foreword in memory of maurizio gaetani rivista italiana di paleontologia e stratigrafia (research in paleontology and stratigraphy) vol. 125(1) march 2019 the first issue of volume 125 (2019) of rivista italiana di paleontologia e stratigrafia is dedicated to the memory of maurizio gaetani (1940-2017), world-renowned professor of geology and stratigraphy at the university of milan, to honour his fruitful life of research and his grandeur as a man, a friend, a mentor and an editor of this journal. many friends, colleagues and scholars of maurizio gaetani joined us in the task to produce a volume focused on the many challenges sought by maurizio in his glorious carrier, from palaeontology to stratigraphy, and from geological mapping to palaeogeography in territories stretching from the southern alps and the balkans, himalaya and karakorum, caucasus and iran, the country that inaugurated his career in the sixties and concluded it in the two thousands: “in a perfect circle”, as he loved to say. in the first paper of the volume, roberto rettori, a scholar and friend of maurizio, with his phd student valerio gennari introduce a new taxon of biseriamminoid foraminifer, globigaetania angulata gen. n. sp. n., from a middle permian succession of nw iran. the discovery of the new taxon sheds light on the evolution of the palaeozoic biserial microgranular foraminifera. as underlined by angiolini & muttoni (2018, permophiles 66), maurizio “never wanted a taxon named after him. we did it anyway because of his great standing as a scientist”. the second paper is by a turkish colleague of maurizio, ercan özcan and co-authors, who want to honour maurizio describing the palaeobiogeographic significance of eocene foraminifera. the authors focus on one of the favourite topics researched by maurizio: palaeogeographic reconstructions and the correlation of peri-mediterranean deposits with those from the south asia. the following paper is by a colleague and a lifelong friend of maurizio, giulio pavia, who, with sixto fernandez-lopez, dedicates to maurizio an impressive monograph on bajocian lissoceratinae (ammonitida). besides presenting in detail their systematic descriptions, the authors discuss their phyletic relations and palaeobiogeographic significance, shedding light on these almost neglected ammonites. eduardo garzanti, a scholar and friend of maurizio, and mohammad ghassemi, one of the iranian colleagues of maurizio in iran, with their co-authors, discuss the provenance of the exotic forewordii and remote karakum dune field of turkmenistan, a central asian desert bound by recent orogenic belts. the composition of the dune sand indicates that the amu darya river, the largest river in central asia, represents the major sediment source for the karakum desert. eugen grădinaru was a friend of maurizio for a long lime and they were working on a paper on triassic brachiopods from north dobrogea, when maurizio passed away prematurely. eugen struggled to finish the manuscript in time to honour his memory, completing maurizio’s perfect circle: he started with a paper on brachiopods in 1964 and his last paper is again on brachiopods. in between, is a rich collection of publications on very different subjects. alfréd dulai, a brachiopod palaeontologist colleague of maurizio, presents additional data on a miocene brachiopod fauna from tetti borelli (piedmont, n italy). there are two new taxa introduced in this systematic and palaeoecological study: one dedicated to maurizio (borellithyris gaetanii n. gen. n. sp.) and one dedicated to a friend of maurizio, giulio pavia (eucalathis giulioi n. sp.). in the paper lead by john powell, many friends and colleagues of maurizio – in particular alda nicora who followed the steps of maurizio during adventurous field trips from the balkans to the himalaya – gathered to produce an excellent and multidisciplinary stratigraphic study of the upper permian to lower triassic succession of the dead sea in jordan, with a focus on the permian-triassic boundary and the pattern of the lower triassic recovery. silvia frisia was a student of maurizio in milan during the earlier part of her career, before moving to australia. silvia, with patrick meister, writes a paper that focuses on a long-lasting debate about the origin of the huge platform of the triassic dolomia principale, contributing to this debate with new data. daniela basso, a scholar and friend of maurizio, and co-authors describe for the first time coralline algal assemblages from the qom formation in nw iran, and their bearing on the interpretation of palaeoenvironmental evolution through time, providing new data on the stratigraphic distribution of several taxa. elisabetta erba, a friend and a colleague of maurizio in milan, presents with her coauthors a discussion of a carbon isotope shift in the jurassic basinal succession of the southern alps, proposing the identification of the “gaetani event” to celebrate maurizio gaetani’s pioneer researches in the southern alps. leopold krystyn and marco balini, respectively a friend and a scholar of maurizio, with two iranian colleagues, describe norian ammonoids from the nayband formation of central iran and their significance in constraining the timing of the collision of the iran plate with eurasia, a subject of study very dear to maurizio. giovanni muttoni, a scholar of maurizio in the late eighties-early nineties, was successfully introduced by maurizio to the world of palaeomagnetism during his phd: with his colleague dennis kent, a friend of maurizio, giovanni presents a straightforward synthesis of the discussion about the transformation from pangea b to pangea a in the permian. andrea tintori, one of the first scholars of maurizio and one of his lifelong friends, remembers maurizio with a paper on the palaeoecology of a species of saurichthys from the triassic succession of northern grigna mountain, a place they visited together many times in the past. based on a very interesting finding, he suggests that the record represents scavenging on floating carcasses and not active predation. marco balini, alda nicora and andrea zanchi with several co-authors dedicate to the memory of maurizio, for them a mentor, a colleague and a friend, a very comprehensive geological study on the structural and stratigraphic setting of the lower to middle triassic sedimentary succession of aghdarband in kopeh-dag, ne iran. besides revising the lithostratigraphy and bio-chronostratigraphy, they propose a new palaeogeographic reconstruction for the aghdarband basin in the triassic. the diverse subjects of this volume, spanning from palaeontology to palaeogeography, from stratigraphy to diagenesis, from palaeomagnetism to regional geology, testify to the vastness of the geological interests of maurizio. we struggled to finish it in time following his teaching and mentoring, and we are sure that it represents an important testimony of the human and scientific qualities of our mentor and colleague. lucia angiolini, fabrizio berra and cristina lombardo who received from maurizio the precious legacy of rivista italiana di stratigrafia e paleontologia. in memory of maurizio gaetani iii mountaineering has also been one of maurizio’s (and lia’s as well) passion: they were able to pass it to the 3 of us and to all the 6 nephews. mountaineering, whenever in winter or summer, has been a way to experience wonderful settings and to test and train vigour, endurance, determination, perseverance, friendships, all of them fundamental life traits. but life was also something more than job and mountaineering; the curiosity for different natural, technological and cultural features has been always encouraged. the whole world was the place for living and playing a role, being aware of our rooted italian culture. cultural differences could make relationships more complex but could not be a dividing-wall: on the contrary they were an option for growing and improving our human qualities. curiosity for nature, from agricultural activities to bee keeping. curiosity for technology devices, when helpful for the daily life, even though not fully understood in their operation and functioning. love for the history, a way to know who we are in order to foresee where we are going. overall, and this is something we learnt and found often in maurizio’s life style, he applied a comprehensive and holistic approach to his life and his job activities. each subject had to be seen by different perspectives, eyes, skills, human and scientific perception, to get the final goal of its whole and deep comprehension….. and typically, best results come from team work …... we are sure you will recognise this approach also in his job activities. finally we would like to thank all the persons we have met in our lifetime thank to his life style: some of you, who are reading now this volume in memory of maurizio, have been path-mate for part of our life time. we want also to thanks the colleagues who are continuing his work with the same enthusiasm and passion. we conclude with his last wish: “siate felici”. from maurizio's family it is with great pleasure and emotion that we, as maurizio’s family, are writing this brief note on maurizio, who has been for us a husband and a father. we would like to give a family perspective of a person that has been a colleague and a professor for scientists and students, respectively; the aim is to find some fundamental learning from a man who tried to fully live his life. according to our experiences, different believes and approaches have been at the basis of maurizio’s life. as for the personal perspective, we have always been incited to find out our personal life-track, according to an old but still modern adagio: “find out your own best talents donated by nature, improve them throughout your life and, if possible, make their benefits available to others”. as kids, we have been encouraged kindly but insistently to pursue this approach. the limit was not “a standard target” but commensurated to each personal talent; after each intermediate step, we were asked to target the following one, increasing skills and strengthening attitudes (“non dormire sugli allori”, often used to say). we were also invited to improve our inclinations too, by studying, reading, meeting people, being reflective and autonomous in thinking and actions. not a pre-set path but the best personal talent-matching one, found thanks to a deep self-understanding. in so doing, all of us could try to do his best in his-own specific field, different for each child, even-though nobody of us undertook the path of maurizio’s personal interest like geology, paleontology and stratigraphy. as for the latter, to go on the field for him was not a task but a pleasure and a way to discuss with colleagues and students, aiming at investigating the earth changes. we all spent several months in himalaya and karakorum to help in measuring mountains and carrying down rock samples and fossils. the talent fulfilment was the target and a severe self-regulation was the way, but a gentle compassion was given when some deviation occurred. independence was given and encouraged, but also verified in the frame of the parenthood responsibility. he has been always open to dialogue and discussion. short note nota breve a pliosaurid tooth from the argille varicolori formation near castelvecchio di prignano (modena province, northern italy) cesare a. papazzoni receioed october 25, 2002; accepted may 5, 2aa3 rivista italiana di paleontologia j""ú ]:,,:1'i november 2003 their synopsis of upper cretaceous plesiosaur from europe, bardet & godefroit (1995) did not include any ltalian plesiosaurs. renesto (1993) reported the first discovery of a plesiosaur in italy: a single pliosauridae indet. humerus from the santonian-campanian argille varicolori formation near zavattarello (pavia province, lombardy, northern italy). in this note, another fragment of cretaceous pliosauridae from the norrhern apennines is recognised and described. moreover, in this case a specific taxonomic assignment is possible. the tooth under stùdy was collected in a gully along the right side of the secchia river, some 2-3 km norrheast of castelvecchio di prignano (modena province; fig. 1). it comes from the argille varicolori formation, of cenomanian-lower campanian age according to bettelli et al. (1989a, b). unfortunately, the lacking of any matrix adhering to the roorh means that the age cannor be better determined. palaeontological description class reptilia laurenti, 1268 subclass sauropterygia owen, 1860 order plesiosauria de blainville, 1835 superfamily pliosauroidea (seeley, 1874) welles, 1943 family pliosauridae seeley, 1874 genus polyptycbodon owen, i841 polyptychodon interruptus owen, 1 841 fig. 2a, b key -uords : sauropterygia, pliosauridae, polyp4,cbodon intenup trzs, tooth, upper cretaceous, northern italy. abstract. the first discovery of a creraceous pliosaurid tooth in itaìy is reported. it comes from the cenomanian-lower campanian argille varicolori formarion near casrelvecchio di prignano (modena province, northern lraly). excepting this new specimen, italy's only reported pliosaurid is a humerus from the upper cretaceous of zavattarello near pavia. the tooth morphology allows it to be ascribed to polyptychod.on ;nterrilptus owen, 1841, a species only reported thus far from northern-central europe (england, germany and the czech republic). this suggests the presence of marine reptile remains in the northern apennines may have been underestimated. riassunto. viene segnalato per la prima volta in italia un dente di pliosauride del cretaceo. il reperto è staro rrovato vicino castelvecchio di prignano (modena), nella formazione delle argille varicolori, attribuita al cenomanianocampaniano inferiore (cretaceo superìore). lunico altro resto di pliosauride finora trovato in italia è un omero del cretaceo superiore di zavattarello (pavia), attribuiro tentarrvamenre alla famiglia pliosauridae. la morfologia del dente studiato permerre un'attribuzione a polyptychodon interruptus owen, 1 841, considerata l'unica specie di pliosauride sicuramente presente in europa durante il cretaceo. finora essa era stata trovata soltanto in inghilterra, germania e repubblica ceca. il nuovo ritrovamento suggerisce che i resti di rettili marini dell'appennino serrentrionale siano forse più frequenti di quanto finora stimato. lntroduction plesiosaur remains from the upper cretaceous of europe a.re rare, and usually fragmentary. nevertheless, they have been recovered from several localities in different countries including belgium, the czech republic, england, germany, the netherlands, poland, portugal, and sweden (milner 1987;barder & godefroit 1995). in c/o dipartimento del museo di paleobiologia e dell'orto botanico, via università, 4 41100 modena; e-matl cpapazzoni@tiscali.it 564 1 841 polyptychodon interruptus orven, p. 19, p|.72, fig. 3, a. 1851t polyptychodon interruptus owen owen, pp. 209-212, lacertilia pi.2, fig. 16,17,p|.8, fig.3. crocodilia, pi.26, fig. 1(?),2-7,8 (?), pl. zr, fig. 1-3. 18516 polyptychodon interruptus owen owen, pp. 55-58, tab. 10, fig. 7-9,tab.11, fig. 1(?),2-l,8 (?), tab. 14, fig. 1-3. 1856 polyptychodon interrhptlls owen von meyer, pp. 3-7 , pl. 2, f ig. 5-8, 10-13, 15,1,6 (?),17. 1861 polyptychodon inteltuptus owen owen, pp.2a-22, trb. +, fig. 3. 1.987 polyptychodon intetruptus owen milner, p. 272, pl. 57, fìg. 2. material. a single tooth crown. locality. in the vicinity of castelvecchio dì prignano (modena province), on the right sìde of the secchia rìver (fig. t) history. collected by mr. enzo grazioli, this specimen was mentioned (but not described) by rompianesi & sirotti (1995) as an ichthyosaur tooth. repository. collection of the "istituto di paleontologìa", universitv of modena and reggio emilia (ipum 30142), donated by mr. pietro rompianesi. description. the maximum height of the tooth is 32 mm (fig.2a). the basal diameters are 14.5 mm (transverse) and 16.5 mm (longitudinal), therefore the crown is laterally compressed (fig. zb). the enamel is very thin, and dark brown in colour. there are alternating long and short longitudinal ridges, a feature which owen (1851a) retained as diagnostic of the species polyptycbodon interruptus owen, 1841. for 8-9 mm at the crown tip, the enamel is fairly smooth. one of the ridges could reach the tip on one side, but the enamel is lacking on it, so we cannot verify this condition. it is noteworthy that the ridges are formed by enamel folding, so they leave impressions on the outer surface of the dentine. remarks. this tooth was previously regarded as belonging to an ichthyosaur, given the presence of location-map of castelvecchio di prignano, with the site (asterisk) where the tooth was collected. some platypterygiws remains very close to castelvecchio di prignano (rompianesi 8c sirotti 1995). during the revision of the ichthyosaur remains from the northern apennines (sirotti & papazzoni 2002) the taxonomic attribution of this tooth has been reconsidered. because the tooth morphology matches that of pliosaurids, and appears very similar to the holotype teeth of p interruptus, it is here regarded as referable to that species. apart from some authors (e.g. white 1940), who included it within the family polycotylidae, the genus polyptycbodonhas been usually regarded as a member of the family pliosauridae (milner 1987; bardet 8c godefroir 1995). according to bardet er godefroit (1995), polyptychodon rs the only vaiid genus of pliosauridae presently recognized in europe. the species p interrwptus has been reported from e,ngland (cenomanian-?campanian), germany (cenomanian), and the czech republic (turonian). the only other valid species of this genus, p. budsoni, was described from north america (\velles & slaughter 1'963). the ipum 30142 tooth is the first record of p interrwptws o'wen, 1841 in italy, thus extending the known geographical range of the species. analysing the tooth morphology in p. interruptus according to massare (1987), one notes it bears an acute but rounded apex, with a natural smooth surfrce; moreover, the longitudinal sculpture is not pronounced, the cross-section is slightly compressed, and there are no cutting edges (fig. z). this suggests the species belongs to the general guild of piercing-smashing predators, which probably ate both soft prey (cephalopods, small fishes) but also somewhat harder ones (ammonites, large vertebrates ) . c. a. papazzoni lcm fig. 2 polyptychodon interruptus owen, 1841. tooth fron.r castelvecchio di prignano (modena province) cat. no. ipum 3a142. a) l.ateral view; b) proximal (basal) view. conclusions the italian record of cretaceous marine reptiles is very poor, so even the single tooth ipum 30142 is oí some importance, because: pliosaurid tooth from northern ltaly 565 a) it represents the first time the species polyptycbodon interruptus owen, 1841 has been recognised in italy; b) this discovery together with that of renesto (1993), substantiates the presence of the pliosauridae in italy during the late creraceous. moreover, this suggests the cretaceous mrrine reptile fossil record in the northern apennines region has probably been underestimated. acknou/edgemerzts. partìcular thanks to the late prof. a. sirotti, who introduced me into rhe study of vertebrates and made an initial description of this tooth. manl' thanks to prof. n. bardet (mus. nat. hist. net. paris) and dr. m. \xi maisch (tùbingen university), who dreq'my attention to the pliosaurid narure of the tooth under study. thanks to mr. p rompianesi, who provided the material and showed me the locality where it was discovered. the author also thanks d. naish (university of portsmounth) and l. noé (sedgwich museum, cambridre) for the critical reading of the manuscript. refer bardet n. & godefroit p. (1995) plesiosaurus houzeaui dollo, 1909 from the upper campanian of ciply (belgium) and a revieq' of the upper cretaceous plesiosaurs from europe. bull. inst. roy. sci. nat. belg. (sc. tène), 65: i 79i 86, bruxeìles. bettelli g.,bonazzi u. & panini f. (1989a) schema introduttivo alla geologia delle liguridi dell'appennino modenese e delle aree limitrofe. mem. soc. geol. 1t.,39 119871:91-125, roma. betteiìi g., bonazzi u.,fazzini p, gasperi g., gelmini r. & panini f. (1989b) nota illustrativa alla carta geologica schematica dell'appennino modenese e delle aree limitrofe. mem. soc. geol. [t.,39 119871:487-498,1 geol. map, roma. blainville h.m.d. de (1s35) description de quelques espèces de reptiles de la californie, précédée de i'analyse d'un s)rstème général d'erpétologie et d'amphibiologie. nouv. ann. mus. hìst. nat. paris, 3: 233-296, parls. laurenti j. n. (1/68) josephi nicolai laurenti austriaci viennensis medicum, exhibens synopsin reptilium emendatam cum experimentis circa venena et antidota reptilium austriacorum. 21'l pp., \wien. massare j.a. (1987) tooth morphology and prey preference of mesozoic marine reptiles.j. vert. paleont.,t (2):121137, lawrcnce. mever h. von (1856) saurier aus der kreide-gruppe in deutschland und der schweiz. palaeontogr.,6:3-1.3, cassel. milner a.c. (1987) reptiles. in smith a.b. (ed.) fossils of the chalk. tbe pale ontological as s ociation ; 266-28a, london. owen r. (1841) odontography, vol. il 79 pp.h. bailliere publ., london. owen r. (1851a) a history of british fossil reptiles. section ii. fossil reptilia of the creraceous formations: 155-274, london. ences owen r. (1851b) monograph of the fossil reptilia of the cretaceous formations. mon. palaont ,loc., 5, part i: 1118, london. owen r. (1860) on the orders of fossil and recenr reptilia, and their distribuiioi in time. rep. br. ass. aht. sci.,29: l5l-166, london. owen r. (1861) monograph of the fossil reptilia of the cretaceous and purbeck strata; including supplement no. iii. cretaceous pterosauria and sauropterygia. mon. palreont. soc., 1,2: 1-25, london. renesto s. (1993) a cretaceous plesiosaur remain (reptilia, sauropterygia) from the argille varicolori of varzi (pavia, lombardy, northern italy). rio. it. paleont. strat., 99 (1): 101-106, milano. rompianesi p & sirotti a. (1995) vertebre di ittiosauro nei "terreni alloctoni liguridi" di prignano (modena).,4ri soc. nat. mat. modena, 125 119911 3-9, modena. seeley h.g. (1874) note on some of the generic modifications of the plesiosaurian pectoral arch. quan. j. geol. soc. lond., 30: 436-449, london. sirotti a. &-papazzoni c.a. (2002) on the cretaceous ichthyosaur remains from the northern apennines (italy). boll. soc. paleont. ital.,11 (2-3):237-248, modena. \íelles s.p (1943) elasmosaurid plesiosaurs with a description of new material from california and colorado. mem. univ. calif., 13 (3): 125-215, berkeley. \x/elles s.p & slaughter b.h. (1963) the first record of the plesiosaurian genus polyptychodon (pliosauridae) from the new vorld./. paleontol.,37: 131-133, ti.rlsa. vhite t.e. (1940) holotype of plesiosaurws longirostris blake and classification of the plesiosaurs. j. paleontol., 74: 451-467,\tlsa. preface 3..4 ������� ����� ���� �� ���� � � ��������� � � ���� ����� ��� ������ ������ ���� � � ���� � ��� ������ !���"#� !����#� �$ $���% !���&!'��� &"% ��(��)��"# �&(�!&#�!* �#�%��� �$ #�" *�&!� �"���#�+ ,&#��"� �" #�� �����#�'�"� �&"%��!� -���&. �&��" ��''��+ ���"�/ �� &!� ��!* ,�&% $�! #�� � �!#�"�#* #� �(���� #��� '����'#��" �$ & �!� &� & �� ����"# �$ #�� ������� ������ � � ���� � � ��� � ������������/ �� �$$�! & � �+ '�&� #�&"0 #� �%�#�!�&� '����##�� �$ #�� 1��!"&� &"% �� �'�&��* #� #�� �%�#�! �" '���$� !�$/ �&�!�2�� 3&�#&"�� 4�� ���# �&�� (��" 4�!!��% &� #� 4��#��! #�� ������ 4���% $�"&��* (� '�� ��#�% �" ��'� & ���!# #���� (�# �&� �#�&%��* �$$�!�% &%��'� &"% �� �!#/ ��! #���! �"�&��&(�� ��� �" #�� !����4 &"% �%�#�", !�'��� �$ #�� ������� ��"'�!� & !�'�&#��" �� �5#�"%�% #�6 �/ �!���!+������ �/�/ ��!"�!� �/ %� �& &!�"# %� �!��"� �/�/ ���"�",� �/ �!&*�!� �/ 3&��(�!#�� �/�/ ��4���� �/�/ �&!#�"�� �/ ��'0$�!%� 7/�/ �&,�� �/ &,��&'�22�� �/ ��&��!� &� 4��� &� #� �#��! &"�"*���� !����4�!�/ �! $���% 4�!0 !���&!'��� �" #�� "�!#��!" �&"&+ 0�� &!�& (�,&" �" �&#� 899: &� & 1��"# !�1�'# (�#4��" #�� ���&!& �� &!#��"# �$ ��"�� &"% #�� ;"���!��#* �$ ���!�"'�� �&�"�* $�'���% �" ,����,�'&� ��!��* &"% !�+ ,��"&� ,����,* -<<�� ���� � ��� � � � � � � �����== �!�1�'#./ �� $�!�# ��!��*� �" �&#� 899:+�&!�* 899> ��% #� #�� �%�"#�$�'&#��" �$ & ����+�����#�'�"� $����� �&�+ �&�+(�&!�", ��''�����" ��'&#�% ?@ 0� ���#� �$ ���& ����&,� �" #�� "�!#��!" �&"&0�� -�$&!. %� !�����"/ �� �"#�!��# $�! #�� &!�& �"'!�&��% (* #�� %��'���!*� �" �&#� 899>� �$ &" &����# '�� ��#� ���&" '!&"��� &� 4��� &� ����!&� ��#�� 4�#� 4�%�� !�&% &"% &(�"%&"# �%�4&" &"% �'�����&" #����/ ��"'� #�� %��'���!* �$ $����� �&�+ �&��� &���&"#�!� ���,�'&� &"% &!'�&����,�'&� !��&�"�� #�� !�1�'# (�'��� �$ ��'� 4�%�! �"#�!��# &"% �# �&� (��" �"�&!,�% #� �"'��%� $�!#��! 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'�"#�"�+ ����* !���%�%� &��", #���� *�&!�� ��"'�!� $!��"%��� &"% �"�&��&(�� ��,��#�' �� �!#/ ����!&� $!��"%� &"% '����&+ ,��� �" ���&!& �&�� (��" �$ ,!�&# ��� �" �&!���� �&"+ "�! %�!�", #���� #�" *�&!� 4�!0 &"% %���!�� & � �'�&� ��"#��"6 �"%!�& ��%� -����.b �&"��� 7��&""�� &"% 3�,������ ����''��!& -3���'��"'�./ ���", #�� �&�# *�&!�� "���!��� '����&,��� �&�� (��" �$ !�'���� ��� $�! %��'�����" &"% &����#&"'� %�!�", �&!���� �&��� �$ #�� !�1�'# &'#���#���6 �/ �22&!���� �/ ��"��"�#�� e/ ��!+ ������ �/ ��"'�##�� �/ �� !�&"�� �/7/ ��&!0�� 3/ ��"#�"�� �/ ��22�"�� �/ �!�����&!�� �/ �&"$&"�� 3/ ��''&!����� �/ 3��#+ #�� �/ �&"%�''�� �/ �&!'�''�� �/ ��"�##�� �/�/ ���!� �/ �&���!�"�� �/�/ ��''�##� �22&!���� �/ �!!�� �/ e�%&��� �/ �&"&,&/ ��"&��*� & � �'�&� ��"#��" $�! #�� �� �� �$ #�� ����&,� �$ ���&� $�! #���! 4&!� ��� �#&��#* &"% $!��"%+ ��� / �� ������� �� � ����!� � �� ���: dicembre 2ool nota breve-shom note turriglomina ? anatolica, n. sp (foraminiferida) from the cretaceous of north-\testern anatolia (turkey): remarks on the evolution of mesozoic meandrospirids demir altiner 1, roberto rettori 2, louise,tte zaninetti : se ross.ana martini 3 receroed may 18,20a1; accepted september 2a,2001 key -rt ords. foramìnifers, nerv species, taxonoml', cretaceous, turkey. riassunto. nel presente lavoro viene descritta una nuova specre dì foreminifero, tuniglomina? anatolica, n. sp., del cretacico inferiore dell'anatolia nord-occident;rle. la nuor';r specie è caratterizzata da uno stadio ìniziale ad alvolsimento meendrospiroìdeo seg;uito da una seconda parte rettilinea ad andanrento elicoidale. la morfologia di questo nuovo taxon è simile a quella del genere triassico turrig/omina zaninetti. l'attribuzione generica della nuova specie è comunque dubitativa a ceusa del fatto che il trend evolutivo di questo gruppo di foraminiferi non è documentato dal triassico al cretacico. abstract. a new species of foraminifer, turrig/omína ? anatolicd, n.sp., ìs erected frorn the lower cretaceous of north-\ffestern anatolia, turkey: the species is characterized by a well developed meandrospirid stage folloned by a rectilinear, heljcoidal stage. the morphology of the nen taxon ìs sin-rilar to that of the tiassic genus turriglomina zamnetti; hon'ever, the generic attributjon is doubtful as the evolutjonary path of meandrospirids is not documented from triassic to cretaceous. lntroduction. iìr the evolution of triassic foraminifers, rhe genus tkrriglomina zaninetti in limongi er al. (1982) (rype species: turritellella mesotridsica koehn-zaninerri, 1968) is a well-known taxon composed of several species ranging stratigraphically from anisian to norian. this genus is considered to be linked in evolution to tiassic representatives of meandrospira loebltch & tappan (rettori, 1995). although turriglomina is not reported from the jurassic, we describe from the creraceous of anatolia a meandrospirid foraminifer, similar to the triassìc genus turriglomina, earlier recorded and figured by altiner (1991) as meandrospiranella sp. the new form is described as turriglomina ? anatolica, n. sp. flowever the generic attribution remains doubtful as the evolutionary link between the triassic and cretaceors me.lndr o sp ira -twrriglom ina linea ge s is not do cumente d. the new species is recorded from a succession located 25 km north of the bilecik city (fig. 1). the jurassic to lower cretaceous sequence in this area is composed of four formations, widely exposed in north\(/estern anatolia u'hich is located in the sakarya continent, one of the tecronic enriries in the neo-tethyan evolurion of turkey (altiner et al., 1991;. the hettangian-pliensbachian bayirkóy formation is composed of continental to marine siliciclastics intercalated with fossiliferous rosso ammonitico facies (fig. 2). after a gap in sedirnentation, a carbonate regime starts in the succession comprising the nodular limestones of the ^ llcallovran-oxtordian taseibayiri forrnation in the lowest part. the major unit of the carbonate succession is the kimmeridgian-valanginian gúnóren limestone, mainly consisting of coral-rich reefal and high-energy limestones. the calcareous and argillaceous pelagic sediments of the latest valanginan-aptian sogukgam limestone overlies the gùnòren limestone with a condensed level rich in crinoids at the bottom. turriglomina ? anatolica, n. sp., is recorded from this condensed level. the upper boundary of the sogukqam limestone is not exposed in the studied area. systematic palaeontology order foraminiferida eichwald, 1 830 suborder miliolina delage & hérouard, 1896 superfamily cornuspiracea schultze, 1 854 1 middle east technical universìtv, department of geoìogical engineering, 06130 ankara, turkey, e-mail: demir@lmetu.edu.tr 2 dipartimento di scienze della terra, piazza unìversità, 06100 perueia, italy', c-m;ri1: rrettori@)unìpg.it. 3 dép:rrtement de géologie et paléontologie, 13 rue des maraîchers, 1211 genèr-e,{, suisse, e-mail: louisette.zaninetti(aìterre.unjqe.ch 178 d. altimer. r. rettorí. l. zaninetti €; r. martini family meandrospiridae saidov;r, 1981, emend. zaninettr et al., 1.987 subfamily turri glomini nae zaninetti i n limongi et al., 1987 genus turriglominazantnettrin limongi et al., 1982 type species: turritellella mesotriasíca koehn-zaninettr, 1968 turriglomina ? anatolica, n. sp. pl. 1, fìgs. 1-1ó 1991 lleandrospiranellt sp. altiner, pl. 13, figs.6-8. 199ir lleanttrospiranella n. sp. altiner et al., p. 32 (not figured). origin of the name. the name of the nerv species is after the region anatolìa, whcre turríghmìna ? anato/íca n. sp. has been fìrstl1' recorded. holotype. the holotvpc (p1. 1, fig. 1) is a slightlv oblique longitudinal section, passing through the proloculus anc{ shon'ing both the helicojdal and meandrospirid parts of the tesr. paratypes. they rre figured in pl. 1, figs. 5, /, 13, 15, 16. !,'pe localin-. 25 hm north of bilecik, north-western anatolia (tlrrkc1.) (fig. t). type level. condcnsed level ;rt the bottorn of the sogukgam limestone, sample g-5, referable to the hedbergella sigalildelrìoensìs zone (latest valanginian to earliest barremian) of robaszvnski tr caron ( 1 995). material and repository. the nen specìes is present in four samples (g-5, g-9, g-10 and go-11a) from tn'o measurcd sections (g and go); the metcrial rs cleposited in the micropaleontological coìlection of the marine micropaleontologl'research unit, middle erst technicel universìt1-, ankara. microfacies and microfaunal association. the microfacies containing turriglomina ? anatolica n. sp. are wackestones to packstones rich in crinoids, bryozoa and pelagic elenents such as planktonic foraminifers, thin shelled bivalves etc. the microfaunal àssemblage connins globuligerina boteriz,ica (subbotina), hedbergella srgali (mou1lade), globochaete alpina lombard, meandrospira faz,rei (charollais, brònnimann & zaninetti), vidaliùa spp., ophthaìmidìum sp., textularia sp., spirìllina spp., patellina turriculata dieni & massari, patellina spp., lenticulina sp., gavelinella sp., epistomind sp., artd ataxophragmiid foraminifers. description. the test of the new species consists of two parts. the meandrospirid spherical portion is characterized by a globular proloculus, followed b,v a second undivided tubular chamber, rvhich is rounded in section. the chamber describes, as in the genus meandrospira loeblich ee tappan, 2 to 211, (aìmost 3 in the holotype) planispirally coiled involute r.r'hor1s, composed of several tight meanders, 5 to 6 in each whorl. the diamcter of the tube incre:rses with gron'th, in order to reach in the second whorl, twice the diameter of the tube in the first whorl; however, the number of the meanders reueins constant. in the second part of the test, the tubular chan'rber fie. 1 geographic location (a and b) and geological map (c) ot the studv area (after altiner et :r1., 1991, modified). describes a high spire, helicoidal and close-coiled in order to form a columella ; the number of n'horls varies from 5 to 7, and the diameter of the tubular chamber remains constant. the wall is black. complct and porcelaneous, sample no. vxce t":r* ^ i .^ 1\ g kks: sogukgam limestone (tatest valanginian-aptian) jkbgr cúnóren limestone (klmmeridgian-valanginian) jbt: ta g g'bayiii formation (calìovjan-odoidia n) jb: bayirkòy fo.rnation (henangian-pliensbachian) new, species of turriglomina sample no 179 fig.2 measured stratìgraphic sections (g end go; after altjner et al., 1991, modified) and type leveì ol turriglornina ? anato/ica n. sp. a.e th'ck-bedded, coarse-grained contjnental or shallow marine sandstone ól) f) ,'\w v 1--l coral brachiopcd ec,\inoid ^@ooosponae cretac€ouss.îraller globdligenna spicuie pianktonicforamrnifers often recrystallised. the aperture is simple, at the end of tube. in the studied material, we have recognized two types of specimens characterized bv different dimensions of the proloculus, and interpreted rs megalospheric (pl. 1, figs. 1, 6-7,11., 13) and n-ricrospheric generations (p1. 1, figs. 4, 12, 14).in the megalospheric forms, the dimensions of the proloculus approximately correspond to the diameter of one u..horl around the proloculus of the microspheric specimens. for the orhcr p.ìrameters of the test, the two different generations do not exhibit a noriceable change in the overall size and general rnorpholoe,v. dimensions (in microns). height of the test: 260400; width of the rectilinear portion: zo-1oo; diameter of the meandrospirid portion: 120-150; di:rmeter of the @ ó@ u t^ t o'ó proloculus: ,10-50 (megalospheric), 1o (microspheric) ; diarneter of the tubular chamber in the first whorl: 1o; diameter of the tubular chamber in the last planisoirai whorl: 20; height of the tubular chamber in the helicoidal portion: 2o-30; thickness of the wall: less than 10. remarks. altiner (1991) attributed the spccimens figured in pl. 2, 3 and 5 ro rhe triassic gews meandrospiranella salaj. however, rh;s.rttribution can not be correct since the rectilinear porrion of turrig/omìna ? anatolica n. sp. ìs helicoidai and does not describe a meandrospirid uncoiled sra€ie as in meandrospiranel/a. turriglomìna scandonei zaninettr et al. and tur rig/omina mdgnd (urosevic) are the two triassic species characterized by a well-developed meandrospirid stage (limonei et al. 1987; urosevic 1977; zminetti et al. 1987; zaninetti et al. 1.99a; bérczt-makk 1993; rettori 10 (t \ll-l cù (j o co ycú o c'. o_ .: h 10 f:rl i::::_ lrmestone l_,'l i h.igh-€nergy llmestone z77vs t//4(\ cross-bedded sanostone i:rt reefal ilmestone w <><> nodular imesìon9 argillaceous condensedlìmestone limestone wfth crinoid sand vj \.j _ò -) 1^ td c-) ls ffi dasyciad r\ b€lemnite ó crinoid go 480 altimer, r. rettori, l. zaninetti €; r. martini 1995; martini et al. 1995); turriglomina ? anatolica n. sp. differs from twrriglomina scandonei in having smalier dimensions, a iower number of meanders in the spherical portion, and a thinner wall. it is drstinguished from turriglomina magna by r much regulrr coiling of the meandrospirid part, and a well developed helicoidal stage. the other triassic species of twrriglomina, turriglomina mesotriasica (koehn-zaninetti), turriglomina conica (he-yan) and turriglomina carnica (dager) characterized by their small dimensions and a reduced meandrospirid stage, can be clearly distinguished from all the other representatives of the genus turriglomina, and also ftom twrriglomina ? anatolica, n. sp. finally, the ladinian-carnian turriglomina robusta bérczi-makk from hungary does not exhibit the peculiar features of the genus; for this reason it can not be compared to tuniglomina ? anatolicd, n. sp. from the lower cretaceous of algeria, sigal (1952) cited a species called meandrospira cljffiensis characterized by a meandrospirid initial part followed by î rcafilrncrr sfîqe floweveq from the two specimens illustrated and given as drawings by sigal (1952, pl 12, two figs.), it is not possible to recognise the type of coiling of the rectilinear portion, which is described by the author as uncoiled (dérowlé). for this reason, meandrospira djaffaensis can not be compared to turriglomina ? anatolica, n. sp. phylogenetic considerations. the stratigrrphic range of turrìglomtnd, composed of several species, is anisian to norianl the genus is so far completely unknown from the jurassic platform or basinal facies. in the triassic, twrriglomina, because of its me:rndrospirid early stage, has been considered to be linked to meandrospira and derived from this genus lzaninerti et al.. 1987; rettori. le95r. similarly. the discovery of turriglomina-llke forms in the cretaceous of anatolia su€igests, as in the evolutionary lineage 6 9 10 77. 1.2 observed in the tiassic meandrospiridae, rhat turriglomina ? anatolica, n. sp., could also have been derived from a meandrospira ancestor. this is in concordance with the fact that the genus meandrospira (species ,m. favrei) is also known from the lower cretaceous (valanginian) of several western tethyan localities, including the type locality of t. ? anatolica, n. sp., in north -western anatolia. in this hypothesis, the meandrospira hneage should be continuous from the triassic to the cretaceous, with the evolutionary potential to produce, at any time, specimens with a helicoidal stage. these specimens, resulting from an iterative evolution, are morphologically identical to turriglomina zaninetti in limongi et al., and can not be distinguished from this genus. for this reason, turrìglomina is here considered as a polyphyletic taxonomic unit, and the genus is used with reservation for the cretaceous species from anatolia. aclenouledgements. the authors are grateful to tubitak (the scientìfic and technical research council of turke,v), to the cnr (consiglio nazionale delle ricerche, italia) and to the fnsrs (grant n' 2050577.97) ior supporting this study. prof. a. koqyigit, prof. a. farinacci, dr. u. nicosìa and dr. m.a. conti helped in r.arious stages of tbe fields.ork. thanks are due to a. cherchi (unìversìtv of cagliari) i"r her heìp in the ìn'pr.rernent ol-rlrc rn:nu.cript. plate 1 all specìmens ,rre from the lorvermost prrrt of the sogukgam limestone, north--ù/estern anatolia, turke1.. fìgs. 1-1,1, 1.5,1.6 turriglomina ? anatolica, n. sp. 1 holon-pe. longitudinal section. sampje g5/5. 2,3 longitudinai sections. sample go 1'1 a/1. .+, 8, 1 4 '1: longitudinal scction; 8: longituclinal tangenti,rl ,rnd oblique section; 1,1: subequatorial section ol the meandrospiroid initial part. sam ple g10/1. 5,7,13,15, 16 paratypes. 5: longitudinrl section; 7: longitudìnal oblique sectìon; 13: equatoriai section of the meandrospiroid initial part; 15, 16: tangential sections of the n-reandrospiroid inìtial part. sampjes g5li,g5l9, g5/5, g5/9, g5/8 longitudinal section. sample g1o/2. longitudinal tangential and oblique section. sample g9l1. oblique section. sample g1o/3. longitudinal oblique sections. samples g11al2, go 14a/12. pì.1 neia species of turriglomina 3 o o c\i 482 d. altimer. r. rettori. l. zaninetti g r. martint re,fe,rences altiner d. (1991) microfossil, biostratigraphy (mainly foraminifers) of the jurassic-lower cretaceous carbonate successions in north-\lestern anatolia (turkey). geologica rom., 27 , 167 -273, roma. altiner d., kocygit a., farinacci 4., nicosia u. & conti a. (1991) jurassic-lower cretaceous stratigraphy and paleogeographic evolution of the southern part of north-lwestern anatolia (turkey). geologica rom., 27, 13-80, roma. berczi-makk a. (1993) turriglomina zaninetti in limongi et al. (foraminifera) species in the triassic formations, aggtelek-rudabanya mts. (northern hungary). acta geol. hungarica, 36, 297 -314, budapest. koehn-zaninettt l. (1969) les foraminifères du trirs de la region de i'almtal (haute-autriche). jb. geol. bundesanst. sonderbd., 14, 155p, \x/ien. limongi p, panzanelli-fratoni r., ciarapica g., birilli s., martini r. & salvini-bonnard g. (1987) turriglomina zantnettr, n. gen., un noveau nom pour uturritellella" mesotriasica koehn-zaninetti, 1968 (foraminifère, trias moyen) arc. sc. genèae, 40 (1),13-22. loeblich a.r. & tappan h. (1987) foraminiferal genera and their classification. van nostrand reinhoìd company inc., new york. martini r., rettori r., urosevic d. & zaninetti l. (1995) le genre piallina rettori & zaninetti (foraminifère) dans des calcaires à turriglomines du trias (carnien) de serbie orientale (domaìne carpatho-balkanique). rez. paléobiol., 14 (2), 411-415, genève. rettori r. (1995) foraminiferi del trias inferiore e medio della tetide: revisione tassonomica, stratigrafia ed interpretazione filogenetica. publ. dép. de géol. et paléont. genète, 1.8, 147 pp. robaszynski f. & caron m. (1995) foraminifères planctoniques du crétacé: commentaire de la zonation europe-méditerrranée. bull. soc. géol. france, 166(6), 681-692, paris. sigal j. (952) apercu stratigraphique sur la micropaleontologie du crétacé. congr. géol. internat. xix, monogr. régionales, algiers, ser. 1 (algeria),26, 17. ijrosevic d. (.1977) stratigraphic position of some foraminifers in tiassic sediments of the carpatho-balkanides. ,4nn. géol. penins. balk., 41, 227 -231, beograd. ztninetti l., ciarapica g., martini r. & rettori r. (1990) paléoécologie des turriglomines (foraminifères) dans 1e trias de 1' apennin méridional (bassin de lagonegro) hahe. archs. sc. genève. 4l{2). 295-j05 genève. zamnetti l., ciarapica g., martini r., salvini-bonnard g. & rettori r. (1982) turriglomina scandonei, n. sp., dans les calcaires recifaux du trirs moyen (ladinien) en apennin méridional. rea. paléobiol., 6(2), 1,77-182, genève. cyclostratigraphy a methodological approach mario sprovieri 1, angelo bonanno 2., salvatoremazzola 2b e. bernardo patti z' recebecl july 1 5, 2aa1; accepted judnuatt 9, 2aa2 keyusords : cyclostratigraph,v, insolation curve, spectral anal,vsis, astronomical calibration. rittssuntoin questo lavoro vengono descritte e commentate sinteticamente le metodologie pìù usate di analisi ciclostratigrafica per la calibrazione astronomica di sequenze sedimentarie. dapprrma sono descritte dir.erse soluzioni numeriche della curva di insolazione e, in particolare, della più nota curva astronomica la9o1r,ry. vengono quindi, in ordine, descritte le metodologie di calibrazione astronomica i) su base lito-ciclostratigrafica e ii) sulla base di metodoiogie di analisi spettrale applicate a segnali faunistici e geochimici che rìspondono ìn fase alle dìverse forzanti astronomiche. abstract. aim of this study is r synrhetic description of the most used methodologies of cyclostratigraphic analysis for the astronomical calibration of sedimentary sequences. the different numerical solutions of the insolation cune and, in particular, the most used astronomìc cur-ve la901r,ry are analtzed. then. a detailed description of the methodologies for the astronomic calibration of different sedimenrar)sequences i1 on the basis of a iitho-cyclostratigraphic approach and ii) on the basis of spectral analysis applied to faunal and geochemical climate-sensitive records, is proposed. introduction a;-^^*,.-. ^1rr of this volume is declicated to the reconstruction of an astronomically calibrated time scale from about 1 1 ma to about 1,3.7 ma, on the basis of cyclostratigraphic analyses of different land-based sections. in the last years, a restricted group of researchers provided to the international scientific community a reliable astronomical calibration of the late neogene geological record te.g. hilgen l99la, 199 lbl hilgen et al. 1995; shackleton et al. 1995; lourens et ai. 1996; hilgen et al. 2000). they also developed the procedures to tune sedimentary sequences, characterised either by homogeneous or cyclically-repeated lithologies, to different numerical solutions of the insolation curve. the most used numerical and cyclostratigraphic techniques applied for calibration of sedimentary records are here synthesized. fwo different kinds of sedimentary records are generally used for astronomical calibration: i) sequences characterised by lithologic alternations organised in well-recognisable cluster patterns and ii) sequences characterised by homogeneous lithology and studied by means of spectral methodologies applied to selected climate-sensitive records. when possible, the combination of these two cyclostratigraphic approaches represents the most reliable system to calibrate geological time series with the astronomical curves. the insolation curve the insolation on the earth depends on its orbital parameters. until 1988, the solution adopted for paleoclimate computation consisted of the orbital elements of the earth, complemented by the computation of its precession and obliquity (berger 1928,1988). in 1988, laskar proposed a solution for orbital elements of the earth, which was obtained in a new manner, making use of vast analytical computation and numerical integration (laskar 1988). in this solution the precession and obliquity parameters, necessary for paleoclimate computations, were integrated at the same time, insuring a good consistency of the solutions. ljnfortunately, for various reasons, this latter solution for the precession and obliquity was considered reliable only for the last 3 ma (berger & loutre, 1,991).later on, laskar et al. (1993) produced a new solution (hereafter named la 90) that is a slight improvement of the previous one. this solution allows evaluation of orbital parameters, precession and obliquity, up to the last 20 ma. the orbital solution lago is obtained by numerical ' dipartimento (cfta) via archirafi 3(),9a1.23 palermo, itall', e-mail: marios@unipa.ir ' istituto irma cnr via luigi vaccara, 61,, 91,a26 mazara del vallo, italv, e-mail: 2" bonanno@irma.pa.cnr.tt;2b mazzola@irma.pa.cnr.it; 2 b p atti @)irma.pa. cnr. it 180 m. sprooieri, a. bonanno, s. mazzola & b. patti integration of an extended averaged sysrem that represents the mean evolution of the orbits of the planets. all the 8 main planets of the solar system are taken inro account, as well as the main lunar and relativistic perrurbations. the use of numerical integration for computing the solution of the secular system is one of rhe reasons for the good quality of this solution, which was checked by comparing with the ephemeris over a short time scale (laskar 1988). the la 90 solution is in quasi-periodic form over 10 mr, but these representations are s1owly convergent, which prevents good accuracy of the solution. the reason for this slow convergence is due to the presence of multiple resonance in secular periodicity of the inner solar system (laskar 1990). because of these resonance the motion of the solar sysrem is chaotic in the iong term, and not quasi-periodic. the lyapunov exponents of the solar system, that quantify the average grow of small initial error in the calcuiation, was esrimated to l/5yr-r. this implies that it is not possible to give any precise solution for the motion of the earth over more than about 10 mr, and most probably, ephemerids can only be given with good precision for no more than 20 mr. the solution of laskar et al. (1993\ can be "recalibrated" at different ages because it gives the possibility to vary the values of the dynamical ellipticity and totai dissipation by the sun and moon. lourens et ai. (1996) carried out a careful correlation between plio-pleistocene mediterranean land-based sedimentary sequences characterised by well-recognisable lithologic rhythms and well calibrated from a cyclostratigraphic point of view, and the different solutions of the insolation curve la 90. the best fit between the geological record and the insolation curve has been obtained using the numerical solution of laskar et al. (1993) with unitary values of both the dynamical ellipticity and the total dissipation by the sun and moon la9o.11.11. successive works (hilgen et al. 1995; sprovieri et ai. 1999: hilgen et al. 2000) confirmed these results for the neogene stratigraphic records. in the papers of iaccarino (2002), the lago 1r,r; solution has been used ro calibrate the studied sedimentary records. astronomical calibration of cyclic land-based sedimentary sequences the studies of hilgen and co-workers (e.g., hilgen 1991,a, 1991b; hilgen et al. 1995, 2oo0) on mediterranean sections demonstrated that the high frequency alternations of different lithologies, recognised throughout several land-based sedimentary sequences, can be directly related to the precessional forcing and that the alternation of lonqer sedimentary intervals characterised by absence/presence of high frequency lithologic cycles can be related to the eccentricity periods of t i ^^ | /^^ i adout tuu ancl +uu kvr. these papers demonstrated that the same kind of cyclic pattern with clusters on different scales superimposed on the basic sedimentary cycle can be recognised in the mediterranean from the tortonian up to rhe plio-pleistocene. alternation of homogeneous grey marls and sapropelitic layers represents the classic high frequency cyciicity recognised in most of the mediterranean records. alternativeiy, these two lithologies can be substituted by similar variants (e.g., grey marls tone /carbonate layers, light coloured/ grey coloured marls rone, etc.). individual sapropelitic layers or substitutive sedimentary expressions correspond to precession minima or the equivalent summer insolation maxima, while smalland large-scale lithologic clusrers can be related to i '^^ ^^^ | r^^ ^^^the 100.000 and 400.000 years eccentricity maxima cycles. this simple sedimentary response to insolation forcing remained in practical identical over the last 1o myr, with the same phase relations between sedimentary and orbital cycles described for the plio-pleistocene. the assumption of a constant phase relationship between lithological cycles and astronomic forcing for all the neogene, ailow us to identify a rigorous stepwise methodological approach for calibrating sedimentary sequences, characterised by lithological alternated records. firstly, a detailed lithologic description of the studied sections allows the identification and the characterization of the sedimenr^ry rhythms along the records. secondly, the studied successions musr be accurately dated with classic bio-magnetostratigraphy. in particular, polarity chrons musr be identified on rhe basis of cande tr kent (1995;ck95). then, the ages of ck95 and shackleton er al (1995a; schps95) for the poiarity reversals, and especially for the youngest ones, have to be adopted as starting point for the actual tunirg. thirdly, a first-order tuning between the sedimentary records and the insolation curve is obtained by a first-order correlation of large-scale sapropel clusters ro the 400,000-year eccentricity maxima and a direct correlation of the smal1-scale sapropel ciusters to the 10o.oooyear insolation cycles maxima. fourthiy, the astronomical calibration is completed by tuning individual sapropels to precession minima and the corresponding summer insolation maxima. according to hilgen (1991a, 1991b) and lourens et al. (1996), a phase lag of -3 ky between mid-summer northern hemisphere insolation and lithologic cycies has to be considered for the final tuning. during the tuning exercise, larger-scale eccentricity cycle patterns must be taken into account, because cyclostratigraphy, a methodological approach 181 they provide a direct check on the validity of the calibration and prevent accumulations of errors due to tuning of the individual cycles (i.e., several 20 kyr cycles could be missing or not recognisable). the resulting tuning is probably accurate to the 1eve1 of the individual precession/insolation cycle in certain intervals but may be off by one cycle in others. the final calibration provides astronomical ages for the sedimentary cycles, as well as for the bio-events recognised throughout the studied record. it should also provide refinement of duration of individual polarity chrons. astronomical calibration of homogeneous sedimentary sequences when lithological rhythms are not present rn a sedimentary sequence, the acquisition of detailed time series of climate sensitive records (oxygen isotopes, geochemical elements, planktonic and benthic foraminifera species, nannofossils species, etc.), known to be forced by astronomic periodicities, can be used to calibrate the studied sequence. in this case application of spectral analysis allows the recognition of a set of periodicity directly correlatable to astronomic parameters. the first step for comparing proxy records to the insolation curve is an appropriate sampling rate of the section (selected for a proper analytical resolution of the highest frequency band) and the selection of the tuning straregy thar enables the establishment of the phase reiationships among the different chosen signals and the astronomic record for the different frequency bands (generally the three milankovitch frequencies of precession, obliquity and short and long-eccentricity). the different records have to be converted in the time domain using the classic procedures suggested by martinson et al. (1982) and the numerical algorithms of the inverse conjugate. identification of a set of tie points (already independent dated biohorizons and/ or magnetostratigraphic intervals recognized along the section) allows the definition of a first approximation of the sedimentarion rate of the studied succession. it is necessary to transform the available signals from the spaceto the time-domain and to identify in the power spectra, estimated for the selected signals, the classic milankovitch periodicities. the filtering of the original signals in the longand short-eccentricity frequency bands and the direct comparison with the same cyclicity of the insolation curve allows a first-order astronomical calibration of the sequence. then, filtering of the signals in the precession frequency bands and correlating them with the 19-23 kyr cycles present in the astronomic curve, produces a detailed short-time calibration of the studied record. cross-spectral analysis between the paleoclimate signals and insolation should confirm with high coherency values for the milankovitch frequency bands the correctness of the obtained calibration. at this point all the stratigraphic events recognised throughout the succession can be dated. shackleton et al. (1997) suggested that application of numerical techniques of complex demodulation could be used as useful tool for calibrating sedimentary sequences, characterised by high variance concentrated in the precession frequency band. the complex demodulation method is a tool for examining the instantaneous amplitude and phase of that portion of the variability that is in a particular frequency band of the signal spectrum. for the mathematical procedure, reader is referred to shackleton et al. (1995b) and pisias & moore (1981). there are several applications of such a method for investigating the amplitude modulation in the precession band, or in the obliquity band (pisias & moore 1e8 1) . the complex demodulation can be used as a method to assess the correctness of a timescale. shackleton et al. (1995b) pointed out thar ir is extremely difficult to use cross-spectral analysis to investigate whether signal and forcing share the same amplitude modulation (e.g., the eccentricity modulation of the precession signal), so that this aspect of the validity of a timescale must be evaluated by complex demodulation or other means like band-pass filtering, etc. however, demodulation works only if the paleoclimatic record is modulated (as the insolation curve). then, application of this numerical technique must be proceeded by a verification of the temporal relationship between the selected climate sensitive-record and the primary forcing. in the following papers we preferred to use the similar method of the band-pass filtering technique. in the papers presented in iaccarino (2002), the spage,os software package (bonanno et ai. 1996) has been used for application of all the above-described methodologies of spectral analvsis. acknozoledgmenrs. \we thank e. helpful crìtìcal comments. this research cofin 98. erba and l. lanci for theìr has been 5upporred b1 mur.t 182 m. sprovieri, a. bonanno, s. mazzola & b. patti references berger a.l. (1978) a simple algorithm to compute long-term variations of daily or monthly insolation. contrib., inst. astr. geophys. geogr. lemaitre, unfu. cathol. de louvain,y.18, 17 pp., louvaine. berger a. (1988) milankovitch theory and climate. rev. geopbys ., 26: 621-657 . berger a. ec loutre m.f. (1991) insolation values for the climate of the last l0 m.y. quat. sci. rev., l0:297-317, amsterdam. bonanno a.,mazzoia s., sprovieri m. & patti b. (1996) spageos, a system for the anal1'sis in time-and in frequency-domain of paleoceanographic and sedimentological data. paleopelagos,6: 371-384, roma. cande s.c. & kent d.v (1995) revised calibration of the geomagnetic polarity timescale for the late cretaceous and cenozoic. j. geopbys. res., 1oo (84): 6093-6095, \x/ashington d.c. hilgen f.j. (1991a) astronomical calibration of gauss to matuyama sapropels in the mediterranean and implication for the geomagnetic polaritv time scale. earth planet. sc. letters, 1a4: 226-244, amsterdam.. hilgen f.j. (1991b) extension of the astronomically calibrated (polarity) time scale to the miocene/pliocene boundary. eartb planet. sc. letters,107: 349-368, amsterdam. hilgen f.j., krijgsman 'il{, langereis c.g., lourens l.j., santarelli a. & zachariasse \mj. (1995) extending the astronomical (polarity) time scale into the miocene. eartb planet. sc. letters,136: 495-51q, amsterdam. hilgen f.j., krijgsman \{1, raffi i., turco e. &'zacharjasse wj. (2000) integrated stratigraphy and astronomical calibration of the serravallian/tortonian boundary section at monte gibliscemi (sicily, italy). mar. micropal., 38: 1,81-211, amsterdam. iaccarino s.m. (ed.) qaa4 integrated stratigraphy and paleoceanosraohv of the mediterranean middle miocene..--.-*_'"b'_r_'/ "' rip. it. paleont. strat., l08: 175-353, milano. laskarj. (1988) secular evolution of the solar system over 1o million years. astron. astrophys., 1.98: 341.-362, vashington d.c. laskar j. (1990) the chaotic motion of the solar system: a numerical estimate of the size of the chaotic zone. icarus, 88: 266-291, orlando. laskarj.,joutel f. & boudin f. (1993) orbital, precessional, and insolation quantities for the earth from -20 myr to *10 myl astron. asrroplrys.,27a: 522-533, \ù/ashington d.c. lourens l.j., antonarakou a., hilgen f.j., van hoof a.a.m., vergnaud-grazzinic. & zachariasse \(j. (1996) evaluation of the plio-pleistocene astronomical timescale. pale o c e an o grap hy, l l : 39 1. 4 13, \lashin gton d. c. martinson d.g., menke \( & stoffa p (1982) an inverse approach to signal correlation. j. geophys. res., 87: 4807-48 1 8, washington d.c. pisias n.g. & moore t.c. (1981) the evolution of the pleistocene climate: a time series approach. earth planet. sc. let ters., 52: 450-458, amslerdam. shackleton n.j., crowhrust s., hagelberg t., pisias n.g. & schneider d.a. (1995a) a new late neogene timescale. application to leg 138 sites. proc. odp resuhs, 138 73-101, college station (tx) shackleton, n.j., hagelberg, t.d. & crowhurst, s.j. (1995b) evaluating the success of astronomical tuning: pitfalls of using coherence as a criterion for assessing pre-pleistocene times c ales. pal e o c ean o grdp hy, ia : 69 3 69 7, vashington d.c. shackleton n.j. ec crowhrust s. (1q97) sediment fluxes based on an orbitally tuned time scale 5 ma to 14 ma, site926: appìication to odp leg 154 sites. in: shackleton n.j., curry \í8., richter c. and bralower t.j. (eds.) proc. odp resuhs., l54: 69-82, college station (tx). sprovieri m., bellanca a., neri r., mazzola s., bonanno 4., patti b., & sorgente r. (1999) astronomical calibration of late miocene stratigraphic events and analysis of precessionally driver paleoceanographic changes in the mediterranean basin. mem. soc. geol. it., 54 7-24, roma. rivist;r italian;r di paleontologra e stratisrafia key uords; anornoepus ichnogenus, lower jurassic, hettangirn, sinenrurian, l-avini di marco tracksite. riasslnúo. 11 sito p;rleontologico dci lavini di m:rrco, presso rovereto (trcnto) riìppresenta un yasto tida/ flat fossile di età giur;lssico infcriore (formezione dci calcari grigi membro jnferiore; hett:rngiano sinemuriano inf.). un larso insieme di improntc riferibili a dìnos;ruri, vi ò steto scoperto pochi anni fa cd esso è attualmente oggetto di estensivi studi icnologici e paleobiologici. lc inrpronte oggetto di questa nota sono riferibili al piede dcstro e sinistro di un medesimo individuo posti jn posiz-ionc affiancat:r e subparallela in ur.r attessiarnento di erresto e accucciamcnto. le orme sono attribuibili ad attometepus hitchcock 1848. tra le orme finora docunrentrte le mrggiori effinìtà corrispondono t moyenisauropus tlodai ellenbcrger 197'1 (.recte anontoepus detdai olsen & gelton 198'1) del lcsotho mentre diffenscono dalle forme attribujte ad anontoepus identificate nell'emisfero settentrionale. questo potrebbe confcrnr.rre la p:rcntcla gondwanica per l'icnofeuna clì rovereto già supposta nello studio di altri titxf,. abstract. the palaeontologic site at lavini di ìúarco, near to rovereto (trento), reveals a wjde fossil tidal flat of early jur:rssic :rge (calcarì grigi formations lorver member; he ttangiln to lol.er sjnemurian ). an extensive sct of dinosaur prints were discovered a few vears ago and are non the subject of ichnologrcal and paleobiological stucljes. the prints rvhich arc dcscribed ìn present short note are believed to represent the right and ìett foot of thc same individual, set in a side-by-side, sub-parallel, sirting posrur.. thc prints can be classified. as anomoeplts hitchcock 1848. ar-nongst recogniscd ichnospecies, most of thc chrrrchterìsric' of the prints here describecl poìnt to ùloyenìsauropus tìodai elenberger 1974 (recte artotnoepus dodai olsen & gelton 198a) oi l.esotho. 81' contr:rst, the prints found rt lavini dì marco diffcr {rom the anornoepus for-rnd in the northcrn emisphere. these characters would seem to confirm the gonclr.anic origìns of the rovcreto ichnofauna as previouslrsupposed from thc studlof other tax:r. lntroduction. the lower jurassic tracksite at lavini di marco, near ro\rcreto (northern italy) is a large , fossil tidal flat aprile 2001 (lanzinger & leonardi 7992; leonardi & lanzinger 1992;leonardi & avanzini 1994; avanzini er ^1.1997). the dinosaur footprints were discovered a few years ago and have been the subiect of extensive ichnological and paleobiological studies (lanzinger & leonardi 1992; leonardi ec avanzini 1994; dalla vecchia 1994; leonardi ec mietto in press). the paleoenvironment at lavini di marco, as deduced from sedimentology and geochemistry, is a relatively arid coastal zone lacking perennial freshwater reservoirs (avanzini et à1. 1997). there is evidence of fluctuations in vegetation density and difference o\rer time documented by the occurrence of paiynomorphs, among them corollina sp. and porcellispora sp., (avanzinl et al. luuu). regional scale stratigraphic and sedimentological characteristics allow reconstruction of the early jurassic settinf, of a coastal environment that probably stretched southwàrds. the coastal belt was à tongue of tidal flats ltrento platform). bordered to the west and to the east by basins developed as a consequence of the foundering of the lombardy basin to the west and the belluno tough to the east. trackways and footprints nost commonly occur on six layers within one inter-to supratidal interval charecterized by vlriable lateral thickncss and facies (lower member of calcari grigi formation, hettxngian to lower sinen-rurian in age). detailed studies of the tracks are ongoing. with the present state of knowledge. tridactyl footprints are represented by theropod of grallator hitchcock, 1858 and ewbrontr:s hitchcock, 1845 ichnotaxa. other tracks are attributed to primitive sauropods (parabrontopodus lock\ey, farlow & meyer, 1994 and breztiparopus dutuit & ouazzou, 1980, ichnonota breve short notf first report of sitting anomoepus tracks in european lo\íer jurassic (lavini di marco site northern italy) marco avanzini'i, gerard gierlinski':'!e{ giuseppe leonardi)t':r'' recei.':ed scprember 23, 2aa0: acceptetl january 1 5, 2aal 'i museo tridentino di scienze naturalì, r.ia calcpìna 1'+, 38100 trento, itaìv avanzini(rr-ìmtsn.tn.it ':': panstwory instvtut geologicznr'-, ul. rakon'iecka '+, pl 00-9/5 -warszawa, poland )ijiji museo trìclentino di scienze nrturali, via calepina 14, 18100 trento, italy; present address: via modiglìani 2,e00/0 lv{onterusciello 1\a7. irr'r gi.leorrrji,,ìibcro.ir t32 m. artanzinì, g. gierlinski & g. leonardi genera) and small ornithischian dinosaurs (leonardi & mietto, in press). this association is similar ro orhers found in the lower li.rssic. systematic lchnology. the footprinrs rhat are the subject of the present report are imprinted on the surface of stromatolitic bindstone level (layer 106). this layer was dolomitized in the earll' stages of diagenesis under semi-arid climatic conditions, where evaporation exeeded precipitation. in particular the trampled layer was characterized by a succession of plastic semiliquid mud and partially lithified sediments, capped by elastic cyanobacterial laminites (avanzini 1998). the footprints are poorly preserved, as commonly occurs for footprints impressed in carbonate sediments. poor prese rvation is also due to karst corrosion of the surface. order ornithischia seeley, 1888 ichnofamily anomoepodida lull, tso+ ichnogenus anomoepus hitchock, 1848 description. the prints belong to the right :rnd lcft foot of the same individual, and are arranged in a side by side, and diverging slightly distally. unfortunately the left footprint is partly eroded, but three digit impressions and the outline of the heei are visibie. the right footprint, which is better preserved, is clearly trid.rctl-ì, and there is marked elongation close to the axis of the third digit, which can be interpreted as the impression of the metatarsal area (fig. 1). pes: tridactyl with digits ii-iv impressed. low values of projection of digit iii beyond ii and iv(2.5 cm). rì'ihe pes is longer thrn wide. digit iv is longest. footprint length, 12.3 cm; with the metapodium, 24.9 cm. footprint width, 1 1.i cm. divarication of digits ii-iii, 34'; iii-i! 23"; iiiv 55'. ler-igth of digit ii,7.5 cm; of iii, 9.5 cm; of i! 10.0 cm. phalangeal pads are not well defined. metatarsalphalangeal pads of digits iii and iv made rwo isolated posterior impressions. on the distal end of digits are preserved impressions of stumpy and rounded claws. the metapodium print js 12.6 cm in length and 4.5 cnr in width. the proximal end shows a marked asymmerry with the lateral part being larger than the medial. manus; close to the right foot, the impression of three parallel furrows set in front of an elongated depression can be recognised and in a cenrral position between the rear limbs there is another print in which two probable short, and robust digits are visible. these incomplete prints are ,1.5-5 cm long and 4-5 cm wide. it ls not corrìpletely clear whether or not these tracks can be attributed to the front limbs. however, some of these (i.e. the central print) show characteristics which indicate that the prints may have been left by the hands, when the subject was exploring the ground before stopping. comparison. the size and general morphology of these footprints assign thern to ichnotaxon,4nomoepus hitchcock, 1848. the n-rain features of ichnogenus anomoepus àrei broad pes with toes tending to be broadly splayed (divarication ii-iv zo'), relatively large width of digits, lon' values of projection of digit iii beyond the other toes, digit ii, iii and iv subequal in length but the pedal digit iv is always slightly longer than others (olsen & galton 1984; demathieu 1990; thulborn 1990; farlow & lockle;. 1993). the manus impression r'hen present, has four or five subequal toes.and is sometimes rot:lred outrvards (olsen & galton 1984). anomoepus trackwàys are usually bipedal. some tracks shows that the animal sometimes switched from bipedal to quadrupedal progression and even ro a stationarv crouching position. in resting tracks all four feet, along with the metatarsal and sometimes the rschiadic callositv and the belly x1. impressed. the size of the feet, the relative length of the digits and the metatarsal lengths of the specimen described, are similar those of some african specimens as i.e. moyenisauropus dodai ellenberge r 1.974 (recte anomoe, pus dodai olsen 8r g:rlton, 1984) : l:14 cm, 24cm with heel, \f = 05-115 cm, digit i (non functional), digit ii : z.o (with metatarsal-phalangeal pad : 1o cm), digit iii : 9 cm (u.ith rnetatarsal-ph:rlangeal pad :1o.5cm), iv: 10 cm. projection of digit iii: 4.5 cm (ellenbcrgcr 1971).the total angle of divarication ii-iv is similar to that described by ellenberger (1 971) for the typc of this ichnospecies; in track 70", in resting pose 55", (ii-iii, 30', iii-iv 4o') . typical north american rnaterial differs from our specimen in having shorter digits end genfig l -anomoepussp.{ootprintsatlavinidimarcorcvealevidenceofcrouching(a).specificallvnotethemetatiìrsal printsarranecdinaside b1' side, and dìverging slightly distallv (b). the right footprint (d), shows the main features of ichnogcnus,4 nontoepus: broad pcs *,ith digit li' iii and iv subequal in length and broadll'splaved, relativclv large r-idth of digits end iowvalues ofprojection of.1;gi, rtt beyond the other toes. intcrpretetive drawing shows other prints on the same surface (c). somc of tl.r. r.r-,,"t h".-. b.", left ùy the hands. scale bar : 10 cm. s ittìng anomoepus tracìes ùa: i' | .r / ,--\-\cd,é ) i .-l iaij"c '.--l \ \/ \ t'1.-j' ,.1 lz\ / t --/ \j \ i-j\j' ' ,\,y i\j ... ,'{,r ._r_ \ \'-ll.. tu i't,,, c; o , -,.-...(] ? i_ \\{ * \!\:\.. ..1j 131 m. avanzini, g. gíerlinslei €: g. leonardi fig. 2 ncrv spccimen of anontoepus pìenleot-skii gicrljnski, 1991 (nluz. pig 1662.1l1) from the hettansian of gili;rnrlas (poland), which q'as ìeft during a plantier;rde gait. scale b:rr -5cm. erally smaller dimensions, but divarication angles ii-iii and iii-iv are sin'rilar ro rhose of anomoepus scambus hitchock, 18,+8 (35',23'). among the northern hemisphere anomoepodicls, the most similar form is anomoepus pienkow,s,èii gierlinski, 1991from the lower jurassic of poland. fiowever, polish anomoepus rracks have a more robust ge neral structure u.ith the shorter metatarsal imprints, which appared in the new discovered specimen muz. pig 1662.ii.t (fig. 2). discussion. anomoepus is a distìnctive classic earlv jurassic track type first reported from the connecticut valley (hitchcock 1848; lull 19a4, d53; olsen 1980a; olsen & galton 1984) but now also known from europe, africa and australia (olsen & galton 1984; thulborn 1994; lockley 8c hunt 1995; lockley 8c meyer 2ooo). the early jurassic assemblage from the newark supergroup and glen canyon group of north america is predominantly composed of late triassic survivors (mainly large sized grallator, and exrremely. rare rhyncbosauroides), with the addition of anomctezzs (welles 1921; olsen 1978, 1980 a-c; clark & fastovsky 1986; olsen & padian 1986; olsen ec baird 1986, silvestri 1ee6). the ichnoge n:us anomoepzs is an abundant ichnite in the lower jurassic stormberg group of lesot}ro (ellenberger 1972,1971; olsen er galton 1984; colbert 1986; haubold 1986) and early jurassic prints, attributed to small ornithischian dinosaurs (anomoepus gracil lintus) have been reported in the carnarvon gorge, quecnsland (thulborn 1994). anomoepus is :rlso present in central europe. the polish ichnofauna described by gierlinski (1991, 1994, 1995, 1,996a-b, 1.997), gierlinski & sawicki (1998) and gierlinski & pienkowski (1999), comes from the norrhern slopes of the holv cross mountain, located about 150 km sourh of \fars:rvi from the zagtje formation, skoloy formation and przl,sucha ore-bearing form;tìon. floristìc remains and sequencc strarigraphy correlation indicate a hettangian age of all these formations. thel' represcnr various continental and marginal-marine environmenrs. fluvial and lacustrine scdiments dominate in the contincnt:rl los'er ztga,je form:rtion, n,hile the ncarshore and deltaic facies arc present in the sholoy formation, uppcr zagale formation and przl-such.r orc-bearing formation. various ornithischi;rn) s.ìuropod and the ropod rracks occur in these sediments (gicrìir-rski er pienkon'ski 1999; lockley et meyer 2ooo). the presence of early ornitischians in europe of is particularll' significant ro inrerpretation of the lavini di marco ichnoassociation. gierlinski (1991, 1999) contends that the polish ichnospecies of anomoepus pienkctvsleiì differs fron'r the classic connecticut forms in beine lrrger .rnd having a larger manlìs print. however, anomoepus pienko.usleii, gierlinski, 1991, differs also from thc prints described in this reporr. taking inro accounr the distribution of anomoepus, and considering the concept that early jur:lssic faunas are cosmopolitan (hunt et al. 1996), it is interesting to note that the anc.tmoepus obse rved at lavini di marco is the most similar to the forms documented in lesotho, slightly different from thc polish specimens and substiantialiy distinctive among the other northern hemisphere anomoepodids. this fact could suggest that the ichnofauna of rovereto may have gondwaniln origins, as previously postulated by leonardi and mietto (in press) from the study of other taxa. it is also note$/orthy that the hettangian tracks of poland, robust anomoepodids and sauropod tracks (gierlinski 1992; gierlinski & pienkowski 1999; gierlinski & sawichi 1998), remain stronger similarities ro rhe distinctive italian assemblage than to other laurasian ichnofaunas. ackno.o/edgements. \we are very srateful to j. o. farloq h. frey and h. g. lockley for the scientific and linguistic revision of the manuscript and uscful suggestions thar have improved substantially this paper. 1: sitting anomoepus tracks re,fere,nces r35 avanzini m. (199s) anatomy of a footprint. bioturbation as a key to r.rnterstand dinosaur walk dynamics. ichnos, 6 (3): 129-139, chur, new york. avanzini m., frisia s., keppens e. & van den driessche k' (1997) a dinosaur tracksite in an early liassic tidal flat in northern italy: paleoenvironmental reconstruction from sedimentology and geochemistry' palaios, i l: 5j8-55 l, tul'r. avanzini m., leonardi g., n{ietto p. 8c roghi g. (2000) icnofaune dinosauriane nel giurassico inferiore della piattaforma di tento: aspetti stretigrafici, paleoambientali e paleogeogrrrficr. b)a rìunione esti''-a soc. geol. italiana. (abstract volume), 42-'14, trieste. clarkj.m. & fastovsky d.e. (1986) vertebrate bìostratigraphy of the glen canyon group in northern arizona. in: padian k. (ed.) the beginning of the age of dinosaurs: pp. 285-302, cambridge universitv press, cambridge. colbert e.h. (19s6) historical aspects of the age of dinosaurs. faunal changes across the triassic-jurassic boundarv. in: padian k. (ed.) the beginning of the age of dinosaurs: pp. 9-20, cambridge university press, cambridge. dalla vecchia f.m. (1994) jurassic and cretaceous sauropod evidence in the mesozoic carbonate platforms of the southern alps and dinarids. in: lockley m.g., dos santos vf. meyer c.a. & hunt a.p (eds.) aspects of sauropod paleobiolog,v. gaia, la: 65-73 lisboa. den"ratl'rieu g. r. (1990) problems in discrimination of tridactyl dinosaur footprints, exernplified bv the hettangian trackways, the causses, france. ichnos, l: 97-11a, chur, new york. dutuit j.-m. er ouazzou a. (1980) décour.erte d'une piste de dinosaure sauropode sur le site d'empreintes de demnat (haut atlas marocain). mém. soc. géol france, n.s., 139:95-102. paris. ellenberger p. (1972) contribution à la classification des pistes de vertébrés du trias: les types du stormberg 'd'afrique du sud. (i partie). paleoztertebrata, mém. extraord. 1972, pp. i -1 i 7, montpe llier. ellenberger p. 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(1996a) avialian theropod tracks fron-r the early jurassic strata of poland. zubia, 1'4: 79-87, logrofro. gierlinski g. (1996b) dinosaur ichnotaxa from the lower jurassic of hungar. geol. quart., a0(1): 119-128, \farszawa. gierlinski g. (1997) sauropod tracks in the earlyjurassic of poland. acta palaeont. pol., 42 (a): 533-538, \farszawa' gierlinski g. (1999) tracks of a large thyreophoran dinosaur from the earlv jurassic ol polend. acta palaeont pol., ++ (2) : 23 1 -23 4, \(/ars zawa. gierlinski g. & pienkowski g. (1999) dinosaur track assemblages frorn the hettangian of poland. geol quart., $q): a3-a5, \ilarszawa. gierlinski g. & san'icki g.(i998) new sauropod tracks from the lower jurassic of poland. geol. qwart., 42(4'): 177480, lvarszawa. haubold h. (1986) archosaur footprints at the terrestrial triassic-jurassic transition. in: padian k. (ed.) the beginning of the age of dinosaurs: pp.189-2a1, cambridge {j nio ersity pre ss, cambridge. hitchcock e. (1845) an attempt to name, classify, and describe the animals that made the fossil footmarks of new england. proc. 6th annual meeting, am. ass. geol. naturalists: pp. 23-25, new haven, connecttcut hitchcock e. (1848) an attempt to discriminate and describe the anirnals that made the fossil footmarks of the united states, and especially new england. mem. am' acc. art. science,3 (2): 129-256, new har-en. hitchcock e. (1858) ichnology. a report on the sandstones of the connecticut valle1', especially' its fossil footmarks. v of 220 pp., boston. hunt a.p, lucas s.g., huber p.& lockley m.g. (1996) faunal evolution in l,,lte triassic, nonmarine tetrapods. in: aspects of triassic-jurassic rift basin geoscience, state geol. nat. hist. suro. connecticut, nat. resources center, dep bwlletin 26, p.57, connecticut. lanzinger m. & leonardi g. (1992) piste di dinosauri nel gir-rrassico inferiore ai lavini di marco (trento). in: aa.w dinosaurs. il mondo clei dinosauri: pp. 88-94, museo tridentino scienze natur:rli, trento. leonardi g. & avanzini m. (1994) dinosauri in it:rlia. ie s cienze, quaderni, 7 6: 69 -81, milano. leonardi g. & lanzinger m. (1992) dinosauri nel trentino: venticinque piste fossili nel liassico di rovereto (trento, italia). paleocronache, 1992(1): 13-24' milano. leonardi g. & mietto p (in press) dinosauri in italia: le ornre giurassiche dei lavini di marco (trentino) e gli altri resti fossili italiani. v of 496 pp. accademia editoriale, pisa. lockley m.g., farlowj.o. & mever c.a. (1994) brontopodus and parabrontopodzs ichnogen. nov. and the significance of wideand narrow-gauge sauropod trackr'vays. gaia,8: 1-10. lisboa. 136 m. aoanzini., g. gierlinski g g. leonardt lockley m.g. & hunt a.p (1995) dinosaur tracks and other fossil footprints of the wesrern united states. v of 33g pp., columbia university press, new york lockley m.g. & meyer c.a. (2000) dinosaur tracks and other fossil footprints of europe. y. of 323 pp., colun.rbia university press, new york. lull r.s. (1904) fossil footprints of the jura-trias of north america. mem. boston soc. natural history 5(11): pp. 461-557, boston. lull r.s. (1953) triassic life of the connecticut valley. bull. connecticut geol. nat. hist. suroey, 81: 1-336, hart_ ford. olsen pe. (1978) on the use of the term newark for triassic and early jurassic rocks of easrern north america. n ew s i e tter s tratigraphy, 7 (l : 9 a -9 5, berlin (usa). olsen pe. (19boa) a comparison of the verrebrate assemblages from the newark and hartford hasins (early mesozoic, newark supergroup) of eastern north an.rerica. in: jacobs l.l. (ed.) aspects of vertebrare history: pp.35-53, museum of northern arìzona press, fìagstaff. olsen pe. (1980b) triassic and jurassic formations of the newark basin. in: manspeizer \m (ed.) field studies of the newjersey geology and guide to field trips: pp.239, geol. dept. newark college of a.ù 5., rutgers univ. newark, nj. olsen pe. (1980c) fossil great lakes of the newark supergroup in the newjersey. in: manspeizer \xl (ed.) fieìd studies of the new jersey geology and guide to field trips: pp. 352-398, geol. dept. neutark college of a t" 5., rutgers univ. newark, nj. olsen pe. & baird d. (19s6) the ichnoge nus atreipus and rts significance for triassic biostratigraphy. in: padian k. (ed.) the beginning of tl-re age of dinosaurs: pp.61-gg, cambridge university press, cambridge. olsen pe. and galton pm. (1984) a review of the reptile and amphibian assernbiages from the stormberg of southern africa, with special en-rphasis on tl-re footprints and tl,re age of the stonlberg. palaectntologia africana,25: 871 1 o, johannesburg. oisen pe. & padian d. (1986) earliest records oí ijatrachopus f rctm the southwestern united states, and a revisior-r of some ear.ly mesozoic crocodyìomorph ichnogenera. in: padian k. (ed.) the beginning of the age of dinosaurs: pp. 259-274, cambridge university press, cambridge. silvestri s.m. (1996) ichnite record of terrestriai tetrapocl abr-rndance and diversitv through a critical period in earth histor1., jacksonvald syncline, newark basin, penrrsylvania. state geol. nat. hist. surrt. connecticut, nat. resources center, miscel. reports 1: p.43. thulborn a. (1990) dinoseur tracks. v of +10 pp., chepmen and hall, london, new york. tlrulborn a. (1994) ornitl.ropod dinosaur tracks from the lower jurassic of queer-rsland. alcheringía, 18: 247-258, melbourne. \x/elles s. p. (1971) dinosaur footprints from the kayenta formation of nortl-rern arizona. plateau, 44: pp.27-3g, flagstaff. luglio 2000 a late triassic ostracod assemblage from the quattervals nappe (austroalpine, northern italy) sylvie crasquin-soleau 1, fabrizio berra 2 s. robemo rettori3 receit,ed february 15,200a; accepted april 3,200a key-u,ords: late triassic, ostracods, stratigraphs austro:ìpine. p:ìaeon toìogy. riassunto. la successione norica della falda quattenals (austroalpino centrale, italia settentrìonale) registra una evoluzione stratigrafica da facies di piattaforma carbonatica di n.rare basso soggetta a fenomeni d:i dolomitìzzazione precoce (dolomia princiaplehauptdolomit) r'erso facies dì transizione (formazione di pra grata), rappresent:rte da alternanze di dolomie e calcari scuri e brecce dolomitiche di pendio. la successione evolve, poi, verso facies di bacino intrapiattaforma (calcare dì quattervals), rappresenrate da calcari scuri con subordinate intercal:rzioni di brecce contenentì clasti con facies di piattaforma e lìvelli paracongiomerati. le facies bacjnali e transizionali consistono prevalentemente di materiale risedimentato daila piattaforma; sono, inoltre, presenti live lli di materiale detritico fine c orizzonti microbialitìci . le facies bacinali contengono abbondante materiale bioclastico, all'interno del quale è stato possibile riconoscere un'interessante associ:rzione ad ostracodi, costituita da numerosi individui appartenenti a poche spccie, che denota condizioni ambicntali poco favorevoli carxtterizzate da variazioni di salinjtà e scarsa ossigenazione. nonostante la intensa tettonica alpina e la diffusa ricrisrallizzazione, i'analisi del materiale raccolto alla base del caicare di quattervals, ha permesso dì identificare, insieme a forme eià note, lc due nuove specie di ostracodi rbombocythere dimorpbica e kerocythere quattercalsi. abstract. the up to 1200 m thìck norian succession of the quatteruals nappe (central austroalpine , italy) documenrs a tectonically-driven passage from intertidal facies (dolornia prìncip:lehauptdolomit), represented by early-dolornitized bedded ro massive gray dolostones, to intraplatform basin sediments (pra grata formation and quattervals limestone), consisting of resedimented dark limestones and thin bedded laminated limestones, alternatìng (maìnly in the pra grata fonlatìon) with intrafonnational breccias contaìning abundant shallow-water facies. the transitional and basinal facies contain an interesting ostracod assembìage, beside bioclastic layers containing shallow-water and upper slope skeletal graìns. thc ostracod fauna is rich in specimens belonging to few genera, denoting restricted environmental conditions characterized by variations of salinity and lon oxygenation. dcspite the intense deformation and recrystallization related to the alpine tectonics, the paleontological analysis of the ostracod assemblage from the base of the quattervals limestone, allowed the identìfication of the two new species of osracods rhombocythere d,imorphica and kerocytbere quatten:alsì in association with other already known forms. introduction and geological setting. despite the strong recrystallization related to the alpine tectonics, an interesting ostracod assemblage has been found in the norian carbonates of the quatterwals nappe (upper austroalpine, eastern t.ombardy, italy; fig. 1). the sedimentary succession of the quattervals nappe is exclusively represented by a late tiassic (mainly norian) up to 1200 metres thick carbonate succession (hess, 1953; somm,1965;berra,1.995) (fig. 2). the lower formation of the quattervals nappe is represented by dolomitized inner carbonate platform facies referred to the dolomia principale-hauptdolomit. this formation passes upwards to the pra grata formation (hess, 1953), characterised by alternations of limestones and dolostones. upwards, it is covered by the n'ell-bedded dark limestones of the basinal quattervals limestones, having at the top about 60 metres of marls (crappa mala beds; furrer, 1983). above, a recovery of shallow-water carbonate platform is documented by the murter dolomite (furrer, 1983). the top of the norian succession is represented by subtidal facies belonging to the murter plattenkalk. the stratigraphic boundary to the overlying rhaetian kossen formation was documented by somm (1965) both in the quattervals nappe and terza unit. stratigraphy. in the studied area, only the lower part of the quattervals nappe succession is preserved, from hauptdolomit to the upper quattervals limestone. the thickness of the hauptdolomit, whose base is lacking, is about 400-500 m and it is mainly represented by irregui cnrs esa 2073, upmc, dtp. géologie sédimentaire, t. 15-25, e.4, case 104, 4 place jussieu, /5252 e-mail: crasquin@ccr. jussieu.fr 2 regione lombardia, ufficio progetti speciali per la geologia e la sismica, yìa f. frlzi 22, e-mail: fabrizio-berra@regione.lombardia.it 3 dipartimento scienze dellaterra, piazza dell'università,06100 perugìa, italy; e-mail: simocir@unipg.it paris cedex 05, france; 20124 milano, itaiy; 182 s. crasquin-soleau, f. berra & r. rettori fìo i lar cy.clic alternations of fine grained dark-grey dolostones and doloarenites, both in centimetric and plurimetric layers, with locally intraformational breccias (somm, t965;berra, 1995). doloarenites mainly consist of often normal grading intraclastic packstones, rarely bioclastic or oolitic; ripples and cross laminations, locally burror.ed, are also frequent. stromatolitic horizons (up to more thf,n i m thick) are also common. emersions are documented by mud cracks, authigenic evaporitic crystals and fenestral fabric. in the upper hruptdolomit, the occurrence of sedimentary dykes and breccia bodies documents the beginning of the tectonically induced platform drowning leading to the developmenr of the quatterwals basin (berra, 1995). the hauptdolomit of the quattervals nappe was deposited in a shallow water basin under low dysoxic conditions with local and short-term emersions. bioclasts are represented by pelecypods and gastropods and rarely by dasycladacean algae and foraminifera. the overlying pra grata formation whose thickness increases from 30 to 200 m souths/ards (berra, 1995) is characterised by irregular alternations of limestones and dolostones. their abundance varies along the succession: dolostones prevails at the base, while limestones become more important toward the top. limestones consist of fine grained, biociastic, commonly thin bedded calcarenites, often with millimetric marly horizons. locally slumps are present. in the upper part, intercalations of paraconglomerate beds up to 3 metres thick often occur. the dolostones consist of generally legend i i m. motto/allgàu fm. i i fraele/r\ossen rm. [-tl ^,.^*^-..^r^ t :*^^+^ -l vudlletvals llmestone t--l ^| | t.ra urata fm. t-----^ ,| .l l)olomia del crisrallohauptdolomit f_.l bu.n-n.,i s su-ntins.itn 2km light-gray chaotic breccia bodies (up to 10 m thick; sometimes amaìgamated, containing carbonate platform derived clasts. centimetric to decimetric doloarenite beds are common. alternations of dolostones and limestones are indicative of the existence of an adjacent submerging carbonate platform of dolomia principalehauptdolomit. muds and fine calcarenites were exported by currents or storms from the surrounding carbonate platform, as documented by microfacies often rich in platform-derived shallow water allochems. dolornitic breccias were directly derived from the dismantling of dolomitised and lithified carbonate platform facies, probably exposed along scarps (berra, 1995). the large number of breccia bodies and the occurrence of slumps indicate that pra grata formation deposition occurred on a developing slope. both the lower and upper limits of the formation are transitional. the quattervals limestone (600-650 m thick) is almost entirely represented by dark, bedded mudstones and wackestones, with marly joints and frequent intercalations of graded packstones yielding platform-derived crtins lsnmm lgas'r.'"' lgq(ì f)..'.^--ld-*... .. -_, jcrra, t11j). t à.rjeongtomefate in up to 4rn thick beds, and more rarely breccias, are intercalated. the upper limit with the crappa mala beds (obere mergel; somm, 1965) has not been obser-ved in the studied area. the quattervals limestones were not deposited on perfectly flat-lying sea-bottoms: slumps, overfolds and slump scars document deposition on a low-anglei slope. microfacies analysis shov'ed rwo generically different deposits: intraclastic-bioclastic limegcological n-rap with the location of thc position of the sampling site. the structural sketch refers to an area slightl,v larger than that of the geological raap. l'.' jqt' ,*\t a. \j cdefopel ì,aì aoro de covo t a:i ó" ,.,ts o late ti'iassic ostracctcl assemblage 2 -val trenzeira tagliata ffi h [.-.---l li-------\------#,ht1 /, l-'i-r-il .f^1 ./ 183 trio ) stones directly derived from sedimentation of platform material and limestones derived from decantation of fine grained material and represented by mudstones and finegrained wackestones. smal1 foraminifers, sponge spiculae, rare ammonoids and calcified radiolarians are present. the dark colour of the sediments and the absence of bioturbation document unfavourable conditions for the development of diversified benthic communities on the bottom of the quattervals basin. the evolution of the stratigraphic succession allows reconstruction of an asymmetric intraplatform basin controlled by syndepositional extensional tectonics, with a north-south width of at least about 8 km and an east-west width of some tens of km (berra, 1995). the sedimentary input was twofold, with breccias mainly from southern high and limestones from sweeping of the two margins. the southern margin of the basin was tectonically-controiled, whereas the northern was a flexural-type margin: the depocentre was asymmetric, and situated near the southern margin (berra, 1995). facies analysis demonstrates that the quattervals basin was a partly isolated sea-way; water exchanges with a wider marine basin occurred only for the surface water, while the bottom conditions were dvsoxic to anoxlc. 3 diga del gallo 4-m.la /s ennsso dei corno palaeontology. despite the intense and polyphasic alpine tectonics and the strong recrystallisation locally destroying the original features of the sediments, ostracods occurrence has been documented in ail the three forrrations considered here. ostracods are particularly abundant in the fine-grained calcareous facies of the pra grata formation and quattervals limestone, generalll. associated with rare specimens of norian-rhaetian foraminifers referable to gandinella falsofriedll (salaj, borza 8c samuel) . the present paper is focused on a rich ostracod assemblage from a calcareous outcrop stratigraphicaily referred to the pra grata formation (coord. gauss boaga 1588, 510-5159, 250). the assemblage has been recorded from well-bedded black limestones consisting of recrystallised packstones-wackestones. the ostracod-bearing facies are mainly represented by dark resedimented fine-grained calcarenites and by dark microbialites, where ostracods are trapped between the organic laminae. the sedimentological features indicate that the ostracods probably lived within the mic,robialitic mats, which developed in subtidal conditions on low-oxygenated bottoms. i -v delcantone (tl o 3 l#l t | | lquohervolslimestone9f-?r-r.a sompling slteta (urot'nr:::.., posirion) ha\n poroconqlomerotes |-;.=n slumps ffi breccios t----f------l " l-r----'--i i"\< | * -'.. -........... 2200 re [:"ff:i:"j1"3::;") 'cablecar/chairrirt a ksíx ht"il"?"',;,$:,i# '",:::," f{| o-g) saîilirgto:eic.s --g-.form top and slopes, and which are present throughout the studied succession (maurer et ai. 2002). most samples derive from the seceda core, which \il.as recovered in proximity to an age-equivalent carbonate platform (fig. 1b). additional samples were taken from outcrop sections around the well site that cover a short stratigraphic interval missing in the core. foraminiferal assemblage and diversification. in the analyzed thin sections foraminifera afe present as part of the debris shed from the adjoining carbonate platforms and slopes by turbidity currents (maurer et a1. 2oo2). throughout the core we encountered platform-derived benthonic foraminifera in association with calcareous hyaiine (lagenina) foraminifera of open-marine environment (fig. 2). the assemblages attributed to a shallow-water environment are characterized by the constant presence of unilocular, tubular foraminifera referable to the genus earlandia plummer, in association with microgranular multilocular foraminifera with elongated morphology belonging to the genera endotebanella yachard et ai. endotriadella vachard et al., and agglutinated foraminifera such as ammobacwlites cushman, reo\itax de montfort and palaeolituonella bèrczi-makk. less frequent there occur trochospiral and streptospiral types of the genera endoteba and glomospira. furthermore porcelaneous foraminifera are represented by ophthalmidium sp., arenovidalina chialingcbiangensis lho, hoyenella gr. sinensis (ho) andagathammina sp. throughout the whole succession v/e noticed the presence of originally aragonitic foraminifera of the family duostominidae brotzen with the genera variostoma kristan-tollmann, d uo stomina kristan-tollmann and krikoumbilica he. the sample 69.90 marks the appearance of the originally aragonitic family involutinidae; upsection there follows a diversification of these types, represented by the lenticular genvs aulotortus \feynschenk and the cone-shaped genvs lamelliconus piller (fig. 2, 3, 4). elongated calcareous hyaline foraminifera include the genera nodosaria lamarck, pseudonodosaria boomgaart, frondicularia defrance, astacolws de montfort and lenticwlina lamarck. the distribution of foraminifera along the studied section (fig.2) shows phases of increase and decrease in the diversity (: richness) of the microfauna both at generic and specific levels. a general increase is given from the base of the section until sample 81.00. in the upper part of the curionii zone follows a decrease in the fig. 2 distribution of foraminifera found in 224 thin sections in the seceda core along with a lithological log of the 20-103 m core interval. the range of ammonoid zones and current candidates 2 (sensu krystyn 1983) and 3 (sensu brack & rieber 1993) for the definition of the anisian/ladinian boundary are indicated on the left. important macrofossils found in surrounding outcrop sections are indicated on the right (after brack & rieber 1993, brack et al. 2ooo, brack et al. 2oo1). the lithoiogy of the core is explained in inset on fig. 3. important macrofossil occufrence 393middle triassic foraminifera from dolomìtes t. i al. a i t t, i i .t' t daonella pichlei protrachyceras sp . daanella taramellii daonella maussoni eooptott achyceras d. recubariense chiose,beras chleserse fuevadlfes secédensis, daonella cl. golana stap pa n i ce ra s evo i ut u m, ncinites dalamiticus daonel{a elongata group daonella cernercensis, aplococeras sp. i rr .t daanèlla lommeli group, daanella tyralensis ì: èè ìì ff,i 2 i =q j 394 presence of foraminifera and a total absence in the interval between samples 72.94 and 70.64. above this level a new increase in diversification is present, dominated by the first appearance of the representatives of the family involutinidae bùtschli from sample 69.90 upwards. in the gredleri and arcbelaus zones the microfauna does not show any major decrease in diversification. only the interval 64-61m, which is characterized by very fine-grained (<1.77 micron) turbidite beds, lacks foraminifera (fig. 2). correlation of this interval with outcrop-equivaients revealed the presence of foraminifera in coarser grained portions of turbidites closer to the platform (fig.3). in the outcrop we also sampled the interval of section 10, illustrated in fig. 3' and missing in the core due to local tectonic unconformities. the outcrop samples show the same assemblages as the surrounding samples in the core close to this interval. in general v/e noticed a low faunistical diversification compared to the foraminiferal assemblages in the anisian or carnian, although there is a good state of preservation in the analyzed samples. stratigraphic implications. the well constrained biostratigraphic age of the section allows a more precise calibration of the first and last appearence of some of the described species. due to an accurate core-to-outcrop correlation a direct comparison of the microfauna with the occurrence of age-diagnostic macrofossils in the outcrop is possibie (fig. 2). the sample 95.06 shows the presence of meandrospira dinarica kochansky-devidè & pantic (fig' 4)' a well known species in the anisian of the tethyan domain (rettori 1995, cum bibl.). this species occurs in the reitzi zone, but does not show up anymore further up in the section; it is therefore likely to have died out in the reitzi zone. the species arenooidalina cbialingchiangensls fio, so far known as middle triassic in age (rettori 1.995, cum bibl.) is limited to the reitzi zone as weli and disappears above the sample 93.09. the species variostoma aha kristan and hoyeneìla grsinensis are present in the reitzi zone and both do not superate the curionii zonein age in the studied succession. palaeolituonella meridionalis bèrczi-makk disappears at the base of the gredleri zone. the biostratigraphic most important event occurs from leve1 69.90 m upwards with the appearance of the genera larnelliconus and aulotortus (fig.2, 4). these involutinidae are represented by the species l oentroplanus (oberhauser), l. mwltispirws (oberhauser), ,4. aff. a. friedli (kristan-tollmann) and a. gr. sinuosus \leynschenk. based on ammonoid data their appearence can be pinpointed at a level slightly older than the base of the gredleri zone (base late ladinian sensu krystyn 1983). in the buchenstein beds in the dolomites this level approximately matches the first occurrence of budwror.,ignathus hwngaricus (kozur & vegh), a conodont marker (maurer 1999). paleoecology from an ecological point of view some considerations can be made on the calcareous hyaline foraminifera (lagenina) of open-sea environment' in particular we noticed that the elongated, cylindrical morphotypes (ì)1odosaria, frondicularia and pseudonodosaria) are most frequent at the beginning or at the end of the anoxic intervals (e.g. levels 92.22 m and 64.49 m in fig. 2, level 36 m in section 2 on fig.3). this is in agreement with the fact that these morphotypes are adapted to disaerobic and low-energy environments (murray 1'991; tyszca 1993). the planispiral and biconvex strictly coiled forms such as biconvex lenticwlina, are in contrast more typical for better oxigenated substrates (murray 1991.; tyszca 1993). the fauna typical for restricted, low-energy shallow-marine habitats (i.e. backreef and lagoon) is characterized by porcelaneous milioiid foraminifera (haunold et a1.. 1997; davaud & septfontarne 1995; travè et al. 1996; fornos tt ahr 1'997) and include the genera ophthalmidium kubler & zwingli, arenooidalina ho, hoyenella rettori and agathammina neumayr. in the studied section these types are more frequent than in older anisian formations in the dolomites, where only the genus ophthalmidiwm is present (senowbari-daryan et a1. t993). in recent environments miliolid foraminifera are not limited to restricted shallow-marine areas, but occur also in forereef and slope settings (in the bahamas, e.g. rose & lidz 1'977; in the gulf of aqaba, reiss & hottinger 198a). the here described genera may therefore have colonized the same range of environments in the triassic. f. maurer & r. rettori fìg. 3 correlation of rhe 52-66 m core interval (section 1) with equivalent portions in outcrop sections 2,3 and 10, where 10 is the most proximal and 1 the most distal section relative to the nearby carbonate platform. the foraminifera found in the outcrop samples are indicated. note the occurrence of nodosaria at the beginning and end of the anoxic facies intervals (e.g. 33.8 and 36 m in section 3)' the stratigraphic column in the core and the sections 2 and 3 is dìsturbed bv local tectonic unconformities. the 35-40 m interval shown in section 10 most probably represenrs the interual missing in the core. the occurrence of the macrofos srl daonella rnoussoni mérran, collected durirg this ,trd1,, i, indicated. the layer nith accretìonary lapilli represents a unique marker horizon that can be correlated over the entire buchenstein basin within the dolomites (gianolla 1991, maurer 1999). due to the local unconformity at seceda, the lapilli-la1,er disappears in section 3 and in the core. for general correlation of core and outcrop see brack et al. (2000). i=, r@phaxsp i-= : :]-l --:)-f>:1 '/ lapilli layer -t=r iri::é:-;i#-*"componenls (oniy frg.2) ì camonate tenses baf ma*;ng laminaled, anoxic facies a6relonary lapilli €oonale nocu€s with chefr | m cneí nodrlea sno palenes i:=-:=_-:: : r=ri -=-l middle hiassic foraminifera from dolomites \ 'ab i seceda l : core-outcropcorrelation j:.r.3èji: earlandja g@cilis, e. amplr + duastamina sp. + eadandia gmcilis, e. ampliearlandía gftcilis, e. ampli!' mumtis = eadandzancils e amplr ? muahs opntnatnidiumsp = g/orcspi€//a sp. ! ealandîag6cìlís, e. afrpli= nuélis, qphthalnidium sp. = ealandia oècilís e. am1l; = = = = = = = = = = = = = = = = = = = = = = = = = ? eadandia_g?cilis .e ampl| muftxs, uuastomtna s0 , aulototlus sp eadandìa gftcilis, e ampli+ frurelis, reophax 9p , duostamina sp . nodosara sp + earlandta^g@cilts e amplimu@ils uuostomtna sp auloto ft us gt. si t1 uosu6 + ea.landia gftcilís. e amplimuftlis. duostóúina sp. 395 i duostamina 9p., eajo/ 2 leba all e. dadau\) + earlandia afrp1ímuelís reophax sp. lunucammtna sp. i lunucak*ìre sp. =t murts à = duostamína spo, n] !l ol oì (rl ::) ? earland;a ampl'mara, s -daonella moussoni iuff ayeé f"plei.a ve.de ì ealandia /nplinunlis reoplèx,ép. o (corej i___ì :=-=' :key for sections dolom[ized _ /b.e,cia(ontyf,g 2....,..i]%-'ntonto'tny -.:k r ^ r ble@ia wilh car borale reophax sp 396 f. mawrer & r. rettori fig. 4 illustration of some species of foraminifera found in the seceda core. [ meandrospira dinarica kochansky-devidè & pantic, level 95.06 m, magnification x66. 2 palaeolítuonella mínima he,level 95.06 m, magnification x106. 3 palaeolituonella meridionalis bètcztmakk, ler.el 68.92 m, magnification x8o. 4 lamelliconus mubispinzs (oberhauser), level 69.22 m, magnification x50. 5 awlotortus ex gr. sinuosus.weynschenk, level 6'1.83 m, magnification x,15. 6 aulotortus afl. a. friedli (kristan-tollmann), level 67.7a m, magnification x55. 7 lamelliconus.oentoplanus (oberhauser),level 69.90 m, magnification x58. the agglutinated and microgranular elongated genera reophax, endotriadella, endotebanella and ammobaculites are also paft of shallow-marine environments; they are suggested to live infaunal with detritalscavenger feeding habit (jones ec charnock 1985). the elongated, unilocuiar genus earlandia, whrch is quantitative the most abundant in the samples, and the originally aragonitic genera duostomina, vnriostomd and krihowmbilica of the family duostomini dae are generally common in triassic carbonate platforms. in the gredleri and arcbelau.s zones the lamellar, aragonitic foraminifera of the family involutinidae appear as additional shallow-water inhabitants. these types, represented by the genera lamelliconus andawlotortus) live in various shallow-marine environments ranging from the reef to restricted lagoon (piller 1928). discussion and conclusions. a significant event in the faunal evolution of foraminifera recorded in the buchenstein beds is the appearance of the family involutinidae bùtschli at the base of the gredleri zone (base late ladinían sensu krystyn 1983). lenticular morphotypes are considered the ancestral forms of this family at the base of its evolution, followed by conical types (salaj et al. 1983; gazdzicki 1983; di bari & laghi 1994). for instance, oravecz-schef{er (1.987) describes the appearance of lenticular involutinidae, assigned to the genera aulotorfas weynschenk and triadodiscus pilier, followed by the conical genlrs lamelliconws piller, in the carnian of the transdanubian central range. in contrast, we encountered first the conical genus lamelliconws (first appearance at level 69.90m) and then the lenticular morphotype, representedby awlotortus (fitst appearance at level 67.70m; fig. 2). we attribute this phenomenon to a lack of lenticular morphotypes due to the general poor percentage of foraminifera in the sampies, rather than to a different evolutionary trend of the family involutinidae in the buchenstein basin. the appearance of involutinidae at the base of the late ladinian as recorded here is in agreement with the general description of the evolution of this family in the late ladinian/ early carnian (e.g. salaj et a1. tls:). note, however, that the conicai morphotype lamelliconws has been described already in the anisian eros limestone of hydra (greece; rettori et al. 1,994), which has been dated as pelsonian in age based on conodonts (angiolini et al.1.992). it is not clear what favourized the appearance of involutinidae in the gredleri zone.in view of mirquez er trifonova (2000), their occurrence depends on the relative percentage of aragonite solved in the sea water. in this respect, a rise in aragonite in the sea water could have favoured the evolution of involutinidae in this part of the tethys. in addition, it could have contributed to the rise in carbonate production on the adjoining platforms (e.g. mmrer 1.999). the repeated alternation of oxic (:bioturbated) and anoxic (=laminated) facies in the buchenstein beds of the seceda core is a feature that is restricted to the area of the dolomites and may have been caused by episodic closure of the seaways of the buchenstein basin bordering the open ocean and related variations in oxygenation of the sea floor. equivalent basinal limestones in the lombardian alps (bagolino, e.g. brack ec rieber 1993) and hungary (oravecz-scheffer 1987) show bioturbated facies throughout their deposition. a comparison of the foraminifera found in the iaminated facies of the reitzi zone in the seceda core with the equivalent bioturbated interval in the transdanubian central range (oravecz-scheffer 1987) reveals no major differences in faunal composition. lagenina are represented by similar genera in both areas, indicating that these foraminifera were probably not affected by oxygenation-variations middle triassic foraminifera from dolomites 397 on the sea bottom. the fauna described by oraveczscheffer (1987) is poor in shallow-water foraminifera, probably related to a lack of turbidites in that succession. the dataset presented here is limited to the source area oî the turbidite sediments, which may not cover the whole shallo.w-water habitat of the foraminifera. the middie triassic buchenstein beds show a continuous record of foraminifera in the late anisian and early ladinian over a time interval of five ammonoid zones. the presence of shallow-water, benthic foraminifera in turbidite deposits in this biostratigraphically well defined succession gives the opportunity to narrow down the biostratigraphic range of some species. this is a goal that can almost not be achieved when these species are found in place in their shallow-marine habitats, which normally lack age-diagnostic macrofossils. overall, the fauna is characterizedby a lower diversification compared to older anisian or carnian foraminiferal assemblages. however, it has to be mentioned that the presented dataset is limited to the turbidite source area, which might not have covered the whole foraminiferal habitat. acb.notuled.gements. the authors appreciate useful comments by reviewer l. zaniner.ti (geneva) and the journal editor m. gaetani (mìlano). fm thanks the austrian academy of sciences and the vrije universiteit industrial associates program in sedimentology for financial support. the preparation of thin-sections by marco stefani (ferrara) and felix heller (innsbruck) is acknowledged. hans rieber (zurich) kindly provided the determination of the macrofossils. references angiolini l., dragonetti l., muttoni g. & nicora l. (1992) triassic stratigraphy in the island of hydra (greece). rie. it. paleont. strat. 98: 137-180, miiano bosellini a. (1984) progradation geometries of carbonate platforms: examples from the triassic of the dolomites, northern kaly. sedimentologt 31.: 1-24, oxford. bosellini a. ec ferri r. (1980) la formazione di livinallongo nella va11e di s. lucano (ladinico inferiore, dolomiti bellunesi). ann. unio, ferrara, n. s. sez. x,6/5: 63-89, ferrara. brack p & rieber h. (1993) towards a better definition of the anisian/ladinian boundary: new biostratigraphic data and correlation of boundary sections from the southern aips. eclogae geol. heb. 86/2: 415-527, basel. brack p & muttoni g. (2000) high-resolution magnetostratigraphic and lithostratigraphic correlations in middle triassic pelagic carbonates from the dolomites (northern kaly). palaeogeogr, palae oclim. paleoecol. 1 67 : 361-380, amsterdam. brack p, schlager'wi, stefani m., maurer f. & kenter j. (2000) the seceda drill hole in the middle triassic buchenstein beds (livinallongo formation, dolomites, northern italy) a progress report. riot. it. paleont. strat. la6: 283-292, milano. brack p, muttoni g. & rieber h. (2001) magnetostratigraphy and biostratigraphy of the middle triassic margon section (southern alps, itaiy): comment. earth and planet. sci. lett., 193/l-2: 255-257, amsterdam. davaud e. er septfontaine m. (1995) post-mortem onshore transportation of epiophytic foraminifera: recent example from the tunisian coast. /. sed. res. 65: 136-142, ruisa. di bari d. & laghi g.f. (1994) involutinidae bútschli lforaminiferida) in the carnian of the northeastern dolomites (italy). mem. scienze geol. 46: 93-118, padova. fornos j.j. & ahr \ilm. (1997) temperate carbonates on a modern, iow energy, isolated ramp: the balearic platform platform, spain. j, sed. researclt 67/2: 364-373, tulsa. gianolla, p (1991): eruzione freatomagnetica a grande magnitudo nel ladinico delle dolomiti (nota preliminare). rend. soc. geol. it. 14: 65-70, roma. jones \w & charnock m.a. (i985) morphogroups of agglutinating foraminifera. their life positions and feeding habits and potential appiicablility in paleoecological studies. re,uue de paléobiologie 42:311-320, geneve. gazdzicki a. (1933) foraminifers and biostratigraphy o{ 398 f. maurer & r. rettori upper triassic and lower jurassic of the slovakian and polish carpathians. palaeont. polon. 14: \09-1'69, \farszawa. haunold t.g., baal c. & piller \(e. (1997) benthic foraminiferal associations in the northern bay of safaga, red sea, egypt. marine micropal.2g q-a):185-210, amsterdam. krystyn l. (1983) das epidaurus-profil (griechenland) ein beitrag zlrr conodonten-standardzonierung des tethyzalen ladin und ljnterkarn. in: neue beitrîge zur biostratigrrphie der tethys-trias (ed. by zapíe h). \ehrrltpnr i-rrtzt t<<. komm. ()sterr. akad. wiss, 5: 2ll258, vienna. mírquez, l. & trifonova, e. (2000): tasas evolutivas de algunos subórdenes de foraminíferos triísicos del írea occidental del tethys. rer,. espan. micropaleont. 32/1': 1'le, madrìd. maurer f. (1,999) 'wachstumsanalyse einer mitteltriadischen karbonatplattform in den westlichen dolomiten (sùdalpen). eclogae geol. hek;.,92 (3):361-378, basel. maurer f., reijmer j.j.g. e schlager, \( (2002) quantification of input and compositional variations of calciturbidites in a middle triassic basinal succession (seceda, dolomites, southern alps). geol. rund,schau, berlin, in p res s. murray j.\v (1991) ecology and distribution of living benthic foraminiferids. lee & anderson eds., london. oravecz-scheffer a. (1982) triassic foraminifers of the tansdanubian central range. geoiogica hwngarica, ser. palaeontologica 5a 1-331, budapest. piller \il (1978) involutinacea (foraminifera) der trias und des lias. beitr. palàont. òsterr.,5: 1-164, vienna. reiss, z. & hottinger, l. (1984) the gulf of aqaba. ecological micropaleontology. ecological studies 5a: 1-351, snrinper-verlrp. berlin. rettori r. (1995) foraminiferi de1 trias inferiore e medio de1la tetide: revisione tassonomica, stratigrafia ed interpretazione filogenetica. publ. du départm. de géol. et paléont. de gene'oe 18.1-147, geneve. rettori r., angiolini l. & muttoni g. (1,994) lower and middle triassic foraminifera from eros limestone, hvdra island, greece. journ. of micropal. 13/1: 25-46, london. rose, pr. er lidz, b. (1977) diagnostic foraminiferal assemblages of shallow-water modern environments: south floriada and the bahamas. sedimenta, 6: 1-55,l'{iami. salaj j., borza k. ec samuel o. (19s3) triassic foraminifers of the \il/est carpathians. geolog. ustat dinyza stwra, pp. .l 1-)l /. drailslava. senowbari-daryan b., zùh1ke r., bechstàdt t. & flúgel e. (1993) anisian (middle triassic) buildups of the northern dolomites (italy): the recovery of reef communities after the permian/triassic crisis. facies 28: 181-256, erlangen. tyszca i. 0993) response of middle jurassic benthic foraminiferal morphogroups to dysoxic/anoxic conditions in the pieniny klippen basin, polish carpathians. palaeogeogr., palaeoclimatol, palaeoecol. ll0/1: 55-81, amsterdam. travè a., serra-kiel j. &.zamarreno i. (1996) palaeoecological interpretation of transitional environments in eocene carbonates (ne spain). palaios 11/2: 141-16a, tulsa. trifonova e. (1992) taxonomy of bulgarian triassic foraminifera. i. families psammosphaeridae to nodosinellidae. geol. balc. 22/ 1: 3-50. sofia. trifonova e. (1993) taxonomy of bulgarian triassic foraminifera. il families endothyridae to ophthalmidiidae. geol. balc. 23/2: 19-66,sofia. trifonova e. (1994) taxonomy of bulgarian tiassic foraminifera. iii. families spiroloculinidae to oberhauserellidae. geol. balc. 24/2: 21-7a,softa. viel c. (o791 litostratigrafia ladinica: una revi:ione. ricostruzione paleogeografica e paleostrutturale dell'area dolomitico-cadorina (alpi meridionali). ri'0. it. paleont. strat. 85 (1), pp. 85-125, milano. zaninettt l. (1976) les foraminifères du trias. riz,. it. paleont. strat.,82 (\): 1-258, milano. facies control on the composition of serpukhovian and early bashkirian foraminiferal assemblages in the middle tien-shan mountains. central asia olga orlov-labkovsky received january 1, 2002; accepted march i t, 2a03 ke1 zaords: facìes, foraminifera, statistics, mid-carboniferous, tien-shan, central asia. absrract. serpukhovian-lower bashkirian deposits are widely developed in the middle tien-shan mountains of uzbekistan and adjacent countrìes of central asia. these deposits formed in a sedimentary basin exhibiting four distinctive facies that differ in foraminiferal diversity and population densit,v. the facies types, named for mountain ranges containing representàtive sections, are called 1) talassìc, for inner shelf, shallowwater marine carbonates; 2) ugamic, for carbonaceous deposits accumulating on an open, shallow-water, outer carbonate shelf platform; 3) karzhantauìc, for interbedded volcaniclastics and shallowwater marine carbonates deposìted on a eroded surface; and 4) paltauic, for basinal beds containing thin-bedded, graded and lamìnated organic limestones and interbedded turbidites. a statistical program (sorenson's coefficients of species similarity) was used to compare assemblages ìn eight foraminiferal zones from coeval fades across the basin. highest similarity coefficients occur in the early serpukhovian and are probably related to a marine transgression that flooded the basin. regression and volcaniclastic sedimentation account for lower coefficients in the remainder of the serpukhovian. increased foraminiferal diversity and abundance in the earliest bashkirian were probably caused by the opening of new connections to adjacent paleotethyan basins only to be followed by more restricted environmental conditions and lower similarity coefficients later in the early bashkirian. riasswnto. successioni di età serpukhoviana-bashkiriana inferiore sono molto diffuse nelle montagne del tien-shan centrale in uzbekisran e nelle regioni adiacenti dell'asia centrale. esse si sono formate in un bacino sedimentario in cui è possibile individuare quattro associazioni di facies sulla base della diversità nei foraminiferi e la densità delle loro popolazioni. le associazioni di otto zone a foraminiferi sono stare confrontate tra di loro mediante un programma statistico (coefficiente di similarità specifica di sorenson). le maggiori similarità si rrovano nel serpukhoviano inferiore e sono probabilmente collegate alla trasgressione marina che interessò il bacino. la successiva regressione e sedimentazione vulcanoclastica spiegano le basse similarità per il resto del serpukhoviano. la ripresa della diversità ed abbondanza in foraminiferi nel successivo bashkiriano basale furono probabilmente dovute all'apertura di nuove connessioni con l'adiacente paleotetide. tuttavia gìà durante il bashkiriano inferiore condizioni più confinate produssero più bassì coefficienti dì similarità. introduction serpukhovian lower bashkirian deposits are widely developed in the middle tien-shan of uzbekistan and adjacent countries (fig. 1). they are part of a continuous marine carbonate sequence, over 1200-m thick that sras formed in different areas on a carbonate shelf in serpukhovian early bashkirian time. regional stratotypes (type sections) of the lower carboniferous and the lower bashkirian are concentrated in the western part of mountain ranges between the chatkal and talass alatau mountains, including the mashatian, keltimashatian and koikebiltaustian substages of the serpukhovian and the seslavian and uzunbulakian substages of the lower bashkirian (fig.2). the foraminiferal assemblage zones, established for these substages (orlova 1994; orlov-labkovsky 1.999), are correlated with urals-russian platform {oraminiferal zones in table 1. basin from the chatkal to talass alatau ranses and in the karzhantau range exposes four distinct facles types. these are the talassic (shallow-water carbonate sediments), ugamic (an open shallow sea accumulation of carbonaceous sediments) , karzhantauic (a volcanogenic terrigenous shelf sequence with interbedded shallowwater carbonates) and paltauic (carbonaceous terrigenous sediments of the basin) facies-types of sections. the foraminifers in each facies differ in species and generic diversity and density of populations. materials and method the means used to correlate facies to foraminiferal associations include determining the characteristic or dominant species, and the areal distribution of the fauna department of zoology, tel-aviv university, tel-aviv 69978,israel, olgaorl@post.tau.ac.il euroasiatic scheme in urals and russian platform (kagamonov & donakova 1990) itratigraphical scheme in the middle tien-shan (orlova, 1994, 1995) il a brachiopods foramìnifers conodonts ammonoids e a a ! o tr é-, î; € :l choristile.s bísu lcati/òrmís chu'islile.s k,shemischensis meekella eximia pseudosta/fèl la oraegorsbi idiognothodu.s s íttuostt,s bilinguites canceìloceras +l: ìèitr;ì pseutlostal/èìla antiqu0 ii è{ qkt \ :ì< iù= è (n f::s \€ : !ì sè ì*:ì i {r -qs .tí |pirifer hisulccttus oroducttts roductifbrmìs plectostulfel lct longísarla plecto,stoffellu .otundu '). noduliferus 'd. cotugalus reticuloceras ba,\hkoríocerqs plectosta//èlìct iakhensis plectostaf/el ia vqrvartensls i a 5 ls. vcntriz f p ct) c a i f' = c beleutella mugna plectostaf/èlla korsdklensis g. po,stbilineatus fayettevìllect delepinoceras )lectostc(fella miro tbtusa eostallèlla lffke.\tanica gnathodus bilinedtus bollandensis s =î ;triatlera qngusl0 eo,sìgmoilina explicota loebìichiq ninima plectostqf/èllu. orimilivcr +. è.ò ! i -'< 'f = latíproducftrs re(le.tlrilr,\ neoarchaedísctrs regularís biseriellu purva z a jig0nîoprodltcîus lrgqnleus enclolhyranopsís c ra.\,so parugnaîhodus nodosus h.vpergonialiles ferganoceras f''i =:è ìi:l 188 o. c)rlov labkouley and its ecological features (rauzer-chernousova 1960' 1967; fomina 1969). the database consists of 1663 specimens (303 species). in order to evaluate the similarity of foraminiferal associations within the same facies or between different facies for each foraminiferal zone, we used quantitative sorenson's coefficients of species similarity css (odum 1971; southwood 1928). among the ten studied sections, five were representative; therefore, the coefficients were calculated only for these sections: koikebiltau and mashat (talassic ft), akkuiluk (ugamic ft), llya 60l7 (karzhantauic ft) and paltau (paltauic ft) (fig. 2). the sorenson's coefficients of species similarity is calculated with the followine formula: tab. 1 comparison of serpukhovian and bashkìrian faunal zones between the mìddle tìen-shan and ur:l' ru..i:n pl:rform (modified after orlova 1994, 1995). css:2cl(a+b) where "c" is the number of species common to two sections in a foraminiferal. zone, and "a' and "8" are the total number of species in each section within the foraminiferal zone. this index is designed to equal 1 (or i 00%) in cases of complete faunal similarity and o (or 0%) if the sites have no species in common. one of the great advantages of this method is its simplicity. the index is particularly useful for evaluating the similarity of two species lists (magurran 1988). it has been applied, for example, in analysis of the biogeography of devonian rugose corals (naimark et al. 1998), and of the paleobiogeographic affiliation of carboniferous faunas of the qaidam basin, china (li 8*. zhang 1999). r \ t baikinsít0 100 ?0s 30g km kau,a,khs?àn almaq. carboniferows foraminiferal assemblages from c entral asia t89 fig. 1 location of studied serpukhovian/bashkirìan sections, middle tien-shan: talassìc ft: 1 mashat, 2 koikebiltau, 3 djianisusai; ugamic ft: 4 akkuiluk, 5 yakhak; paltauic ft: 6. paltau; karzhantavic ft z river uya 60//, 8 river uya 61/8,9 river uya 69/9,10 river djegirgen. ft facies types. facies types marine sedimentation began around the devonian carboniferous boundary in middle tien-shan and persisted into the serpukhovian and early bashkirian (fig. 3). differences in lithological sequences testify to differences in topographic relief, tectonic movements and sedimentation within the basin (osipova et. al.1971,; fomina 1969). talassic type the talassic type contains shallow-water carbonate sediments from an inner marine shelf (koikebiltau and mashat sections). the serpukhovian inter-val is characterizedby high-energy, near-shore to offshore facies, consisting of light gray, thick-bedded, ooid, crinoidal, algal foraminiferal and brachiopodal limestone banks. these beds formed shoals. during the early bashkirian near-shore, moderately agitated shallowwater environments prevailed, forming graiz, bedded, fine-grained and nodular limestone to ooid, algal foraminiferal limestone chert concretions abound at the top of sequence. foraminifers such as endothyranopsis, omphalotis, globoendothyra, forshia, eostffilla (ikensieformis) and others represent eurobiontic and stenobiontic populations. the most diverse and highest density populations are observed in this facies. ugamic type tl"' i r^^*:^ *'^: is open, shallow marine carbonate sediment and represents accumulation on an outer carbonate shelf platform (akkuiluk and yakhak sections). the carbonates are typically gray, bedded, fine-grained limestone. continuous sedimentation and a relatively constant stratal thickness characterize the sequence. shoal and intershoal sediments as well as up ro 1 m thick lenses of calcareous breccia also occur. chert, both concretions and bedded, appears in the middle part of the section. fossils include conodonts and foraminifers as well as rare brachiopods. the foraminiferal association is characterized by a gradual increase in diversity during the serpukhovian and into the lower bashkirian. the high species diversity of the foraminifers contrasts with the low abundance of specimens except in offshore, shallow-water shoals that have high population density and species diversity. the population density increases from the bottom to the top of the sequence. karzhantauic type the karzhantauic type is a volcaniclastic and shallow marine carbonate sequence, deposited in the karzhantau range ({jya and djegirgen river basins). serpukhovian strata consist of massive-bedded tuffaceous conglomerate and tuffaceous sandstone which are replaced westward by finer-grained terrigenous and detrital limestone. continuous beds of limestone occur at the top. foraminiferal diversity and abundance are high. bashkirian, tuffaceous sandstone passes to the west into detrital foraminiferal limestone and interstratified tuff to the east. upward, strata of trachytic tuffaceous conglomerate, foraminiferal sandstone and fossiliferous sandy limestone occur. the foraminifers have low diversity, but a high population density. talas,t alatau @ *pn* c i.* chirehik 15 s l$km ch*tka 190 o. orlov-lableot,sky f;) a^"'.1..;^^h."r of the main sections for the various facies and formations in the serpukhovian / bashkirian sections in the middlel !b'_ tien-shan. paltauic type the paltauic type includes basinal carbonaceousterrigenous sediments, characterized by thin-bedded, graded and laminated organic limestone with turbidite interbeds (paltau section). the limestone contains admixtures of sand and clay in the middle of the section. the sequence terminates with a limestone conglomerate and calcareous shale. the foraminiferal assemblage is characterizedby a gradual but insignificant increase in diversity and has a moderate population density. foraminifers inch)de b is eriella p arua (n.tchernysheva), n eoarchaedis cus regwlaris (suleimanov) and m onotaxinoides priscus (brazhnikova & yarzeva). some species, such as ompbalotis omphalota (rauzer-chernousova er reitlinger) and eostffilla ihensis vissarionova have thick shells and show traces of transportation. they appear in the late visean, but disappear in the early serpukhovian. other fossils, such as conodonts, ammonoids, algae (koninleopora) and brachiopods are also present but not abundant. facies associations and similarity of foraminiferal zones seroukhovian lower bashkirian foraminifers are moderately to highly abundant and highly diverse (from 40 to 160 species) depending on depositional environments within the basins. assemblages within foraminiferal zones i viii (fig. 3) show a gradual change in eostaffellid species and genera and other taxa. the number of species and genera found from the late visean is sharpiy reduced. the serpukhovian is characterized not only by the evolution of the family eostaffellidae, but also by the appearance of sma1l foraminiferal genera such as rectoendotbyra, b iseriella, globivaktulina, m onotaxinoides, representatives of the family asteroarchaediscidae and others. the sorenson's coefficients of species similarity (css) values for foraminiferal zones i-viii are presented in table 2.they are based on comparisons between sections in the same facies ikm in table 2\ and from different facies (ka through pu in table 2). in order to interpret the significance of the coefficient values, i separated them into five categories (fig. 4): highest css at 85 -100 %; high css at 70 -85 "/"; moderately high at 55-70 "/". moderate ar 4055 o,o afld low at 25 -40%.the highest css records are from zones i (75% of total records) andii (25%); the high css are from zones ii (20%),iii (10%), ry (10%), vi (30%) and vii (30%); the moderately high css are from zones iii tr i (1 è ,j) -è {t} \ facics typr' talassic ugamic paltauic karzhantauic àn scctiorr koikebiltau mashaf akkuiluk paltau uya (60/7) g, u} d n f d v) lx pseudostaffella pruegorsnyai f ll : :\ a f. .j) limcstonc n:edium and drin bedded at botlòm" lìne grairred. ngdular arrd deirital rvith chelt concretioix. limestone mediurr bedded finc grained nodular and detrilal with shells. alsalfbraninilì:rs. ioids che{ rùncretiòns. oe !:4 à limestone drin and medium bedded. nodular jine and micro grained ciayeyìnarly dehita liilltl f i : e tmchyardesite rvith tuflaceous sandstone and sandstone ylil pseudosraljùlla 0t1tiqu0 yii pltctostqffelkt fongi''culu p i e ctostajfè i la ronmdo rirgsionc lhrtki llftilii tr 8"di -imeslone mediunr :edded, dotrital. ilgaf lìlraminifcral. roclular fine grained lith concretions md intcrlaycrs of :ììen tu{lrceous sandsto ne with limestone sandy. detritài, furaninìfèru|, nodular, ooids ard iúerlayen of trachye vl plectostufftlla .j nklrnsis pleuostof: .fèllu ,-onuriansis z o f trl ,"/) 6'a =-u a y plectosnlfltlla karsaklensis 1l llfit e. t! g z j tuffaccous conglonìemle and lnè iàceous sandstone with iinrestone sandy, defitài, lbraminilèral, nodular and r:oids and thin bedrled with cnrsion surfacesforaninifirs brachiopo85-100 <40>55 .25>40 categorè c arb oniferous foramintferal as s emblages from c entral asia references t93 bensh f.r., nigmadzanov i.m. & iskandarkhodgaev t.a. 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(1978) ecological methods. chapman & hall. a halsted press. 524 pp., london. foreword proceedings of the 8th international brachiopod congress brachiopods in a changing planet: from the past to the future milano, 10-14 september 2018 rivista italiana di paleontologia e stratigrafia (research in paleontology and stratigraphy) vol. 125(3) november 2019 the 8th international brachiopod congress took place in the prestigious venue of the university of milano, italy, in september 2018, after the previous edition held in nanjing, china, in 2015. 150 participants from universities and research institutes from all over the world attended the meeting, from argentina, armenia, austria, belgium, canada, china, czech republic, denmark, france, germany, hungary, iran, israel, italy, japan, new zealand, poland, russia, slovakia, spain, sweden, united kingdom, and united states of america. besides oral and poster scientific sessions, pico-presentations of young researchers, and two prestigious plenary lectures, the congress was preceded and followed by three field trips (spain, united kingdom and sicily), and by two mid-congress day excursions at castell’arquato and grigna mountains. during the congress, all the topics and the recent advances in the study of brachiopods, marine invertebrates that have dominated the palaeozoic seas and had an important role in the phaneorozoic benthic communities, have been touched: from systematics and evolution to biostratigraphy, palaeoecology, palaeobiogeography, mass extinctions, and the biology of recent taxa. one of the novelties of this edition was a session dedicated to brachiopod shells as biomineral archives of fundamental importance to understand the evolution of marine calcifiers during climate and environmental changes in recent and deep time. so, also biomineralization and geochemistry were among the topics of the event. the high scientific level, activity and wide and interdisciplinary interests of the brachiopod researchers’ community are testified by the numerous contributions received for the proceedings of the 8th international brachiopod congress, 16 of which are published open access in issue 125/3 of rivista italiana di paleontologia e stratigrafia. the first paper published in this issue is by donald macfarlan, who describes a large lower jurassic terebratulid collection from new caledonia and new zealand. he recognizes seven genera and twelve species, eight of which are new, and discusses their stratigraphic and geographic distribution, enhancing the biostratigraphic potential of these taxa and offering new insights on the tjb and toarcian faunal changes. forewordii the paper by aleksandra bitner investigates a brachiopod fauna collected in the tonga islands in the south-west pacific ocean, identifying species never reported before from those islands. the author also compares the affinity and the biodiversity of the tonga fauna with those from fiji, new caledonia, new zealand and french polynesia, providing new data on the distribution of recent taxa. hui-ting wu and co-authors describe a brachiopod fauna from a mixed nearshore clastic–carbonate setting in the changhsingian longtan formation and the griesbachian yelang formation of the liuzhi section (guizhou province, south china). they classify 33 species and 16 genera and discuss the end-permian extinction magnitude in different localities and palaeoenvironments, with a particular emphasis on shallow water clastic vs carbonate settings. emma taddei ruggiero and co-authors discuss and propose a solution to a long-standing nomenclatural problem related to the definition of the species terebratula sinuosa. following an error of linnaeus, this species was considered by subsequent taxonomists a synonym of terebratula terebratula. the authors now provide stratigraphic and morphological evidence that t. sinuosa deserves the full rank of a species and that the stratigraphic distribution of t. terebratula should be amended and limited to the pliocene-pleistocene. studying internal molds and steinkerns, jisuo jin and co-authors investigate pentamerida brachiopods from lower silurian rocks of the michigan basin to clarify the taxonomy of several virgianids. after proposing a new genus and revising the biostratigraphic and palaeobiogeographic distribution, and the type of shell deposits of several virgiana palaeocommunities, they discuss the factors controlling their composition, providing very valuable palaeoecological and palaeobiological data. fernando lavié and co-authors describe and illustrate a collection of micromorphic linguliform brachiopods from the middle-upper ordovician of the argentine precordillera. they recognize nine linguliform families and propose two new acrotretid species: conotreta andina n. sp. and biernatia rhapsody n. sp., providing new and interesting biostratigraphic data. elizabeth harper and coauthors provide quantitative data concerning patterns of shell breakage predation and repair in rhynchonelliformean brachiopods collected from a new zealand fiord. the authors observe that only a few individuals show signs of having been able to repair damage and that the proportion of individuals showing unrepaired, and hence presumably fatal, breakages was higher. the authors provide new and important data on brachiopod ecology and in general on predation. daniel stadtmauer and susan butts describe the skeletal microstructure and morphology of a unique permian brachiopod genus, pirgulia, from the permian sosio limestone of sicily. in this taxon, the ventral valve forms a cone that fully encloses the lobate structure which replaces the dorsal valve. based on a functional morphology approach, the authors reconstruct the lifestyle and the dynamics of water flow of this heteromorphic brachiopod, providing very valuable palaeoecological information. zhiwei yuan and co-authors provide a detailed palaeoecological analysis of an in situ preserved brachiopod palaeocommunity from the lower carboniferous of guizhou, south china. the authors also erect a new spiriferide species, weiningia ziyunensis n. sp., describing in detail its population dynamics and explaining why this was a dominant species in the palaeocommunity, forming dense clusters attached to living and dead shells. attila vörös and co-authors present a compilation of brachiopod genus-level diversity through the phanerozoic and compare the traditional extinction events, the so-called ‘big five,’ with the episodes of synchronous termination of multiple orders, named ‘clade extinctions.’ their compilation reveals that three of the five episodes are recognized both as mass and clade extinctions. the paper proves how these episodes are relevant to shape the history of the clade, as for instance in the early jurassic, when spire-bearing brachiopods disappeared. yuchen zhang and co-authors describe the upper ordovician atrypide genus rongatrypa, expanding its distribution from the kazakh terranes, from which it was previously known only, to south china, and providing important palaeobiogeographic implications. a detailed analysis of its ontogenetic variation provides an interesting explanation for its allometric growth pattern. proceedings of the 8th international brachiopod congress iii geochemical studies, underscores that brachiopods are excellent archives of past and present oceanographic conditions. bernard mottequin and gabriela cisterna reinvestigate in detail histosyrinx vautrini, the type species of histosyrinx, a poorly known carboniferous spiriferinide genus from north africa. the study of new material from lybia and algeria allows to improve the knowledge of the variability of the internal characters at the subfamily level and to compare histosyrinx with allied genera, some of which still inadequately known. gabriela cisterna and co-authors open up about the opportunity to elucidate some aspects of the late palaeozoic ice age demise by performing a comparative study of the eurydesma faunas and associated brachiopod assemblages from the bonete (east argentina) and wasp head (east australia) formations. providing a very interesting palaeoecological investigation, the authors demonstrate that the faunas show similar composition, but differences in the relative abundance of taxa. they conclude that several factors controlled the faunal composition, arguing for more in-depth analyses in the next future. the guest editors of the 8th international brachiopod congress proceedings volume lucia angiolini, gaia crippa, claudio garbelli, renato posenato ulrich jansen discusses and interprets the biodiversity changes of lower devonian brachiopods from the rhenish massif, in the framework of a regional palaeoenvironmental evolution. in the paper, a new genus, ingentistrophia n. gen., and two new species, fascistropheodonta? wiltzensis n. sp. and pachyschizophoria amygdala n. sp., are proposed. patterns of extinction and speciation are interpreted in terms of the interplay of sea-level fluctuations, subsidence rates and siliciclastic input in a very interesting reconstruction. florencia leone and juan benedetto describe the morphology and ontogeny of the rhynchonellide clarkei antinsiens from the upper silurian tarabuco formation of bolivia and compare the ontogenetic stage of the species with harringtonina australis and anabaia paraia. they demonstrate that, throughout the silurian, these species represent an evolutionary lineage characterized by increasing peramorphic characters, providing an interesting case study of heterochrony in brachiopods, with implications on the current taxonomic classification of the studied taxa. uwe brand and co-authors propose an updated brachiopod-based oxygen-isotope thermometer, that can be applied to most rhynchonelliformea, both recent and fossils, from cold to warm and shallow to deep water settings. based on a very large dataset, the authors record a -1 ‰ offset of brachiopod shells oxygen isotopes from 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(2020) the occurrence of eocenochelus (testudines, pleurodira) from sardinia supports palaeogeographic reconstruction of the proximity of the island to continental western europe during the eocene. riv. it. paleontol. strat., 126(3): 833-846. rivista italiana di paleontologia e stratigrafia (research in paleontology and stratigraphy) vol. 126(3): 833-846. november 2020 abstract. in this paper, we describe pleurodire turtle material from the island of sardinia, italy, originating from two eocene localities of the cixerri formation. the more complete among the two specimens bears strong resemblance with the continental western european eocenochelus eremberti and is tentatively referred to the same species, while the second, less complete specimen is only provisionally referred to the genus eocenochelus. the new sardinian turtles add to the so far scarcely documented fossil record of pleurodires in italy, while it ranks as among the very few paleogene vertebrates known from the island. the identification of the widespread eocene western european genus eocenochelus in the fossil record of sardinia supports recent palaeogeographic reconstructions of the island, according to which the sardinia-corsica massif was located rather close to mainland europe and was subjected to a significant, counterclockwise rotation during the paleogene. received: july 8, 2020; accepted: september 10, 2020 keywords: pleurodira; testudines; paleogene; biogeography; palaeogeography. introduction pleurodires are currently confined solely to the southern continents (rhodin et al. 2017); however, this turtle clade achieved a significantly broader distribution during its mesozoic and cenozoic past (see gaffney et al. 2006, 2011; cadena 2015; pérezgarcía et al. 2017b; ferreira et al. 2018; joyce & georgalis g.l., zoboli d., pérez-garcía a., pillola g.l. & delfino m.834 chelys bolcensis was initially introduced as platyarkia bolcensis by bergounioux (1953); however, he provided no description, definition, indication to a previous description, or figure of the material and as such, this 1953 binomen has to be considered a nomen nudum. neochelys bolcensis was formally described by the same author the following year (bergounioux 1954), but nevertheless, currently this taxon is considered to be a junior synonym of n. capellinii (e.g., broin 1977). the validity of emys nicolisii is at flux with the taxon likely representing a nomen dubium or at least a junior synonym of n. capellinii, though it has also been recently considered as a valid species as well, neochelys nicolisii (lapparent de broin 2003; pérez-garcía & lapparent de broin 2015; lapparent de broin 2018). in a broader frame of a revision of the fossil turtles of italy (abbazzi et al. 2008; chesi 2009; chesi et al. 2007a, 2007b, 2009; colombero et al. 2014, 2017; cirilli et al. 2016; georgalis et al. 2020), here we describe two shells of eocene age from sardinia, which are assigned to the genus eocenochelus. we address their taxonomic affinities and compare them with other finds of the genus from continental europe. biogeographic implications of the new pleurodire remains are discussed, along with their impact on the palaeogeographic reconstruction of the western mediterranean during the eocene. material and methods the fossil material described herein is curated at the collections of the mdlca. the first of the two specimens (the most complete, mdlca 14006) was briefly mentioned in abstracts and papers during the past two decades (righi & delfino 2003; delfino & rook 2008; pérez-garcía & chapman 2017; pérez-garcía et al. 2017b); however, it has never been formally described or figured. the second one (mdlca 3018) is presented here for the first time. taxonomy follows joyce et al. (2004) and pérez-garcía et al. (2017b). institutional abbreviation. mdlca, museo sardo di geologia e paleontologia “domenico lovisato”, university of cagliari, italy. geological settings the turtle remains described herein originate from the cixerri formation, a continental succession cropping out discontinuously in southwestern sardinia (pecorini & pomesano cherchi 1969). the most complete specimen (mdlca 14006) was collected in a sand quarry in the locality medau is lapparent de broin 2013, 2015; cadena 2015; pérezgarcía 2016; pérez-garcía & chapman 2017; pérezgarcía & smith 2017). their most recent records in this continent are from the early neogene (ristori 1895; georgalis et al. 2013; georgalis & kear 2013). a number of paleogene taxa from western europe have been identified as pertaining to the lineage of erymnochelyinae, which is now represented by the extant taxon erymnochelys madagascariensis (grandidier, 1867), which is exclusive to madagascar (ferreira et al. 2018); nevertheless, erymnochelyinae were once much more widespread during their cenozoic past, distributed over europe and africa (ferreira et al. 2018). erymnochelyines were widespread in the eocene of central and western europe, achieving a diverse radiation comprising multiple species, pertaining to two genera, the freshwater neochelys bergounioux, 1954, and the littoral eocenochelus pérezgarcía, lapparent de broin & murelaga, 2017b (bergounioux 1954; broin 1977; pérez-garcía et al. 2017b). despite their abundance and high levels of species diversity, european erymnochelyines became ultimately extinct at the end of the eocene (pérezgarcía et al. 2017b). among european erymnochelyines, the genus eocenochelus, species of which have been found across the eocene of france, spain, belgium, and the united kingdom, is the most phylogenetically proximate genus to the extant erymnochelys baur, 1888 (pérez-garcía & chapman 2017; pérezgarcía & smith 2017; pérez-garcía et al. 2017b). in italy, the fossil record of pleurodire turtles has been so far limited to continental remains coming from the province of veneto, in the northeastern part of the country. the remains from veneto are represented by several specimens, including complete, articulated skeletons, and are already known since the late nineteenth century (zigno 1889, 1890). they originate from two middle eocene localities and have been originally referred to their own species: emys capellinii zigno, 1889 and neochelys bolcensis bergounioux, 1954 from the famous lagerstätte of monte bolca, and emys nicolisii zigno, 1890 from avesa (zigno 1889, 1890). all of them seem to pertain to the podocnemidid erymnochelyine genus neochelys (bergounioux 1954; broin 1977; kotsakis 1978; pérez-garcía & lapparent de broin 2015). neochelys capellinii is in fact the type species of neochelys and is currently almost universally considered a valid species (lapparent de broin 2003; pérez-garcía & lapparent de broin 2013, 2015; cadena 2015). neothe occurrence of eocenochelus (testudines, pleurodira) from the eocene of sardinia 835 fenus, near flumentepido (municipality of carbonia). the other specimen (mdlca 3018) was collected in the locality of murecci (municipality of gonnesa, fig. 1). the geographic distance between these outcrops is around 7 km in straight line. the cixerri formation is a terrigenous formation mainly represented by sandstones, subordinated siltites, and conglomerates with very rare intercalations of lignitiferous clays and lacustrine limestones (pecorini & pomesano cherchi 1969; barca & costamagna 2010; costamagna & schäfer 2013). the conglomeratic levels are more frequent in the upper part of the succession. generally, these are represented by rounded, polygenic and heterometric clasts originated from paleozoic, mesozoic and early eocene lithotypes cropping out currently in the sulcis-iglesiente subregion. furthermore, several authors reported the presence of pebbles with ?aptian–albian microfaunas originated from lithotypes of the pyrenean domain, totally absent in the sardinian area (cherchi & schroeder 1976; cherchi 1979; barca & cherchi 2002). the formation crops out extensively in southwestern sardinia (sulcis subregion) and in the cixerri valley, with a maximum thickness of about 300 m (pecorini & pomesano cherchi 1969; funedda et al. 2009; barca & costamagna 2010). the nature of the sediments indicates a continental depositional context with a prevalence of fluvial facies (pecorini & pomesano cherchi 1969). more precisely, the cixerri formation represents the molassic post-pyrenean phase sedimentation in sardinia (cherchi 1979; barca & costamagna 2010) and deposed in the middle to lower, distal part of a foreland alluvial plain fed by iberian relieves (costamagna & schäfer 2013). presently, only a portion of this alluvial plain is preserved, because a great part of this was cut in half by the following detachment of the sardinia-corsica massif from the southern european margin (costamagna & schäfer 2013). furthermore, barca & costamagna (2010) reported fluvial facies alternated with siliciclastic deposits containing marine ichnofacies and tidal stratal pattern in the monastir area (campidano di cagliari subregion) suggesting at least temporary coastal/river mouth area. marine influences are documented in eastern and southeastern sardinia since the thanetian (matteucci 1985). however, the general trend in facies distribution during the cixerri formation time indicates a provenance of the sediments from the west to the east and suggests that the previous “short living” marine to lagoonal basin identified in the sulcis area (starting with the ?late thanetian, middle–late ypresian “calcari a macroforaminiferi” auct.), showing a clear transgression, running from the southwest to the east in the present day position, was bounded by emerged land towards the east and filled by continental deposits during the early lutetian lignitifero auct formation and cixerri formation molasse times. it is difficult to give more detailed palaeogeographical information, but the characters of the transgressive-regressive cycle and the tectono-sedimentary setting play in favor of a relfig. 1 schematic geological map of the nw sulcis basin, sardinia, showing also the position of the fossil localities which have provided the remains of turtles studied in this paper. 1) undifferentiated aquitanian–holocene deposits; 2) cixerri formation (middle lutetian–early oligocene); 3) lignitifero formation (late ypresian– early lutetian); 4) miliolitico formation (early ypresian); 5) variscan metamorphic basement and carboniferous–mesozoic deposits. georgalis g.l., zoboli d., pérez-garcía a., pillola g.l. & delfino m.836 atively short time for the deposition of the cixerri formation. according to barca & costamagna (2010) and costamagna & schäfer (2013), the cixerri formation is settled in a more limited time span (middle lutetian–early rupelian). these authors used the presence of volcanoclastic pebbles at the top of the continental succession to discriminate another formation (flumentepido formation; barca & costamagna 2010). the cixerri formation rests conformably to unconformably over the late ypresian–early lutetian lignitifero formation or it is posed unconformably over the variscan metamorphic basement (costamagna & schäfer 2013, and references therein). upwards, it passes sharply through an unconformity to the calcalkaline volcanics of the oligocene–miocene cycle (about 22.8±1.3 ma) in the sulcis area (lecca et al. 1997) or to the terrigenous of the late oligocene to aquitanian ussana formation in the campidano area (pecorini & pomesano cherchi 1969). in addition, in the cixerri valley, the cixerri formation is clearly intruded by the cenozoic volcanic rocks dated at 29.9–27.5 ma (savelli 1975; savelli et al. 1979); more recently these rocks were recently dated to be 29.30±1.2 to 26 ma (funedda et al. 2009). the fossiliferous content of the cixerri formation is rather poor; in addition to the herein described turtle remains, only scarce plant remains (in particular palms), freshwater gastropods (lymnaea cf. orelongo), and isolated bone fragments from lacustrine levels were reported from the villamassargia, gonnesa, and perdaxius areas (taricco 1924; maxia 1959). summarizing, the age of the cixerri formation is bracketed between the early lutetian (which is the age at the top of the lignitifero formation) and the local first occurrence of intruded mid-late rupelian volcanic products. according to these data, the turtle remains from the cixerri formation (sensu barca & costamagna 2010) can be referred to the middle lutetian–early rupelian; this time range slightly encompasses the so far known chronologic distribution of eocenochelus, that is known exclusively up to the late eocene. but, the reasons exposed above (tectono-sedimentary setting) suggest that the age of the sardinian pleurodire remains is, in all likelihood, middle eocene. systematic palaeontology testudines batsch, 1788 pleurodira cope, 1864 pelomedusoides cope, 1868 podocnemididae cope, 1868 erymnochelyinae broin, 1988 (sensu pérez-garcía et al. 2017b) erymnochelyini pérez-garcía, díaz-berenguer, badiola & canudo, 2019 eocenochelus pérez-garcía, lapparent de broin & murelaga, 2017b eocenochelus eremberti (broin, 1977) eocenochelus cf. eremberti figs. 2-3 material: mdlca 14006, an almost complete carapace and plastron, missing only tiny fragments of the anteriormost and posteriormost portions of the carapace (fig. 2-3). locality: medau is fenus, near flumentepido (municipality of carbonia), sardinia; cixerri formation, probably middle eocene. description. mdlca 14006 is an almost complete shell, preserving most parts of the carapace and plastron (figs. 2-3). the carapace misses only tiny fragments of the anteriormost and posteriormost portions, especially at the posterior right portion, near the marginals, as well as near the level of the first pleurals. three prominent breakages are apparent on the carapace. also, certain portions of the carapace are slightly deformed. the carapace is oval, being 372 mm long and 306 mm wide (at mid-length and mid-width respectively). its posterior part is relatively flattened dorsally; there are no signs of a medial or lateral keels. the posterior margin of the carapace is dorsally directed. it is not possible to determine whether a cervical scute was present (in the case of being present it should be extremely narrow, which is unlikely). the limits between several plates are not clearly visible and these often coincide with the fractures. the plates of the carapace, which has been possible to identify, are represented by: a nuchal, six neurals, a suprapygal, a pygal, eight right and eight left costals, and at least nine right and nine or ten left peripherals. the nuchal is located immediately anterior to the neural series. it is covered by the vertebral i and the first pair of marginals. there are six neurals. the dimensions of all the neurals are the occurrence of eocenochelus (testudines, pleurodira) from the eocene of sardinia 837 more or less similar between them, with the exception of the last one, which has significantly smaller dimensions than the others. their shape is hexagonal, with their smaller sides situated in their anterior positions and their formula is: 4-6a-6a-6a-6a-5. the last neural does not contact with the suprapygal, fig. 2 eocenochelus cf. eremberti from sardinia. mdlca 14006, almost complete carapace and plastron. photographs in dorsal (a), ventral (b), and left lateral (c) views. fig. 3 eocenochelus cf. eremberti from sardinia. mdlca 14006, almost complete carapace and plastron. interpretative drawings in dorsal (a), ventral (b), and left lateral (c) views. georgalis g.l., zoboli d., pérez-garcía a., pillola g.l. & delfino m.838 due to the medial contact of the last three pairs of costals. the costals at the central area of the carapace are larger than laterally and they are decreasing both in width and height, towards both the anterior and posterior position in the series. peripherals i-ix can be distinguished in the right series; the sutures of peripherals x and xi are hidden by fractures and areas in which shell portions are missing. from the left series, peripherals vii and viii are not recognizable. the remains of the scutes on the carapace afford some information about all the elements that originally constituted its external appearance. the scutes are represented by two series, each arranged on one side, of 12 marginals and four pleurals and by a single series of five vertebrals. the medial region of both first marginals is not preserved. the marginals ii-vii have a rather similar shape and size between them, and are longer than wide. among pleurals, pleural ii is the largest, followed by pleurals i, iii, and iv. the vertebrals ii and iii are the largest, both showing a similar size; they are followed in size by vertebrals i, iv, and v. the plastron is 34.6 cm long and 26.4 cm wide (at the level of the mesoplastra). its central part is slightly concave (it is unclear whether it a product of deformation or if it was also the case of the specimen when in life). the anterior plastral lobe, 10.9 cm long, is rounded at the front and widens towards a posterior direction. its outer edge folds slightly upwards. the posterior plastral lobe, 11.7 cm long, has an anal notch with an angle of 107o; its distal margin folds slightly upwards. the left bridge is 12.7 cm long. it is damaged by displaced fractures, in which the posterior area was collapsed. the right side bridge is well preserved, being 12.0 cm long. several variably pronounced (and sometimes dislocated) fractures are visible, mainly with a transversal or oblique direction. in the plastron, the sulci that delimit the scutes are relatively evident, while several areas corresponding to the sutures between some plates are not clearly visible. the plates in the plastron are represented by the entoplastron, the epiplastra, hyoplastra, mesoplastra, hypoplastra, and xiphiplastra. the entoplastron is of rhomboid shape and is located in the anterior portion of the plastron. the epiplastra are trapezoidal. the hyoplastra are longer than the hypoplastra. the mesoplastra are situated laterally, placed at the lateral limits of hyoplastra and hypoplastra. scutes are represented by a single gular and a pair of extragulars (sensu hutchison & bramble 1981), humerals, pectorals, abdominals, femorals, and anals. the element size formula of medial surface of the plastron is as follows: abdominals > pectorals > femorals > anals > gular ≥ humerals > extragulars. the gular has small dimensions. the femorals have a trapezoidal shape and the sulcus with the anals is slightly posteriorly arched. the anals are rhomboid. their posterior extremity is semioval and its margins form an angle of 107o. cf. eocenochelus sp. fig. 4 material: mdlca 3018, an incomplete carapace and plastron, missing the most of its anterior halves of both carapace and plastron (fig. 4). locality: murecci, (municipality of gonnesa), sardinia; cixerri formation, probably middle eocene. description. mdlca 3018 is a partial shell, lacking the anterior portion of both carapace and plastron (fig. 4). the preserved portion of the shell is filled with matrix and is locally heavily fractured and incomplete. the surface is partly covered by a hard concretion. therefore, most of the skeletal elements are not visible, though their presence can be supposed on the basis of the general shape of the shell. however, its morphology does not significantly differ from that of mdlca 14006 and thus it will be only briefly commented here. the carapace has a preserved midline length of 33.6 cm, while its maximum preserved width is 28.8 cm. the carapace preserves the neurals from the second (maybe a little portion of the second) to the last, but only the third, fourth and fifth can be seen with relative confidence (a posterolateral suture of the sixth could be present); the costals from the second to the last; the peripherals from the fourth to the last one. pygal and suprapygal could be also present but this cannot be deciphered with certainty. the plastron has a preserved midline length of 26.5 cm, while its maximum preserved width is 23.2 cm. the plastron preserves partial hyoplastra, the hypoplastra, and the xiphiplastra. the entire left mesoplastron and possibly part of the right mesoplastron are present but their sutures cannot be seen due to the presence of concretion and fractures. the occurrence of eocenochelus (testudines, pleurodira) from the eocene of sardinia 839 discussion taxonomic allocation mdlca 14006 can be identified as a member of pleurodira considering characters as the present of a pair of laterally located mesoplastra, and that of a single gular. the absence of a trapezoidal anterior plastral margin allows to exclude its attribution to neochelys (pérez-garcía & lapparent de broin 2015), the only genus of pleurodira that was hitherto identified in the fossil record of italy (see introduction above). the development of a long medial contact of the extragulars, due to the presence of a reduced gular, has been recognized as exclusive of erymnochelyini (pérez-garcía et al. 2017b, 2019). this specimen can be referred to the genus eocenochelus on the basis of the identification of an exclusive character combination within this clade including: absence of a medial keel on the carapace; vertebral i heptagonal, with short latero-anterior margins, and wider than the nuchal; vertebral ii approximately as wide as vertebral iii, both scutes lacking wide lateral protrusions anterior to the limit between the pleural sulci; first pair of marginals overlapping no more than half of the length of the lateral nuchal margins; posterior plastral lobe narrower than the anterior one; absence of clear extragular protrusions; relatively long gular scute, reaching the anterior margin of the entoplastron or overlying its most anterior region; very short dorsal expansion of the plastral scute borders (see characters in pérez-garcía et al. 2017b, 2019; pérez-garcía & smith 2017). among eocenochelus spp., this sardinian specimen shares several characters with the type species of the genus, eocenochelus eremberti, which constitute an exclusive combination for this form: sinuous lateral margins of the vertebral scutes; relatively long epiplastral symphysis, less than half the entoplastral length; relatively wide posterior plastral lobe; subrounded lateral margins of the posterior plastral lobe; relatively wide anal notch, its length being equivalent to half this width fig. 4 cf. eocenochelus sp. from sardinia. mdlca 3018, incomplete carapace and plastron; photographs and interpretative drawings in dorsal (a, b), and ventral (c, d) views. georgalis g.l., zoboli d., pérez-garcía a., pillola g.l. & delfino m.840 or less (see pérez-garcía et al. 2017b, 2019). it is worth noting that in terms of size, the sardinian material approaches mostly the smallest species of the genus (i.e., eocenochelus lacombianus pérez-garcía, lapparent de broin & murelaga, 2017b), which has a plastron length of around 30 cm and not respective elements of the largest species, e. eremberti, which reaches even around 50 cm in plastron length (see pérez-garcía et al. 2017b, 2019). the presence of six neurals instead of seven differs from the known condition for the middle eocene e. eremberti, being compatible with the one that characterizes the late eocene eocenochelus farresi pérez-garcía, lapparent de broin & murelaga, 2017b (pérez-garcía et al. 2017b). however, other characters allow to exclude its attribution to this form (e.g., the combination of characters indicated as shared with e. eremberti as well as exclusive characters for e. farresi, such as the presence of a very short epiplastral symphysis, long lateral overlap of the humeral scutes onto hyoplastra, and an almost as long as wide u-shaped anal notch), as well as to the early eocene e. lacombianus (e.g., the combination of characters indicated as shared with e. eremberti as well as the development of concave lateral margins of its notably narrow posterior plastral lobe, which represents an exclusive condition for e. lacombianus) (see pérezgarcía et al. 2017b). the differences between the shell mdlca 14006 and those hitherto known for e. eremberti cannot be ruled out as due different ontogenetic stages (e.g., the smaller size of the sardinian specimen), and to an intraspecific variability for this species greater than that until now known (e.g., the difference in the number of neurals). since other shell characters used for the characterization of e. eremberti (pérez-garcía et al. 2017b, 2019; pérezgarcía & smith 2017) are unknown in mdlca 14006, and also considering that the age of mdlca 14006 cannot be confirmed as compatible with the known stratigraphic range of distribution for that species (lutetian), we attribute mdlca 14006 to eocenochelus cf. eremberti. most of the characters available in the partial shell mdlca 3018 are compatible with those observed in mdlca 14006 (e.g., the morphology of the preserved neural plates and vertebral scutes; the width/length ratio of the anal notch and the morphology of its lateral margins), and the differences observed between them (e.g., the morphology of the posterior plastral lobe) are recognized as compatible with the intraspecific variability observed in various members of pleurodira, including eocenochelus eremberti (pérez-garcía & smith 2017; pérez-garcía et al. 2019). despite the relative low number of characters provided by mdlca 3018, it is attributed to cf. eocenochelus sp. considering its potential compatibility with the same taxon represented by the almost complete sardinian shell described above, as well as the fact that their remains are found in the same formation and, in particular, in two localities that are situated relatively close to each other. the well-preserved specimen (mdlca 14006), comprising both carapace and plastron, ranks as one of the most complete shells of eocenochelus known to date, being surpassed in completeness only by a recently described shell of e. eremberti from spain (pérez-garcía et al. 2019). palaeobiogeographic and palaeogeographic implications the present identification of eocenochelus in the eocene of sardinia provides interesting biogeographic implications. so far, eocene tetrapods from sardinia were represented by pan-trionychids, crocodylians, the marsupial amphiperatherium filhol, 1879, and the perissodactyls atalonodon monterini dal piaz, 1929, and “lophiodon” sardus bosco, 1902 (bosco 1902; dal piaz 1929; kotsakis 1985; kotsakis et al. 1997; zoboli et al. 2019). with the exception of atalonodon monterini, the crocodylians, and the pleurodire turtle material described herein, all tetrapod fossils come from the late ypresian–early lutetian lignitifero formation. the eocene pan-trionychids from sardinia are represented by indeterminate remains (kotsakis 1985) that cannot be identified beyond the family level (georgalis & joyce 2017). the crocodylians are represented by a single small tooth from the ypresian of escalaplano (southeastern sardinia) that cannot be taxonomically more precisely referred (zoboli et al. 2019). the marsupial remains have not been yet formally described and were only provisionally referred to the genus amphiperatherium (kotsakis et al. 1997), a genus that is widespread in the paleogene of continental europe. “lophiodon” sardus has been either referred to the lophiodontid genera lophiodon cuvier, 1822, and paralophiodon dedieu, 1977 (bosco 1902; esu & kotsakis 1983; palombo 2009), both genera that are widespread in the paleogene of europe, including the occurrence of eocenochelus (testudines, pleurodira) from the eocene of sardinia 841 the iberian peninsula (holbrook 2009). anyway, a revision of the type material of “l.” sardus should be necessary to confirm the taxonomic position and validity of the taxon. however, the described fossil material of this perissodactyl is currently lost (kotsakis et al. 2008). the other large mammal, atalonodon monterini, was collected in the lithotypes of the miliolitico formation (early ypresian) cropping out near the terras de collu coal mine (gonnesa). atalonodon is also currently considered a lophiodontid (e.g., prothero & schoch 1989; holbrook 2009), whereas alternative relationships within perissodactyls have also been suggested (palombo 2009); however, whether it indeed represents a sardinian insular endemic genus or is instead referred to a mainland european one remains yet to be demonstrated. the presence of related faunal elements among sardinia and continental western europe should not appear at strange. rather than that, current geodynamic reconstructions suggest that, during the early paleogene, the sardinia-corsica massif was rather close to continental western europe, with an important, about 45°, counter-clockwise rotation subsequently taking place during the eocene (advokaat et al. 2014). nevertheless, faunal distinction, at least to the species or even genus level, is the case for certain vertebrate taxa that occurred in the early eocene of continental western europe and sardinia, therefore implying the existence of some sort of temporary geographical or palaeoecological barriers between the sardinia-corsica massif and the iberian-occitanic area that allowed endemic radiations flourishing in these nearby different landmasses during the late paleocene–early eocene time span (matteucci et al. 2000; palombo 2009). the rotation of the sardinia-corsica massif continued also in post-eocene times, as an even more prominent and well documented rotation (counter-clockwise of about 50°) took place, mostly during the early miocene (gattacceca et al. 2007); that being said, the total rotation of the sardinia-corsica massif was of about 95° counterclock-wise and was completed by around 15 ma (gattacceca et al. 2007). it is further worth noting that during the late oligocene the sardinia-corsica massif was still connected with the european mainland as documented by the presence of the continental ruminant “amphitragulus” quercyi fig. 5 palaeogeographic reconstruction of the western mediterranean during the eocene, showing the position of the sardinia-corsica massif. silhouettes represent the shared continental tetrapod lineages that occurred in both areas during the late ypresian–lutetian. abbreviations: ba, balearic islands; be, betic; ib, iberia; ka, kabylias; cpa, calabrian-peloritan arc; sa, sardinia; co, corsica; aq, aquitania. map modified after omodeo and rota (2008) and advokaat et al. (2014). georgalis g.l., zoboli d., pérez-garcía a., pillola g.l. & delfino m.842 (mennecart et al. 2017) and early miocene insular taxa with clear mainland affinities (de bruijn & rümke 1974; van der made 2008; zoboli & pillola 2017; mennecart et al. 2019). the new material described herein adds the pleurodire turtle eocenochelus as another shared faunal element between sardinia and continental western europe. rather interestingly, in this case, such shared faunal resemblance reaches almost the species level, as the sardinian material is identified as very close to eocenochelus eremberti, if not conspecific. that species was originally established from the middle eocene of northern france on the basis of both cranial and postcranial remains (broin 1977), while additional, more or less coeval, material from belgium (pérez-garcía & smith 2017) and spain (pérez-garcía et al. 2019) has recently been referred to it, demonstrating a rather broad geographic distribution during the middle eocene. the other two named species of eocenochelus are known from only from shell material from southern europe and have much more confined geographic distributions, as only one specimen has been attributed to each of them so far: eocenochelus lacombianus from the early eocene of southwestern france, and eocenochelus farresi from the late eocene of northeastern spain (pérez-garcía et al. 2017b). the new shell material from sardinia allows the recognition of a form rather close to eocenochelus eremberti. the depositional environment of the two sardinian localities that yielded the new pleurodire remains indicates a freshwater environment (in which, eocenochelus, adapted for life in littoral environments, would have arrived), demonstrating the non-marine nature of the sediments. such close or even conspecific affinities between continental western european coastal areas and sardinian eocenochelus do not necessitate direct land connection between these land masses in order to be explained. instead, eocenochelus has been suggested be a taxon adapted for coastal habitats (pérez-garcía et al. 2017b) and thus, even short marine distances between western europe and sardinia during the eocene would certainly not be a significant barrier for these pleurodires (see also pérez-garcía & chapman 2017). after all, pleurodires appear to have been rather capable of marine dispersals, with many taxa also inhabiting near shore environments (sánchez-villagra et al. 2000; gaffney et al. 2006, 2011), with such dispersal events having been repeatedly suggested for cenozoic european taxa (georgalis et al. 2013; georgalis & kear 2013; pérez-garcía & chapman 2017; pérez-garcía et al. 2017b, 2019). the 95° counterclock-wise rotation of the sardinia-corsica massif significantly shifted the position of the fossils of eocenochelus, that have never lived in the central tyrrhenian area, and therefore provides an interesting biogeographic case of “viking funerary ship” (mckenna 1973) in a mediterranean setting. finally, the occurrence of eocenochelus in the sardinian fossil record further adds to the growing evidence of the rich and diverse extinct reptilian fauna of the island that once comprised many different lineages that are currently globally extinct or extirpated from europe (portis 1901; abbazzi et al. 2004, 2008; venczel & sanchíz 2006; chesi et al. 2007a; delfino et al. 2008, 2011; georgalis et al. 2017, 2019; tschopp et al. 2018; zoboli et al. 2019). conclusions the first pleurodire material from the island of sardinia is described herein, comprising of two shells from two different, probably middle to late eocene, localities. the most complete shell is referred to the erymnochelyine podocnemidid eocenochelus cf. eremberti, while the incomplete specimen is only tentatively referred to as cf. eocenochelus sp. furthermore, the most complete sardinian specimen represents one of the most complete known shells of the genus eocenochelus, bearing more resemblance with the type species, eocenochelus eremberti, fossils of which have also been found in spain, france, and belgium. the biogeographic importance of the new eocenochelus remains is discussed, as these represent another shared related faunal element between sardinia and continental western europe. taking into consideration the former proximity of sardinia with continental western europe and the rotation of the sardinia-corsica massif from the eocene onward, we envisage that eocenochelus directly dispersed to sardinia somewhen during the eocene. acknowledgments. the shell mdlca 14006 was prepared by f. landucci (department of earth sciences, university of florence); d. righi (university of modena and reggio emilia) prepared its preliminary interpretative drawing and description in the frame of her master thesis. md thanks h. tong for her comments on mdlca 14006 and for having suggested its erymnochelyine affinities. project supported by fondi di ateneo dell’università di torino (2018-2019), the occurrence of eocenochelus (testudines, pleurodira) from the eocene of sardinia 843 generalitat de catalunya (cerca program) and spanish agencia estatal de investigación (cgl2016-76431-p, aei/feder, eu) to md. glg acknowledges support from a postdoctoral grant from the university of torino and forschungskredit of the university of zurich, grant no. [fk-20-110]. dz was supported by grants p.o.r. sardegna f.s.e. 2014-2020 asse iii “istruzione e formazione, obiettivo tematico: 10, obiettivo specifico: 10.5, azione dell’accordo di partenariato: 10.5.12 “avviso di chiamata per il finanziamento di progetti di ricerca anno 2017”. apg acknowledges funding by the ministerio de ciencia e innovación (pid2019-111488rb-i00). glp research is supported by fondi di ateneo cagliari university. glp and dz work was also supported by grants from fondazione di sardegna (grant number: f71/17000190002). the quality of the manuscript was improved by the useful comments of the reviewer walter joyce (university of fribourg). references abbazzi l., angelone c., arca m., barisone g., bedetti c., delfino m., kotsakis t., marcolini f., palombo m.r., pavia m., piras p., rook l., torre d., tuveri c., valli a. & wilkens b. 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(2019) an overview of the crocodylian fossil record from sardinia (italy). annales de paléontologie, 105: 123-137. _goback _goback _goback _goback _goback _goback _goback _goback _goback _goback _goback _goback _goback riri'ra irllirna di paleonrologir e strarigrr l;-;l *;;,r short note nota breve metaxytherium medium (mammalia: sirenia) from upper miocene sediments of the arenaria di ponsano formation (tuscany, itaty) giovanni bianucci s{ \talter landini receioed march 19,2aa2; accepted marcb 2g,20a3 key raords: mammalia, strenia, metaxvtberium med,ium, systematics, late miocene, tuscany, italy. abstract, records of metaxytherium medium (mammalia: sirenia) from tortonian (late miocene) sediments from the arenaria di ponsano formation (tuscany, ftaly) are described. they consist of fragmentary specimens, including several partìal cranial elements representing at least three skulls, two humeri, fraqments of vertebrae and some incomplete ribs. the neu, tuscan records confirm the wide diffusron of metaxytherìum in the mediterranean during the miocene. this sirenian's occurrence in the arenaria di ponsano sediments is in accordance with the shelf environmenr indicated by other fossils. the low sea botton was at least parrially covered by segrass meadows, the food source of this dugongid. riassunto. vengono descritti resît di metaxytheriam medium (mammalia: sirenìa) provenienri dai sedimenti tortoniani (miocene superiore) dell'arenaria di ponsano (toscana, italia). si tratta di reperti fr.rmmentari rappresentati prevalentemente da diversi elementi craniali appartenenti almeno a tre distinti crani, due omeri, frammenti di vertebre e alcune coste incomplete. questa segnalazione conferma i'ampia diffusio ne di metaxytherium nelmedtterraneo duranre il miocene. il ritrovamento di sirenia nei sedimenti dell'arenaria di ponsano è in accordo con l'ambiente di piattaforma indicato anche dagli altri fossili. il basso fondale marino era almeno in parte ricoperto da praterie algali, la sorgenre di cibo di questo dugongide. introduction metaxytberium is an extinct dugongid sirenian that shows a wide geographical and biostratigraphical distribution. in fact, metaxytherium rernains have been collected in several early-late miocene localities of the mediterranean region (see below; summary in domning & thomas 1,987), northern france (corrreau 1928), north and south america (muizon br domning 1985; domning 1988; aranda-manreca et al. 1994), and also in pliocene sedimenrs of the mediterranean basin (domning & thomas 1987). the miocene specimens from mediterranean and adjacent areas were collected mainly from central and tvestern paratethyan deposits of austria and switzerland (domning & pervesler 2001), from spain (pilleri er al. 1989), and from many localities of southern italy and sardinia (moncharmot zeí t< moncharmont 1986; carone 1997; bianuccr et al. 2003). new specimens referred to metaxytheriwm mediwm, collected rn 1965 by dr. angelo varola from tortonian sediments of the arenaria di ponsano formation near volterra (tuscany, central ltaly) (fig. 1), are described in this paper. the sirenian remains examined here belong to an undescribed collection of marine mammals, containing also odontocete and mysricere specimens kept in the museo di storia naturale e del territorio (msnt) of the university of pisa. anatomical terminology follows domning (1988). systematic description. class mammalia linnaeus, 1258 order sirenia illiger, 181 1 family dugongidae gray, 1821. subfamily halitheriinae (carus, 1868) abel, 1913 genus metaxytberiun de christol, 1840 metaxytherium medium (desmarest, 1822) hooije r, 1952 (figs.2-5; pls. 1,2) dipartimento di scienze della terra, via s. maria, 53,i-56126 pisa (italy). e-maìl: bianucci@dst.unipi.it; landini@dsr.unipi.it. 568 referred specimens. several fragments belonging to at lerst three animals; among these the most significant parts are two skull roofs (msnt i10140; msnt i10095), one right zygomatic process of the squamosal and the associated jugal (msnt 10068), right (msnt 110141) and left (msnt 110138) humeri probably of the seme anìmal, a fragment of a right scapula (msnt i10148), and some incomplete r-ertebrae and ribs. description frontal the dorsal surface of the preserved posrerior portion of the frontals of msnt i1 0095 is flat and narrow and it is laterally delimited by rounded and slightly laterally concave temporal crests (fig. 2;pi. 1,, fig. 2) . the crests are separated by a distance of at least 27 mm. parietal temporal crests on the parietal roof of msnt 110140 correspond to the "type e" morphology described by domning (19s8) for metaxytheriwmflorìdanum (fig.3; p1. 1, fig. 1). they are prominent and lyriform and they come closest to the midline (12 mm) about fig. 2 skull roof (msnt 110095) in dorsal vìew of metaxytberium meclíum from the arenaria di ponsano formation. fig. 1 geographic location of the arenaria di ponsano fonnarion wlrere rhe.irenirn specimens examined in this paper were collected. the numbers refer to the fossiliferous outcrops (moclìfied from menesini 1967a). 55 mm anterior to the nuchal crest. the median valley between these temporal crests is concave and about 14 mm deep. in msnt 10095, the temporal crests are less prominent and there is not a deep median valley. the crests of this specimen are of the "type a' of domning (1988). the parietal roofs of the po:-rsano specimens are narrower than in any of the specimen s of m. floridanum. internally, the parietal of msnt i10140 exhibits an evident bony falx cerebri that runs anteroposteriorly from the frontoparietal suture to the internal occipital protuberance. supraoccipital this bone is entirely preserved in msnt 110140 and partially in msnt 110095. in lateral view, the supraoccipital of msnt i10140 forms an angle of about 125'with the parietal roof. the nuchal crest is relatively rounded and the external occipital protuberance rises above plane of parietal roof, and below it, the median ridge separates two evident rugose semispinalis muscle insertions. the supraoccipital-exoccipital sutures form an angie of about 145". jugal the jugal (msnt i10148) is laterally broad and triangular, and it has straight posteroventral and anteroventral margins forming an angle of about 100" (fig. 4; pl. 1, fig. 3). the ventral tip of the jugal is located under the orbit. squamosal only an anterior fragment of a left zygomatic process is preserved (msnt 10068). the squamosal-jugal suture is straight. scapula a fragment of a right scapula (msnt i10148) is preserved (p1.2, fig. 6). the glenoid fossa is moderately concave and oval in shape; its greatest diameter is 48 mm and the smaller is 36 mm. flumerus the epiphyses of the two humeri are worn and the tubercles. heads and trochleae are not well g. bianucci & v/. landini supraoccipita i meolan ridge nuchal crest temporal crest ..--l: fro ntopa rieta i sutu ra nuchal crest external occip ita i protu berance med ia n ridge pa rieta i median valley 5cm internal medial ' p rolu bera nce bony falx cerebri temporal cresls supraoccipital fig. 3 skull roof (msnt i1al4a) oí metaxytherium mediunt ftom the arenaria di ponsano formation. a, dorsal vieu'; b, ventral r.iew; c, lateral vien'. preserved (fig.5; pl.2, figs. 1,2). the srrong right humerus (msnt i10141) is more than 205 mm long at it exhibits a prominent and elongated deltoid crest. velldeveloped tubercles are separated by a deep and wide bicipital groove. the head apparently is relatively small (but it is worn) and hemispherical. the trochiea is anessef t!bercle }{ttmer of mearytherium medium fron the arenaria di por.rsano formation. a, rìght humerus (msnt i10111) in medial vieu'; b, the same in anterior vien'; c, left humerus (msnt i10138) in anterior view. metaxytherium medium from miocene of twscany 569 sq uamosal fi,1 portion of right squamosal and jugal (msnt 10068) in lateral vìew of metaxytherium medium from the arenarìa di ponsano formation. teroposteriorly flattened and the oleocranon fossa is wide. the left humerus is almost as anreroposteriorly elongated as the right, but its diaphysis is unusually thin and apparently lacks the deltoid cresr. this bone might have suffered a pathological malformation (fornaciari, pers. com.). vertebrae only vertebral fragments are known. a small portion of an atlas, a corpus of a cervical vertebra, and some incomplete neural arches and transverse processes of thoracic vertebrae are preserved. ribs several fragments and some almost complere ribs are preserved (pl. 2, figs. 3-5). comparison. the' lragmentary sirenian remalns from the arenaria di ponsano are referable rc metaxytberium medium because: some well-preserved skeletal elemenrs, such as the skull roof and the humerus, show close resemblances to those of metaxytherium: temporal crests of "type e" as in msnt i1o140, and a very narrow parietal roof as in msnt 10095, are mosr frequent in m. medium, among the species of this genus; m. mecjiu;,a is the only known sirenian species in the late miocene of the mediterranean and easrern north atlantic. age and stratigraphic occurrence. all specimens were collected in the arenaria di ponsano, a miocene formation that consists of sandstone and marls ourcropping in three distinct areas between volterra and siena towns (tuscany, italy) (fig. 1). the main outcrop is in the area of ponsano farm, about 10 km southeastern of volterra. the sirenian fossils here examined were collected in this ponsano area where a succession over 500 m thick of sandy sediments representing the upper portion of the formation are exposed (giannini ec tongiorgi 1959 mazzanti et al. 1981; foresi et al. 1997). these sediments were attributed by foresi et al. (.1997) to the lr,leogloboquadrina acostaensis zone. this zone is referable to the early tortonian (late miocene) and to an ase interval between about 10.5 and 8.14 ma. fìo q 570 g. bianucci 8t tà/. lanjini in the 43 recognized fossiliferous outcrops of this area were collected also fishes (menesini 1967a), mollusks (tavani & tongiorgi 1963), echinoids (menesini 1967b), cirripeds (menesini 1963, 1966), foraminifers (bartolani 1966; foresi et al. 1,997) and calcareus nannopiancton (bartolani & pirini 1969). discussion and conclusions. specimens referred to metaxttherìum and here described represent the first miocene sirenian occurrence from tuscany and they confirm the wide diffusion of this genus in the mediterranean area during the miocene. from a paleoecological point ofview, the occurrence of sirenian remains contributes for a better definition of the environment already indicated by other fossil records. particularly, the ichthyofauna indicates a subtropical and shallow marine environment with possible estuarine conditions (menesini 1967a). the analysis of the foraminifers indicates deposition in the circalitoral-infralitorai zones and. relativelv to the upper part of the succession, a deltaic condition lforesi et al. 19871. these data are confirmed by the sirenians, considering that, as pointed out by domning (2001) and domning tc pervesler (2001), a1l fossil dugongids apparently have been exclusively marine shallow-water and (eccept for hydrodamalines) tropical. besides, the presence of sirenians indicates that the low sea bottom was at least partially covered by segrass meadows. infact domning (2001) hypothesized lor metaxytherium spp. a diet based on leaves and rhizomes of smal1 and mid-sized seagrasses in water deeper than 1 meter. aclenou;ledgemenrs. special thanks are due to d.p domning (laboratory of paleobiology, department of anatomy, howard university, ril/ashington, dc) for very useful suggestions and for a critical reading of a first version of the manuscrìpt. \ùfe also thank a. varola (museo dell'ambiente, università di lecce), who collected and restored the specimens here examined, c. nocchi (museo di storia naturale e del territorio, università di pisa) for her help in the cataloging of the ponsano fossil collection, and c. sorbini (dipartimento di scienze della terra, ljniversità di pìsa), for fruitful discussions and suggestions. the editor (m. gaetani) and c. sagne (muséum national d'histoire naturelle de paris) made costructive comments on the manuscript. metaxytherìum nediurn frorn the arenaria dj ponsano formarion. vies.s; 2, skull roof (msnt i10095) in leteral (a) and dorsal (b) metaxytherium medium from miocene of tuscany 5cm plate l l,skull roof (msntil0l+0) ìndorsal (a), ventrel(b) ,letcral (c) andposterior(d) vìen's; 3, portion of right squnosal end jugal (msnt i10068) in lateral vieu.. 571 572 g. bianucci &-'v/. landint plate 2 metaxytheriunt mediun from the arenarja di i'onsano form:rion. i, right humeru' (msnt i10141) in humerus (msnt 110138) in anteriorvieu,; 3,,1,5, ribs;6, frrqnent r{ rirht sc.rpul.r (msnt i10148) in fossa (b). mcdial (a) and anterjor (b) views; 2, left lateral vier' (e) end distal vien-of glenoid aranda-manteca f. j., domning d. p e barnes l. g. (1994) a new middie miocene sirenian of the genus fuletaxytherium fron'r baja california: relationships and paleobiogeograohic in-rolications. in: berta a. er deméré t.a."_'ò'_-ò_*r'__-'^_'r (eds.) contributions in marine mammal paleobiology honoring frank c. tmhitmore, jr. prctc. san diego sctc. nat. hrst., 29: 191-204, san diego. bartolini c. (1966) i foraminiferi dell'arenaria di ponsano. palaeont. it., 61: \7 -64, prsa. bartolini c. & pirini c. (1969) découverte de nannoplancton calcaire dans ìe grès de ponsano, miocéne moven, toscane, italie. in brònniam p& renz h. h. (eds.), proc. first int. conf. planlet. microfossils: 81-bb, geneva. bianucci g., landini'm & varola a. (2003) ner.v records of metaxytherium (marnmalia: sirenia) from the late miocene of cisterna quarry (apulia, southern ltaly). boll. soc. paleont. it., 42 (1-2):59-63, modena. carone g. (1997) metaxytherium medium (desmarest) 1822 (dugongidae, sirenia, mammalia), delle arenarre tortoniane di cessaniti (calabria, kalia). atti soc. it. sci. nat. mus. ciis. st. nat. milano, 137:91-fi0, milano. cottreau j. (1928) le l[etaxytherium cw,uieri du golfe de la r ^ 7 | t\i otre. hnn. i'dteontot., i /: j-lu, l'jrl\. domning d. p (1988) fossil sirenia of the \il/est atlantic and caribbean region. i. metarytherium floridanwm hay, 1922. jour. veft. paleortt, 8(4):395-426, northbrook. domning d. p (2001) sirenians, seagrasses, and cenozoic ecological change in the caribbean. palaectg., palaeocl., pal eoecol., | 6b: 27 -5a, a mstcrdem. don.rning p d & perr.esler p (2001) the osteology and relationships of metaxytberium lerahuletzl deperct, 1895 (manimalia: sircnia). abh. senck. natur. ges.,553: 189, frankfurt. domning p d & thon-ras h. (1982) metaxytherium serresii (mammalia: sirenia) from the early pliocene of libye and france: a reevaluation of its morpholog,v, phyletic position, and biostratigraphical and paleoecoloeical significancc. in: boez n.t., el-arnauti a., gaziry a.\{1, de 573 heinzelin j. & boaz d. d. (eds.) neogene paleorttologt and geologt of sahabi:205-232, new york. foresi l. m., pascucci v & sandrelli f. (1997) larenaria miocenica di ponsano (toscana, italia): evoluzione paleoambicntale e bio-cronostratigrafica. boll. soc. pal. it., 36: t3-na, modena. gi.rnnini f. & tongìorgì m. 11059.1 stratigrafia neogenic.t toscana. 1 larenaria elveziana di ponsano (yolterra). boll. soc. geol. [t.,78(2): 83-100, roma. mrzzanti r., mazzei r., menesini e. & salvatorini g. (1981) larenaria di pons:rno: nuove precisazioni sopra l'età. in: ix cont,egnct della società paleontologica ltaliana: 1351 60, pisa. menesini e. (1963) cirripedi miocenici de lle 'arenarie di ponsano" (volterra, prov. di pisa). atti soc. tosc. sci. nat., mem., ser. a, 70(1): 15-22, pisa. menesini e. (1966) i balani miocenici delle'arenarie di ponsano" (volterra, prov. di pisa). palaeont. i t., 6a: 99 -129, pisa. menesini e. (1967a) i pesci miocenici delle 'arenarie di ponsano". atti soc. tosc. sci. nat., mem., ser. a, 74(1): 122, pisa. menesini e. (1967b) gli echinidi n-riocenici delle 'arenarie di ponsano" (volterra, provir-rcia di pisa). palaeont. (t.,60: 143-167, ptsa. moncharmot zei m. & moncharmont ij. (1986) i1 metaxytherium medium (desmarest) 1822 (sirenia, mammalia), delle arenarie tortoniane (miocene sup.) di s. domenica di ricadi (catanzaro, lr.lit). mem. sc. geol.,39 285-341, padova. muizon c. de & domning d. p (1985) the first records of fossil sirenians in the southeastern pacific ocean. bull. mus. natl. hist. nar., section c,'[ ser., 7:189-21.3,paris. pilleri g., biosca j. & via boada l. (1989) the tertiary sirenia of catalonia. v of 98 pp. ostermundigen, brain anatomy in.titute. varnnrala. tavani g. & tongiorgi m. (1963) la fauna miocenica delle 'arenarie di ponsano" (volterra, prov. di pisa). i parte. lamellibranchiata. palaeont. 1r., 5b: 1-43, pisa. ?i?a references key'oords; taxonornr-, gatropods, naticids, pliocene, northcrn itaìr.-. riassuttto. viene descritta e figurate euspira ntagctto r nuor':r specie di naticidae rinvenuta in depositi argiijoso-sabbìosi plioccnici csposti nelle località di r:io rosello (emilia, provincia di piacenza) e vjllalvernia (piemonte, pror,ìncie dì alessandria). l:r specie, di piccola tagììa. e car.rtterizz.rtr d.r sutura canalicolata, ombelico :rbbastenza ampio e profondo, nonché dell'epertura prolungata abapic:rlmente. si tratta di un elemento poco comunc, probabilmente inlralitorale e leeato a frcies di transiz-jone trl paleobioccnosì analoghe a queile rrru.ììi medìterranee sfbc e dc. abstract. the nen. n:rtìcic1 specics euspira magenesz is describecl rnd figurcd. thc t1-pc-nr:terial u'as recovered fron plioccne deposits exposcd along the strearr rìo rosello (emilia resjon, pi;rcenza province) and ncar villalvernia (piedmont, alessandria pror,:ince). the snrall-sizecl euspira magenesi is featured b1. channeled sllturej open, rather n-ide umbilicr.rs trnd :bapicrìly prrtlucrd .ìperrure. it is :rn urlcorrìmon infrelittoral element ìike1v related ro ccorones betu.een palcobiocoenoses sìmihr to the modern mediterranean sfbc and dc. introduction in the course of field work aiming to recover narlcids from the pliocene outcrops along the stream named rio rosello near sariano (piacenza province), ren specimens of a small polinicine spcc;es were found. the material was obtained from a lenticular body of clayey sand that crops out on the right bank of rio rosello, about 280 m south west of case badini di sopra (fig. t). the calcareous nannofossil contents of the sandy clay resulted to be mostly ren'orked and gave a general pliocene age (p maior.rno, wrirren communication, 2000) . according to s. raffi (oral communicrtion, 2001), this lithotype belongs to the monte zago unit of piacenzian age as treated by roveri et al. (1998). an 'l additional specimen was recovered later on in piedmont, from tr layer of fine sand exposed on the right bank of the stream rio vaccaruzza northeast of villalvernia (fig. 1). the sandy layer pe rtains to the uppermost p.1rr of the pliocene formation known as argille di lugagnano (lugagnano clay) . for further information on this latter loc.ility, reference can be made to brambilla (1976). the species, that fits in with the characters of the genlrs euspira agasúz in j. sowerby, 1837, after thorough examination of literature data and of number of neogene naticids in the bellardi-srcco collection (university of torino), proved to be previously undescribed and is discussed and figured herein. systematic account the suprageneric rrr.rngemenr is th:rt adopted in recent major revisions of the family naticidae (kilburn 1976;marrncovìch 1977; majima 1989). we follow majima (1989) and bouchet & waren (1993) in ranking euspira agassiz in j. sowerby, 1837 as a full genus of the subfarnily polinicinae. the bulk of the studied material is housed in the museo g. cortesi di castell'arquato (mgc); paratype 551/gf (unfigured) is stored in the museo di ecologia e storia naturale di marano sul panaro, modena, paratype 23957 (unfigured; in the collezione malacologica of the museo di geologia e paleontologia g. cappellini, bologna and paratype mzb 18569 (unfigured; in the laboratorio di malacologia, bologna. symbols for shell dimensions are: nw number of whorls; pd, diameter of the protoconch; h, height of the shell; d, maximum diameter; ha, height of the aperture; msa, mean spire angle. rivista italiana di paleontologra e strtrtigrafia uiccnidte iuu nota breve-short note euspira magenesi, a ne\t species of the naticidae (gastropoda) from the pliocene of italy luca pedriali': & elio robba'}'i recehed july 5, 20al: accepted septetnber 21, 2a01 " vja s. pertini 29,i11a16 san martino, ferrara, itali,'. 'r': dipartimento di scienze gcologichc e geotecnologje, unìr'ersità degìi studi di mìlano-bicocca,piazzt clella scienza 4,i2a126 milano, italr'. e maìl: elio.robbacaunimib.it 484 l. pedriali €' e. robba family naticidae forbes, 1838 subfamiiy polinicinae finlay er marwick, 1937 genus euspira agassiz in j. sowerby, 1837 the eocene natica glaucinoides j. sowerby, 1812 is the type-species by subsequent designation (bucquoy, dautzenberg & dollfus 1883, p. 1a3). the synonymy of natica glaucinoìdes with natica labellata lamarck, 1804, stated by cox (1930) and accepted later on by several authors, was rejected by vrigley (1949) who regarded the two species as distinct and redescribed sowerby's taxon. according to kabat (1991), laguncula benson, 1.842, bensonia gray, 1847, lunatìa gray, 1847, ampul' lonatica sacco, 1 890, labellinacca cossmann, 1'918, dallitesta mansfield, 1930, scarlatia schileyko, 1,977 and pseuclopolinices golikov er sirenko, 1983 are synonyms of euspira. fig. 1 location map of the typelocrìir; o[ [uspira mage't,si sp. n. (bottom, right) and of the orltcrop near villalvernia (bottom, left). the diagnostic characters o{ euspira are 1) shell thin to moderately thick, globose or globose-elon gate, 2) protoconch low-turbiniform of r.5-2.5 smooth whorls, 3) sprre depressed to moderately elevated, somewhat stepped in some species, 4) sutllre we akly impressed to channeled, 5) umbilicus open rnd deep. narrow to wide, 6) funicle weak to absenl, 7) parietal callus moderately thick, slender, n'ith distinct anlerior lobe slightly overhanging the adapical part of the umbilicus, 8) umbilical callus indistinct to rroderately developed, merging into the anterior lobe of the parietal callus or demarcated from it by a weak groove. 9) corneous operculum. euspira is a distinctive cosmopolitan genus useful to accomodate polinicine species with the open (not slitlike) umbilicus virtually devoid of funicle and with strongly reduced to absent umbilical callus. six well known species from the italian pliocene do belong to euspira, namely natica fusca de blainville, 1825, lwnatia grossularia marche-marchad, 1957, l"latica guillemini payraudeau, \826, nerita helicina brocchi, 1.81.4, natica macilenta philippi, 1844 and natica pulchella risso, 1826. sacco (1891) created several varieties oí euspira catena (da costa, 1778) that, at least partly, belong here. they will be dealt with in a forthcorning revision o1 pliocene naticids (pedriali & robba, in preparation). f19.1 fig. 3 plate 1 euspira magenesi sp. n. holotype. rio rosello. mgc 567; a) apical n'horls arrorv), b) top view, c) protoconch, d) apertural side, e) abapertural side' f) base, ì) umbilical cavitl4 l) and m) sculpture of the umbilical wall euspira magenesi sp. n. paratype. rio rosello. mgc 568; apertural side. euspira magenesì sp. n. paratype. villalvernia, mgc 523; apertural side. (the protoconch-teleoconch transition indic;rtcd by spire, g) detail of the channeled suture (arrow), h) euspira //tlrga//est 486 l. pedriali & e. robba euspira magenesi sp. n. pli t, fig. l-3 derivation of name. the spccìes is nlmcd after paolo mirgcnes s'ho provided the type-meteriai. holotype. rio rosello: mgc 562 (p1.1, fie. 1a-m). paratypes. rio rosello: 5 spms., mcc 568-5/2; 1 spm., 551/gf; 1 spm.,23957; 1 spm., mzb 18569; villalvernia: 1 sprn., mgc 573. preservation. txcept for minor danages of rhe perisromc ìl some specimens, the preservation is fair. type-locality. rieht benk of the srream rio rosello, 280 m south west of case badìni di sopra ne:rr rhe village of sari;rno (piacen 7,x) . horizon. lcns-shaped intercalation oi clavel, send in the monr,. z.rgo unit, lon cr pìrienzi.rn. diagnosis. depresscd-globose, thin shcll u.ith moderatelv high to depressed, somervlrar stepped spire. protoconch o[ l.l5-2 smooth s.horls. sutures channeled. umbilicus wide and decp, bounded bv a sharp :rngulation. umbilìcal n.all excar-ated, sculpured with spiral cords crossed b1. coarsc growth lines. aperture hrgh, projuccd abapicallr: parietal callus with slender anterjor lobe reaching the basal fasciole. funicle wanting. umbìlicel callus modcratel). d"""'oo.o, attachcd to the adapical part of the umbilicus and nerging ìnto the anterior lobe of the pariet:rl callus. description. shell small, only slightly exceeding z mm in height, thin, depressed-globose, with moderately elevated, obtuse spire that is about 2a 'k oî the shell height. protoconch low-turbiniform of 1.75-2 apparently smooth whorls, faintly but distinctly demarcated from the teleoconch because of change in the shell texture; the diameter averages 0.93 mm. spire somewhat stepped, whorls convex, meering rt deeply and broadly channeled sutures. body whorl large, somewhar depressed, rather quickly expanding and produced abapically toward the aperture; periphery sliehtly above midìine. umbilicus wide and deep exposing earlier whorls, clearly demarcated from base by a sharp enguhtion; umbilical wall descending steeply ro a narro.w and shallow spiral groove, rhen excavared to form a broad spiral depression sculptured with uneven, low spiral cords brossed by coarse gros/rh markings resulring in xn irreg;ular square-reticulated pattern. basal fasciole poorly differentiated. aperture ovarely d-shaped, slìghtly obiique, markedly produced abapically, nearly as high as twice its width. parietal callus thin to moderarely rhick, reaching the basal fasciole; anrerior lobe slender, with pointed edge slightly extending over the adapical part of the umbilicus. funicle absent. umbilical callus moderately developed, triangular to semicircular, occasionally with faint transverse furrow; it attaches to the ad;rpic.rl part of the umbilicus and merges into the anterior lobe of the parietal callus. operculum unknown, supposed ro be corneous. outer surface smooth except for dense, thin grosrth lines that are coarser over rhe basal fasciole; a faint spiral striation is noted on lower base. nicely preserved specimens exhibit a yellowish background witl-r fine, distant reddish collabral lines, in some insrances restricted to parr of the body n'horl; surural channel reddish-brown, with adaxial blackish band. dimensions (mm): n\l pd h d ha msa mgc 567 (holotl.pe) 4.00 0.92 1.03 4.05 3.31 1oo" mgc 568 4.25 0.e9 1.31 4.16 3.4r 96" mgc 569 ,t.50 0.95 mgc s70 4.75 8.oo mgc 571 4.00 3.58 1.02 2.63 1os. mgc 572 .t. 10 3.65 1.13 3.a7 115" mgc 573 4.75 a.96 7.a5 7.11 6.41 109. remarks. the present new species fully matches the characters of the genlrs euspira agassiz in j. sowerby, 1837 as summarized above, and shares the most significant features wrth euspira glaucinoides (j. sorverby, 1812). the type-species has markedly globose shell with higher spire, impressed suture and somewhat smaller tumbilicus devoid of inner spiral sculprure. euspira mtlgenesi is easily distinguished from the others euspira species in combining 1) somewhat stepped spire,2) channeled sutures, 3) depressed body whorl, 4) rvidely open umbilicus, 5) sculptured umbilical wall and 6) high, abapically produced aperture. the miocene north european euspira nysti (d'orbrgny, 1852) and euspira gottschei (kautsky, 1925) are closely related, but have lower and broader, more distinctly dshaped aperture (cf. janssen 1984, pl. 55, figs. 4, 5). euspira fusca (de blainville, 1825), euspira grosswlaria (m.irche-marchad, 1957), euspira guìllemini (payraudeau , 1826), euspira beìicina (brocchi, 1814), euspira macilenta (philippi, i844) and euspira pulchella (risso, 1826), occurring in pliocene deposits of italy, attain a larger size, have more globose, more robust shell with higher spire and smaller umbilicus. the channeled suture is clearly developed in the recent north atlantic euspira montagui (forbes, 1838). this species, also small-sized, differs from ewspìra magenesì rn having more globose shell, larger protoconch (1.2 mm in diameter) with spirally sculprured first whorl, lower and broader aperrure and unsculptured umbilical wall. the spiral sculprure of the umbilical wall occurs rather commonly in species of the polinicinc genus glossaulax pilsbry, 1929 (cf. majima 1989) whereas it is unusual in euspira. it is noted in euspira catena (d^ costa, 1778) and appears to be particularly coarse in pliocene specimens from belgiurn. in the norrheasrern pacific area (cf. maricovich 1977), the miocene to recent polinices (euspira) le.raisii (gould, 1842) and the pliocene to recent polinices (euspira) draconis (daii, 1903) exhibit a spirally sculprured umbilical wall. all these species differ lrom euspira mdgenesi in several respects and none of them has either channeled sutures or an abapically produced aperture. euspira magenesi is an uncommon taxon so far recovered from piacenzian deposits of piedmont and western emilia regions. the rather fragile shell and the smalì size likely have contributed to the failure to find specimens in the past. furthe r work n-ray expand the distribution and refinc the stratigraphic range that, on the basis of the present material, appears to be restricted to the upper piiocene. it is difficult to infer the paleoecological meaning of euspira magenesi due to the scanty records. in the type-localitv, it belongs to a fossil association characrerized by the abundant occurrence of circomphalus foliaceolamelloszs (dillwyn, 1817) and callista italica (defrance, 1818), and interprered (s. rrffi. oral communication, 2001) as an infralittoral ecotone between pliocene counterparts of the modern mediterranean biocoenoses sfbc (biocoenosis of fine \vell sorted sand) and dc (biocoenosis of the coastal detritic) . the mollusk association at villalr-ernia comoares more closely to a modern sfbc biocoenosis. accordingly, euspira megenesi is likely to be regarded as a tolerant sand-related, infralittoral element that has dwelled in sfbc-influenced pliocene serrrnss. acknouledgtnents. this paper has benefìted from critìcal reading bv s. raffi, university of bologna and from revier.ing b,v a. varen, swe dish nluseurn of natural histor1., stockholm. f. campanino, museo di geologi,r e paleontolo{ìia, llniversitv of torino, provided àccess ro the bellardìsacco collection. m. grìgis, ninove (belgium), donated specimens of euspird cdtena from the pliocene lillo formation of belgiun-r. g. della bella, monterenzìo, bologna, ailon ed access to his collccrìon. x4. mariani, civica stazione idrobiologica, mjlano, essisted u'ith photocopies of some essentiri refcrences. nannoplancton anall.ses bv p maìorano, llnivcrsitv of bari, are also acknou'ledged. the scanning electron micrographs were made b1.a. rizzi, unìr'ersity o{ milano. the computer-assernbled platc w;.ls prepared by f. facchini, san martino, irerrara. the research u-as granted bv the unri.ersrtv of milano-bicocca. 487 references bouchet p &'waren a. 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accepted: february 11, 2011 1 dipartimento di scienze della terra, università di cagliari, via trentino 51, i-09127 cagliari, italy. e-mail: corradin@unica.it 2 steinmann institute of geology, mineralogy, palaeontology; university bonn, nussallee 8, d-53115 bonn, germany. e-mail: sakaiser@ uni-bonn.de 3 dipartimento di scienze della terra e geologico ambientali, alma mater studiorum-università di bologna; via zamboni 67, i-40127 bologna, italy. e-mail: mariacristina.perri@unibo.it 4 dipartimento di scienze della terra e geologico ambientali, alma mater studiorum-università di bologna; via zamboni 67, i-40127 bologna, italy. e-mail: claudia.spalletta@unibo.it key-words: biostratigraphy, taxonomy, devonian/carboniferous boundary, conodonts, protognathodus. abstract. the current definition of the devonian/carboniferous boundary is the first occurrence of the conodont siphonodella sulcata. due to difficulties in identification of the early siphonodellids, such as s. praesulcata and s. sulcata, investigation of protognathodus which enters in the latest devonian and extends into the mississippian, was undertaken to determine use as a better indicator of the base of the carboniferous. during the d/c boundary interval, protognathus is represented by four species: pr. meischneri, pr. collinsoni, pr. kockeli and pr. kuehni. although pr. kockeli can be abundant in boundary interval sections, none of the four protognathodus species has a high potential as a tool for redefining the d/c boundary, based on regional variation in first occurrence data, restricted stratigraphic ranges and global distribution, poorly understood facies occurrences, as well as general rarity of the taxa. riassunto. la base del carbonifero è definita dalla prima comparsa del conodonte siphonodella sulcata nella linea evolutiva si. praesulcata-si. sulcata. poiché la identificazione dei primi siphonodellidi, quali s. praesulcata e s. sulcata, presenta difficoltà, è stato esaminato il genere protognathodus, la cui distribuzione stratigrafica si estende dalla parte più alta del devoniano al mississippiano, al fine di valutare se fosse un indicatore migliore per la definizione del limite basale del carbonifero. durante l’intervallo di tempo comprendente il limite devoniano/carbonifero il genere è rappresentato da quattro specie: pr. meischneri, pr. collinsoni, pr. kockeli and pr. kuehni. benché pr. kockeli sia a volte abbondante in alcune sezioni che comprendono il limite, nessuna delle quattro specie di protognathodus presenta elevate potenzialità per la ridefinizione del limite stesso, a causa della variabilità regionale dei dati di comparsa, di ridotte distribuzioni stratigrafiche e geografiche a livello globale, imputabili forse ad una dipendenza dalle facies non ancora ben chiarita, e, soprattutto, ad una generale rarità delle specie. introduction the base of the carboniferous system is defined by the first appearance datum (fad) of the conodont species siphonodella sulcata, within the s. praesulcata-s. sulcata lineage (paproth et al. 1991). the global stratotype section and point (gssp) is located in the la serre trench e’ section, montagne noire, france. flajs & feist (1988) published a biometric study of s. praesulcata and s. sulcata based on the la serre faunas, demonstrating that transitional forms are very common. despite these taxonomic uncertainties, the fad of s. sulcata was chosen to define the base of the tournaisian. further studies on the stratotype section have revealed a series of problems, such as lack of other important stratigraphic guides and the existence of reworking (e.g., flajs & feist 1988; ziegler & sandberg 1996; casier et al. 2002; kaiser 2009). additionally, difficulties in discriminating s. praesulcata from s. sulcata arose quite immediately (e.g., ji 1987a; flajs & feist 1988; wang & yin 1988). a redefinition of the devonian/carboniferous boundary (dcb) was reputed nec16 corradini c., kaiser s.i., perri m.c. & spalletta c. essary, and in 2008 the international commission on stratigraphy established a working group with the goal to propose a new criterion for defining the boundary and finding a new gssp. a taxonomic overview and revision of the early siphonodellids is in progress (kaiser & corradini 2011), but also other conodont genera should be considered for evaluating their potential for identifying the dcb. this paper addresses the genus protognathodus, a taxon with stratigraphic potential around the dcb and well known since ziegler (1969) introduced the “protognathodus fauna” to define the youngest devonian strata, between the “costatus” and the “s. sulcata-p. kockeli” zones. later, ziegler & sandberg (1984) defined the base of the upper praesulcata zone by the entry of pr. kockeli. four species of protognathodus are known in the time frame across the dcb: pr. meischneri ziegler, pr. collinsoni ziegler, pr. kockeli (bischoff) and pr. kuehni ziegler & leuteritz. the goal of this study is to summarize the current knowledge of these species, and take in to account their original taxonomic diagnosis and definitions, precise biostratigraphic ranges, and geographic distributions, in order to evaluate the possible use of one of these taxa to define the base of the carboniferous. phylogeny and stratigraphy of the early protognathodus genus protognathodus comprises six species: pr. meischneri ziegler, pr. collinsoni ziegler, pr. kockeli (bischoff), pr. kuehni ziegler & leuteritz, pr. praedelicatus lane et al. and pr. cordiformis lane et al. morphological features and stratigraphic distribution of these forms allow discrimination of two groups: an early group, including pr. meischneri, pr. collinsoni, pr. kockeli and pr. kuehni, occur in the uppermost famennian and lower tournaisian; in these species the anterior margins of the cup are opposed. a late group, ranging in upper tournaisian, includes pr. praedelicatus and pr. cordiformis, which have larger cups that have slightly offset anterior terminations. lane et al. (1980) suggested that pr. praedelicatus is derived from pr. kockeli, but there is a short stratigraphic gap between the two groups, corresponding to part of the lower crenulata zone. the appearance and evolution of the genus protognathodus is easy to follow in the latest devonian (fig. 1) as documented by ziegler (1973). protognathodus meischneri, the oldest species of the genus, evolved from bispathodus stabilis (ziegler, 1969) by a variation in the position and shape of the basal cavity (cup). in protognathodus the basal cavity extends to the posterior end of the element and is more rounded and inflated; pr. meischneri has an almost symmetrical and unornamented cup. this happened during the late famennian, contemporaneous to the first occurrence of representatives of the genus siphonodella or slightly before, as suggested by the occurrence of a single specimen of pr. meischneri in the upper expansa zone in sardinia (corradini et al. 2003: tab. 2). in the very early part of its range pr. meischneri gave rise to pr. collinsoni, characterized by the occurrence of scattered nodes on the upper surface of the slightly asymmetrical cup. both of these species range to within the upper duplicata zone (over 1992). protognathodus kockeli, characterized by an asymmetrical cup covered by at least one longitudinal row of nodes, evolved from pr. collinsoni just after the fig. 1 phylogeny and stratigraphic distribution of early representatives of genus protognathodus across the d/c boundary plotted against selected events and aligned to the biozonation schemes after sandberg et al. (1978), ziegler & sandberg (1990) and kaiser et al. (2009). sketches are based on the holotypes figured in ziegler (1973). protognathodus (conodonta) and its potential as a tool for defining the devonian/carboniferous boundary 17 hangenberg event. it ranges up to the lower part of the lower crenulata zone in north america (sandberg et al. 1978), whereas elsewhere it is limited to the sandbergi zone. protognathodus kuehni is distinguished by robust transverse ridges on the upper surface of the cup running radially from the margins to the carina, that can be suppressed. it evolved from pr. kockeli and ranges from around the present dcb to the sandbergi zone (lane et al. 1980). repository of the figured specimens. dstc: dipartimento di scienze della terra, università di cagliari; ic: dipartimento di scienze della terra e geologico ambientali, università di bologna, alma mater studiorum; ipum: museo di paleontologia, università di modena e reggio emilia; smns: staatliches museum für naturkunde stuttgart. systematic palaeontology protognathodus synonymy lists herein only include figured specimens that could be clearly assessed. for suprageneric classification, the scheme proposed by sweet (1988) is followed. the four species considered are discussed in stratigraphic and evolutionary order: pr. meischneri, pr. collinsoni, pr. kockeli and pr. kuehni. taxonomy is focused only on p1 elements, as the multielement apparatuses of the different species are unknown. translation of the original diagnosis in german was provided by sandra kaiser; it is essentially in agreement with the previous translation by ziegler (1973). class conodonti branson, 1938 order ozarkodinida dzik, 1976 family idiognathodontidae harris & hollingsworth, 1933 genus protognathodus ziegler, 1969 type species: gnathodus kockeli bischoff, 1957 remarks. the apparatus of protognathodus was tentatively reconstructed as seximembrate by chauffe & nichols (1995). however, no distinction between members of the apparatuses of the various species was provided and the authors described p2, m, s0, s1 and s2 elements as “vicarious elements of protognathodus collinsoni, pr. kockeli and pr. meischneri.” protognathodus meischneri ziegler, 1969 pl. 1, figs 1-3 1969 protognathodus meischneri n.sp. ziegler, p.353, pl. 1, figs 1-13. 1973 protognathodus meischneri ziegler – ziegler (in ziegler), p. 421-422, schmidtognathus pl. 2, fig. 3. 1973 protognathodus meischneri ziegler – szulczewski, p. 43, pl. 2, fig. 8. 1976 protognathodus meischneri ziegler – dreesen et al., pl. 2, fig. 10. 1982 protognathodus meischneri ziegler – higgins & wagnergentis, pl. 34, fig. 10. 1984 protognathodus meischneri ziegler – hou et al., pl. 4, fig. 9. 1984 protognathodus meischneri ziegler – wang & yin, pl. 3, fig. 17. 1985 protognathodus meischneri ziegler – austin et al., pl. 4.9, fig. 14. 1987b protognathodus meischneri ziegler – ji, pl. 1, figs 1-2. 1988 protognathodus meischneri ziegler – garcia alcalde & menendez-alvarez, pl. 1, fig. 8. 1988 protognathodus meischneri ziegler – schönlaub et al., pl. 4, figs 4. 1988 protognathodus meischneri ziegler – wang & yin, pl. 22, figs 1-2 (only). 1990 protognathodus meischneri ziegler – gagiev & kononova, pl. 3, figs 27-28. 1992 protognathodus meischneri ziegler – over, fig.7.14. 1993 protognathodus meischneri ziegler – nemirovskaya et al., pl. 2, fig. 13. 1995 protognathodus meischneri ziegler – chauffe & nichols, pl. 2, figs 27-28, 33, 36. non 1997 protognathodus meischneri ziegler – caridroit et al., pl. 1, fig. 4. 1997 protognathodus kockeli (bischoff) – dzik, fig. 8 c. 1999 protognathodus meischneri ziegler – sanz lopez et al., pl. 1, fig. 15. 2003 protognathodus meischneri ziegler – corradini et al., p. 235, pl. 4, fig. 4. non 2004 prothognathodus meischneri ziegler – bardasheva et al., pl. 13, fig. 44. original diagnosis (ziegler 1969 german): a species of the genus protognathodus with a generally symmetrical platform, whose surface has no ornamentation. remarks. pr. meischneri differs from its ancestor bispathodus stabilis for the more posterior position and greater expansion of the basal cavity; it is distinguished from all other species of protognathodus by the absence of ornamentation on the upper surface of the cup. stratigraphic range. according to ziegler & sandberg (1984) pr. meischneri first occurs at or near the base of the lower praesulcata zone; however, a single specimen (pl. 1, fig. 2) have been found from the upper expansa zone in sardinia (corradini et al. 2003: tab. 2). the last occurrence is within the upper duplicata zone (sandberg et al. 1978; kaiser et al. 2009). protognathodus collinsoni ziegler, 1969 pl. 1, figs 4-8 1969 protognathodus collinsoni n. sp. ziegler, p. 353, pl. 1, figs 14-18. 18 corradini c., kaiser s.i., perri m.c. & spalletta c. 1973 protognathodus collinsoni ziegler – ziegler (in ziegler), p. 415-416, schmidtognathus pl. 2, fig. 4. 1973 protognathodus collinsoni ziegler – szulczewski, p. 42, pl. 2, figs 9-10. 1974 gnathodus kockeli bischoff – gedik, pl. 7, fig. 6. 1980 protognathodus collinsoni ziegler – ebner, pl. 16, figs 3, 5. 1983 protognathodus collinsoni ziegler – wang & ziegler, pl. 8, fig. 16. 1984 protognathodus collinsoni ziegler – hou et al., pl. 4, fig. 12. 1984 protognathodus meischneri ziegler – wang & yin, pl. 3, fig. 16. 1984 protognathodus kockeli (bischoff) – luppold et al., pl. 4, fig. 1. 1985 protognathodus collinsoni ziegler –austin et al., pl. 4.8, fig. 20. 1987 protognathodus collinsoni ziegler – kalvoda & kukal, pl. 4, fig. 4 (only). 1988 protognathodus collinsoni ziegler – wang & yin, pl. 22, figs 5-6 (only). 1988 protognathodus kockeli (bischoff) – wang & yin, pl. 22, figs 9, 13, 17. 1988 protognathodus collinsoni ziegler – flajs & feist, pl. 9, fig. 6 (only). 1989 protognathodus collinsoni ziegler – ji et al., p. 90-91, pl. 18, fig. 8-9 (only). 1990 protognathodus kockeli (bischoff) – gagiev & kononova, pl. 3, figs 29-30. 1990 protognathodus collinsoni ziegler – gagiev & kononova, pl. 4, fig. 4. 1992 protognathodus collinsoni ziegler – over, fig.7.15. 1992 protognathodus collinsoni ziegler – ji & ziegler, pl. 3, fig. 22. 1993 protognathodus meischneri ziegler – nemirovskaya et al., pl. 2, fig. 20. 1995 protognathodus cf. collinsoni ziegler – chauffe & nichols, pl. 2, figs 30, 31-32, 38. 1995 protognathodus kockeli (bischoff) – chauffe & nichols, pl. 2, figs 35, 37. 1998 protognathodus collinsoni ziegler – kalvoda & kukal, pl. 4, fig. 4. 2001 protognathodus collinsoni ziegler – perri & spalletta, pl. 1, fig.4 2003 protognathodus collinsoni ziegler – corradini et al., p. 235, pl. 4, fig. 3. 2009 protognathodus collinsoni ziegler – kaiser, pl. 1, figs 1, 3. original diagnosis (ziegler 1969 german): a species of the genus protognathodus with a weakly asymmetrical platform, which has single nodes on the surface. the nodes can be both on the right side and left side (outside or inside) of the carina. remarks. pr. collinsoni is characterized by one or more nodes scattered on the cup, that can occur on one or both sides of the carina. a few specimens with the cup covered by nodes, identified as pr. kockeli, should be referred to pr. collinsoni, as the nodes are not arranged in a distinct row diagnostic of pr. kockeli. stratigraphic range. from the lower part of the lower praesulcata zone (ziegler & sandberg 1984) to at least into the upper duplicata zone (over 1992). protognathodus kockeli (bischoff, 1957) pl. 1, figs 9-19 1957 gnathodus kockeli n.sp. bischoff, p. 25, pl. 3, figs 27-32. 1959 gnathodus kockeli bischoff voges, pl. 33, p. 281-282, fig. 27 (only). 1967 gnathodus kockeli bischoff – van adrichem boogaert, p. 179, pl. 2, figs 17-18. non 1968 gnathodus kockeli bischoff – manzoni, p. 659-660, pl. 62, fig.2 1969 gnathodus kockeli bischoff – ziegler, p.354, pl. 1, figs.19-25; pl. 2, figs 1-5. 1969 gnathodus kockeli bischoff – schönlaub, p.330, pl. 1, figs1-2. 1970 gnathodus kockeli bischoff – ziegler & leuteritz (in koch et al.), pl. 8, figs 1-3, 5. 1970 gnathodus kuehni n. sp. – ziegler & leuteritz (in koch et al.), pl. 8, fig. 4. 1973 protognathodus kockeli (bischoff) – ziegler (in ziegler), p. 417-418, schmidtognathus pl. 2, fig. 5. 1973 protognathodus kockeli (bischoff) – szulczewski, p. 44, pl. 2, figs 11-13. 1974 gnathodus kockeli bischoff – gedik, p. 13, pl. 7, fig.5. 1980 protognathodus kockeli (bischoff) – ebner, p. 16, figs 4, 6. 1980 protognathodus kockeli (bischoff) – lane et al., pl. 3, fig. 1. 1984 protognathodus kockeli (bischoff) – hou et al., pl. 4, figs 10. 1984 protognathodus kockeli (bischoff) – luppold et al., pl. 4, fig. 2; pl.6, fig. 4 (only). 1984 protognathodus kockeli (bischoff) – wang & yin, pl. 3, fig. 15 (only). 1985 protognathodus kockeli (bischoff) – austin et al., pl. 4.9, figs 13. 1987 protognathodus kockeli (bischoff) – kalvoda & kukal, pl. 4, figs 2-3 (only). 1988 protognathodus kockeli (bischoff) – garcia-alcalde & menendez-alvarez, pl. 1, fig. 7. 1988 protognathodus kockeli (bischoff) – flajs & feist, pl. 9, figs 8-10. 1988 protognathodus collinsoni ziegler – flajs & feist, pl. 9, fig. 7. 1988 protognathodus kockeli (bischoff) – schönlaub et al., pl. 4, figs 1-2, 5. 1988 protognathodus praedelicatus lane et al. – schönlaub et al., pl. 4, fig. 10. 1988 protognathodus kockeli (bischoff) – weyant., pl. 2, fig. 10. 1988 protognathodus kockeli (bischoff) – wang & yin, pl. 22, figs 8, 10-11, 14-15 (only). 1988 protognathodus kuehni ziegler & leuteritz – wang & yin, pl. 22, fig. 19. 1989 protognathodus kockeli (bischoff) – ji et al., p. 91, pl. 18, figs 3-5 (only). 1989 protognathodus collinsoni ziegler – ji et al., pl. 18, fig. 7. 1989a protognathodus kockeli (bischoff) – clausen et al., pl. 5, figs 3, 5. 1990 protognathodus kockeli (bischoff) – gagiev & kononova, pl. 4, figs 5-6. 1993 protognathodus kockeli (bischoff) – nemirovskaya et al., pl. 2, figs 15-16 (only). 1993 protognathodus kuehni ziegler & leuteritz – nemirovskaya et al., pl. 2, fig. 19. 1992 protognathodus kockeli (bischoff) – over, fig.7.16. 1994 protognathodus kockeli (bischoff) – korn et al., pl. 5, figs. 4-11; pl. 7, figs 8, 10 (only). protognathodus (conodonta) and its potential as a tool for defining the devonian/carboniferous boundary 19 ? 1995 protognathodus kockeli (bischoff) – chauffe & nichols, pl. 2, figs 29, 34. 1997 protognathodus kockeli (bischoff) – chauffe & guzman, pl. 2, figs 7, 15. 1997 protognathodus kockeli (bischoff) – dzik, fig. 8 a-b. 1999 protognathodus cf. kockeli (bischoff) – sanz lopez et al., pl. 2, fig. 1. 2001 protognathodus kockeli (bischoff) – perri & spalletta, pl. 1, fig. 5. non 2000 protognathodus kockeli (bischoff) – wang et al., pl. 1, fig. 11. 2002 protognathodus kockeli (bischoff) – buggisch & michl, pl. 4, fig. 125 (only). 2003 protognathodus kockeli (bischoff) – corradini et al., p. 235, pl. 4, figs 1-2. non 2006 protognathodus kockeli (bischoff) – dzik, figs 116 m-n. 2008 protognathodus kockeli (bischoff) – habibi et al., p. 772, figs 4.6. 2007 protognathodus kockeli-prothognathodus kuehni – kaiser, pl. 2, figs. 3, 5-6. 2009 protognathodus kockeli (bischoff) – kaiser et al., pl. 1, figs 10-11; pl. 2, fig. 16. 2009 protognathodus kuehni ziegler & leuteritz – kaiser et al., pl. 1, figs 8-9. 2009 protognathodus kockeli (bischoff) – kaiser, pl. 1, fig. 2. original diagnosis (bischoff 1957 german): a species of the genus gnathodus [protognathodus] with the following features: approximately a hemispherical platform with one or two lines of coarse nodes on the inner and outer side of the platform. the lines run parallel to the blade. emended diagnosis: a species of protognathodus with a row of coarse nodes parallel to the carina on one or both sides of the cup. remarks. the coarse, nodose, ornamentation and the presence of at least one row of nodes parallel to the carina on one half of the cup characterizes pr. kockeli; scattered unarranged nodes may occur on the other side of the cup. some large specimens bear one row of nodes on both sides of the cup, sometimes together with other scattered nodes. some specimens that bear unarranged coarse nodes on the cup have been referred to this species; however, these should be referred to pr. collinsoni. transitional forms between pr. kockeli and pr. kuehni are well known (i.e.: bischoff 1957: pl. 3, fig. 31; voges 1959: pl. 33, fig. 26; higgins et al. 1964: pl. 5, fig. 27; kaiser 2007: pl. 2, fig. 4); these specimens have a more rounded cup than typical pr. kockeli and bear nodes, sometimes transversally elongated. protognathodus kockeli is by far the most abundant and most widely documented species of protognathodus. it has a wide geographic distribution, although in some regions only co-occurs with lower carboniferous faunas (see discussion in the next chapter). stratigraphic range. the species first appears in the latest famennian and is the marker for the base of the upper praesulcata zone (ziegler & sandberg 1984). protognathodus kockeli ranges at least up to the sandbergi zone in europe (kaiser et al. 2009) and into the lower crenulata zone in north america (sandberg et al. 1978). protognathodus kuehni ziegler & leuteritz, 1970 pl. 1, figs 20-22 1962 gnathodus n. sp. b collinson et al., text-fig. 3. 1969 protognathodus n. sp. a ziegler p.355, pl. 1, fig. 26. 1970 protognathodus kuehni n.sp. ziegler & leuteritz (in koch et al.), p.715, pl. 8, figs 6-16. non 1970 protognathodus kuehni n.sp. ziegler & leuteritz (in koch et al.), p.715, pl. 8, fig. 4. 1973 protognathodus kuehni ziegler & leuteritz – ziegler (in ziegler), p. 419-420, schmidtognathus pl. 2, fig. 6. 1984 protognathodus kuehni ziegler & leuteritz – wang & yin, pl. 3, fig. 13. 1984 protognathodus kockeli → kuehni – luppold et al., pl. 4, fig. 3 1987 protognathodus kuehni ziegler & leuteritz – feist & flajs, pl. 2, fig. 1. 1988 protognathodus kuehni ziegler & leuteritz – flajs & feist, pl. 9, figs 11-12. 1988 protognathodus kuehni ziegler & leuteritz – schönlaub et al., pl. 4, figs 3, 7. 1988 protognathodus kockeli-prothognathodus kuehni – schönlaub et al., pl. 4, fig. 6. 1988 protognathodus praedelicatus lane et al. – schönlaub et al., pl. 4, figs 8-9. 1990 protognathodus kuehni ziegler & leuteritz – gagiev & kononova, pl. 4, fig. 7. 1994 protognathodus kockeli (bischoff) – korn et al., pl. 7, fig. 9. non 2000 protognathodus kuehni ziegler & leuteritz – wang et al., pl. 1, fig. 10. 2007 protognathodus kuehni ziegler & leuteritz – kaiser, pl. 2, figs 2, 7. 2009 protognathodus kuehni ziegler & leuteritz – kaiser et al., pl. 2, fig. 15 (only). original diagnosis (ziegler & leuteritz in koch et al. 1970 german): a new species of the genus protognathodus ziegler [...] with a weak asymmetrical platform, whose surface has robust transverse ridges. these ridges run unordered from the edge radial to the carina. the carina can be, as some specimens reveal, suppressed. remarks. the occurrence of transverse ridges on the cup characterizes pr. kuehni and distinguish it from other representatives of the genus. transitional forms from pr. kockeli are known: they bear both transverse ridges and nodes and have a more asymmetrical cup than pr. kuehni. in lateral view the carina does not rise above the ornamentation and in some specimens can be suppressed in the posterior part of the cup. protognathodus kuehni has a wide geographic distribution, documented in europe, south china, siberia and the central usa; however, outside europe, it is very rare (see discussion in the next chapter). 20 corradini c., kaiser s.i., perri m.c. & spalletta c. stratigraphic range. from the base of the sulcata zone (kaiser et al. 2009) or just below (ziegler 1973) to within the sandbergi zone (lane et al. 1980). stratigraphic and geographic distribution representatives of genus protognathodus are relatively widely distributed in uppermost devonianlowermost carboniferous rocks around the world (figs 2-3; tab. 1). however, their abundance is often associated with peculiar environmental conditions. in deeper-water sediments the occurrence is very irregular and often the different species first occur together, or not in stratigraphic order. in several localities they have been reported only from a higher part of their stratigraphic range (see below for examples and discussion). in north america “the major habitat of the protognathodus fauna was in nearshore or lagoonal settings where non-argillaceous microbial or algal micrite was deposited” (c. sandberg, pers. comm. march 30, 2010), and thus protognathodus was regarded as a shallow-water genus. in the woodford shale in oklahoma, occurs in association with offshore taxa. in southern europe protognathodus is thought to reflect depositional settings at the continental rise and lower slope due to the microfacies characteristics (kaiser 2005) . more data from different palaeogeographic settings, however, are needed for a more complete evaluation. the standard biofacies model favored by sandberg (1976) and ziegler & sandberg (1984), especially the assessment of the protognathodid biofacies as an indicator of a shallowing, has to be reconsidered (see discussions in kaiser et al. 2008). more likely, the occurrence of the protognathodus fauna can be related to biotic opportunism during a rise in sea level in the latest devonian, as evidenced in many sections in the rheinisches schiefergebirge. in the base of the carboniferous gssp at la serre trench e’, representatives of the genus protognathodus are rare in boundary beds and their presence is not regular throughout the section. protognathodus kuehni is reported only in the topmost part of the section (flajs & feist 1988; kaiser 2005, 2009) where the protognathodus fauna is abundant. in the same region, but in a different tectonic unit, in the puech de la suque section (kaiser et al. 2009), pr. kockeli enters just above the hangenberg shale equivalent, slightly before the joint first occurrence of pr. meischneri and pr. collinsoni. a single specimen of pr. kuehni has been found higher in the section, together with si. quadruplicata (kaiser et al. 2009: tab. 6). similar late first occurrences of pr. kuehni, and often also of pr. kockeli, are well documented in various geographic areas: poland (szulczewski 1973; dzik 1997), cantabrian mountains (higgins & wagnergentis 1982; garcia-alcalde & menendez-alvarez 1988), sardinia (corradini et al. 2003; corradini 2008) and algeria (weyant 1988); in the western united states no protognathodids have been found in the devonian (c. sandberg, pers. comm.). plate 1 figs 1-3 protognathodus meischneri ziegler, 1969 1) ic p139 966502; sentiero per cresta verde section, carnic alps, sample scv 4; 2) dstc 30116; monte taccu north b section, sardinia, sample mt z; 3) ipum 27648; monte taccu north b section, sardinia sample mt 2a. figs. 4-8 protognathodus collinsoni ziegler, 1969 4) ic p149 966564; plan di zermula c section, carnic alps, sample pzc 1; 5) ic p149 966569; plan di zermula c section, carnic alps, sample pzc 1; 6) ic p149 966565; chiarsò area, carnic alps, sample chrb; 7) ipum 27647; monte taccu north a section, sardinia sample mt x; 8) smns 67400; la serre trench e’ section, montagne noire, sample 70. figs 9-21 protognathodus kockeli (bischoff, 1957) 9) ic p150 966570; plan di zermula c section, carnic alps, sample pzc 4; 10) ic p190 212112; plan di zermula a section, carnic alps, sample pza 2ai; 11) ic p149 966567; plan di zermula c section, carnic alps, sample pzc 4; 12) ic p174 967133; passo di monte croce carnico section, carnic alps, sample pmc 1; 13) ic p139 966503; sentiero per cresta verde section, carnic alps, sample scv 4; 14) ipum 27646; monte taccu north b section, sardinia sample mt 1a; 15) ipum 27645; monte taccu north a section, sardinia sample mt x; 16) ic p149 966568; chiarsò area, carnic alps, sample chrb; 17) smns 67375; trolp section, graz palaeozoic, sample 16; 18) ic p149 966566; plan di zermula c section, carnic alps, sample pzc 1; 19) dstc 30117; monte taccu north a section, sardinia sample mt x; 20) smns 67374; trolp section, graz palaeozoic, sample 16; 21) smns 67401; puech de la suque section, montagne noire, sample ps 16. figs 22-23 protognathodus kuehni ziegler & leuteritz, 1970 22) smns 67400a; milles section, french pyrenees, sample mi 9 top. 23) smns 67710; grüne schneid section, carnic alps, sample gs 6c2. protognathodus (conodonta) and its potential as a tool for defining the devonian/carboniferous boundary 21 22 corradini c., kaiser s.i., perri m.c. & spalletta c. in the french pyrenees, at milles (cygan & perret 1998; kaiser et al. 2009) abundant protognathodids (mainly pr. kockeli) enter at the base of bed 9, just above a siliciclastic bed interpreted as an equivalent of the hangenberg black shale (kaiser et al. 2009). here two specimens of pr. kuehni occur only in the upper part of this bed (kaiser et al. 2009: tab. 7), representing less than 1% of the whole association. in other sections in the same area (e.g., pont de saubette and moustarde) pr. meischneri, pr. collinsoni and pr. kockeli have their first occurrence all together, and pr. kuehni is not reported (perret 1988); finally, in the garcet section (perret mirouse & majesté-menjoulas 1998), where the dcb is located in a continuous calcareous section with a good recording of siphonodella, no protognathodus have been found across the boundary. in the carnic alps, the grüne schneid section exposes a continuous condensed calcareous sequence across the boundary (schönlaub et al. 1988, 1992; kaiser et al. 2006). protognathodus meischneri and pr. collinsoni enter in association and are followed, in sequence, by pr. kockeli and pr. kuehni (kaiser 2007). the latter species is documented only by two certain and five questionable specimens from four samples collected in a 15 cm thick interval and represents about 1% of the association (kaiser 2005: 118), while pr. kockeli, pr. meischneri and pr. collinsoni are quite abundant in some beds. in the kronhofgraben (schönlaub et al. 1992; kaiser 2005) and plan di zermula a sections (perri & spalletta 2001; kaiser 2005; kaiser et al. 2009), where the equivalents of the hangenberg black shale, corresponding to the middle praesulcata zone, are present, no protognathodids were recorded below the shales and pr. kuehni is very rare in only one level, a few centimetres below the first occurrence of si. duplicata. in both sections pr. kockeli enters just above the equivalents of the hangenberg black shale, together with si. sulcata. in the rheinisches schiefergebirge, the type area of the protognathodus fauna of ziegler (1969), several sections spanning the dcb have been studied and documented (i.e. clausen et al. 1987, 1989b; korn et al. 1984; kaiser 2005 and references therein). protognathodids are always present, but in many sections they are very rare and their occurrence is limited to a few beds. the only sections in which protognathodids are abundant (e.g., seilerschurf iii and schurf 0, clausen et al. 1989b) are characterized by much more shaley sedimentation. this is even more evident if we consider the occurrences and abundance of pr. kuehni, which in most sections is represented only by one or two specimens from samples slightly below the entry of si. duplicata, whereas it is abundant only in the seiler-schurf iii section (clausen et al. 1989b). in the nanbiancun ii section, guangxi, south china (ji et al. 1989; wang 1995; wang & yin 1988) pr. kockeli first occurs in bed 52 (sulcata zone after ji fig. 2 geographic distribution of protognathodus species. pa laeogeographic map redrawn after scotese (2001). protognathodus (conodonta) and its potential as a tool for defining the devonian/carboniferous boundary 23 et al. 1989; upper praesulcata zone after wang 1995), whereas pr. meischneri only enters in bed 54 (lower duplicata zone after ji et al. 1989; upper praesulcata zone after wang 1995). protognathodus kuehni has a single occurrence in bed 56 (lower duplicata zone after ji et al. 1989; basal sulcata zone after wang 1995), but this finding should be confirmed, since the only specimen figured as pr. kuehni actually belongs to pr. kockeli. in levels across the dcb in the nanbiancun ii section, protognathodids are uncommon, representing only 3-7% of the fauna (wang & yin 1988: 109), and from the seven sections spanning the boundary in the nanbiancun area, pr. kuehni is reported only from nanbiancun ii. in the muhua section, guizhou, south china, all four species of protognathodus have been found together in a “grey dense limestone” lens (wang & yin 1984: 233), immediately below the first occurrence of si. sulcata. according to ji et al. (1989), the species occur in different levels: pr. meischneri first occurs in level 21-1 (middle praesulcata zone), pr. collinsoni in level 21-2 (upper praesulcata zone), pr. kockeli in level 22-1 (upper praesulcata zone), and pr. kuehni in level bed 22-2 (upper praesulcata zone). in dapoushang section, guizhou, south china (ji et al. 1989) protognathodids occur in limestone beds from the lower praesulcata zone to the lower duplicata zone, but it is difficult to consider these data fig. 3 geographic distribution of pr. kockeli. a) carboniferous occurrences; b) devonian occurrences. 24 corradini c., kaiser s.i., perri m.c. & spalletta c. because of inconsistence between text and figures (i.e. page 31-34, 90-91; text figs 6-7; pl. 18). the protognathodids are relatively abundant in the limestone beds just above the equivalent of the hangenberg black shale (bed e), but the abundance drastically decreases at the base of the sulcata zone (ji et al. 1989: fig. 7) no protognathodids have so far been found in australia (j.a. talent, pers. comm.). conclusion 1. the original diagnoses (in german) of the four early species of protognathodus are clear. a translation in english was provided very soon after they were established (ziegler 1973). the great majority of the specimens of protognathodus can be easily assigned and only a few transitional specimens are known, tab. 1 localities and relevant literature for reports of late devonian/early carboniferous protognathodids. data are grouped in two lists concerning figured and not figured elements. f: famennian, t: tournaisian; x: finding in neptunian dykes. p r. m ei sc hn er i p r. c ol lin so ni p r. k oc ke li p r. k ue hn i p r. m ei sc hn er i p r. c ol lin so ni p r. k oc ke li p r. k ue hn i bechar – algeria (famennian–tournaisian) (weyant 1988) t cantabrian mt – spain (famennian–tournaisian) (van adrichem boogaert 1967) t cantabrian mt – spain (famennian–tournaisian) (garcia alcalde & menendez alvarez 1988) t t cantabrian mt – spain (tournaisian) (higgins & wagner gentis 1982) t t t cantabrian mt – spain (famennian–tournaisian) (sanz lopez et al. 1999) f f t f czech republic (famennian–tournaisian) (kalvoda & kukal 1987) t t f f f f–t graz – austria (famennian–tournaisian) (ebner 1980) f f f–t t t iran alborz mts. (famennian–tournaisian) (habibi et al. 2008) t iran -tabas (famennian–tournaisian) (bahrami et al. in prep. pers. com.) t t montagne noire france (famennian–tournaisian) (feist & flajs 1987) t f–t f–t t montagne noire france (famennian–tournaisian) (flajs & feist 1988) t t t f f f montagne noire france (famennian–tournaisian) (kaiser et al. 2009) f–t t t f f pyrenees france (famennian–tournaisian) (perret 1988) f f–t f–t sardinia (famennian–tournaisian) (corradini et al. 2003) f–t t t t southern alps (famennian–tournaisian) (gedik 1974) t southern alps (famennian–tournaisian) (kaiser 2007) f–t t f–t t southern alps (famennian–tournaisian) (perri & spalletta 2001) t t t southern alps (famennian–tournaisian) (schönlaub 1969) t ? t southern alps (famennian–tournaisian) (schönlaub et al. 1988) f f–t f ardennes – belgium (famennian) (dreesen et al. 1976) f f f germany (tournaisian) (buggisch & michl 2002) t t t germany (famennian–tournaisian) (clausen et al. 1989a) f t t germany (famennian–tournaisian) (clausen et al. 1989b) f–t f–t t germany (famennian–tournaisian) (kaiser et al. 2009) t t f-t f-t f germany (famennian–tournaisian) (korn et al. 1994) f–t germany (famennian–tournaisian) (luppold et al. 1984) t t t t germany (tournaisian–visean) (voges 1959) t germany (famennian) (ziegler 1969) f f f f germany (famennian–tournaisian) (ziegler & leuteritz in koch et al. 1970) x t f–t f–t f–t great britain (famennian) (austin et al. 1985) f f f holy cross mt. – polonia (famennian–tournaisian) (szulczewski 1973) x x x polonia (tournaisian) (dzik 1997) t t polar ural russia (famennian–tournaisian) (nemiroskaya et al. 1993) t t t f f usa – illinois (tournaisian) (collinson et al. 1962) t usa – midcontinent (famennian) (chauffe & nichols 1995) f f f usa – missouri and illinois (tournaisian) (chauffe & guzman 1997) t usa – oklahoma (famennian–tournaisian) (over 1992) t t f f f t usa – wyoming (tournaisian) (lane et al 1980) t omolon block far east russia ne siberia (famennian–tournaisian) (gagiev & kononova 1990) f f–t t t t f tajikistan tajikistan (tournaisian) (bardasheva et al. 2004) t t guangxi china (famennian) (wang & ziegler 1983) f f guilin – china (famennian–tournaisian) (wang & yin 1988) f f f–t guizhou china (famennian–tournaisian) (hou et al. 1984) t t t f f guizhou china (famennian–tournaisian) (ji et al. 1989) f f–t f f f guizhou china (famennian–tournaisian) (wang & yin 1984) t f f f jianghua – china (tournaisian) (ji 1987b) t north gondwana south china laurussia figured not figured protognathodus (conodonta) and its potential as a tool for defining the devonian/carboniferous boundary 25 mainly in the upper part of the lineage (pr. kockeli-pr. kuehni). the diagnosis of pr. kockeli is slightly amended to include specimens, actually rare in collections, with one single row of nodes on only one side of the cup. some authors seem to have applied a very personal taxonomic approach in the attribution of specimens to the four species, often made on the basis of criteria clearly in contrast with the diagnoses (cf.: wang & yin 1988; chauffe & nichols 1995). 2. in north america protognathodus is often abundant in shallow water environments. in southern europe the occurrence of protognathodus is thought to reflect depositional settings at the continental rise and lower slope related to biotic opportunisms. more data from different palaeogeographic settings, however, are needed for a more complete evaluation; the standard biofacies model needs to be assessed. 3. protognathodids are rare in the great majority of devonian/carboniferous boundary sections where they comprise a minor component of the conodont association; while pr. kuehni is very rare (often <1%), pr. kockeli is relatively abundant. only in a few sections, mainly in the type area, protognathodus species have a local range corresponding to their known global stratigraphic distribution, whereas often they have a very restricted stratigraphic distribution. in some sections, the different species of the genus have a coincident first occurrence, or evolutionary younger forms enter below their evolutionary ancestors. 4. outside central-southern europe protognathodus is quite rare, and in some geographic areas such as in australia, completely absent, or enters only in the carboniferous, often together with si. duplicata (i.e.: western usa). 5. pr. kockeli is the most abundant and widely documented species of protognathodus, used as the marker of the upper praesulcata zone (= kockeli zone after kaiser et al. 2009, the last devonian biozone). it has a wide geographic distribution, but in many regions it occurs only in the carboniferous (fig. 3). 6. protognathodus kuehni is, in general, an extremely rare taxon, with a restricted range in many sections, and, outside the type area, it often occurs only in higher stratigraphic levels. as pointed out by alberti et al. (1974: 272) recoveries are irregular, even in the type area: “whether the pr. kuehni is found depends on sample quantity and some luck: sample 1008-330 yielded one specimen of each of the four species of protognathodus, while in sample 1008-294 [same bed] among 170 protognathodus specimens no pr. kuehni was found.” in conclusion, protognathodus can be of help in stratigraphic works across the devonian/carboniferous boundary, but its potential as a tool for defining the dcb is low, and a decision should consider the rarity, at least of pr. kuehni, pr. collinsoni and pr. meischneri, and the restricted stratigraphic ranges of the taxa. acknowledgements. jim barrick, gil klapper, jeff over and charlie sandberg kindly provided information on the occurrence and distribution of protognathodus in north america. thomas becker informed us on his finding in morocco and ali bahrami on data from iran. we are deeply grateful to hanna matyja and jeff over for critical revision of the manuscript. r e f e r e n c e s adrichem boogaert h.a. van (1967) devonian and lower carboniferous conodonts of the cantabrian mountains (spain) and their stratigraphic application. leidse geol. mededel., 39: 129-192. alberti h., groos-uffenorde h., streel m., uffenorde h. & walliser o.h. 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(1959) conodonten aus dem unterkarbon i und ii (gattendorfia und pericyclus-stufe) des sauerlandes. paläontol. z., 33: 266-314. wang c.y. (1995) on the devonian-carboniferous boundary in neritic facies area of south china: a viewpoint of integrated stratigraphy. j. china univ. geosci., 6(2): 184-191. wang c.y. & yin b. (1984) conodont zonations of early lower carboniferous and devonian-carboniferous boundary in pelagic facies, south china. acta palaeontol. sinica, 23(2): 224-238. wang c.y. & yin b.a. (1988) conodonts. in: yu c.m. (ed.) devonian-carboniferous boundary in nanbiancun guilin: 105-148, science press, beijing. wang c.-y., zhu m.-k., yan y.-j., wu y.-l., zhao y.-g. & qian y.-z. (2000) lower devonian conodonts from the sawayardun goldmine area, wuqia (ulugqat) county, xinjiang. acta micropalaeontol. sinica, 17(3): 255-264. wang c.-y. & ziegler w. (1983) devonian conodont biostratigraphy of guangxi, south china, and the correlation with europe. geol. palaeontol., 17: 75-107. weyant m. (1988) relationship between devonian and carboniferous strata near the northern confines of the bechar basin, algeria. in: flajs g, feist r. & ziegler w. (eds) devonian carboniferous boundary results of recent studies. courier forschungs-institut senckenberg, 100: 235-245. ziegler w. (1969) eine neue conodontenfauna aus dem höchsten oberdevon. fortschr. geol. rheinl. westfal., 17: 343-360. ziegler w. (ed.). (1973) catalogue of conodonts. 1, 504 pp., stuttgart (schweizerbart’sche verlagsbuchhandlung). ziegler w. & sandberg c.a. (1984) palmatolepis-based revision of upper part of standard late devonian conodont zonation. geol. soc. america, spec. pap., 196: 179-194. ziegler w. & sandberg c.a. (1990) the late devonian standard conodont zonation. courier forschungsinstitut senckenberg, 121: 1-115. ziegler w. & sandberg c.a. (1996) reflexions on frasnian and famennian stage boundary decisions as a guide to future deliberations. newsl. stratigr., 33: 157-180. rivista italiana di paleontologia e stratigrafia voìume 108 short palaeoecological notes tortonian ostracods pagine 289-296 luglio 2002 on the middle serravallian-basal from the tremiti islands barbara dall'antonia receiaed july 15,20a1; accepted juanuary 9,2aa2 key uords: ostracoda, palaeoecology, systematics, middle miocene, tremiti islands. riasswnto. viene discusso il significato paleoecologico dell'ostracofauna rinvenuta nelf intervallo serravalliano medio-tortoniano basale della successione delle isole tremiti. questa è indicativa di un ambiente batiale per tutto i'interuallo investigato, ma modificazioni nella composizione delle associazioni suggeriscono che le condizioni al fondo subiscono alcuni cambiamenti. in particolare, nel serravalliano medio gli ostracodi indicano i'esistenza di un ambiente a carattere termosferico profondo soggetto, probabilmente, a deboli influenze psicrosferiche. nel serravalliano terminale-tortoniano basale registrano l'inizio di una graduale transizione verso condizioni tipiche della termosfera superìore. la specie bradleya (?) saxolensis russo, 1966 viene discussa ed attribuita d, genere agrenocythere benson, 19l2. abstract. the palaecological significance of the ostracod faunas from the middle serravallian-basal tortonian of the temiti islands is briefly discussed. the fauna is typical of the bathyal envìronment throughout the investigated interual. compositional changes recorded in the assemblages, however, indicate that bottom-water conditions varied. sfithin the middle serravallian, ostracods point to the existence of a lower thermospheric environment with possible feeble psychrospheric influences. in the latest serravallian-basal tortonian, the start of a gradual shift towards upper thermospheric conditions is recognizable. the species bradleya (?) saxolensis russo, 1966 is herein redescribed and placed into the genus agrenocythere benson,1972. introduction deep-sea ostracods are a valuable tool for the diagnosis of the physical conditions of deep water-masses and have been often used to detect and investigate global and local palaeoceanographical changes (benson & sylvester-bradley 1971; benson 1.973, 1975, 1.976, 1978, l990; benson et ai. 1,984; vhatley & coles 1991; majoran 6c dingle 2001). some ostracod genera are p^rticularly useful for recognizing past thermospheric and psychrospheric conditions. among these is the genus agrenocytbere benson, 1972, which is thought (benson 1,972, 1976) to be virtually restricted to the upper psychrosphere (oceanic .waters beneath the permanent thermocline with temperature from 4'c to 8-10'c). oblitacytbereis benson, 1977 is likewise regarded (benson 1973, 1.976, 1.977) as typical of the lower thermosphere (waters with temperature from 1o'c to 1,6-2a'c overlying the psychrosphere in the open ocean or filling restricted basins). the aim of the present note is to discuss briefly the palaecological significance of the ostracod fauna from the middle serravallian-basal tortonian of the tremiti islands. in addition. some few, essential information on the langhian-early serravallian assemblages from the tremiti islands succession, which have been recently studied by the present author (dall'antonia 2ooo, ph.d. thesis 2001, in review), are incorporated. the sampling and bio-chronostratigraphical framework used for the ostracod study are essentially those of iaccarino et al. (2001), although herein, the foraminiferal zonal scheme has been modified according to the new proposals of sprovieri er al. (2002). this note supplements the more extensive palaeoecological analysis performed on the benthic foraminifera from a coeval composite section of the tremiti islands by russo et al. qa04, notwithstanding that the sampling and temporal framework utilized are quite different. material and methods the middle serravailian-basal tortonian [mmi 7 (p partimlabiata) zone-base of the mmi 10 (g. obliquws obliquws) zone] succession of the tremiti islands consists of whitish indurated hemipelagic marls intercalated with grey or reddish marls belonging to the cretaccio formation. the succession is well exposed in the san nicola island and is documented entirely in section 9 andpartialiy in sections 5,6,7 and 8 (fig. 1) of iaccarino et al. (2001). all together, 59 samples from the dìpartimento di scienze della terra, università di pisa, via s. maria 53,56726 pisa, italy, e-mail: barbarad@dst.unrpr.it 294 b. dall'antonìa above-mentioned sections have been used for the ostracod study. systematic analysis revealed the presence of assemblages of deep-water character in all the examined samples. only in a single sample (sample 172 of section 8) a few ;'uvenile specimens pertaining ro the shallow-water genus awrila pokorny, 1955 have been recovered. these specimens have been regarded as allochthonous and harre been consequently disregarded in the palaecological considerations. collectively a total of 69 taxawere identified, of these 11 are yet undescribed. synthetic range charts for ostracod species are reported in tab. 1. some species temporarily disappear from the record within the interval under discussion, but are already presenr below and/or reappear higher in the miocene (langhian-messinian) succession of the tremiti islands. local first occurrence (fo) and last occurrence (lo) records are, rherefore, indicated in tab. 1. since biozonal resolution is inadequate to enable a {fig. 1 geological map of the tremiti islands with location, lithology and biostratigraphical correlation of the middle serravallian-basal tortonian sections used for the ostracod study (modified after taccarino er al. 200 l;. )tab. 1 ostracod range charts for the middle serravallian-basal tortonian of the tremìti islands. detailed correlation between samples of the various sections to be made, no rigorous quanritarive analysis has been carried out on the ostracod faunas and only longterm compositional changes have been investigated. fig. 2 gives the relative abundance (percent value) of the most significant (major component or palaeoecologically noteworthy) taxa recorded in the studied interval. palaeoecological significance of the ostracod fauna the ostracod faunal composition is typical of bathyal environment throughout the investigated interval. both qualitative and quantitative changes in the assemblages, however, indicate that bottom-water conditions altered during the middle serravallian-basal tortonian interval. in the middle serravallian [mmi za (p. partimlabiata/g. drury) subzone-lower part of the mmi 7b (p s, nicola island bio-chf onostratigraphy planktonic foram n fe6 post-lrt oc"n" un s i l\,4iocene un ts e ere-tultoc"neunts ffi 5,6,7,8,9 locaton ofthe investigated sectons .r r--==-l l i -"' i t: l\larls of the cretaccio fm. l . _ -_.06+sa_ptes faulls // liostracods from the tremiti islands 291, mayeri) subzone] the major components of the fauna (mean percent value > 2%) are, in order of relative abundance, henryhouella asperrima reuss, cytherella owlgata ruggieri, bwntonia rnwlticostata ruggieri, cytherella postdenticulata oertii, cardobairdia glabra? van den bold, bairdia profwnda (aiello, barra & bonaduce), obliacytbereis (paleoblitaqtthereis) apula daii'antonia, kritbe iniqua abate, barra, aieilo ec bonaduce, cytlterella sp. 4, cosa ciampol ruggieri,,4 wstraloecia posterocurua barra & bonaduce, xestoleberis prognattl bonaduce er danielopol and cytherella russoi sissingh. the deep thermospheric genus oblitacythereis is represented mostly (99%) by the above-mentioned o. (p) apwla and to a lesser extent by o. rugiedl (russo) . such a faunai composition characterized by the common occurrence of typical deep water faxa, i.e. australoecia, cardobairdia, cytherella (smooth species), henrythowella, krithe, xestoleberis prognata and notably fenurs australoecía barra & bonaduce, 200 1 bairdia barra & bonaduce, bunlonia dedonenis ruooieri. 1954 g!99!9!rde eelra r]ln q9!'_991d , l! e6_8__ cytheteila vulg?!?. ruggeí 1.962 henryhowella aspeffima (reuss, 1850) .kilthe cmilessa (seguenza, 1880) . cvîhercila sp.4 rectobunton i a m ira nd a bol4_qg_er_9e]!pq 3 àrq!9!!.1919 xesfo/eòer! prognafa bonaduce & oanielc baidia sp. 2 cylheropteron (aversov.l pinarense van den bold aiello. barra & bonaduce. 1996 eucytherc pubera bonaduce, ciampo & l,4asoli, 1976 krithe iníaua abate. bara. aiello & bonaduce. 1993 occultocylhercis cullgr aiello, bara & bonaduce, 2000 1980 sensu paakrithe rotundata aiello,baralabate & borladuc€, 1993 ___4 tsjll999e 99 y!!!!99]li utt { :-18e 4 loxocurnig!.ll!!! quadlj9gm,s (ru.ssier,, 1962) s/vele, athersuch. 1978 argílloecia gonzalesi barg, aielìo & bonaduce, 1996 ct'thetopteron (c.) bîfidun colalongo & pasini, 1980 subsp. 1 retibythere (bathibyiherc ) vandenboldi (rsggie1, 19601 cadobaidia sp.2 cylhercpteron (a.\ lancei carbonnel, 1969 9r4lc.elqigvg!4 r9991!!i9q macrocyprissa sp. aft. m. arcuata (col€]ln€o & pasini, 1980) cailnovalva aquila (ruggieri, 1972) macrcmckenziea masoli & . t977 costa tricostata (reuss, 1850) subsp. 1 sagmatocylherc tenuis (ciampo, 1980) saida ionia ciampo, 1988 mac@cypris sp.2 ary,uoecè 4isjq4qyqlllsrqisjltg[ 1 97?_ _ __ cythercplercn la .) sp. 1 292 b. dall'antonia oblitacythereis ([or explicative remarks on rhe palaeoecology of these taxa see dall'antonia et al. 2oo1), suggests bottom water conditions characteristic of the lower thermosphere. the assemblage, however, includes rare psychrospheric forms. agrenocythere saxolensis (russo) is, in fact, randomly present with mostly juvenile specimens and very low values of relative abundance (1-2%).it is noteworthy that psychrospheric ostracods, comprising both,4. saxolensis and ayenocytbere hazelae (van den bold), are relatively common in the langhian and in the earliest serravallian [upper part of the mmi 5 (o. suturalis-p peripheroronda) zone) of the temiti islands (dall'antonia ph.d. thesis 2001, in review). their occurrence documents that before the time interval investigated herein, deep oceanic water-masses characterized the circulation pattern of the area. psychrospheric forms temporarily disappear from the record (dall'antonia ph.d. thesis 2001, in review) during the early serravallian [mmi 6 (d. altispira ahispira) zone) to re-appear sporadically (as previously discussed) in the middle serravallian [mmi 7a (p partimlabiata/g. drury,) subzone and lower part of the mmi 7b (p mayeri) subzone]. in the above outlined contexr, the occurrence of a. saxolensis within a typical de ep thermospheric fauna seems to be suggestive of a 1ate, very weak influx of psychrospheric water-masses in the area. within the late serravallian [upper part of the mmi zb (p mayeri) subzone-lower parr of the mmi s (n. atlantica praeatlantica) zone], major components of the assemblages do not show significant change and the fauna appears to be typical of the lon'er thermosphere as indicated by the common occurrence of oblitacythereis (p) apula and the absence of psychrospheric taxa. the latest serravallian-basal tortonian interval [upper part of the mmi 8 (lx[. atlantica praeatlantica) zone-base of the mmil a gd. obliquws obliqwus) zonef is characterizedby a gradual shift from deep thermospheric to higher temperature tolerating species. such a compositional change includes: a decreasing incidence of the gews oblitacythereis and notably the lo ol o. (p.) apula at the top of the serravallian ftop of the mmi 9 (p siakensis) zonef; a decrease in the relative abundance of australoecìa posterocurúd and the genus cardobairdia; an increase in the relative abundance of the genus co.rta neviani, 1928 represented by costa ciampoì and costa tricostata (reuss) subsp. 1 fthe genus costa is mainly typical of the shelf (ruggieri 1961; benson 1973),but most tolerant species are able to live in deeper environments (ruggieri 7992, p. 176-178), as are presumably those encountered in the present studyl; an increase in the relative abundance of acanthocytbereis hystrix (reuss) fthe optimum of this species lying between 100 and 200 meters in the modern mediterranean (bonaduce et al. 1976; ruiz &, gonzalezregalado 1996)l; the fo of sagmatocythere tenuis (ciampo) [recent congeneric species live most often in the innerouter shelf (bonaduce et ai. 1,976,1988)]. conclusions ostracods from the middle serravallian-basa1 tortonian of the tremiti islands are indicative of a bathyal environment. vithin the middle serravallian bottom water conditions are essentially those of the lower thermosphere, but the scattered occurrence of agrenocythere saxolensis (mostly as juveniles specimens) seems to be suggestive of the presence of feeble deep oceanic influences. from this time onward, in contrast to indications from the benthonic foraminifera (russo et ai. 2oo2), ostracods are unable to provide evidence of further psychrospheric infiuences in the area and the faunas of the late serravallian appear to be typical of a deep thermospheric environment. in addition, in the latest serravallian-basal tortonian the decreasing frequency of the €lenera oblitacythereis, australoecia and cardobairdia and the increasing incidence of taxa mainly related to shelf environments indicate the srart of a transition from lower thermospheric to upper thermospheric bottom water conditions. appendix taxonomic note on agrenocythere sdxolensis (russo, 1966) the examination of the type material of bradleya (?) saxolensis russo, 1966 and further specimens from the miocene succession of the tremiti islands indicates that this species should be assigned to the genus,4grenocythere benson and that its original description requires amendment. the terminology of the external carapace features employed herein is that proposed by benson (1972).the figured specimens are housed in the ostracoda collection of prof. a. bossio (c.o.b. 165-166), dipartimento di scienze della terra, università di pisa, ita1y. class ostracoda latreille, 1806 order podocopida mùlier, 1894 suborder podocopa sars, 1866 superfamily cytheracea baird, 1 850 family tachyleberididae sylvester-bradley, 1948 subfamily trachyleberidinae sylvester-bradley, 1948 genus agrenocytbere benson, 1972 type species ,4 grenoclthere spinosa benson, 1972 agrenocythere saxolensis (russo, 1966) (fig. 3a-d) 1966 bradleya (?) saxolensís russo, p.2.11, text-fig. 3. pl.14, fig. 3a-d. ? 1979 agrenocythere sp. l ducasse & pe1-pouquet, pl. 2, trg. 7. ostracod; from the tremiti islands 293 c .9 o ct) @ c @(f) \ .9 @(f) o be ls co ezzo -?2 l1 7/,u, 9.-"?o, -oolo* 9 ó>^-,í1v "?soo'% 9r\ *-ju ,"r"r.i_ -o, 294 b. dall'antonia fig. 3a-d sem microphotographs of agrenocythere saxolensis (russo, 1966). (x 80 unless otherwise stated). a, b, c) l! c.o.b 165, section 3iii of iaccarino et al. (2001) sample 37, early langhian: a, external view; b, dorsal view; c, detail oí castrum, x 180; d) rv external view, c.o.b 166, section 3iil of iaccarino et al. (2001) sample 32, early langhian. ? 1993 agrenocythere sp. riha, pl. 1, fig. 1-6. ' material. 22 valves of which many are juveniles. emended diagnosis. a species of agrenocytbere charactenzed by a thin, regular reticulate ornament with quadrangular meshes; castrum distinctive and moderately prominent; bwllar seríes reduced to bìunt rpinet. posterìor end markedly acuminare. emended description. left valve subtrapezoidal, elongate in lateral view. anterior margin broadly rounded with tuberculate perimarginal rib; posterior margin triangular with apex below mid height. both extremities are furnished with short marginal spines. dorsal margin straight, sloping downwards to posterior; ventral margin rectilinear to slightly convex. reticular patrern quire regular, with quadrangular fossae and short, obtuse con.junctive spines. in the posterior area the fossae are rhomboidal and the muri are aligned into opposed crossed system. castrwm moderately prominent and characterized by a regular ballium; parapectus confined to the three through sixballialfossae.therzt,ir consists of a u-shaped upper portion (bearing a prominent specula), which surrounds the peraial fossa and of a l-shaped lower part, which delimits the fossa arcis. the forwm is wide and depressed. the dorsal bullae are reduced to more or less prominent spines. eye-tubercle absent. ventro-lateral rib well developed and ponticulate. in dorsal view carapace s\à/ollen medianly; both extremities laterally compressed. internal features typrcal of the genus. size lmm). lv (c.o.b. 165) l :1.25, h:0.68, h:0.36 rv (c.o.b. 166) l: 1,.04, h = 0.55, h = 0.30 remarks. a. saxolensii strongly differs from a. hazelae (cythereis hazeli van den bold, 1946, p.92, pi. 10, fig. 4a-c) in the less massive and more regular reticular pattern and the structure of the castrum. these features make ,4. saxolensis quite similar to agrenocythere radula (cythere radula brady, 1880, p. 102, pi. 1,9, fig. 4a-b). the two species, however, differ in the lateral outline (,4. saxolensis having a more acuminate posterior end and a ostracods from tbe tremiti islands 295 less convex ventral margin), development and nature of the ventro-lateral rib (a. saxolensis having a clearly ponticulate and more prominent ventro-lateral rib) and, finally, the structure of the castrum. agrenocythere sp. 1 in ducasse & peypouquer (1979) and agrenocythere sp. in riha (1993) strongly resemble the present species in the ordinate and regular reticulate ornament, but quality of the illustrations and lack of description prevent their positive assignment to russo's species. previous records, ? late palaeocene-early pliocene of the north atlantic (ducasse tr peypouquet 1979) reported from leg 48 site 400-4. site 401 (northern continental margin of the bay of biscay) and site 403 (rockall area). in site 403 agrenocythere sp. 1 ducasse & peypouquet was found in association wrth bradleya, poseidonamicus and echinocytherels. according to ducasse & peypouquet (1979, p. 346) this association is indicative of a depth close to that of the site today (about 2700 m). ? early badenian of moravia {riha 1993 1 reported in a mixed association of shallox-water and deepwater ostracods, which is not characteristic of typical psychrospheric conditions (riha 1,993,p.158). the estimated depth of this association is of 200-1000 m (rihe refer benson, r.h. (1,972) -the bradleya problem, with description of two new psychrospheric ostracode genera, agrenocythere and poseidonamiczs (ostracoda: crustacea). smithsonian contr. paleobiol.,v. 12, pp. 1-138, vashington. benson, r.h. (1973) an ostracodal r'iew of the messinian salinity crisis. in: drooger, c. \il (ed.), messinian events in the mediterranean. wrh. le. neder. akad. .'weten s ch., pp. 23 5 -242 amsterdam. benson, r.h. (1975) the origin of the psychrosphere as recorded in changes of deep-sea ostracode assemblages. letbaia, v. 8, pp. 69-83, oslo. benson, r.h. (1976) miocene deep-sea ostracodes of the iberian portal and the balearic basin. marine micropaleontologt, v. 1, pp. 249-262, amsterdam. benson, r.h. (1977) evolution of oblitacythereis from paleocosta (osrracoda: trach;.leberididae) during the cenozoic in the mediterranean and atlantic. smithsonian contr. paleobiol., v. 33, pp. 1-47,rvashington. benson, r.h. (1978) the paleoecology of the ostracodes of dsdp leg 42-a. init. rep. deep sea drill. project,v.42 (t), pp. 777-787, (u. s. government printing office) rvashington. benson, r.h. (1990) ostracoda and the discoverv of global cainozoic paleoceanographical events. in: \whatley, r. & maybury, c. (eds.), ostracoda and global events, pp. 41-58, chapman and hall press, london. benson, r.h. & sylvester-bradley, pc. (1971) deep-sea ostracodes and the transformation of ocean to sea in 1.989, 1993). langhian of pescale, northern apennines (russo 1966) reported associated wrth macrocypris, argilloecia, krìtbe, bythocyprìs, b untonia radiatopora radiatopora (seguenza), bairdia and cytherel/a (russo 1966, p. na-tr). occurrence in the tremiti lslands succession. early langhian fupper part of the mmi 4a (p glomerosa sicana) subzone] and mid serravailian [mmi 7a (p partimlabiata/g. drury) subzone-lower part of the mmi zb (p mayeri) subzonel. ackno'eledgements. the author gratefully acknowledges professor nevio pugiiese and an anonymous referee for their fruitful comrìents to the manuscript. the author is especially indebted to professor antonìo russo for examining the type material of agrenocytbere saxolensis and discussing the morphological features of the specìes with him. ences the tethys. in: oertli, h. j. (ed.), paléoécologie des ostracodes pn 197q. bull. cent. rech. pau-snpa, suppl. 5, pp.63-91, pau. benson, r.h., chapman, r.e. & deck, l.t. (1981) palaeoceanographic events and deep-sea ostracodes. nature, v. 224, pp. 1334-1336, london. bonaduce, g., bismuth, h., g., ruggieri, russo, a. & mascellaro, p (1988) marine ostracods of the upper miocene of the \x/e11 ashtart 1 (gu1f of gabes, southern tunisia). in: hanai, t., ikeya, n., ishizaki, k. (eds.), evolutionary biology of ostracoda. deoelop. paleont. stratigr., 1 1, pp. 1087-1 1oo, tokyo. bonaduce, g., ciampo, g. & masoli, m. (1976) distribution of ostracoda in the adriatic sea. pubbl. staz. zool. napoli, v, 40 (1), pp. 1-154, napoli. brady, g.s. (1880) report on the ostracoda dredged by h. m. s. challenger during the years 1873-1876. rep. sci. res. voyage h. m. s. "challenger" 1873-1876, zoologt, v. 1 (3), pp. 1-184, london. dall'antonia, b. (2000) revision of the ostracode subgenus paleoblitacytherels benson, 1977. rio. ital. stratigr. palectnt.,v. 1a6 (l), pp. 391-398, milano. dall'antonia, b. (2001) unpublished thesis ostracodi miocenici dell'avampaese apulo-ibleo. ph. d. thesis, unit. pisa, pp. 1.-257. dall'antonia, b. (in review) miocene ostracods from the temiti islands and hyblean plateau: systematics and biostrar i graph y. g eobi os. lv on. dall'antonia, b., di stefano, a. & foresi, l. m. (2001) inte296 b. dall'antonia grated micropalaeontological study (ostracods and calcareous plankton) of the langhian wesrern hyblean successions (siciln italy). palaeog., palaeoclim., palaeoec.. v. 176. pp. 59-80. amsterdam. ducasse, o. ec peypouquet, j. p (1979) cenozoic ostracodes: their import.rnce for bathymerry. hydroìogy and biogeography. proc. ocean drilling program, init. repts.,v. 108 (48), pp. 343-363, (u. s. government prinring office) 'washington. iaccarino, s., foresi, l.m.,mazzei, r. & salvatorini, g. (2001) calcareous plankton biostratigraphy of the miocene sediments of the tremiti islands. rev. espafi. micropaleont., v. 33 (2), pp. 237-248, madrid. majoran, s. & dingle, r.v (2001) palaeoceanographical changes recorded by cenozoic deep-sea ostracod assemblages from the south atlantic and the southern ocean (odp sites 1087 and 1088). letbaia, v. 34, pp. 63-84, oslo. neviani, a. (1928) ostracodi fossili d'italia. 1. vallebiaja (calabriano). mem. pont. accad. nuoai lincei, v. 1.1., pp. 1-118, roma. pokorny, \( (1955) contribution to the morphology and raxonomy of the subfamily hemicytherinae puri. acta univ. carolinae, v. geologica 3, pp. 1-35, pragae. riha, j. (1989) ostracod interpretation of palaeodepth of miocene (lower badenian) calcareous clays near brno, czecholos lov akia. c o ur. f or s ch. i nst. s en ckeb erg., v. 7 1, 3, pp. 103-116, frankfurt am main.. riha, j. (993) some notes to the paleohydrology lower badenian (miocene) sea in the vicinity of knihonicka zpn, v. 15, pp. 1.57-1.63. of the brno. ruggieri, g. (1961) alcuni ostracodi quaternari e recenti aooartenenri aì cenere costa neviani. boll. soc. paleont.*rr-^ '''_'__'_ *' b",ital.,y. 1. (2), pp. 1-9, modena. ruggieri, g. (1992) considerazioni tassonomiche su ostracodi neogenici e pleistocenici risultate dalla revisione di vecchi lavori dello scrivente. boll. soc. paleont. ital., v. 31 (2), pp. 175-188, modena. ruiz, f. & gonzaìez-regalado, m.l. (1996) les ostracodes du golfe mio-pliocène du sud-ouest de l'espagne. rev. micropaleont., v. 39 (2), pp. 1.37-1.51,paris. russo, a. (1966) ostracodi langhiani del pescale (appennino settentrionale modenese). boll. soc. paleont. ital.,v. 3 (2), pp. 227-251., modena. russo, b., sgarrella, f. & gaboardi, s. (2002) benthic foraminifera as indicators of paleoecological bottom conditions in the serravallian tremiti section (eastern mediterranean, italy). in: iaccarino s.m. (ed.) integrated stratigraphy and palaeoceanography of the mediterranean middle miocene ria. ital. paleont. strat., v. 108 275-287, milano. sprovieri, r., bonomo, s., caruso, a., di stefano, a., di stefano, e., foresi, l.m., iaccarino, s.m., lirer, f.,mazzei, r. & salvatorini g. (2002) an integrated calcareous plankton biostratigraphic scheme and biocronology for the mediterranean middle miocene. in iaccarino s.m. (ed.) integrated stratigraphy and palaeoceanography of the mediterranean middle miocene riz,. ital. paleont. stratigr., v. 1a8 337 -353, milano. van den bold, wa. (1946) contribution to the study of ostracoda, with special reference to the tertiary and cretaceous microfauna of the caribbean region. dlss., utrecht unio., pp. 1-167, amsterdam. in: ellis, b.f. & messina, a.r. (eds.), catalogue of ostracoda, new york. \flhatley, r.c. & coles, g. (1991) global change and the biostratigraphy of north atlantic cainozoic deep water ostracoda./. micropaleont., v. 9, pp. 1.1.9-132, london. rivista italiana di paleontologia e stratigrafia volume 1ol nota breve short note description of an early ontogenetic evolutionary step in zepid o rb i t o i d e s : lep i d o r b i t o i d e s b i s am ber gensis asymmetrica subsp n., early maastrichtian (central turkey) e,rcan ozcan'i & sevinq ozkan-altiner'i'* receited april 16, 1999; accepted januatt 10, 2001 ke1'ruords: foramjnifera, nepionic arrangenent) lepidorbitoides, early maastrichtian, anatolia (turkey) riassunto. lepidorbitoides bisambergensis è caratterizzrto per ayere una camcra embrionale quadrìserìale senz-a nessuna cameretta che si origini dircttamentc dalla deuterocor.". è r.r. specre comune e molto diagnostica nelle successioni fl-vscioidi del maestrichti:rno inferiore dell'anatolia. alcune popolaz-ioni dr questa specre presentano una disposizione marcatamente asimmetrictr della prima c:mcrr, in visjone orizzontale, nei primi stadi ontogenetìci. tale asjmmetria è ."^ ,1" ..-" marcata differenza neila dimensione dellc canlerette ausiliaric chc si trovano sia nella protoconca che nella deuteroconc? e sono anche enfatizz,ate dallo sviluppo dispari di camerette nella serie che si origina da queste camerette ausiljarje sul lato della protoconca. questc popolazioni asirnmetrjche a c:rusa della compars;r cli un;r nuova camerette trusiliaria si trovano comlìnemente rr orrzzonti stratigrafici sottostanti quelli a disposizione simmetrica, che le sostitniscono progressivamente. gli esemplari asimmetrici "quadriseriali" rappresentano lo stadio filogenetico iniziale de l. biambergensis e sr troyàno negli orizzonti stratigrafjci corrispondenti alle zone t g. ha"r,anensís and g. aeg'ptiaca (?). si ritiene meritino uno stàtus tassonomico sottospecifico e vengono descritti con il nome dr lepidorbìtoides bisambergensis asymmetrica subsp. n. abstract. lepidorbitoides bísambergertsis is characterìsed by heving a 'quadriserial' enrbrl-o without any chamberlet directly arisine from the deuteroconch and is a very diagnostic and common specìes in lor.er maastrichtian flvsch successions in anatolia. some populations of this species present iìn earlv ontogenetic morphologic feature which js characterised bv distinctly asl.mmetric early chamber arrangement recognised jn the horizontal sections. this asymmetry is mrinly caused by the pronounced djfference in the size of auxiliary chamberlets which rest on both, protoconch and deuteroconch and also enhanced by the dcvclopmcnt of uncqu.rl rumbcr of chamberlets in the series arising fron.r these auriliary chamberlets on the protoconchal side. these asymmetric specimens are commonlv identified in stratigraphic horizons belor. the svmmetric ones after the introductìon of a new ruxiìia11ch:mberlet and proeressivell' replaced by symn.retric ones ìn the vounger popuìations. asymmetric 'quedriserial' specirlens representìng thc c:rly phl,logenetic stage of l. bisambergensis described ìn the stratigraphic horizons corresponding to g. ha'oanensls and g. deg)pt;dca (?) zones are thought to deserue a particlrlar taxonomìc status and are attributed rc lepidorbitoides bkambergensis asymmetrica subsp. n. lntroduction. lepidorbitoides bisambergensis ís an early maastrichtian species characterised by a 'quadriserial' nepionic chamber arrangement (specimens having two auxiliary chamberlets of van gorsel 1975 and 1928 and without any chamberlets arising directly from deuteroconch). although papp (1954, 1956) considered this species late campanian in age, same horizons which bear sìderolites calcitrapoides above l. campaniensis populations were dated as early maastrichtian by van gorsel (1975). caus (1988) rnd òzcen & ózkan-altiner (1999a) also considered this species eariy maastrichtian in age. l. bisambergezsis, which possesses two auxiliary chamberlets, has both symmetric and asymmetric variants as recognised in horizontal sections. asymmetric z. bisambergensls is distinguished from the symmetric ones basically by having an auxiliary chamberlet comparatively smaller than the other. the e.ìrly ontogenetic a5ymmetrv in some specimens of l. bisambergensis and its stratigraphic position below the symmetric ones were first recognised by van gorsel (1975, 1978). this author illustrated the embryos with tl.o auxiliary chamberlets; one ì/ith the dimensions smaller than the other and from which only few and small subsequent chamberlets are formed. although these asymmetric specimens were considered as the primitive stages of development of z. bisambergensls, the variation in early chamber arrangement, their biometric aspects and stratigraphic position have not been fully discussed. ' department of geological engineering, akdeniz university, 07200 topqular, antall'a, turke,v. ''r marìne micropaleontology research unìt, department of geological engineering, middle east technical universìty, 06531 ankara, turkev 5'ìì.ìi.ì.-;t/)--ihan] bartin 138 e. ózcan & s. òzkan altiner fig. 1 location of the sarnplìng sites (top) and generalized colurnnar sections of the measured sections han and hay-\fi/ (below). stars indicate the lepidorbitoiles and other orbitoidal foramìnifera bearing horizons. anatolian l. bisambergensis malnly represented by symmetric specimens have been identified in numerous horizontal secrions and its early ontogenetic biometric aspects we.re previously described by òtr^n (1995), ozcan & ozkan-aitiner ( 1997) and ó.crn & ózkanaltiner (1999a, b). the analysis of early onrogeneric features of lepidorbitoides specimens from secrions fian and hay-v measured in northwest central anatolia has revealed the different developmental stages of asymmetric and symmetric l. bisambergensis. .nli the specimens in the lower part of the secrion han have invariably asymmetric nepionts. these specimens have been attributed.to l. bisambergensis asymmerrlca subsp. n. (ózcan ec ozkan-akiner, 1999a). however, rhe complete description of the new raxon is here presented for the first time. lepidorbitoirles specimens secrioned in the stratigraphically lowermost orbitoidal foraminifera bearing horizon below the asymmetric ones have only one auxiliary chamberlet ('biserial' nepionrs of van gorsel 1975, 1978\ and were attributed to l. campaniensis. asymmerric specimens which are replaced by symmetric ones in the younger horizons seem to represent an important evolutionary step after the introduction of a second auxiiiary chamberlet during early maastrichtian and thought ro deserve an independenr subspecific status. asymmetric and symmetric l. bisambergensis populations are diagnostic for gl. bar.,anensis and g. deg/ptirtca zones. chamberlets which arises directly from deuteroconch (adauxiliary chambers of van gorsel, 1975 and 1978) are first observed in the populations consisring of asymmetric l. bisambergensis. material and age of the samples. asymmetric l. bisambergensis specimens have been identified in sections han and hey-v rneasured in the r icinity of town of hanònù (kastamonu) near the black sea coast and near the town of haymana (ankara) in northwest central anatolia respectively (fig. t). section han which commences with condensed, planktonic foraminiferal clayey limestone or shale lithologies at the base consists of thick, coarsely turbiditic horizons of mainly studied sections han hay.w =uj o o ag tu o uj i a é= of l thick. 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#� +3%'%��.-37(%' �#� �*'%+ �#� �-$' �#��#� �!.3)*'% #� k(%(+* �#� �-;;+ �# � �%3,'* # ��cc�� � �'+&+$/ -,0 5-&-'+,*+&+$/ +" *.' �55'% (+!',' +%+)� ',-&&.+1(,(0 &+!-&(*/� �.-0# 2 � ���� g�h� �e���ee� �+,0+,# rivista italiana di paleontologia e stratigrafia volume luz numero 3 pagine 363-366 dicembre 1996 paleoenvironmental control on the morphology of nummulites fabianii (prever) in the late priabonian parasequences of the mortisa sandstone (venetian alps, northern italy) enrico trevisani1 e. ce,sare andrea papazzoni2 key-utords: facies analysis, numtnulites, paleoenvironments, late priabonian, venetian alps, nonhern italy. riassunto. viene descritta la distribuzione dt nummalites fabiani.i. fabianii e nurnmulites fabianii retiatus nelle arenarie di monisa, un'unità prevalentemente silicoclastica del priaboniano superiore affiorante a sud dell'altipiano di asiago (prealpi venete). lanalisi di facies ha permesso di riconoscere una organizzazione ciclica in parasequenze tbickening-coarsening upward prodotte dalla progradazione di sistemi costieri. nelle facies di base e di tetto di ogni ciclo sono presenti abbondanti macroforaminiferi, fra i quali numerosi esemplari della specie nummulites fabia.nii. la distribuzione dei due norfotipi corrispondenti alle "sottospecie" n. fabianii fabianii (prever) e n. fabianii retiatus roveda è risultata strettàmente legata alle condizioni paleoambientalì,. n, fabianii reti.dtus, forma relativamente appiattita, si trova nella facies marnosa di offsbore a base ciclo, carattertzzata da bassa energia idrodinamica e bassa intensità luminosa. n. fabi.anii fabianii, forma più bombata, è invece tipica della facies biocalcarenitica di tetto ciclo, caratteristica di un ambiente meno profondo con energia idrodinamica ed intensità luminosa relativamente elevate. dalle osservazioni riponate non si evidenzia alcun legame diretto tra la morfologia e la posizione stratigrafica. lutilità strarigrafica comunemente riconosciuta alle due "sottospecie" (utilizzate per distinguere priaboniano inferiore e superiore) deve pertanto essere attentamente verificata. abstract. the distribution of nurnmulites fabianii fabianii and nummulites fabi.anii reti.dtas in the mortisa sandstone is discussed. the monisa sandstone is a terrigenous unit of late priabonian age occurring south of the asiago plateau (venetian alps, northern italy). the facies analysis allowed to recognize thickening-coarsening upward cycles interpreted as shallowing upward parasequences corresponding to depositional regressions. in each cycle five different facies were recognized. the first and the last facies of every cycle contain larger foraminiferal assemblages with numerous specimens of nummulites fabianii. two different morphotypes of this species were attributed to the "subspecies" n. fabianii fabianii (prever) and n. /abianii retiatus roveda. their distribution was influenced by the paleoenvironmental conditions. the flat "subspecies" n. fabianii retialrs occurs in the base-cycle marly facies, deposited under low-energy, lowlight conditions. the more inflated n. fabianii fabianii characterises the shallower top-cycle limestone, deposited under high-energn highlight conditions. our observations suggest there is no obvious link between the morphology and the stratigraphic position, so the biostratigraphic significance of the two "subspecies" (currently used to divide the priabonian in a lower and an upper part) remains uncertain. introduction. this note deals with the problem of the biostratigraphic significance of the so-called subspecies n. fabianii fabianii and n. fabianii retiatus. they have been used since now to divide the priabonian in a lower (with -m fabianii fabiani) and an upper (with n. fabianii retiatus) part, but some authors (herb & hekel, 1973; barbin, 1988) pointed out the possible paleoenvironmentai control on their distribution. in the frame of the igcp 393 ("neritic events at the middle-upper eocene boundary") the problem of the biostratigraphic usefulness of these "subspecies" has to be carefully considered. in this paper we cannot discuss the problem exhaustively: the data presented here are only the first step. additional studies on more morphological features could help to better understand ìf n, fabianii fabianii and n fabíanii retidtus are real subspecies or simply morphogroups ecologically controlled. our study was made on n, fabianii fabianii and n. fabianii retiatus fabianii populations from the mortisa sandstone unit. this is a terrigenous unit of late priabonian age deposited on the eastern margin of the lessini shelf. it crops out south of the asiago piateau, along the piedmont zoîe between the astico river and the brenta river (fig. 1). its thickness decreases from 'sfest to east: about 60 m near calvene and about 10 m in the bassano del grappa area. the mortisa sandstone unit has been studied since the past century and dated as lower oligocene (oppenheim, 1900, 1901). more recently the age'was corrected ' m,r."o di storia naturale di ferrara, via de' pisis 24, i-44100 ferrara, italy. 2 dipartimerto di scienze della terra (paleontologia), università di modena, via università 4, i-41100 modena, italy. les sini mortisa la calvene@ "1 lusiana o ovalle di sopra niarostica a i ì padova bassa bassano del grapp km -. =05 364 e. treaisani ct c. a. papazzoni as upper eocene-i-ower oligocene (frascari ritondale spano, 1967). the regressive features of the unit and its variable thickness 'were pointed out by frascari ritondale spano (1969). frascari ritondale spano & bassani (1973) considered the mortisa sandstone as sublittoral or deltaic deposits forming the basal member of the calvene formation. trevisani (1993) remarked the cyclic organizatron of the mortisa sandstone and interpreted the cycles as shallowing upward parasequences. facies analysis. the mortisa sandstone consists of thickening-coarsening upward cycles (t-c-u), where five different facies have been recognized (fig. z). the a facies is represented by bioturbated marls, less frequently marly limestones, with nummulites; the b facies by bioturbated coarse siltstones to fine sandstones; the c facies by sands and sandstones vrith storm layers; the d facies by conglomeratic sandstones to sandy conglomerates with storm layers; the d facies by biocalcarenites with nummulites. the thickness of each cycle increases westward, from about 10 m in the bassano area to over 40 m near calvene. on the basis of facies analysis these t-c-u cycles are interpreted as shallowing upward parasequences which are the result of depositional regressions produced by rhe progradation of shallow water siliciclastic systems. these progradational phases were interrupted by abrupt relative sea level rise, resulting in a backshift of the various facies belts toward more internal positions. these parasequences ^ppear to be part of a depositional sequence bounded in their lower part by an erosional limit underlying the pradelgiglio formation. therefore, in terms of sequence stratigraphy, the mortisa sandstone fig. 1 l-ocation map of the study area and sites of the examined srretigraphic sections. l) mare section; 2) crosara section; 3) pradipaldo section (from trevisani, 1993) represents the lowstand phase of this depositional sequence. for further details see trevisani (1993). paleontologic analysis. although the mortisa sandstone is prevailing sili ciclastic, in some parts 'we found assemblages of larger foraminifera. in particular, they are abundant in the a facies at the base of the cycles, which corresponds to an offshore environment below wave base level, and in the e facies at the top of the cycles, corresponding to bioclastic sandy shoals. also in the uppermost part of the d facies the larger foraminifera are often abundant. among the larger foraminifera the specimens of nummulites fabianii (prever) were recognized to show different morphotypes in different facies: in the a facies the tests are distinctly flatter than in the e facies. these morphotypes correspond to the so-called subspecies n. fabianii fabianii (the more inflated ones) and n. fabianii retiatws (the flatter ones). the distinction between the two "subspecies" relies mainly on external features such as the ornamentation and the more or less inflated profile. roveda (1959, l97o) reported that n. fabianii retiatus has an ornamentation with larger mesh than n. intermedius (:n. fichtel) and a transverse lamina weaker than that of n. fabianil. moreover, n. fabianii retidtus is flatter than n. fabianii fabianii. the profile is well approximated by the diameter/thickness (d/t) ratio. from the data reported by roveda (1970) this ratio resuits about 2.0 for n. fabianii fabianii and about 2.7 for n. fabianii retiatws. in the upper eocene section of possagno, some fifteen km north-east of bassano del grappa, flerb & :ts : d c b -+2 a {rt hd td i fig. 2 modal cycle (thickness variable from 10 to 40 m) of the monisa sandstone. a-e) different facies recognized; td) transgressive deposits; hd) highstand deposits; 1 and 3) marine-fl ooding surface (parasequence boundary); 2) surface of maximum flooding (modified from trevisani, 1993). hekel (1973) found a lithologically controlled distribution of the two "subspecies": the limestones contain à{ fabianii fabianii and the silty-argillaceous sediments n. fabianii retíatul barbin (1988) made a comparison between the test flattening of -m fabianii fabianii from buso deila rana, near priabona, and the n. fabianii retiatus descrrbed by roveda (1959), concluding that n axial sections of nummulites fabianii (x 25). a) n. fabia' nii rctiatus; sample mv 9, mare section, lower part of the a {acies; b) n. fabianii fabianii; sample mv 21, mare section, e facies. n. fabianii motpbology in the mortisa sandstone 365 fabianii retiatus is an ecologically controlled morphotype of n. fabianiifabianii. our samples from the mortisa sandstone were examined to test the possible environmental control on the morphology. the d/t ratio of the specimens of n. fabianii from different facies was measured. in the e facies, represented only by hard limestones, diameter and thickness were measured on thin sections only where the proloculus was visible and the centrai pillar well recognizable. nevertheiess, a correction was made for the possible obliquity of the section, reducing all the measured values by about 10%. the correctness of this assumption was tested on two samples where both free specimens and thin sections were available. the mean d/t ratio of the n. fabianii from the a facies amounts to about 2.8, whereas it is about 2.7 for the samples colleoed in the e facies. in particular, the sample mv 9 (fig. 3a), from the base of the a facies in the mare section, shows a d/t ratio of 2.89 (twenty-four isolated specimens measured); the sample mv 21 (fig. 3b), from the e facies of the same section, about 45 m above mv t has a d/t ratío o[ 2.05 (six axial sections measured on two thin sections; thickness vaiues corrected). therefore, the e facies is characterizedby n. fabianii fabianii, whereas n. fabianii retiatus occurs in the a facies. discussion and conclusions. in living larger foraminifera the test shape depends mainly on light intensity (influencing algal symbiosis) and water energy (reiss & flottinger, 1984; hallock ec glenn, 1986; hallock et al., 1986, i99l). the recent nummulitid operculina ammonoides shows flatter tests on soft substrates than on hard ones at the same depth (pecheux, 1995). in the mortisa sandstone the a facies contains flat species of nummulitids: together with n. fabianii retiatus there are n. chaoannesi de la harpe and operculina spp. this agrees well with the sedimentoiogical evidences of a low-energy, lowjight, relatively deep paleoenvironment with soft substrate. on the other hand, in the e facies n. fabianii fabianii is associated with the inflated species n. incrassatus de la harpe and l'.1. variolariu.s (lamarck). f{ere a high-energy, highjight, shaliower paleoenvironment (probably with hard substrate) is inferred. in conclusion, the late priabonian n fabianii shows two different morphotypes, probably controlled by the paleoenvironmental conditions (light intensity, water energy, substrate) . nummulites fabianii fabianii prevails on hard substrates, with high light intensity and high water energy, whereas numrnulites fabianii retiatus occurs in opposite environmental conditions. f;. 1 366 e. trevisani & c. a. papazzoni the data here reported are not conclusive to ascertain if we are facing two morphogroups of the same taxon, produced by different paleoenvironmental conditions (like recent operculina arnmonoi.de) or really two distinct subspecies with stratigraphic significance. nevertheless, we underline that the sample mv 9 (with n fabianii retiatu) is well below mv 21 (with n. fabianii fabiani). therefore, the two "subspecies" do not appe r one after the other but are present at the same time, at least in the upper part of the priabonian. thus, our observations suggest that the ,'subspecies" of n. fabianii could be ecological morphotypes, but a more comprehensive study is needed to solve the problem of their biostratigraphic value. acknotaledgements. '!le thank prof. a. sirotti (modena university) for his help in determining the larger foraminifera. many thanks to prof. m. gnaccolini (milan university) and prof. l. hottinger (basel university) for their useful suggestions to improve the manuscrrpt. references barbin v. (1988) réflexions srr nurnmulites fabianii et nummulites fabianii retiatus, leur signification stratigraphique et leur origine phylétique. rezt. paléobiol., v. spéc. 2, n. 2, pp. 609-614, genève. frascari ritondale spano f. ( 1967) osservazioni stratigrafiche e rertoniche sui dintorni di calvene nel vicentino. giorn. geol., v. 3a (966), pp. 307-340, bologna. frascari ritondale spano f. (1969) serie paleogeniche nell'area pedemontana a sud dell'altipiano di asiago (vicenza). mém. b.r.g.m., v. 69, pp. 773-182,parìs. frascari ritondale spano f. ec bassani p. (1973) ricerche geologiche nei dintorni di bassano del grappa (vicenza). mus. tiident. sc. nat., v. 19, pp. 65-106, trento. hailock p., forward l.b. e{ flansen h.j. (1986) influence of environment on the test shape oí amphistegina. journ. foram. res., v. 16, n. 3, pp.224-231, washington. hallock p. & glenn e.c. (1986) larger foraminifera: a tool for paleoenvironmental analysis of cenozoic carbonate depositional facies. palaios, v. 1, n. l, pp.55-64, tulsa. hallock p., róttger r. 8c 'wetmore k. (1991) hypotheses on form and function in foraminifera. in lee j.j. er anderson o.r. (eds) biology of foraminifera. academic press, pp. 47-72, london. herb r. & hekel h. (1973) biostratigraphy, variability and facies reiations of some upper eocene nummulites from northern italy. ecl. geol. helo., v. 66, n. 2, pp. 419-445, basel. oppenheim p. (1900) ueber eine riesige perna (paclryperna n. sbg. laoerdana n. sp.) aus dem alrtertiàr venetiens und die ùbrigen perniden des gebietes. zeitschr. deutsch geol. ges., v. 52, pp. 232-243, stuttgart. oppenheim p. (1901) die priabonaschichten und ihre fauna. palaeontograpbica, v. 47, pp. 1-348, stuttgarc. pecheux m.j.-f. (1995) ecomorphology of a recenr large foraminifer, opercalina ammonoides. géobios, v. 28, n. 5, pp. 529-566, lyon. reiss z. 8c hottinger l. (1984) the gulf of aqaba. ecological micropaleontology. y. oí 354 pp., springer-verlag, beriin. roveda v. (1959) nummulites retiatus, nouvelle espèce de nummulite réticulée des abruzzes (italie). rea, mioopaléontol., v. 1., n. 4, pp. 207-207, paris. roveda v. (1970) revision of the nummulites (foraminiferida) of the n. fabianii.fichteli group. rio. it. paleont. strat., v. 76, n, 2, pp. 235-324, milano. trevisani e. (1993) analisi di facies e ciclicità delle arenarie di mortisa (priaboniano superiore, prealpi venete). mem. sc. geol., v. 45, pp. 57-65, padova. receitled marcb 18, 1996; accepted, october 3, 1996 il 12 novembre 2001 si è spento a roma ardito desio, alla bella età di 104 anni. la sua scomparsa ha avuto larga eco anche tra il grande pubblico, che lo conosceva per la sua attività di esploratore, geologo e per la spedizione alpinistica al k2. in questa sede 1o voglio ricordare per lo speciale legame con la rivista italiana di paleontologia e stratigrafia e per quella parte della sua multiforme attività che ebbe come oggetto la paleontologia e la stratigrafia. laureatosi in scienze naturali a firenze ne| 1920 con di stef"ri qr^n nîrtc .'l"ll" nrime ricerche scientifiche si svilupparono nei campi della paleontologia e della stratigrafia, come al.veniva a quel tempo trattando di successioni sedimentarie. sediin memoria ardito desio (1,8e7-2001) ardito desio nel giorno del suo 100 compleanno. ardito desio the day of bts 1aa birthday. ardito desio died in rorne on not,ember 12, 2001, at the imposing age of 104 years. his departure created a large echo also among the lay pwblic ruho knew bim as dn explorer, a geologist, and for tbe successfwl climbing expedition to the k2 in karakorwm. here i ,ízant to remember him for bis special ties to tbe ri'uista italiana di paleontologia e stratigrafia and for that part of his eclectic professional lfe that deah u,itb p ale onto logt an d stratigrap hy. he became doctor of natwral sciences in 1920 in florence, studying under prof. di stefani and tbe majority of his early scientific researclt deah witb paleontologt and stratigraphy, follouting tbe taditional approach of tbe time to the study of sedimentary succession. sedimentologt and brittle structurmentologia e geologia strutturale de1 fragile dovevano ancora nascere! ricordo tra queste 1o studio delle ammoniti cretacee del bacino di firenze, e dopo il trasferimento a milano, 1o studio della successione dell'albenza, divenuto in seguito il classico terreno di esercitazione degli allievi geologi di milano. a milano si era trasferito nel 1,925 come conseryatore presso il museo civico di storia naturale, gloriosa istituzione voluta da a. stoppani. in quella sede si tenevano anche 1e lezioni di geologia e mineralogia per il politecnico e la nascente università. a milano fervevano grandi iniziative. da secoli, dal tempo del ducato di milano, la città universitaria della lombardia era pavia. ma nei primi anni del'900, la borghesia illuminata del tempo si unì alle iniziative statali e sorsero a relativamente breve distanza di tempo il politecnico, i'università statale e la libera università bocconi. desio, sempre moìto attento a cogliere ie nuove opportunità, vinse la cattedra di geologia nella nuova università e fondò l'istituto di geologia. accanto a questo tuttavia non smise una intensissima attività di terreno, in particolare in libia, ma anche laddove 1'espansionismo del regime aveva bisogno di conoscenze geologiche. gli anni'30 furono ricchi di esplorazioni. e la geologia esplorativa nel campo del sedimentario ha innanzitutto bisogno di stabilire una buona stràtigrafi^, al geology were still to come! among bis early rporks i r.oant to remember tbe study on the cretaceows ammonites of the florence basin and, after bis move to milan, the study of the albenza swccession tbat later became a classic area for practicingfield worh by the geologt stwdents of milan. he transferred to milan in 1925 as curator of the mwseo ciolico di storia natwrale, a gloriows institution established by a. stoppani, where the lectures of geolog, and mineralogt for the polytechnic and the developing unioersity were also held. milan was bwstlingruith great initiatioes. paoia had been the city hosting the only wniversity in the region of lombard.y for centwries, since the time of the dukedom of milan. hozaeper, at the beginning of the 1900's tbe enlìghtened bourgeoisie joined the effort of the state to create, in a sbort interval, the polytechnic, tbe state university and the free university bocconi. desio, always ready to seize new opportunities, won the chair of geolog with tenure in the new uniaersity and founded the institwte of geologt. in addition, be neaer stopped to uorh aert intensely in the field, especially in lybia and wheret,er geological knowledge was required by tbe ltalian expansionistic regime. tbe thirties u)ere rich with explorations, and exploratioe geologt first needs a solid stratigraphic backgrownd that in twrn is based on sound paleontology. desio carried owt some good, strictly palememorte la quale non vale niente se non c'è una altrettanto buona paleontologia. qualche studio prettamente paleontologico desio riuscì anche a farlo su reperti raccolti in libia, ma ne mancava il tempo tra tante iniziative. nel 1938 discusse con lui la tesi di laurea carla rossi poi maritata ronchetti, che da allora divenne i1 suo punto di riferimento in questo settore. la passione editoriale lo portò a fondare gli annali del museo libico, in cui furono pubblicati gli studi sulle ampie collezioni di fossili che nel frattempo aveva accumulato nelle sue esplorazioni a cufra, ne1 bacino di murzuch e lungo le coste della tripolitania e della cirenaica. ne furono pubblicati quattro volumi. le collezioni di fossili depositate a tripoli andarono purtroppo disperse, ma molte copie degli annali sono ancora depositate nella biblioteca di scienze della terra dell'università di milano. e venne la guerra. nonostante questa, nel 1942 si concrerizzàrono due iniziative. l'istituzione del corso di laurea in scienze geologiche di cui desio fu un propugnatore e l'acquisto della proprietà della rivista italiana di paleontologia. questa rivista, sorta nel 1895 a perugia, dapprima si era limitata a recensire articoli pubblicati altrove, ma poi gradualmente aveva preso a pubblicare anche ricerche originali. ne1 1923 1a sede venne trasferita presso i'istituto geologico della r. università di pavia e nel 1.927, come recita la carra in bollo conservata negli archivi della rivista, i1 procuratore generale del re ne dichiarava il prof. paolo vinassa de regny, allora professore a pavia, direttore responsabile. nel 1942 vinassa de regny, che aveva anche acquistato la proprietà, vendette la testata ad ardito desio, i1 quale ottenne dal procuratore generale del re impe ratore la nomina a direttore responsabile. e così la rivista approdò a milano, gestita presso l'istituto di geologia, ma proprietà personale di desio, che rimase direttore responsabile sino a1 1995. carla rossi ronchetti, coadiuvata poi da fausta guaitani e carla albanesi, ne curava la redazione. nel 1947 su iniziativa di desio che fu il primo presidente, venne fondata la società paleontologica italiana con sede presso il museo civico di storia naturale di milano. la rivista divenne i'organo ufficiale della nuova società. poiché, sulla base della sua esperienza professionale trovava difficile scindere paleontologia e stratigrafia, desio cambiò nome alla testata che divenne rivista italiana di paleontologia e stratigrafia. negli anni '50 però, anche perché desio era intensamente impegnato nelle spedizioni in karakorum e hindu kush, la sua spinta propulsiva per sosrenere 1a società paleontologica si affievolì. verso la fine degli anni '50 assunse la presidenza della società e. montanaro gallitelli, ed il nuovo consiglio si diede un proprio periodico, il bollettino de11a società paleontologica italiana, con redazione presso l'istituto di paleontologia dell'università di modena. il primo numero di questa nuova rivista uscì nel 1960. ontological research, but with so many initiatioes his time was limited. in 1938 carla rossi, later married roncbetti, defended with him her tbesis and became from tben on bis referral point in this area. a passion for pwblishing browght hìm to fownd the annali del mwseo libico, in r,uhich tbe large fossil collections assembled during his expeditions to cufra, the murzucb basin and along the coast of tripolitania and cirenaica were published in fowr oolumes. [jnfortwnately, the fossil collections stored in tripoli zuere lost, but ser.,eral copies of the annali are still leept in the library of tbe earth science department of the unioersity of milan. tben came the war. even so, t'tso initiatives materialized in 1942, the creation of the degree in geological sciences, one of wbose supporters r.uas desio, and tbe purchase of the rivista ltaliana di paleontologia. this publication, born in perugia in 1895, was limited initially to tbe review of papers pwblisbed elsewhere, bwt gradually began to publish orìginal stwdies. in 1923 it zuas tra.nsferred to the geological institute of the royal uni,sersity of pavia and in 1927, as recorded by a document zoith official seal preserved in the archiaes of the rivista, tbe general prosecwtor of the king declared prof. paolo vinassa de regny, then professor in paoia, editorin-chief of tbe riz,ista. in 1942, p vinassa de regny, who had also purcbased the title of the rioista, sold it to ardito desio, wbo obtained from the prosecutor of the king emperor the appointment of editor-in-chief. so the rioista uds transferred to milan, operated from the institwte of geologjt, but ns the personal property of desio, who remained director in charge wntil 1995. carla rossi ronchetti, helped by fausta guaitani and carla albanesi, utas the managing editor. in the 1947, under the leadership of desio who was the first president, the società paleontologica ltaliana uas founded, based on tbe museo cipico di storia naturale of milan. the rioista became the officìal journal of the new, society. becawse of his personal experience in whicb paleontologt and stratigraphy were strictly tied, desio changed tbe name of tbe journal in ri.c,ista ltaliana di paleontologia e stratigrafia, in tbe fifties hozaeaer, being desio deeply inpolved in the geological expeditions to karaleorum and hindu kush, bis interest in tbe paleontological society decreased. tberefore, towards the end of fifties, e. montanaro gallitelli became the new president of the society and the new board created its orpn bollettino della società paleontologica, with editorial offices at the institwte of paleontologt of the (jnioersity of modena. tbe first number of the new journal u,as issued in 1960. finally, tbe last act. approacbing bis tenure as professor emeritus, desio decided to donate the right of title of the rioìsta to the (jniversity of milan, who accepted the gift in 1976. desio establisbed some clawses, arnongwhich was the rule tbat the director bad to be a professor of paleontolog,t or geologt of the unioersity of milan, and rivista i taliana paleonto logia e stratigrafia infine i'ultimo atto. awicinandosi al fuori ruolo, desio decise di donare la proprietà della testata all'università di milano, la quale accettò la donazione nelt 1976. desio pose alcune clausole, tra cui i'obbligo che il direttore fosse un professore di paleontologia o di geologia dell'università di miiano e che questa si impegnasse, ove necessario, a sostenere frnanzíartamente la rivista. la redazione rimaneva comunque nelle salde mani di carla rossi ronchetti. infine alf inizio degli anni'90, riducendosi finteresse alla redazione da parte della "signora", anche desio decise di passare la mano e nel 1995 indicò chi scrive come nuovo direttore della fuvista. questo piccolo scorcio di storia, che tuttavia abbraccia 3/o di secolo, è parte integrante della storia della paleontologia e della geologia del sedimentario in italia. io credo che noi ele generazioni più giovani di studiosi e studenti debbano essere grati a chi ha saputo non solo tenere viva la fiamma, ma alimentarla e consentire che questo periodico scientifico potesse festeggiare il secolo di vita ed essere incluso nella lista dei periodici isi. le riviste universitarie non hanno un futuro facile, complicato dai costi, dalie necessità della internazionahzzazione e dalla crescente messa on-line del1e pubblicazioni scientifiche da parte dei gruppi editoriali commerciali, ma penso sia nostro dovere cercare di mantenere e sviluppare quanto è stato sinora costruito da persone come ardito desio. e' il modo migliore per ricordarli. that tbe unioersity was committed, if necessary, to support the riuista financially. howe,t,er, the editorial office remained in the experienced hands of carla rossi ronchetti. eventually, at the beginning of the nineties, 'tuhen the interest of the "signora" in the editorìal process started to roane, desio also decided to pass tbe baton and in 199) proposed tbe rpriter of this piece as netu i dttnr-tn-( htel nf thp rivista. tbis short excerpt of history, tbat ner,tertheless spans three quarters of a centwry, is an integral part of the history of paleontologt and sedimentary geologjt ìn ltaly. i belieoe that we and tbe younger generations of researchers ancl students need to be grateful to the person who not only kept the flame bwrning, but also nourisbed it ancl made it possible for this scientific publication to celebrate its centennial and be included ìn the list of isi perioclicnls. (jni.,.,ersity magazines clo not face an easy future, complicated by costs, by the need to become more international and the inueasing number of scìentific journals nou, available on-line. hotoeter, i beliez,e it is our duty to try to maintain and der,,elop zt,bat zoas built to date by people like ardito desio. it is the best way to remember them. maurizio gaetani renesto 145..160 � ��� �������� � �� �� � � � � ��� ������ ������� ������� ��������� ����� ����� � ��������� ���� ��� ����� ������ ������� ��������� ��� � � �� �� �� ��������� ��� �� �� � ��� �� �� ������� � ��� ���� ������ �������� ������ ���� �� ���� �� � �������� �� �� �������!� ��� � ��� "������� �����#���� ���� ����� ��! ���!���� ��!$� ��� � ������% ���� ���% � ������ !����� � #� ���� �!��� � �&��� �� �' � ����� ��(� �� $����� �� ��!��#� % �$� ���!���� )�� !����! � �� *++, '��� �� �� !��� �' $� ������ ����� ��� "���(#� ����(��� �' �)��� �� $���� �' �� �� ������� �� �� �������!� ���� �� $� ��� � ��� "������ ����� ��� $��� � ���% �$� �� ������ �' $� ��� !������ �&��� �� ������ � $� ��������� � $� ����� ������� �� �� ���� �-!�� �� � � #�������� � ������� � ������� � !� �������� � �"# �$� � ����� � &��)� '��� !��.�� ��!��� ��� �' $� ������ ����� ��� �� $� �)��� ��� �' $� ��� � ��� "������ ����% /���� � $� ��!& �' ��� �' $� �&���� )$�!$ !�� ���� ������ �! !$���! ��� � $� ���!��� ��.�� '�� ������� �� $� '�) ������.� !������ ������ � ������ ��� ������ ��� )� $ �% & ����$�� )$�!$ �&���� �� ���� &��)� ��� � �� ���� �' !������#�� ��(�% �$�� ���!���� �� ��� �!������ �����'�!�� #�!���� � �����.�� $� &��)�� �� �' $� �� ������ �' �% & ����$�� �� #�!���� $� ��!�� ������� � ���!���� ������� $�� �% ������� � !% �"�$� �� ������ � !��-�� ��!� �' ����� ��! ���!��� !$���! ���(� #� ��(� �� � ���#�#��� ���$�! �''���� �� ��� )� $�� $� ����� ������� � �� $� ��� � ��� "������ ���� �� �!!���� �� $� !��.�� "������! �����% ����� ���% ����� ��!�� � �� �!$��� �� 0���� !����� �� �� �� ����� � ���� �!��� � � ���.� � 0���� � � �� !������ � �� ��!!��� �� ������'���� ���.����� � ���� �����(���� � ������ "���(#� �� ���(���� ������� � ���1�����!� ����� ��� �������!� �� ���% � !��� ��� �� ������!� ���� �!$��� �� ��� !������� !����� ��� �1� ��#�(���� �� ��� ���� ������� � � ��!�� ��� ��� !�� ���� �� ��� ����� ����(� ���� ���!�� ������� � ������� � !� �������� � �"�$� �� ���2 �� � � �'� '������� � �.�((��� ���� "���(#� ����(���% 3�� ����� �1��������� �2 0���� !����� ���� � ���.� �� !������ !$� !�� ���� �� ������� ��� � �� !��� ��� ������ �!� � ��.���� ���!�'�!� ��� ������� �� � �.�� �� ��� � !�����.� � ����� ��� � ��� �((��� �� �����#��� ����� ����(� ���1�� �������� � �% & ����$��% �� �� ��.���� � �2 ��� �!������� � �����'�!� �.� ���!$�4 �� � ��� ��� � ����� ���##� � ����� ��� !���� ���.����� � �� !����!��(� ����1�� ������� � �% & ����$��� �� ���� !��� �� ������ �� ������(� � ��� ��!�� � ���!�� � ������� � ����� �����(���� � ������ � ��������� ������ ����� � � ������� � ������� � !� ���# ����� � �"�$� �% 5��� � ��������!� �� �!������ � !��������(� � ���!�� ����� ��!$� ���� � ���� ������ ����1���� �� ��� � ��� "������� !��� ���((� � � ��������� �''���� �� !$� 0��� � �� !�#�.��� � ��� � �.���� !�� ���� �.� �� �.������ � ��!!$�� ��'�!$�� !��� �2 � � � ���� ��� � ��� � �� ������ !��.� �� ��!��� "������!�% ������������ �$� ��!$ .�� �#�� � '���� �' ���������� � ��� "������ ���#�� �� � ��� �� 6�� �� ��!���� �)� (��� ��� � �� ����� $� ��� ����� �� �� $� )��� '�� $� �� �� �������! �.�� �' �� $� ��!�� ��!�.����� �� 6$�� ��% �$� �� � �������� � �� ��.����#�� ��� '�� !������� ���� ����� $��� '����� '�� � ���� !����� � ������� ��� �' $� #���������$� �� �.��� ��� �' �� �� ������ ��! 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7�8*% ��� �� �%3% 7;8;� � �$� ��!$������������ � ��� ���� �� $�������� '��� $� �� �� �������! �' ��� � ��� "������ �)� (����� �� )� $ $� ��!��� ��� �' � ��) ���!���% !��$% ' ���% ���% *��% (����� �� ,*= =@7�@:+% �!$��' �% �%� b����� �% a ���� ��� �% �% *+++� � 3�� � �� ��(� � ���� ��(� ���� ����$��� �' ������ '��$ ��� � ��� �� �����!�'�! .���� ��� �� �''�! � �' �� ����� �� #� � ��(� �� ���$�!���!$� #��� $% �� % 4��$% ! �� % *� % *+8 **;�*98% ��!$��( >% 7;8;� � (� ���� � � ))�� �% ��% '��� $� �� � �� �������! �' ��� � ��� "������ �)� (����� � )� $ !����� � �� $� !�����'�!� ��� �' ������� �% !�# $������� ������ �� *+8 7=,�7:;% <������ �% 7;,=� � /�� �����'���� �� �������� >��&����� ;� !�����$� ��� � ���� ����� 6������� ��% �� ����� ��������� ����#��� ����������% ��� *�����) !�$�����% -��%� =@ 7�8+% ������� *%7@+ rir.ista ltaliana di paleontologìa e stratigrefia short note nota breve eo styloceros cf. ptdopltt schkoi korotkevits ch 1,964 (cervtdae, muntiacinae): ne\t element in the neogene mammal assemblage of lo\ter valdarno (tuscany, central italy) laura abbazzi'e{ roman croitor, receii:ed september 16,2002; accepted mdrch 28, 2aa3 key raords: cen idae, muntiacinte, eos4t loceros, sl.stematic plleontologl-, lowcr valdarno, tuscany, early to middle piiocene. abstract. an eostyloceros antler fragment ìs described. tbe specimen makes part of rhe montopoli (lon'er valdarno, tuscany) collection preserved jn the geo-paleontological section of the natural history museum of florence. this site is the t.vpe locality of the homonymous faunal unit of the ltalian biochronologìcrl scheme; ir includes the latest early villafranchian mammal age faunas. the occurrence of an eostyloceros species among this material howeler, is in disagreement with the age of the montopoli local fauna, whìch has been correlated with the gauss-matuyama boundaru and thus dated to about 2.6 ma. the youngest remains of this genus come from sites of eastern europe early pliocene in age (ruscinian mammal age). the possibilìt1' that the anrler ìs an external elenrent to the montopolì assemblage js therefore contemplated. rìassurtto. nel presente lavoro viene riportata per la prima volta la presenz-a della specie eostyloceros cf. pidoplìtschkol korotkevitsch tra ìl materiale rinvenuto nella località montopolì. montopoli, nel valdarno inferiore (toscana), è una località fossìlifera nota fìn dall'800, in cui sono stlti rinvenuti resti di mamrniferi che rappresentano un punto di riferimento per la biocronologia italiana. la fauna locale dì montopoli, calibrata in prossimità del pessaggio tra i magnetocroni gauss e matuyama, è stata infatti utilizzata per definire la parte alta dell'etàl a mamnriferi villafrenchìano ìnferìore. l:r presenza dt eostyloceros costiruisce un dato nuovo per l'italia e contempor.rneamente pone dei problemi biocronologici in quanto le segnalazioni più recenti dei suor rappresententi provensono da localitii dell'europa orientale riferite al pliocene inferiore (e,tà a mammiferi rusciniano). per giustificarc h pre.enza di que.r,, muntiacino tr.r il materiale fossile di montopoli vengono prese in considerazìone due ipotesi: secondo la prima il palco analizzato è rimaneggiato e proviene dall'elaborazione di depositi di origine continent:rle di età pliocene inferiore; sulla base della seconda ipotesi si suggerisce che il reperto non rir rtrto raccolro a monropoìi, mj protengl in rerlrr d.r un'rrer ldiacente in cui affiorano terrenì continentali del pliocene inferiore. location map of the montopoli site. the extension of lower and middle plìocene marine deposits is shown. 50121 firenze itaìia e-mail:labbazzi(!geo.unifi.it 1, 2028 chiqinàu moldor.a e-mail: karpatos@,ìexcite.com 1 2 dipartirnento di scienze della terra institutul de zooìogie al academiej università di fircnze, via g. la pira'1, de $tiinfe din moldova, str. academìeì november o lucco ìî "-r\l\ n\x lj ]) lrvorno o lo km montoooli locol founo aurelion i golefion i c q -c o c i\+-p i c a c-^ a l c. c c) 0)p q c o + oz c) c 0 o o+ -<,/) 0 o_ c) c o o 9 o_ bo u f q 6 o c) op c-.n c c) o o zi o =o_l c o c) o c o n l. abbazzi ?x r. croitor fig.2 chronological scheme. lntroduction a fragment of a shed antler of eostylctceros cf. pidoplitschkol korotkevitsch was found during the inspection of fossil deer material from the locality montopoli (lower valdarno, tuscany, central ltaly, fig. 1), stored in the geo-paleontological section of the natural history museum of florence university. the fossil was apparently recovered by forsyth-mayor in the 1880, as suggested by an original hand-writren nore, but its exact stratigraphic provenance is unknown. montopoli is celebrated (\fleithofer 1893; forsythmalor 1877,1885; azzaroll 1977) for the high numbe r of fossil mammal remains it yielded from middle pliocene shallow marine deposits (de giuli et al. 1983; benvenuti et al.1995). the fossil-bearing beds have been calibrated to the gauss-matuyama boundary and thus dated to about 2.6 ma (lindsay et al. 1980). though the specimens do not all come from the same stratigraphic level, they are altogether considered as a local fauna (de giuli & heintz 1974a) representative of the homonymous faunal unit which defines the l:ìtest part of the early villafranchian mamnr:rl aqe (azzaroli 197a,1977; gliozzi et a1.1997 fig. 2). the unpublished antler fragment, obl'ect of the present work, differs from all other montopoli cervids, which include: eucladoceros sp., pseudodama c[. lyra, procapreolus cusanus and croizetoceros rllmosus (de giuli & herntz 1974b: azzaroh 1977,1992\ the presence of an eostyloceros representative in lower valdarno opens an intriguing biochronological problem, since eostyloceros is reported from various eastern european and asian latest miocene and early pliocene localities (ruscinian mammal age; vislobokova 1983,1990; vislobokova et a|.1993), unless we do not believe it is a ruscinian holdover in middle piiocene times. this however contrasts with the acknowledged disappearance of ruscinian elements that marks the transition to the montopoli fauna. this significant faunal turnover, known as the elephant-equus evenr (lindsay et al. 1980), was due to a global climatic change that gave rise to a generalised cooling and increasing aridity, which occurred at the time of the gauss-metuyxma transition. comparative diffrattometric analyses of the sediments covering the eostyloceros antler and two "pseudod,ama" lyra jaw fragments from the montopoli assemblage were carried out with the intent of knowing more about the possible source of the eostyloceros. the two matrices significantly differ, suggesting a different provenance. the silty-sandy sediment on the eostyloceros antler is characterised by the dominance of calcite and quartz and by a low content of feldspars, while that on lsoth psewdodama mandibles is low in calcite but high in feldspars (n. cipriani pers. com.). moreover, rhe eostyloceros antler is slighlty polished indicaring a certain degree of transport, while the montopoli fossils are generally unabraded, and show evidence of pre-burial weathering and of root etching (mazza pers. com.). these accumulated evidence suggests that the antler of eose,loceros might be extraneous to the montopoli mammal assemblage. tw-o alternatives could explain its occurrence: 1) lower pliocene continental or shallow marine deposits are more widespread than previously supposed, as suggested by recent investigations in nearby areas (e.g. valdelsa and val di pesa basins, benvenuti & degli innocenti 2001; rook et al. 2001; tangocci 2oo1), and therefore the eostyloceros remain could have been reworked from those depo.its: 2) alternatively, the antler might have not been recovered at the montopoli site, but somewhere else in the surroundings, wherein lower pliocene continental deposits outcrop. systematic palaeontology order artiodactyla owen, 1848 suborder ruminantia scopoli, 1777 infraorder pecora linneaus, 1758 family cervidae gray, 1821 subfamily muntiacinae pocock, 1 923 genus eostyloceros zdansky, 1925 fig. 3 eos1'loceros cf. pidoplitschkoi lgf8218rfrom l-or.er valdirr, no. a frontal vien, b rncdial vieq c larcral vien, d ventral vierv of burr. eostyloceros cf. pidoplitschkoi korotkevitsch, 1964 fig. 3 material. a rieht shed antler igf8218v (fig.3, tab. 1) description. the antler is two-pointed, with the bifurcation set close to the burr (fig.3; tab. 1); thc anterior tine is comparatively smaller than the posrenor one, which is broken at abollt 10 cm fron'r its base. the surface is streaked by r.vell devcloped ridges and furrows. close to the bifurcation, on the medi;rl side, the surface is intensely pearled. the cross section of the anterior branch is trianeular-shaped (fig. 3b). the posterior branch h:rs a triangular cross section too, which turns oval in the distal part (fig. 3b). in frontal vierv the bearn is straight. the anterior branch forms an angle of 45' n'ith the postcrior branch. eostyloceros cf. pidoplitschkoi from lower valdarno 577 discussion and comparisons. the antler has a typical muntiacine morphology. the short length, the dichotomous pattern with the splitting close to the burr and the disproportion between the branches are typical eostyloceros traits (fig. 3). \fhile muntiacinae cervids are very common in the european mammal faunas during middle and late miocene (see for example azanza 2000), they decline considerably from the early pliocene, when their geographic distribution shrinks to include only eastern europe and asia, with only one fairly well documented western european genus, paracet"uulus, which characterizes the latest turolianrus cinian f aunas (vis lobokova 1 99 2 ; d ong 19 9 6;'\bbazzr 2001; abbazzi k azanza2a0a). the italian antler has been comoared with muntiacine from latest miocene-early pl'ocene european and asiatic sites: eostyloceros pidoplitschleoi (from lower pliocene deposits of kuchurgan river at sites novopetrovka, yurievka, voinich, and from trifonesti type 1ocality, south-eastern moldova, korotkevitsch 197a), eostyloceros blainvillei (latest miocene of china and moneolia, korotkevitsch 1920; vislobokova 1983, 1990), and muntiacus pliocaenicus (eariy pliocene of ukraine, ko| ' i r^-^.rotkevltscn iy,/uì. the comparison of the leneth of the first segment (lateral distance measured from the bifurc.rtion to the burr, fig. 3) with the antero-posterior diameter of the antler base (measurèd just above the burr, fig. 3) is shown in fig. 4a. in this diagram the points representing e. blaincillei form a quite separate cluster from those of e. pidoplitschkoi, thus xrtesring to a longer 1" segment. muntiacus pliocaenicus is also separated from eose,loceros, being significantly smaller-sized and having a higher bifurcation, like in recent muntiacus. the cloud of points relative to e. pidoplitschleoi frorn ukraine evidences that the 1" segment and the dimensions of antler base are inversely correlated. the lower position of the bifurcation expands the antero-posterior diameter. the lowering of bifurcation is an ontogenetic character, older individuals having a smaller value in the l" segment, as already pointed by korotkevitsch (1920). this feature is shared by other cervid genera as well. focusing on mere morphological differences, the index relating the 1" segment to the latero-medial diameter of antler base (1" segment/lmdbeam of antler basex 1oo, fig. 3) is plotted against the index of flattening of antler base (apdbeam/lmdbeamx100, fig. 3) in fig. 4b. the different morphology ol e, blaìnvillei and muntiacus pliocaenicws due to the higher position of bifurcation and more circular antler base is confirmed. the antler of montopoli differs from that of e, pidoplitschkoi in the sub-circular cross-section of the antler base. on the b:rsis of the accumulated evidence, we can confidentlv refer the itelian antler to the genus eosty/oceros,whlle for its specific attribution we calltiously refer to it "rs to e. c[. oidoolitschleoi. 5 cnr w^3 qrr i \pd-;ili i lnrdbeam l t3xon e. cf. pídoplitschkoi e. pidoplítschkoí e. pidoplitschkoi e. pidoplitschkoi e. pidoplíîschkoi e. pidoplitschkoi e. pidoplitschkoi e. pidopìitschkoí e. pidoplixchkoí e. pidoplìtschkoi e. pidoplítschkoí e. pidoplitschkoi e. pidoplítschkoi e. pidoplùschkoi e. pídoplít.sc'hbi e. blainvillei e. blqinvilleí e. hlqinvillei e. blainvílleí e. b/ainvillei e. cî. pidop/itschkoi m. pliocaenicu.s m. pliocaenicus m. pliocaenicus localilv tuscany trifbnesti yurievka (kuchurgan) voinich (kuchurgan) novopetrovka (kuchurgan) :: clirra apdbeam !4qdbeam i segm. angle 37.0 36.6 32.4 55' 43.5 24.2 39.8 10" j7.0 21.2 2q.0 10.0 20.5 2q.5 3s.6 2-.2 2?.t 30.0 24.0 ll.0 i0.8 2ó.5 .ì5.0 j7.2 24.5 i1.0 200 21.00 25.5 ii.5 12.5 17.0 l).j 21.0 10.5 t().4 21.5 i1.0 2c.b .ì-.5 4l.o -60 o 4 i .0 59.0 __ 4 t.5 17.0 51.5 37 -51.0 40.5 33.0 29.0 90" 21 .5 r8.5 16.0 36.0 20.x iq.6 .r8.0 apdìurr lucesti kuchurgan 51.6 31 .0 43.0 31.3 31.4 25.0 40.0 3 l .5 29.8 26.7 33.5 33.7 38.3 32.0 35.0 30.5 -26.1 29.0 19.6 30.0 27.0 41.0 29.5 36.3 26.2 3 r.0 24.0 52.0 53.5 46.0 57.0 55.5 17.4 ,+8.8 3,{.0 28.0 r 8.5 16.0 20.8 19.6 578 eostyloceros pidoplitscbkoi korotkevitsch is the only species of the genus known from the early pliocene of est europe. the definition of this deer is based on the morphology of antlers, pedicles and frontal bones. the fig. 4 a) scatter diagram of first segrnent against antero-posterior diameter of antler base (apdbeam) of e. pidoplitschboi from trifonesti, yurievka, voinich and novopetrovka, e. blainviliei írom cina, e. cf. pidoplitschboi from lucesti and "montopoli", and m. pliocaenicus from kuciurgan river. b) scatter diagram of first segment/latero-medial diameter of entler base':100 against antero-posterior diameter of antler base/ latero-medial diameter of antler base'tlo0 oí e. pidoplttschèol from trifonesti, yurier.ka, voinich and nor-opetror.ka, e. blaint,illei from cina, e. cf. pidoplitscbkoi îrom lucesti and "montopoli", and m. pliocaenicus îrom kuchurgan rìr'er. measurements of antlers of eostyloceros and muntiacus species. all data are from koroîkevitsch (1970). legend: apdburr : antero-posterior diameter of burr; lmdburr : latero-medial diameter of burr; .rpdbe.rm : rnlero-po\terior diameter of beam just above burr; lmdbeam : latero-medial diameter of beanr just .tbove burr: i 'ce,nì. : fìr.t segment length: rneleangle of bifurcation. type specimen (nr 5634, stored in the national museum of natural history, kiev, fig. 5a), was discovered in the fossiliferous outcrop near trifonesti, (eastern moldova) from the "pliocene bessarabian sands" and the main part of the material ascribed to this species was found in the deposits of the kuchurgan river va1ley (fig. 5b). the other species of eostyloceros, e. blain'uillei zdansky, e. triangularis zdansky, e, propria abdrahmanova and e. maci vislobokova are described'from the latest miocene-early pliocene of china, mongolia and kazahstan, and from the early pliocene of baikal region of russia (zdansky 1925; abdrahmanova 1974; vislobokova 1985, 1990). conclusion the occurrence of eostyloceros in itaiy represents a significant westward expansion of the known pliocene geographic distribution of this genus. unfortunately the unknown stratigraphical context prevents any further conclusions about the occurrence of this genus. the accompanying fauna at kuchurgan, where korotkevitsch (1964) first discovered the finds of e. pidoplisch,éol, is different from that of montopoli, and contains species as muntiacus pliocaenicws korotkevitsch, pli o c ervws k utchwrganicz.r korotkevitsch, parac era ulus australis (de serres), procapreolus cf. cusanus (croizet et jobert) which indicate a ruscinian age (see also korotkevitsch 1965). therefore, also with the support of the sedimentological evidence presented here, the antler fragment from montopoli can be reasonably interpreted as an element stratigraphically incongruous to the rest of the montopoli local fauna, possibly reworked from nearby lower pliocene deposits, or the result of inappropriate recovery. nonetheless, the presence of an eostyloceros representative in the tuscan paleontological collections urges l. abbazzi k r. croitor x tilfbnesti a yilriclla tr voinrch o no\odotro\ka. o e.hlanoillei -ll. pliocaetúcil\ a montoooli o o no\opclrorla u vornrcn a yurie'ka x tilbnesti a moiltopoìi 4t plíoeenieus i t0 130 150 apdbeanr ìnrdbeanrx ì llll b @" w % c bo i ,({ to the adoption of multidisciplinary investigations in the reconstruction of the paleoenvironmental evolution of the neogene of tuscany. acknouleclgements. the authors are deeply obliged to m. benvenuti (earth sciences departmenr, florence) for his suggestions and for the constructive criticism of rhe manuscripr; to n. cipriani (hearth science departnrent, florence) who carried out the diffrattometric analysis; to p. mazza (earth sciences department, florence) eos4tloceros cf. pidoplitschkoi from lott,er valdarno 579 fig.5 a) eostyloceros pidoplixchkoi from trifonesti (moldavia) holotype, right antler on frontal bone n. 5635 (paleontological department of the national museum of natural history kiev), adapted from korotkevitsch (1964), a) frontal view, b) medial view; b) iost1, lo cero s p idop lìts ch ko i f rom kuchurgan (ukraine), left antler on frontal bone n. 1549 (paleontological department of the national museum of natural history kiev), a) medial view, b) anterior vieq c) cross section of anrerior branch, d) cross section of posterjor branch. for his contribution to the identification of the taphonomic characters of montopoli fossils, and for improving english version of text, to l. rook and d. torre (earth sciences department, florence) for critical reading of text. many thanks to t. krakhmalnaya and to e. cìoppi for access to the fossil collections of rhe national museum of natural history of kiev and of the geo-paleontological section of the natural history museum of florence, respectìvely. photos bv f. cozzini (natural history museum of florence ). research supporred by the a.g. side grant of the linnaean society of london and by miur granrs 1d. torret. references abbazzi l. (2001) cervidae and moschidae (mammalia, artiodactyla) from the baccinello v-3 assembìage (late miocene, late turolian, grosseto, ital,v). rir,. it. paleont. strat., 707 : 1a7 123, milano. abbazzil. &. azanza b. (2000) occurrence of paraceruulus cf . australis de serres, 1832 in late miocene (messinian) italian localities. i international congress "the holarctic ungulares of the pliocene and pleistocene", avignone, 19-22 september 2001, abstract book, p. 2. abdrahmanoval.t. (197a) fossil ruminanrs of karabastuz (in russian). materialy po istorii fauny i flory kazahstana, 6: 83-89, alma-ata. azanza b. (2000) los cervidae (artiodactyla, mammalia) del mioceno de 1as cuencas del duero, tajo, calatayud-teruel y levante. mem. museo paleont. univ. zaragoza, 8, 376 pp.,zaragoza,. azzaroli a. (197a) vilìafranchian correlations based on large mammals. giorn. geol.,35: 111-131, bologna. azzaroli a. (1977) the villafranchian stage in italy and the pliopìeistocene boundary. giorn. geologia, 49 61-79,bologna. ltzzarolr a. (1992) the cervid genus psewdodama n. g. in the villaf ranchian of tus cany. pala e o nto gr. i tal., 7 9 : 1 4 1, p isa. benvenuti m., dominici s. & rook l. (1995) inquadramento stratigrafico-deposizionale delle faune a mammiferi viliafranchiane (unità faunistiche tiversa e montopoli) del valdarno inferiore nella zona a sud dell'arno ltoscana). il quaternario 8: ,f57-.{64, roma benvenuti m. er degli innocenti d. (2001) the pliocene deposits in the central-eastern valdelsa basin (florence, italy), revised through facies analysis and unconformitybounded stratigraphic units. rl.rr. it. paleont. strat., 1a7: 265-286, milano. de giuli c. & heintz e. (97aa) gazella borbonica (bovidae, artiodactyla, mammalia) nouvel element de la faune vi1lafranchienne de montopoli, valdarno inferieus, pisa, italia. atti soc. tosc. sc. nat. mem., a,81:227-237,pisa. de giuli c. 8r heintz 8., (1974b) croizetoceros ramosus (cervidae, artiodactyla, mammalia) de montopoli, nouvel element de la faune villafranchienne d'italie. atti soc. tosc. sc. nat. mem., a,81 241-251, pisa. de giuli c., ficcarelli g.,mazza p & torre d. (1983) confronto tra success;oni marine e continentali del pliocene e pleistocene inferiore in italia e nell'area mediterranea. boll. soc. paleont. 1t..22: 323-328. modena. 580 l. abbazzi k r. croitor dong\xl (1996) les cervidae (artiodactyla) rusciniens (pliocene) de languedoc et du roussillon (france). bwll. mus. nat. hist. nat.,18, c (1): 133-163, paris. forsyth-major c.l (1877) considerazioni sulla fauna dei mammiferi pliocenici e post-pliocenici della toscaira. atti soc. tosc. sc. nat., 1.:7-40, pisa. forsyth-major c.i. (1885) on the mammalian fauna of the val d'arno. quat. journ. geol. soc. abstact, 8 pp, london. gliozzi e., abbazzi l., argenti p., azzaroh a., caloi l., capasso barbato l., di stefano g., esu d., ficcarelli g., girotti o., kotsakis d., masini f.,mazzap,mezzabotta c., palombo m. r., petronio c., rook l., sala s., sardella r., zanalda e. & torre d. (1997) biochronology of selected mammals, molluscs and ostracods from the middle pliocene to the late pleistocene in italy. the state of the art. rio. it. paleont. strat..703:369-388. milano. korotkevitsch e. l. (1964) a new species of fossil mwntiacus from the pliocene deposits of the south of ussr. dopooidi akademii naule ulerains,èol àsà, 6: 807-809, kiev. (in ukrainean). korotkevitsch e. l. (1965) on deer from pliocene of the kuchurgan valley and their paieogeographical significance. prirodnaya obstanopka i fauny proshlogo , v. 2: 25-43, kiev lin russian). korotkevitsch e. l. (197q late neogene deer from the north black sea area. nauhoz,a dwmlea, 1-175, kiev. lin russian). lindsay e.h., opdyke n.d. & johnson n.m. (1980) pìiocene dispersal of the horse equus and late cenozoic mammalian dispersal events. nature, 287 :135-138, london. rook l., lbbazzr l., agustij. & tangocci f. (2001) a possible early pliocene sn.rali fauna with a gerbil in val d'elsa (tuscany, kaly) .2"à eeden workshop "late miocene to early pliocene environments and ecosystems, sabadell, 15-17 november 2001. abstract book: 64-65. tangocci f. (2001) stratigrafia dei depositi mio-pliocenici dell'area di poggibonsi (bacino deil'elsa). unpublished thesis, 98 pp, università degli studi di firenze. vislobokova i.a. (1983) the fossil deer of mongolia (in russian). the soviet-mon golian palaeontolo gic al exp e dition. tian s. joint soviet-mongolian paleont. expedition,23: 78 pp, moskva. vislobokova i.a. (1985) a new species oí eostyloceros from ol'khon island (lake baikal). paleont. tourn.4:137-139, moskva. visiobokova i.a. ( 1 990) fossil deer of eurasi a. transactìons of the paleontological institute, 24a, 2a6 pp, moskva. vislobokova i.a. (1992) neogene deer in eurasia. paleont. í er., ol., 24-25, pp. 1 49 -1 54, sabadell. vislobokova i.a., erbaeva m.a. 8{ sotnikova m.v (1993) the early villafranchian stage in the development of the mammalian fauna of northern eurasia. strat. geol. corr., v. 1 (5) : 555-564. 'weithofer k.a. 11893) proboscidiani fossili del valdarno in toscana. mem. carta geol. it. v. 4(2) 1-152, roma. zdansky o. (1925) fossile hirsche chinas. paleontol. sin.,c, 2\3): l-94. peking. rivista italiana di paleontologia e stratigrafia volume luz numero 1 pagrne 131 134 aprile 1996 note brew an exceptional reptile find in the norian (late triassic) lagerstà.tte of endenna (zogno, bergamo, italy) andrea tintori, silvio renesto, cristina lombardo, gianluca manarolla. matteo manenti & massimiliano vendico k^,-",,^.).. p--r;li. norian (late triassic), calcare di zorztno, lombardy q.tronhern italy), new find. riassunta" durante un sopralluogo nella località di endenna (zogno, bg) il 5.10.95 è stato rinvenuto un frammento di un grande rettile parzialmente danneggiato da scavatori abusivi. dopo il salvataggio del blocco già isolato, si è proweduto alla ricerca e al recupero della rimanente pane dell'esemplare. durante questi lavori sono anche stati sorpresi alcuni scavatori abusivi che sono stati segnalati alla competente soprintendenza. il rettile si presenta completamente inglobato in matrice calcarea molto tenace e si prefigura parzialmente arrotolato e con il cranio disarticolato rispetto al resto del corpo. la hnghezza totale supera i 4 m e 1o stato di conservazione è da ritenersi ottimo. sulla base di osservazioni preliminari si può ipotizzare 1a sua àppar-tenenza all'ordine thalattosauria. si tratterebbe quindi del pitì grande rettile acquatico triassico, con i'esclusione degli ittiosauri, finora conosciuto. abstract. durìng a survey àt the fossiliferous site of endenna (zogno-bergamo) of norian age, a huge reptile has been found at october the 5th, 7995. the recovery of the whole specimen took place at the beginning of november. four main blocks, plus minor fragments, are now in the laboratory of the department of eanh sciences of the milano university waiting for the long prepararion. the specimen is somewhat more than four meters long, the head being disarticulated in front of the body. a preliminary survey suggesrs the reptile may be ascribed to thalattosauria. thus, apart from the ichthyosaurids, it would be the largest marine reptile of the triassic. the find. the fossil-site of endenna (zogno, bergamo, italy) is one of the most important sites in the calcare di 7nrzíno (zorzino limesrone) (tintori et a1., 1985). vith the permit of the soprintendenza archeologica della lombardia and under the direction of the senior author, field-works were carried our from 1978 unttl 1983, when works v/ere stopped for logistic causes and started in the nearby zogno2 locality. nonetheless, regular surveys are necessary because private collectors frequently damage the site, anything but exhausted. a couple of them have been catched during the recent works and reported to the competent aurhority. hundreds of fishes have been collected during six years of digging (beltan & tintori, 1980; tintori, 1980, 198t, 1983, !99q, 199l, 1992, 1995; tintori & renesro, 1983; tintori & sassi, t992), together wirh some reptiles (padian, 1981; pinna, 1,979,1,980, 1984;pinna 6c nosotti, 1989; renesto 1.984, 1992, 1,993,1994a,b, c, t995a,b; renesto & tintori, 1995) and several, mostly undescribed, invertebrates (basso & tintori, 1994). on the last 5th of october, the senior author was accompanying dr. w. bausch (erlangen, germany) to a survey for geochemical sampling in endenna. he found a block, measuring about 80x40 cm (block n.3) which showed remains o{ a very large organism, possibly a reptile. coming from level 11 (tintori et al., 1985), the block had been removed and damaged by unauthorized diggers. in this case, splitting was difficult because the big fossil has disturbed the regular lamination and has caused a locally stíonger cementation. the relative posi tion between the block and the resr of the layer was difficult to state, but the possibilities that at leasr part of the fossil rvas stiil in place were good. the problem was that the find was close to the end of the fossillevel outcrop, so we wondered how much of the fossil had been already destroyed by natural erosion. the block was taken to milano only a few day after and, by the end of october, we organized the recover of the rest of the fossil. it rook rwo davs of work (the 3oth of october and the 2nd of norre-t".) for six people, but we had the greatest expected success: the remain of the reptile was still in place and could be completely removed. closely observing the front of layer 11, we could see that the reptile was in its lower parr. the upper part 'was then removed; the underlying thin, shaly level made the rest of the fossiiiferous level easy to take off. taking advantage oí a lateral small karstic canai (kc in fig. 1) and natural fractures the caudal region (fig. 2) milano, via mangiagalli 34,2a1.33 milano, italy.dipanimento di scienze della terra dell' università degli studi di 132 a. tintori, s. renesto, c. lombardo, g. manarolla, m. manenti & m. vendico ll{{ has been evidenced on the upper surface of a slab (fig.1; block +) measuring about 140x105 cm. the rest of the fossil was more problematic to recover: the fracture between blocks 2 (60x50 cm) and 1, (75x30 cm) was irregular, leaving a couple of small fragments in between. on the whole, the reptile has suffered very little damage. the total weight of the blocks is about 250 kg; they have been trasported for about 1 km on hand-barrow up to the car. shape and dimension of the skeleton can be guessed trough the sediment covering the fossil. a part of the posterior leg (knee arriculation) has been already prepared, to help in a preliminary classification. the observation of the fossil suggesrs that the reprile has long trunk and tail, relatively short legs and considerably long skull. its total length should be more than 4 m. the body, lying on its belly, forms an almost complete circle (fig. 1). all these features indicate the skeleton does not belong to an ichthyosaur; in fact, femur, tibia and fibula, showing sharp epiphyses and narrow diaphyses, are very different from those of ichtyosaurs. on the basis of the same morphologic features we can exclude the fossil belongs to the nothosaurs, which have shorrer tail, longer neck and smaller skull. leaving apan rhe ichthyosaurs, to nothosaurs belonged the largest known acquatic reptiles in the triassic, before this find. a specimen of paranothosaurus atnsleri from the ladinian of monte san giorgio (canton ticino, switzerland) reaches 3.80 m and it was the largest one, as far as we know. triassic phytosaurs seem to show more similarities with the new find. they are crocodilelike archosaurs with nostrils back near the eyes. phytosaur remains are known from upper triassic in germany, in usa and in india and fragments are found also in several sires of the middle and far east. phytosaurs are thought ro have dwelled in continental environments, such as lakes and alluvial plains, and to have very rarely visited marine y/aters. this is clearly in contrasr with the fossilization environment of the caicare di zorzíno, which is an intraplatform basin with only small islands nearby (|adoul et a1., 1994; renesto & tintori, 1995; tintori, 1995). nonetheless, an isolated skull of a phytosaur had been already found in endenna, but it is probably a fragmenr of a floating skeleton coming from dry land (renesto, 1995b). the new specimen cannot have been transported: its skeleton is complete and perfectly articulated, apart from the skull. moreover, though the relative dimensions of the skull are comparable to those of phytosaurs, other features have different proportions: tail, for example, is too long. only a last group of reptiles can be compared with the new, large specimen: thalattosaurs. these are possibly archosauromorph reptiles characterized by relatively short legs and long, laterally compressed tail. thalattosaur remains are known from the middle triassic of the formazione di besano (grenzbitumenzone for the swiss authors) and of california and from the upper triassic of british columbia. aiso endenna have yielded two specimens of endennasaurus (renesto, 1984, 1992). thalattosaurs lived in strictly marine environments (fossils are often associated with ichtyosaurs and ammonites) and they likely had a semi-durophagous diet. proportions of the new reprile are well comparable \*" \ fig. 1 schematic draw of the four blocks with the probable reptile outline (dotted area, in black the already exposed posterior leg) as seen from above. kc: karstic canal; s: slumping like deformation of the underlying laminae beside the specimen belly; striped area: empry region. we expect the preparation will rake a very long time: on the lower side, apparently the easiest, at least 18 months are necessary to expose the bones. flowever, it will be convenient to expose the fossil on both sides, at least in important parts, because it is fortunately very slightly compressed; even the head is tridimensionally preserved. 3d conservation is rarely found in our norian locaiities and only regards parts of large specimens with strong structures (the skull of mystriosuchus and the mouth region of large sargodon tomicu). normally, fossils are totally flattened and bones are more or less broken by lithostatic pressure. taphonomic and systematic remarks. the impact of the reptile body on the bottom compressed the underlying laminae, inducing also a uniderectional slumping-like deformation for at least 1 m from the body itself (s in fig" 1); this fa*, together with the absence of further compression of the skeleton, suggests that diagenesis in this level has been remarkably early. laminae in fact, show folds which keep their individuality. an accurare sedimentological-geochemical study on those laminae is trying ro undersrand the first diagenetic stages in an anoxic environmenr with predominant carbonatic sedimentaticn. exceptional repttle find tn endenna fig. 2 the slab (block 4 in fig. 1) with the tail. scale-bar: 15 cm to those of thalattosaurs and their life environment is in agreement with the paleoenvironmental reconstruction of the calcare di zorzrno. in conclusion, for the classification of the new reptile specimen, three hypotheses can be put forward, with decreasing likelihood: it is a new genus of thalattosaur, it is a phytosaur or it is a totally new organism. in any case, this new find will through new light on rhe life environment of the zorzino fauna: before this, no very large reptile had been found. this fact was explained with the presence of geographical and ecological barriers for both marine and terrestrial organisms, which favoured the rising of an endemic reptilian fauna made of mainly small species. now we must reconsider that, or suppose that those infraplatform basins were so rich in iife that aiso large organisms could develop and be sustained. acknouledgements, thank are ciue to the soprintendenza archeologica della lombardia which gave us the special permission ro recover the specimen. the picture is by g.chiodi. the hand-barrow shafts were made of the mast of at brother-in-law old wind-surf: to carlo our sìncere thanks. re,ferences basso d. & tintori a.(1994) new triassic isopod crustaceans from northern italy. palaeontology v. j7, n. 4, pp 801-810, london. beltan l. & tintori a.(1980) the genus sauricbtlrys (pisces, actinopterygii) during the gondwana period. proc. fifth int. gondvtana syrnp., pp. 53-59, rotterdam. jadoul f. masetti d. cirilli s. berra f. claps m. & s. frisia \1994) norian rhaetian srratigraphy and paleogeographic evolution of the lombardy basin (bergamasc a1p$. v. of 38 pp., ii ias regional meeting, ischta, italy. padian k.(1981) notes on a new specimen o{ pterosaur (reptilia, pterosauria) from the norian (upp.. triassic) of endenna, ita|y. rio. mus. cio. sc. nat. bergamo, v. 2, pp. 1,19-l27, bergamo. pnna g.(1,979) il cranio di un giovane placochelide (psepho derma alpinum meyer, 1858) deì norico di endenna (bergamo). atti soc. it. sc. nat., mus. st. nat. milano, v. 120 n. 3-4, pp. 195-2a2, milano pinna g. (1980) drepanosaurus unguicaudatur, nuovo genere e nuova specie di lepidosauro del trias alpino. atti soc. it. sc. nat., mus. st. nat. milano, v. 12l, n.3, pp. 181192, milano. 134 a. tintori, s. renesto, c. lombardo, g. manarolla, m. manenti & m. vendico pinna g. (1984) osteologia d). drepanosaurus unguicaudatus, lepidosauro triassico del sottordine lacertilia. mem. soc. it. sc. nat., mus. st. nat. milano, v. 24, pp. 7-28, miiano. pinna g. ec nosotti s. (1989) anatomia, morfologia funzior'ùe e paleoecologia del rettile placodonte psepboderma alpinum meyer, 1858. mem. soc. it. sc. nat., mus. st. nat. milano, v. 25, pp. 17-49, mllano. renesto s. (1984) a new lepidosaur (reptilia) from the norian beds of the bergamo prealps. rip. it. paleont. strat., v. 90, n. 2, pp. 156-167, milano. renesto s. (1992) the anatomy and relationships oí endennasaurus acutirostris (reptilia, neodiapsida), from the norian (late triassic of l-ombardy) . riv. it. paleont. strat. v.97, (199l), n. 3-4, pp. 409-430, milano. renesto s. (1993) an isolated sternum oî eudirnorphodon (reptilia, pterosauria) from the norian (late triassic) of the bergamo frealps (i-ombardn northern italy). riv. it. paleont. strat., v.99, n. 3, pp. 415-422, milano. renesto s. (199a{ megalancosaurus, a possibly arboreal archosauromorph (reptilia) from the upper triassic of northern itaiy. journ. wrt. paleont., v. 14, n. 1, pp. 3852, lawrence. renesto s. (1994b) the shoulder girdle and anterior limb of drepanosaurus unguicaudatus (reptilia neodiapsida) from the upper triassic (norian) of northern italy. zool. fourn. linn. soc., v. 111, n. 3, pp. 247-264, london. renesto s. (199ac) a new prolacertiform reptile from the late triassic of northern ltaly. rio. it. paleont. strat., v. 100, n. 2, pp. 285-306, milano. renesto s. (1995a) a sphenodontid from the norian (late triassic) of l-ombardy (northern italy): a preliminary note. modern geologl, v. 2q, pp. 149-158, malaysia. renesto s. (1995b) ecology and taphonomy of the reptiles from the calcare di zorzino (norian, late triassic) from northern italy. ext. abstract ii int. symposium on i i th o grap h i c l ift'?. e s ton e s, pp. l27 1.29, m adrid. renesto s. ec tintori a. (1995) functional morphology and mode of life of the late triassic placodont psepbodernta alpinum, meyer from the calcare dí zorzino (i-ombardy, n. italy). rio. it. paleont. strat., v. 101, n. 1, pp. 37-48, milano. tintori a. (1980) teeth of the selachian genus pseudodalarlas sykes, 1971, from the norian (upper triassic) of lombardy. ria. lt. paleont. strat., v. 86. n. i, pp. 19-30, milano. tintori a. (1980 two new pycnodonts (pisces, actinopterygii) from the upper triassic of lombardy (l'.1. italy). riv. it. paleont. strat., v. 86, n. 4, pp. 795-824, milano. tintori a. (1983) hypsisomatic semionotidae (pisces, actinopterygii) from the upper triassic of lombardy (n. kaly). ri,l. it. paleont. strat., v.88, n. 3, pp. 417-442, milano. tintori a. (1990) the vertebral column of the triassic fish saurichtlrys and its stratigraphical significance. ria it. paleont. strat., v. 96, n. 7, pp. 93-102, milano. tintori a. (1991) durophagous vertebrates from the late norian of southern .nips. abstract "7th international symposium on the study of earlt wrtebrates", parc de miguasha, quebec canada, p. 44. tintori a. (1992) fish taphonomy and triassic anoxic basins from the alps: a case history. riv. it. paleont strat., v.97 (1991), n. 3-4, pp. 393-408, milano. tintori (1995) biomechanical fragmentation in shell-beds from the late triassic of the lombardian basin (northern italy). preliminary reporf. ri,o. it. paleont. strat., v. 101, n. 3, pp. 371-380, milano. tintori a. & renesto s. (1983) the macrosemiidae (pisces actinopterygii) from the upper triassic of lombardy (n. italy). rio. it. paleont. strat., v.89, n.2, pp.2a9222, mtlano. tintori a. muscio g. & nardon s. (1985) the triassic fossil fishes iocalities in italy. ritl. it. paleont. strat., v.97, n. 1, pp. 127-210, milano. tintori a. & sassi d. (1992) tboracopterzs bronn (osteichthyes: actinopterygii): a gliding fish from the upper triassic of europe. fourn. wrt. paleont., v. 1,2, n. 3, pp. 265-283, lawrence. received january 23, 1996; accepted february 7, 1996 martinez 531..534 ����� ���� � ���� �� � ��� ��������� �� ������� �� � ����� ��� ���� �� ��� �� � ������� ��� ����� ���������� ���� �� � � ���� ���� ���� � � ��������� ��� �� � � ����� ��������� ������ ��� ���� ��� � ���� ���� � ����������� ����� ����� ����� !"�#�� �!$�% � ������% �#� !$�"�� ���# ��&�� � �!# !' � (!�#�"!�� '�!$ (� �!# � ��)�! �� �� &��)�!*��� ��"��+�� ! � ���� ���������� �*,,�� � (� � +��!#,� ! 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(199a) included in the family syringothyrididae fredericks, 1926 all the punctate, strophic, biconvex sprriferids with ornamentation of simple ribs, smooth fold and sulcus, and high to very high ventral interarea with perideltidial areas. based on the presence or the absence of the syrinx and the ventral median septum, they distinguished three subfamilies within the syringothyrididae. due to the supposed occurrence of a syrinx they included subansiria sahni & srivastava, 1956 in the sublamily syringothyridinae. however as suggested by angioiini et ai. (tlvl) subansiria lacks a true syrinx and thus must be included ir-r the subfamil,v permasyrinxinae \íaterhouse, 1986 together with cyrtelìa fredericks, 1924 and punctocyrtella plodowski, 1968, previously considered synonyms by carter et al. (1994) . as discussed in angiolini et al. (1997), cyrtella and punctocyrtella could be regarded as distinct and separate genera and a new genus pachycyrtella is established in order to includ perigondawanan species previously placed rn cyrtella and characterized by a high and apsacline ventral interarea and thick umbo. genus pachycyrtella n. gen. type species: p omanensìs n. sp. etymology. gcnus named for its thick, heavy umbo (from the ancient greek no,xug) and rs cyrtella like general shape. diagnosis. thick shelled, large species s'ith high :ps.rcìine to orthocline ventraì jnterarea. ventral sulcus usually wide and shallor., dorsal fastigium deep11. furrowed. ornamentation of coarse, adichotomous costae. interior of ventral vaìve q,ith variablv developed umbonal callus, thick delthyrial plate, lone dental platcs with ventral adminicula surrounding the posterior part o{ thc muscìe frcld. interior of dorsal valve with broad sessile cardinal process; socket plates and crural b;rses fused to the cardinal proccss bv a thrck callus. discussion. the new genus pacbycyrtella is erected in order to include species similar to cyrtella but characterized by a high rnd apsacline to orthocline ventral interarea, by r thick umbonal callus, and deep dorsal median furrow. in fact, cyrtella differs from pachycyrtella by its catacline ventral interarea, smaller callus, more transverse shape and shallower furrow on the dorsal fold. furthermore, cyrtella has a characteristic internal secrion with narrow post-delthyrial lcentral r cavity rnd much wider umbonal chambers, very different frorn that of pachycyrtella, characterised by a large central chamber. pacbycyrtella n. gen. differs from permasyrinx \íaterhouse, 1983 by its apsacline-orthocline interarea and the occurrence of a medial sulcus along the fold; from pyramidtbyris hu, 1983, pseudosyringotbyris fredericks, 1916 and tuotalania hu, 1983 by the orientltion of the ventral inte rarea and the dorsal fastigium which in the latter genera is not sulcate. the specimens determined as s. nagmdlgensis lry termier et ai. (t0l+) from the sakmarian of central afghanistan are assigned to rhe new gents pachycyrtel/a because of their apsacline interareas and thick umbones. the same holds true for those determined as cyrtella nagmargensis by archbold & gaetani (1991) from the late sakmarian chumik formation member a of zanskar (india). cyrtella subparallela \flaterhouse 1982 from the early sakmarian elvinia fm. of se bowen basin (e australia) is probably related to pachycyrtella, having an apsacline ventral interarea and a heavl unrbonal thickening. on the contrary, examination of replicas of the type material of cyrtella nagmargensis (bion, 1928) from kashmir kindly shown to me by dr. g. plodowski of senckenberg museum (frankfurt) has pointed out that l. angiolint plate 1 ' all x l. c\lepr s hcn 'pecified r fig. 1 pachycyrte lla omanenis n. gen. n. sp. holotype, dorsal view of complete specimen mpum 8125. fig. 2 pachycyrtella omanenis n. gen. n. sp. holotl'pe, ventral view of complcte specimen mpum 8425. fig. 3 pachycyrtella omattenis n. qen. n. sp. par:type, lateral view of completc specimen mpum 8426. f'ig. a pachycyrtella omanenis n. gen. n. sp. paratype, ventral view of cornplete specimen mpum 8426, bearing a ce mented spccimen oi etherilosia sp. fig. 5 pachycyrtella etmanenis n. gen. n. sp. paratype , ventral vienof complete specimen mpum 8428, bearing a cemented specimen ol etberilosia sp. fig. 6 pacbycyrtella omanenis n. sen. n. sp. p:r:r1-pc, dorsal r-ien of a complete specimen mpum 8129. fig. / pacbycyrtel/a omanenis n. gen. n. sp. paratype, r.entral valr-e interior, specimen mpum 8131. fig. 8 pachyq,rteìla omanettis n. gen. n. sp. paratvpe, dorsal view of complete specimen of a jur-enile mpum 8427. fig. 9 pachycyrtella omanetris n. gen. n. sp. par.rtl-pe, dorsal valve interior, specimen mpum/9/5 fig. 10 pachycyrtella omanenìs n. sen. n. sp. secondary layer showing fìne punct:rtìon, x 52, at a.77 cn from the umbo (mgl 63121). fig. 11 pdcbycyrtella omanenis n. lien. n. sp. section shorving dental plates and delth.rirl pl.rte x,l rr 1.29 cm from the umbo (mgl 63121). pl. 1 nerr' brachíopod from e,trh, i)ertttian of onan t27 t28 l. angiolint the original species nagmargensis belong to rhe genus cyrtella. in fact, the kashmir specimens are characterised by a catacline interarea, transverse shape and internal characters consistent with those ol cyrtella leulibiana (fredericks, 1916), type species ol cyrtella. furthermore, cyrte/la nagmargensis from kashmir is easily distinguishable from the genus punctocyrtella by the orientation and height of its ventral interarea, the ornamentation and the internal characters of the ventral valve. the tibetan c. nagmargensls from the late sakmarian qudi formation of rutog duoma (hu, 1983) belongs to the genus cyrtella, based on the orientation of the interarea and the internal sections. the \x/estern australian c. awstralis thomas, 1971 from the lyons group and callytharra formation is to be retained in the genus cyrtella, according to the orientation of the interarea. c. koopi archbold, 1990 from the sakmarian cuncudgerie sandstone (cunning basin) is difficult to judge being an internal mould. occurrence. early permian of south oman, central afghanistan, zanskar (india), ? e, australia. pachycyrtella omanensis n. sp. 1959 asyrinx haushiensis hudson & sudbury, p. 16-17, pl. 5, fig. 2. 1959 pseudosyrinx sp. hudson tr sudbur,v, p.46, pl.5, fig. 1a-b. 1997 cyrtella aff. c. nagmargezsls angiolini et al., p. 391, fig. 11.16, text fig.8,9,11, tab.5. holotype .a complete specimen: mpum 8425. paratypes. 6 complete specimens: mputrl 8426, mpum 8427, mpum 8,+28, mpum8429, mpum 8,+30, mpum 2960. 2 ventral valves: mpum 8431, mgl6z121. 1 dorsal valve: mpum /925. etymology. species named for its provenance, the sultanate of oman. type locality and age. saiwan (coord. 2a"52'27"n57"36'26"f.),interior of oman, basal bed (ol1a) of the saiwan formation, mid-sakmarian. description. medium to large-sized biconvex shells with maximum width at hinge line. cardinal extremities from angular in .juveniles to truncared in adults. shell substance finely punctate and micropunctate. ventral valve weakly convex, sub-rhomboidal in outline. interarea high, generally slightly concave, oriented at a low angle with the commissural plane (from apsacline to nearly orthocline); perdideltidial areas not well demarcated. delthyrium closed by stegidial plates; hypodeltidial and deltidiai furrows separated by a deltidial ridge. shallow and smooth ventral sulcus arising at umbo, widening anteriorly and protruding anteriorly as a tongue of variable length. ornamentation of simple rounded costae, numbering fi-12 for each flank. costae widening anteriorly up to 3-,1.5 mm in width at the anterior margin. growth lrrmellae of two differenr sizes occur throughout the valve: coarser and widely spaced versus finer, more numerous and denser. micrornamentation of minute pustules. dorsal valve tr.lnsverse, strongly arched. fastigium widening and getting higher anteriorly, bearing a deep and wide median furrow, becoming shallower at the anterior margin. lateral dorsal margins strongly overlapping the ventral ones. ornamentation of 10-12 simple rounded costae on each side of the fold. costae widening anteriorly up to 3-3.8 mm at the anterior margin. growth lamellae as in the ventral valve. ventral valve interior with large and concave delthyrial plare, showing a strongly conc.rve margin towards the hinge line. umbonal callus variably developed, embedding dental plates and adminicula and filling at vcriable degree the central cavity below delthyrial plate and the lateral cavities; anterior sides of apical callus deeply pitted by genital markings. dental plates high, merging with adminicula and concave towards iateral margins. adminicula surrounding the posterior part of the muscle field. teeth stout and coarse. muscle field large, sub-rhomboidal, slightly depressed, dendritic on posterior part and longitudinally striated on anterlor part and divided by a myophragm. dorsal valve interior with a broad, sessil laminated cardinal process; socket plates and crural plates fused to the cardinal process by a callus which is very large in mature specimens. spiralia broad, tightly coiled, with postero-larerally directed apex. adductor scars depressed with a thin median myophragm. discussion. this description is in total agreement with that of ?c. aff. nagmargensis published by angiolini in angiolini et al. (1997, p.391-392) to which the reader is referred for details on the ontogentetic variation, intraspecific variabilitir, and serial sections. the invalidity of the genus asyrinx hudson 6c sudbury. lq59 based on rwo non congeneric specimens has a1read1. been pointed out by angiolini in angiolini et al (1997\. ackno.oledgements. m. balìni, e. garzanti, a. nicore, j.p platel, j. roger and a. tintori are thanked for joint field-lvork. j. carter rnd g. plodowski are thankcd for revìsion and encouragement. reseerch financialll' supported rjthin the peri-tethys progremme and bv italian cnr (grant to m. gaetanì). nero bracbiopod from early permian of oman references 129 angiolini l., bucher h.. pillevuit a.. platei j.p., roger j., broutin j., baud a., marcoux j., & al hashmi h. (1997) early permian (sakmarian) brachiopods from southeastern oman. geobios, 3a: 379-405, lyon' archbold n.\( & gaetani m. (1993) early permian brachiopoda and mollusca from the n\7 himalaya, india. riv. it. paleont. strat.,99 27-56,milano. carterj.,johnsonj.g., gourvennec r. & hong-fei h. (199a) a revised classification of the spiriferid brachiopods. annals of carnegie mwsewm, 63:327-374, pittsburgh. hu c. (1933) new genera and species of spiriferacean brachiopods in the late carboniferous to early permian from duoma district, rutog, xizang (tibet), china. journal of wuhan college of geology,l'9: 105-117, tvuhan. hudson r.g.s. & sudbury m. (1959) permian brachiopoda from south-east arabia. notes et memoires sur le moyenorient. 7: 20-53. parigi. termier g., termier h., de lapparent a.f. & marin ph. 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+�� ,�� � ��� 2���'���** $�$+�� '�� +��� 2��.��$�� 0��,& ;1& �'� �(� �-�� �-2 ��� ��,��'�� �'� ;�&��h �. �'� ���� #�������& �'��%,' � �� ��� "��� �'� ������+%���� �. �'� ��$2 �� ��� �2����� �� ����� .��$ �'� ����� ����� �$& '�� +��� ��%���� 0��,& 61& �'� �-�� ��� � ����%�� ���2����#� " .�� �@&:6h ��� ��&�dh �. �'� ���� #��! �����& ��$2 �� ��� � $��� ������%�%� " ������+%��� � ��, �-�� � �'�(��, � 2��,�����#� ���������, � ��, �-�� �& �'� ��$2 �� (��' '�,'��� �-�� ������ ��� �'��! ��������� +" $��� " $����� �2����� ��� ������2���� �� � %���� + �. �'� � %���� ��� "���& ��$2 �� (��' �������! ��, �-�� ������ ������2��� �� �'��� �. � %���� � �. �'� � %���� ��� "���& �'��� ����� ��� +� .%��'�� �%+��! #���� �� �'��� ,��%2� ��e��, ���� ����%�� �'� � ��� ���! ���+%���� �. �'��� ��$����� �2����� �� �'� �� 2 ��& ��"������� ��$����" ���" ���� �) ��� �"��'�"��������� 1���� 1� �� � �������� ,��$�� 2��"�3 ;@6 �� ���� #-� +"��((� /0 1�((��� 2-;@d species pn o i 1 2 pn o i 2 3 pn o i 1 4 pn o i 1 8 pn o i 1 1 pn o i 2 1 pn o i 2 4 pn o i 2 5 pn o i 2 6 pn o i 2 8 pn o i 2 7 r m 6 r m 7 r m 5 pn o i 4 5 pn o i 3 3 pn o i 3 5 pn o i 3 7 pn o i 4 1 pn o i 4 2 pn o i 4 6 pn o i 4 7 pn o i 4 8 pn o i 4 9 pn o i 5 0 pn o i 5 6 pn o i 5 7 argilloecia kissamovensis sissingh, 1972 bairdoppilata profunda aiello, barra & bonaduce, 2000 bythocypris obtusata producta (seguenza, 1880) cytherella gibba aiello, barra, bonaduce & russo, 1996 cytheropteron (cytheropteron) omega aiello, barra & bonaduce, 1996 henryhowella sarsii profunda bonaduce, barra & aiello, 1999 krithe compressa (seguenza, 1880) krithe exigua abbate, barra, aiello & bonaduce, 1993 paijenborchella iocosa kingma, 1948 parakrithe acuta aiello, barra, abbate & bonaduce, 1993 parakrithe ariminensis (ruggieri, 1967) parakrithe declivis ciampo, 1980 parakrithe dimorpha bonaduce, ciampo & masoli ,1976 acanthocythereis histrix (reuss, 1850) argilloecia minor müller, 1894 cytherella robusta colalongo & pasini, 1980 cytheropteron (cytheropteron) sulcatum bonaduce, ciampo & masoli ,1976 cytheropteron (aversovalva) denticulatum aiello, barra & bonaduce, 1996 eucytherura sp. loxoconcha cf. l. concentrica bonaduce, ciampo & masoli, 1976 argilloecia sp. 1 buntonia sublatissima (neviani, 1906) cytheropteron (cytheropteron) venustum aiello, barra & bonaduce, 1996 parakrithe lata ruggieri & d'arpa, 1993 parahemingwayella tetrapteron (bonaduce, ciampo & masoli ,1976) aurila (aurila) convexa (baird, 1850) aurila (cruciaurila) cruciata (ruggieri, 1950) aurila (aurila) punctata (münster, 1830) bosquetina tarentina (baird, 1850) buntonia robusta ruggieri, 1954 callistocythere flavidofusca (ruggieri, 1959) carinocythereis carinata (roemer, 1838) carinovalva testudo (namias, 1900) cimbaurila cimbaeformis (seguenza, 1883) cistacythereis (cistacythereis) cebrenidos ulicnzy, 1969 costa edwardsii (roemer, 1838) cytherella scutulum ruggieri, 1976 cytherella harrymutvei stambolidis, 1980 cytheropteron (cytheropteron) monoceros bonaduce, ciampo & masoli ,1976 cytheropteron (cytheropteron) ruggierii pucci, 1955 echinocythereis (rhodicythereis) pustulata (namias 1900) eucytherura gullentopsi ruggieri, 1952 henryhowella parthenopea bonaduce, barra & aiello, 1999 leptocythere multipunctata (seguenza, 1884) leptocythere transiens pucci, 1956 loxoconcha ovulata (costa, 1863) palmoconcha subrugosa (ruggieri, 1976) palmoconcha turbida (müller, 1894) pterygocythereis coronata (roemer, 1838) pterygocythereis jonesi (baird, 1850) pseudocytherura calcarata (seguenza, 1880) sagmatocythere versicolor (müller, 1894) semicytherura ruggierii (pucci, 1956) carinocythereis whitei (baird, 1850) cytheropteron (cytheropteron) circumactum colalongo & pasini, 1980 ionicythere reticulata (colalongo & pasini, 1980) loxoconcha rhomboidea (fischer, 1855) paracytheridea hexalpha doruk, 1980 semicytherura rarecostata bonaduce, ciampo & masoli ,1976 xestoleberis communis (müller, 1894) xestoleberis erecta namias, 1900 cytheretta subradiosa (roemer, 1838) hemicytherura gracilicosta ruggieri, 1953 leptocythere ramosa (rome, 1942) neocytherideis subulata (brady, 1868) pontocythere turbida (müller, 1894) semicytherura rara (müller, 1894) eucythere curta ruggieri, 1975 procytherideis retifera ruggieri, 1978 semicytherura incongruens (müller, 1894) cypria ophtalmica (jurine, 1820) pontocypris cf. p. frequens (müller, 1894) procytherideis subspiralis (brady, crosskey & robertson, 1874) ��+& ! ��� �. �'� �������� �2����� ���%����, (��'�� �'� ('� � ����� ����� �%��������& ��"������� ��$����" ���" ���� �) ��� �"��'�"��������� 1���� 1� �� � �������� ,��$�� 2��"�3 ;@@ species pn o i 1 2 pn o i 2 3 pn o i 1 4 pn o i 1 8 pn o i 1 1 pn o i 2 1 pn o i 2 4 pn o i 2 5 pn o i 2 6 pn o i 2 8 pn o i 2 7 r m 6 r m 7 r m 5 pn o i 4 5 pn o i 3 3 pn o i 3 5 pn o i 3 7 pn o i 4 1 pn o i 4 2 pn o i 4 6 pn o i 4 7 pn o i 4 8 pn o i 4 9 pn o i 5 0 pn o i 5 6 pn o i 5 7 cytheridea neapolitana kollmann, 1960 loxocauda decipiens (müller, 1894) semicytherura n. sp. loxoconcha glabra (brady, 1866) neonesidea mediterranea (müller, 1894) semicytherura inversa (seguenza, 1880) aurila (aurila) cephalonica mostafawi, 2006 aurila (aurila) abscisa (terquem, 1878) bythocythere zetlandica athersuch, horne & whittaker, 1983 celtia (celtia) quadridentata (baird, 1850) cytherella circumpunctata ciampo, 1976 echinocythereis vidua barra & bonaduce, 2000 flexus triebeli (ruggieri, 1962) ionicythere n.sp. phlyctenophora affinis (schneider, 1953) propontocypris micropunctigera ruggieri & d'arpa, 1993 ruggieria longecarenata (namias, 1900) semicytherura alifera ruggieri, 1959 tetracytherura irregularis (terquem, 1878) urocythereis sororcula (seguenza, 1880) aurila (aurila) hesperiae ruggieri, 1975 cistacythereis (hiltermannicythere) rubra (müller, 1894) cytheretta adriatica ruggieri, 1952 cytheropteron (cytheropteron) depressum brady & norman, 1889 eucytherura gibbera müller, 1894 eucytherura patercoli mistretta, 1967 grinioneis haidingeri (reuss, 1850) ilyocypris gibba (ramdohr, 1808) leptocythere bacescoi (rome, 1942) loxoconcha alata brady, 1868 loxoconcha rubritincta ruggieri, 1964 metacypris cordata brady & robertson, 1870 paradoxostoma abbreviatum sars, 1866 paradoxostoma ensiforme brady, 1868 protocytheretta obtusa ruggieri, 1962 pseudocytherura miliciae ruggieri & d'arpa, 1992 sagmatocythere napoliana (puri, 1963) semicytherura acuticostata ventricosa (müller, 1894) semicytherura intorta (terquem, 1878) semicytherura paradoxa (müller, 1894) urocythereis praelonga (terquem, 1878) callistocythere crucifera (hartmann, 1953) potamocypris fallax fox, 1967 semicytherura dispar (müller, 1894) aurila (aurila) n. sp. costa batei (brady, 1866) cytheropteron (cytheropteron) latum müller, 1894 microxestoleberis xenomys (barbeito-gongalez, 1971) xestoleberis plana (müller, 1894) cyprideis torosa (jones, 1850) cytherella vulgatella aiello, barra, bonaduce & russo, 1996 heterocythereis albomaculata (baird, 1838) ilyocypris getica masi, 1906 mutilus laticancellatus (neviani, 1928) aurila (aurila) lanceaeformis uliczny, 1969 limnocythere inopinata (baird, 1843) candona (neglecandona) neglecta sars, 1887 heterocypris salina (brady, 1868) basslerites berchoni (brady, 1869) costa punctatissima ruggieri, 1962 sylvestra virgula bonaduce, russo & barra, 1990 argilloecia spissa aiello, barra & bonaduce, 1996 callistocythere n. sp. pseudocandona marchica (hartwig, 1899) potamocypris zschokkei (kaufmann, 1900) ���%2 � �� �'����������� +" ����$+ �,�� $��� �. ��$�! 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true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /fra /enu (use these settings to create pdf documents with higher image resolution for improved printing quality. the pdf documents can be opened with acrobat and reader 5.0 and later.) /jpn /deu /ptb /dan /nld /esp /suo /ita /nor /sve /kor /chs /cht >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [595.000 842.000] >> setpagedevice [rt" italiana cli paleontologia e stratigrafia lampadena ionica: a ne\t teleost from the mediterranean pleistocene. angela girone'! & dirk nolf'i'i recei.oed december 27,2aa1; accepted may 3,2aa2 key uords: lampadena, pleistocene, sourhern ftaly. riassunto. viene descritta la nuova specie lampadena ionica (myctophidae, teleostei) riscontrata in depositi del pleistocene inferiore-medio dell'italia meridionale. in particolare l. ionica è stata riconosciuta dalla biozona a "la,rge gephyrocapsa" a quella a pseudoe milianìa lacunosa ed apparentemente sembra estinguersi prima della fine del pleistocene. sebbene il genere lampadena sia rutrora viventc solo negli oceani atlantico e indo-pacifico, esso è noto nel mediterraneo sin dal miocene inferiore. l. ionica sembra essere i'unica specie di questo genere esìstente neì pleistocene del mediterraneo. sino ad ora è stata rinvenuta associata a fauna bentonica (invertebrati e vertebr,ltì1 caratteristica di ambienti batiali indìcante una profondità maggìore di )uu lnetrt. abstract. the new spectes lampadena ionica (mvctophidae, teleostei) is described from lou.er and middle pleistocene deposits of southern italy. in particular, l. ionica is known from the "large gepbyrocapsa" up to the pseudoemiliania lacunosa biozone. apparentlr.., the species became extinct before the end of the pleistocene. although the genrs lampadena iives only outside the mediterranean today, ìr is known from the mediterranean reaim since the early miocene. z. lon lca seems to be the only species of the genus lampadena existing in the pleistocene deposits of the mediterranean area. the new species has been found only associated nith benthic faunas (invertebrate and vertebrate) indicating an bathyal environment deeper than 500m. introduction the genus lampadena goode & bean, 1896 was known as a fossil in the mediterranean realm from miocene and pliocene deposits but is not represented there in the recent fauna. the miocene species are the extincr l. gracile (schubert 1912) and l. speculigeroides brzobohaty ee nolf, 1996.in the pliocene, the recent z. dea fraser-brunner, 1949 is recorded from zanclean (lower pliocene) deposits of the western mediterranean (nolf er cappetta 1988; nolf et al. 1998). in the recent fauna, the latter species is restricted to the southern part of the indian and pacific oceans, betsreen the latitudes of approximately 2o"s and 5o"s (nafpaktitis & paxton 1968). recent taxonomic srudies on the mediterranean pleistocene otoliths revealed the presence of a new species, lampadena ionica, from two sites in southern italy. the otoliths of the species described here can easi1y be distinguished from the other lampadena species of the mediterranean miocene and pliocene as s/ell as from the living species. otoliths of lampadena ionica n. sp. were collect, ed from silty pleistocene deposits ar rwo locations in southern italy. montalbano jonico a composite succession of marine pleistocene deposits, over 400 m thick, was reconstructed aiong the internal border of the southern apennines foredeep (montalbano jonico area, fig. 1) (ciaranfi et al. 1997). it consists mainly of terrigenous clayey-silts and silty-clays in the basal and middle part, and of sandy-silts, silty-sands, and sand bodies in the upper part. nine volcaniclastic layers (v1-v9) are included at various heights, and a marine conglomerate tops the succession (fig. 1). calcareous nannofossil assemblases indicates a lower to middle pleistocene age, and the successron was proposed to locate the gssp of the lower-middle pleistocene (ciaranfi et ^1. 1,997). the basal part of the section belongs to the "iarge gephyrocapsatt and "small gephyrocapsa" nannofossil biozones, and the middle and upper part to the psewdoemiliania lacunosa nannofossil biozone (ciaranfi et al. 2001; marino 1996). palaecological, taphonomical, and sedimentological analyses suggest a general regressive trend from upper slope to shoreface environmenrs. the otoliths studied were collected from the first 1.2 m of the section, named entalina section, which belongs to the lower pleistocene (iarge gephyrocapsa nannofossil biozone). the benthic invertebrate associa'ldipartimento di geologia e geofisica, unìversità di bari, r.ia e. orabona 4,7a125 bari : email girone@geo.uniba.ir 'r'rinstìtut royal des sciences naturelles de belgique, rue vautier 29, b-1000 bruxelles: email dirk.nolf@natuurwetenschappen.be 494 a. girone & d. nolf tions and the otolith associations are dominated by bathyal species suggesting a palaeodepth of 500-600 m (ciaranfi et al. 2001; girone 2000). archi this section, 9 m thick, is located near reggio calabria, southern italy, (fig. 1) at about 90 m above the sea level and is particularly well-exposed along a ns abandoned quarry front. the section, lower-middle pleistocene in age, mainly consists of well stratified pelitic sediments with a general slight westward sloping of the beds (di geronimo et al. 1997).the deposition occurred during a cold phase, as inferred from the planktonic foraminiferal assemblages. the invertebrate benthic faunas (foraminifers, moiluscs, bryozoans, and serpulids) (di geronimo et al. op. cit.) as well as the fish otoliths (girone 2oo0) testify a bathyal palaeoenvrronment, 5oo to 1ooo m deep. the occurrence of l. ionica in the section is show in fie. 1. systematic palaeontology terminology applied to otolith morphology follows nolf (1985). the classification adopted is that of rosen (1973) subsection ctenosquamata rosen, 1973 sept scopelomorpha rosen, 1923 order myctophiformes regan, 1911 family myctophifdae e gill, 1893 genus lampadena goode & bean, 1896 fio i loc.rrion of monrelb:no jonico and archi sections. type species. lampadena speculigera goode tr bean, 1896. the otolìths of the species of the genus lampadena generally have large and oval-shaped otoliths, considerabiy longer thrn high. they can show a slight or absent posterodorsal notch. only one recent species l. anomala has a small and more round-shaped otolith. lampadena ionica new species fig. 2 a-e etymology, from the montalbano jonico section (basilicata, southern italv), the type locality. type materiaì, holorrpe: a left oroìith 'fig. 2ar /dgcub ag 1) from the entalina section (montalbano jonico composite section) lon er pìeistocene in age; 1o parat,vpes! of which four are figured (fig. 2b-e) (dggub ag 2-s). type locality. montalbano jonico section, southern ltalv. repository. dipartimento di geologia e geofisica, università degli studi di bari, italy (dggub ag1-s). dìagnosis. a species characterised by moderately thin, oval shaped otoliths becoming more elongated with the growth. the rostrum is large and salient with no antirostrum. the posterodorsal angle is r-ide and moderately notched. the sulcus is t-ide. description this species is characterized by oval-shaped otoliths with a wide sulcus. the ostium is somewhat longer than the cauda. the caudal colliculum is narrower than the ostial one, which becomes slender towards the anterior end. the cristae become obsolete in the posterior part of the otolith; the superior one is well marked and more linear than the inferior one. the collampadena ioníca, a nea teleost lampadena lozlca, lvlontalbano jonico, ìou'er pleistocene (large gephyrocapsa biozone): a) holotl-pe; left otolith; b-c) paratypes, leit orolìth': ll-s' p.rr.rrrlrc': r'i1hr orolirh': dl renrrr'ri.*. 495 fig. l licular crest is elongated and separated from the crista inferior by a deep furrow the dorsal and ventral areas are very similar in size; the former is characterized b1:r depression covering most of its surface. this depression is deeper just above the crista superior and becomes more smooth towards the dorsal rim. the ventral area is slightiy conyex and bears a shallou.. groove near to the ventral rim. in large specimens, the dorsal rim is truncated in its posterior part (behind rhe posterodorsal angle) where it reaches its maximum height. in sn-iall specimens the dorsal rim shows a regular curr.e. the ventral rim is curved but not semicircular and bears irregularly spaced obtuse spines. the anterior spines i3 or 4 in number) are more pointed and are closer to each other. they are separated b). deep furrou's on the inner face. the posterior rim is globally rounded, but the lobation shows a marked variability among the available specimens. in several llrger ones, it bears two or three lobes near the posterodorsal angle. the outer face is irregularly convex with the maximal convexity located in the posterior part. the surface of this face shows a low. elongate umbo in the central part and numerous radial lobes in the marginai zone. affinities. otoliths of lampaclena ionica can be distinguished from those of the fossil species l. gracile (schubert, 1912) and l. speculigeroides brzobohaty tt nolf, 1996 from the mediterranean miocene bv their notched and wide posterodorsal angle, their less pronounced rostrum and a more rounded and convex ocrsterior mrrgin. in lhe tn o m iocene species, this nrargin is linear and subvertical (brzobohaty & nolf 1996, pi. 4, figs. 1-6 and figs. 12-1o1. ln the recent fauna, the genus lampadena is represented by eight nominal species and one subspecies (paxton 1979). orcliths collected from the stomach contenr of r pigmv sperm whale (.kogia simus) caught off taiji, japan, are different from all other presently knori,'n lamapadena species and apparentlv represent e ninth species (p1. 1, tig.22), but the entire fishes have not been causht. otoliths of all known taxa have been figured bv various authors and those pictures are scartered through the literature. references to rhese sources are listed in table 1, which also provides a sur\-ey of rhe geographic distribution of each taxon. otoliths of al1 those recent taxa are also figured on pl. 1, which for several of species, provides gro\\rth series that never have been published before. otoliths of l. ionica differ from those of l. luminosa (garman 1899) (p1. 1, fig. 1-4) and l. urophaospaxton,1963 (pl. 1, fig. 16-17) by a wider sulcus, a posterodorsal angle which is conspicuously obtuse and, in adult specimens, a less notched superior part of the postrior rim. in the medium-sized otoliths of the present pleistocene species and the recent l. luminosa, the two lrtter features appear to be very similar, but the medium-sized as q.ell as the large specimens of l, ionìca are less oblongate and have a wider-shaped anrerior parr than i. lumirtosa. the features that distinguish the pleistocene species from the pacific l. urophaos are much nominal species figured otoliths distribution lampadena anomala pan. i 928 lampadena chavesí collet, 1905 lampadena dea fraser-brunner, 1949 lampadena luminosa (garman. 1899) lampadena notialís nafpaktitis & paxton, 1968 lampadena pontifex krefft, 1970 lampadena speculigera goode & bean, i896 lampadena urophaos urophaos paxton, 1963 lampadena urophaos atlantica maul, 1969 lampadena sp. a pl. 1, fig.9-10 cor-strl,a.n & nappaxrtr:s, 1912, fig. 4c, p, 7 nafpakrns & p,qxron, 1968, fig. 10.8, p. 24 pl. l. fig. 5-8 nappa,rrrrrs & p,txron, 1968, fig. 10.7 , p. 24 suele et al., 1995, pi.21, fig. e1-3 pì. 1, fig. 18 napparrrrrs & paxron, 1968, fig. 10.6,p.24 reow, nsra,1992, fig. 36, p. 186 pì. 1, fig. 1-4 napperrrrrs & p,q.xron, 1968, fig. 10.1, p. 24 rrveron & bounrer, 1999, pl. i 19, figs. 1-13 sua.r-e et al., i995, pl. 21, fig. f1-2 pl. 1, fig. 1 i-12 nnpprrrrrrs & paxron, 1968, fig. 10.5,p.24 suers et al., 1995, pl. 2i, fig. g2, not gl(?: bolinichthys) pl. 1, fig. 20-2i krrppr, 1970, fig. 3, p.280 pl. i, fig. 13-15 nerpe.rtrrrs & pe,xron, 1968, fig. 4, p. 12, frg. 10.1 , p. 24 svers er al,, 1995, pi. 21, fig. a1-3 pl. 1, fig. 16-17 kotlyer & p,qnln, 1990, fig. 2, p. 101 nepperrlrrs & paxron, 1968, fig. \0.2,p.24 rlvaron & bounnrr, 1999, pl. 1 19, figs. 14-19 pl. 1, fig. 22 nerparcrrrrs & paxrou. 1968, fig 19fiop1 10.3 atlantic, indian ocean, east and central pacific n and s atiantic, antitropical; southem indian and pacific oceans southem parts ofall three oceans, between 20 and 50"s atlantic, eastern pacifi c southern ocean atlantic, mainly mauritanian upwelling n atlantic, s atlantic, s indian ocean, se pacifrc eastern pacific n atlantic off japan, otoliths from stomach content of sperm whale kogia sinasl fish unknown 496 a. girone & d. nolf table 1 list of recent species of the genus lampadena. the references on iconography and the geographic distribution are also reported. more marked in the atlantic subspecies l. urophaos atlantica maul, 1969 (pl. 1, fig. 19) that, in addition to other features, has a more elongated antirostrum and a more salient rostrum than l. ionica. although the general shape of otoliths of l. ionica shows some similarity to those of the recent north atlantic l. notialis nafpaktitis & paxton, 1968 (p1. 1, fig. 11-12), the pleistocene specimens differ from the latter by having a larger sulcus, a more salient posterodorsal angle and a more expanded anteroventral area. in l. notialis the antirostrum is well developed while in l. ionica it is almost absent. these differences can also be observed in the medium sized-otolith of z. notialis from the south atlantic figured by nafpaktitis er paxton (1968, fig. 10) in which the anterior part of the dorsal area is wider than in l. ionica. otoliths of the i. ionica differs from those of z. deafraser-brunner, 1949 (p1. 1, fig. 18), known as fossil from the lower pliocene deposits of the mediterranean realm, by their marked rostrum and their more obtuse posterodorsal angle which, in the recent species, is near1y right. moreoveq l. dea generally shows a well distinct rostrum and antirostrum (nafpaktitis er paxton 1968). the more obtuse posterodorsal angle is also the feature that distinguishes otoliths of l, ionica from those of the recent l. pontifex (pl. 1, [rg. 20-21) and pl. 1 lampadena ionica, a nezu teleost 497 plate 1 the recent specimens figured in this plate are from the collection irsnb (institut ro1'al des sciences naturelles de belgique). fig. 1-a) lampadena luminosa,leftotoliths.recent: gulf ofmexico.fig.5-b)lampadenacha'",esicollet,1905, leftotoliths.recent:atlantic,off theazores.fig.g-10) lampade naanomalaparr,1928,left otoliths. recent: atlantic (researchvessel knorq station 65). fig. 11-12) lampadenanotìalis nafpaktitis & paxton, 1968, left otoliths. recent: 11) south \west atlantic (research vessel valter herwìg statìon 362), coll. sch\iarzhans; 12) off new south \wales, australia. fig. 13-1.5)lampadenaspeculi.geragoode&bean, 1896, leftotoliths.recent:13)atiantic,offnlllreland;14-15)gulfofbiscaye.fig. 16-17) lampadena uropbaosputon, 1963, leftotoliths:fig. 16) newcaledonia,fig. 1z)california,recent:16) offnewcaledonia;12)offcalifornia.fig. 18 lampadena deafraser-brunner,i949,left otolìths. recent: se pacific. fig. 79) lampadena urophaos atkiltic.lmaul, 1969, right otoliths. recent: off portugal, atlanric. fig. 20-21) lampadenapontifex lkeflì, 1970, left otoliths. recent: atlantic (research vessel atlantis ii sration 59). fig.22) lampadena sp. a,left otolith. recent: from the stomach content of a pigmy sperm whale (kogia simus) caught offtaiji,japan. redraq afternafpaktitis er paxton (1968, fig. 10.3) lampadena luminosa (genmr, 1899) lampodena chavesi collerr, 1905 parr, i928 :.--=--_---1t" lampadena speculigera goooe & bean, 1896 lmm-\ -j tt ,/f-\"" /r l€l+;gl ,,r'.\imto lampadena nolialis nafpaktitis & paxton, 1968 lampadena pontiér krefft, 1970 lampadena urophaos paxron, 1963 lampadena dea fraser-brunner, 1949 lampadena urophaos atlantica maul, 1969 498 a. girone & d. nolf references lampadena sp. a (p1. l, frg. 22). in the recent l, speculigera and l. anomald, rhe general shape of the otoliths is quite different; the otoliths oí l. speculigera have a nearly vertical posterior margin and a narrower sulcus with a very short cauda. according to nafpaktitis & paxton (1968) and coleman & nafpaktitis (1,972), the otoliths of l, anomala can be considered as intermediate between the more elongated otoliths of all other lampadena species and the almost round, smooth-edged otoliths of the genus thaningichthys (myctophidae). otoliths of l. ionica c\ear\y belong ro the lampadena group with elongate otoliths and are quite different from the high-shaped l. anomala morphology. distribution lampadena ionica first appears in the lower pleistocene beds ("large gepbyrocaps4" nannofossil biozone) in the basal part of the montalbano jonico section. the presence of l. ionica in the bathyal assemblages from the archi section (calabria, southern itaiy), referable to the "sma11 gephyrocapsa" and the pseudoemiliania lacwnosa nannofossil biozones (di geronimo et al. 1997;, suggests that this species survived in the mediterranean basin up to the middle pleistocene. most recent species of the gents lampadena appear to be among the deepest-dwelling myctophids; they are merely captured below 600-700 m. l pontifex and l. lwminosa occur also at shallower depths between 275 and 450 m (nafpaktitis et al. 1977). l. ionica was collected from beds which, according to their content of benthic invertebrates and benthic fish assemblages, indicate a bathyal environment of more than 500 m deep (girone 2ooot. in the montalbano sections, l. ionica is absent in the more shallow associations referable to the "small gephyrocapsa" biozone. ac/enou-ledgemen ts. tve wish to thank prof. i. di geronimo for pror-iding the sample of archi section, a. rizzi (university of milano) for his assistance during sem photographs and p hoffman, who helped us in preparing the iconographl' of the present paper. fìnancial support was provided by the phd program of the university of bari ,-,1 l, trc r ^{,l-. f,,,^-.tn communir.. brzobohaty r. & nolf d. 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(1985) otolithi piscium. in schultze, h. p (ed.) handbook of paleoichthyology. g. fisher verlag, 10: 1145, stuttgart and new york. nolf d. & cappetta h. (1988) otolithes de poissons pliocènes du sud est de la france. bull. inst. r. sc. natur. b e lgique, s c. terr e, 58 : 2a9 -27 1, bruxelles. nolf d., mané r. & lopez a. (1998) otolithes de poissons du pliocene inferieur de papiol, près de barcelone. palae oo ertebrata, 27 (l -2) : 1 -1 7, 7 pi, montpeilier. paxton j. r. (1979) nominal genera and species of lanternfishes (family myctophidae) . contrib. science, nat. hist. museum los angeles cownty,322: 1-28, los angeles. radwanska u. (\992) fish otoliths in the middle miocene (badenian) deposits of southern poland. acta geologica po loni ca, 42 (3 4) : 1 4 | -328, \xlar szawa. rivaton j. & bourret p. (1999) les otolithes des poissons de lampadena ionica, a new teleost 499 1'indo-pacifique. doc. sc. techn., inst. rech. déoel., nouméa,2 (2): 1-378, nouméa. rosen d. e. (1973) interrelationships of higher euteleostean fishes. in: greenwood p h., miles r. s. & patterson c. (eds) interrelationships of fishes. zool. journ. linnean )oc.. )j: jt/-jij. lonclonsmale m. j., \fatson g. & hecht t. (1995) otolith atlas of south african marine frshes. ichthyological mon., 1: 1.253, grahamstown. rivista ltaliana di paleontologia e srratigra *.".-*oo, l the castagnone site (cerrina valley, monferrato hills, n\i italy): early pleistocene sedimentary record and biochronology carlo giraudi" alberto mottura" benedetto sala], m. stella siori'& daniele, bormioli, receit,ed may 6, 2002; accepted april 24, 2003 key u^ords: earlv pleistocene, jaramillo subchron, mammal fauna, italv. abstract. geological researches carried out near the castagnone hamìet in the cerrina valle.v (northern monferrato hills, piedmont, n\l italy), have brought to light a post-messinian succession whose sedimentary record starts with a lower complex of pedogenized collur.ial materials and with two superimposed alluvial units (i and ii). the low-er one of these units contains a galerian macrofauna associated with n-ricrotine vole teeth (mimomys satini, mimomys pusillus, ungaromys cl. u. nanus, microtus (allophaiomys) sp.), n-hile the upper one yields only scarce faunal remains. most of this sediments were deposited durine a normal palaeomagnetic phase. the i alluvial unit, due to its biochronologìcal correlation, must be referred to the jaramillo subchron, between 1,070,000 and 990,000 years ago. the ii alluvìal unit, being both unconformable with and younger than the first one, might be best referable to the brunhes chron. or.erall, the bed dipping across the reported succession shos's a progressive syn-sedimentaru tiltìng, with accelerated deformation durìng the i alluvial unit deposition. this tectonic stress over rhe castagnone area is seemingly related to the uplift of the north-easternmosr ridge of the monferrrato hills and appears to have been nearly exhausted before the ii alluvial unit deposition. riassunto. le ricerche geologiche condotte nei pressi dell'abiteto dì castagnone di pontestura, in val cerrina (monferraro settentrionale, piemonte), hanno evidenzixro una successione sedimentaria post-messiniana che inizia con un complesso inferiore cosrituiro cla colluvi pedogenrzz:tr e da due unità sovrapposre formate da sedimenti alluvionali. tlunità alluvionale inferiore (i a.u.) contiene una macrofauna galeriana associata con denti di micromammiferi (mimomys sa.oini, mimomys pusillus, ungaromys cf . u. nanus, microtus (allophaìomys1 sp.), mentre ia superiore (li a.u.) contiene solo scarsi resti faunìstici. tuttr l:r serie sedimentaria risulta deposta in periodi a polarità magnetica normale. per il suo conrenuro in fossili la i a.u. deve essere attribuita al subchron jararnillo, datato tra 1.ozo.ooo-99o.ooo anni fa. la ii a.u. è più recente e discordante sulla i a.u. e sembra attribuibile al chron brunhes. linclinaz.ione decrescenre degli strati che formano la successione sedimentaria indica una deformaz-ione sinsedimentaria progressir.a, parricolarmenre evidente all'interno della i a.u. lo stress tettonico dell'area di castaenone sembra in relazione con il sollevamento del margine nord-orient:le del monferr.rto ed appare quasi completamete esaurito prima della sedimentazione della ii a.u. introduction the early pleistocene sediments located in the cerrina valley (northern monferrato hills, piedmonr, n\f italy) preserve both lithological and fossil record which document palae oenvironmental variations, tecronic evolution and biochronology. the cerrina valley. is in a hi1ly area, mainly formed by cenozoic marine sediments, with elevations of up ro 455 m a.s.l. the monferrato and the relared turin hills form a central relief isolated from the main wesrern a1pine ridge and are surrounded by the alluvial plains of two major rivers: the po (n) and the tanaro (s). the reported pleistocene succession locally and in patches outcrops on the left slope at the end of the cerrina valley (fig. 1). mrjor exposures are preserved near castagnone (pontestura, alessandria) : on rhe local hillslope the sediments are covered by a thin aeolian layer, while in the valley bottom the lower succession merges under the recent alluvium. in the late'zos fossiliferous gravelly-sandy alluvial sediments yrere first located by one of us (c.g.) during preliminary studies in a now disused quarry (peratore quarry: c1 in the text and fig. 1). there, some macrofaunal remains suggested as possible an eariy middle pleistocene age (giraudi 1981), though the severe sediment deformation and the lack of arvicolid finds biased the biochronologic interpretation. from the peratore quarry other fossil remains were collected in ensuing years in î enea., centro ricerche.casaccia", via anguillarese,30l, ooo6o s. maria di galeria (roma). c.p 2400 oo1o0 roma a.d., italy f -mail: qirrudiq/ cr\rccir.ener.il. department of animal and fiuman biology, via accademìa albertina, 17, 10123 torino, itely. e-mail: siori@,dba.unito.ir department of natural and environmental resources, c.so ercole ii d'este, 32,441aa ferrara, ital,v. e-mail: b.sala(g_ìdns.unife.it department of f,arth sciences, via valperga caluso, 35, 10125 torino, italy. e-mail: geobormioli@,ìlibero.it 51u c. girau.di, a. mottura, b. sala, m. s. siori k d. bormioìi sandy sllt aìluv al val ey floor eol aî si r (late dleislocene' or sandy s lr of rhe lrd.2nd 1st alluvial terraces covering the i and i alluvial units e',1:;":':l:';"1':""::1""a l-lneoroc' \ îe.acescarps @'"'uo" still doubtful stratigraphic context (giraudi 1983; sitec 1984). more recentl,v, archaeological finds referable to different levels of the same succession (not examined in this report) led to new extensive and detailed researches. recent studies in a survey of the whole area have made it possible to localize new outcrops of the castagnone sedimentary succession. in particulaq in a newly opened quarry (cascina nuova: ca1), a network of trenches, with an overall extent of about 300 mt and a depth of up to 5 m, were cut for better control of the stratigraphy, along with archaeological, palaeontological and magnetostratigraphic sampling. the following preliminary updating results allow the overall geological framework to be better known. due to the small number of european locations with late early pleistocene faunas, the reported castagfìg. 1 geolosical sketch nap of thc lon er cerrina valle1,. 1: sandy silt of the alluvial vallev floor (huluccrìeli 2: '.rnj1 .ilr of tl-re .ltl-r elluvial terrace (late pleistocene); 3: aeolian silt (late pleistocene) or, local11; sandl' silt of the 3rd, 2nd and 1st alluvial terraces (middle pleistoccne?), w-hich cover the i ancl ii alluvial units; 4: bedroch; 5: terrlce scarps. none succession improves the resolution of the european biochronological scale. the castagnone/pontestura sequence as exposed in the sections of the ca1 quarry the ca1 quarry (fig. t) is in the upper part of the castagnone hillslope, lear the top (193 m) of the local relief. the bottom bed of the sedimentary succession lie s with a slight unconformity over a bedrock of laminated clayey greylyellow sediments with hardened (chalky or silicized) interbedded levels. such sediments are evidence of a brackish water facies with malacofauna (girotti o., esu d., pers. comm. 20oo) and were referred to a late miocene .messinian' formation (c.g.i. 1969). tl bed roc k lau.+ lithostratigraphv and strati graphicel units of the castasnone/pontestura oriented). l: loess cover; il a.u.: pedogenized siltv gravelly sand (a) and silt end sandy silt rvith carbonatic concretions (b); sand (c) u'ith thin ancl (d); l. c.: consolidated silt and cla,v (a); sand, gravel and siìty cìa1'rvith n.ith (a) hardened levels (messinian). succcssion. cross section at the ca1 quarry' (n-s: ìeftright send,v silt, silty sand and cla,v (b); i a.u.: sand and gravej (a); discontinuous calcrete la1'ers (c1); gravelly sand and dark clala discontinuous calcrete layer (b-c); bedrock: laminrted ch1 fio ) the early pleistocene castagnone site 519 the overlying succession starts, from the borom upwards, with the nl-ower complex" unir, rwo ailuvial units, and ends with alayer of 1oess, as below (fig. 2) . the lower complex (lc). it is formed by: a) clayey, massive, light to dark-grey, consolidated and cracked silt of colluvial origin, up to 40 cm thick; b) grey sand, gravel and silty clay of colluvial origin, with patchy rust oxidation, up ro 20-30 cm thick; c) at the top lies a pedogenic, lnore or less continuous calcrete, up to 10-15 cm thick. the first alluvial unit (i au), this unit, about 10 m thick, lies in slight unconformity over rhe lower complex, and is formed by: a) grey-yellowish sand and gravel containing borh macroand micromammal remains; b) massive or poorly stratified grey-yellowish silt and sandy silt, with embedded carbonaric concretions, sized 1 to 15-20 cm, and some faunal remains; c) yellowish and grey unconsolidated sand, with thin and discontinuous calcreres and leaf marks; this sand is interbedded in the middle part of the silt and sandy silt layer; d) a heteropic facies of gravelly sand and dark clay overlies the above sandy silty sedimenrs: rhe coarser fractions which increase ar rhe rop of the unit are fairly rich in fossil macroand microfauna (the reported elepbas sp. remains from c 1 were also in a correlated level: giraudi i983). the largest raw clasrs which occur in the i au are constituted mainly by silicized rvood or cherts. actually, while other fluvial clasts (quartzites, jaspers and other alpine rocks) seldom exceed 6-7 cm in diameter, the silicized wood can atrain here 50-60 cm in length and weigh more than 1o kg; likewise, the largest cherty cobbles may 1 ^^ ^-be up to 20-25 cm rn diamerer. in the same unit, bony and cartilagineous marine fish teeth, echinoderm radiola and silicized inner casts of marine gastropods have been found, due to an intensive redeposition from the surrounding bedrock. the second alluvial unit (ii au). this enrire unir is weathered by a soil. it is 4-5 m thick, with definite unconformity with the former unit. it is formed by: a) basal silty-gravelly sand with diffused yellowish clay chips: these sediments are weathered and contain badly preserved macrofaunal remains; b) sandy silt and silt with interbedded scanty lenses of siìty sand and clay. during the palaeomagneric srudies a marked increase in the total amount of magnetic minerals with respecr to the i a.u. has been observed. the loess cover. in the uppermost slope of the hills, the topsoil whìch caps the above alluvial levels is buried in its turn by a loessic bed, everywhere thinner than z0-80 cm. discontinuous concenrrations of fresh, unpatinated mousterian arrifacrs were embedded in this layer. down the slope, the loessic covering and the underlying alluvial unirs occur in different relationships. as the latter were cur by the subsequent drainage evolution and before the former occurred, the aeolian deposition took place on terraced slopes (mostly erosional morphologies in the ca1 relief). hence there, the loess is directly in contact with the lower i au or with the tertiary bedrock, while the upper ii au is altogether lacking. the dip of the above different units progressively decreases from the bottom upwards: the undulate "messinian" sediments of the bedrock have an areal inclination of about 14o-12", from the base to the top; the lower complex, formed by coliuvia and the calcareous crust, dips 12'to s (down the local slope); the i au dip decreases from 1o'at rhe botrom ro 4o ar the top; finally the ii au is nearly horizontal. two major erosional breaks in the pleistocene sequence (i. e. rhose between the lower complex and i au, and between i and ii au) correspond actually ro unconformable limits where the aggradation/tilting trend was also uncombined. as seen so far, the units of the above successìon are differently preserved along the hillslopes, both vertically and laterally: this fact is due to the mutual overlap and respective thickness and dip. thus, in addition to its greater depth, the i au is also more extensively preser-ved than the upper ii au: the latter, as it is nearly horizontal, is preserved only between the slope elevations of 193 m lthe top) and 185 m. . discussion and interpretation the lower complex was characrerized by a soil strongly affected by the calcrete and carbonatic concretions. these pedogenic features have so far been recorded in the piedmont region only in the "villafranchian, type succession (boano & forno 1996).they are related usually to high evaporarion rates mosrly controlled by rhe temperarure (birkeland 1984) and so indicate an fairly warm but chiefly dry climate. the thin and discontinuous calcreres interbedded in the i au dip s like the sand containing them, and therefore must be nearly coeval: otherwise in . plrce affected by tilting their dipping would be different. the presence of calcareous concrerions and calcretes in the lower and middie part of the i au, not affected by evident pedogenetic weathering, must be due to the rapid ev.rporation above the water table. towards the top of the i aij, a coarser deposition reappears with the energy renewal while the carbonatic precipitation decreases and stops: therefore a ciimatic change occurred during the sedimentation of this unit. regarding the presence of silicized wood and raw clasts of cherts larger than other alluvial clasts in the i au, it should be noted also that today, in the whole cerrina valley basin as well as in the monferrato hills, no other known formatìon contains so many cherts lboth as to va524 specimen: l" a.u. 1150 i 1e 02 1e-03 1 e-04 + 1 e-03 1é-a2 te 01 te+00 te+01 field b (t) fig. 3 acquisitìon cun'e of the isothermal remnant nr:gnctizltion (irm). riety and overall amount) as this unit. therefore' during the i au deposition, fluvial erosion exposed some (cenozoic?) cherty formation which is now entirely eroded or covered by more recent sediments. a comparable redeposition of cherts was not active in the older plio/pleistocene regional bedloads: e.g., in the "villafranchian" ones such pebbles (far smaller in size) rarely occur. thus, during the i au deposition a sudden important suppl;' of silex began in the regional clastic background. silicized woods are frequent in the cenozoic bedrock below the castagnone sequence: their presence in the i au is probably derived from the fluvial incision near this area, as inferable from the size range and minor reworking of such clasts. also the bony and cartilagineous marine fish teeth, echinoderm radiola and silicized inner casts of marine gastropods were reworked from the bedrock. about the ii au, in respect to the i au, a lithological change may be observed mainly in the contents of large siiicized wood, cherts and clastic load mineralogy; the content of reworked tertiary fossils is smaller. these variations suggest possible drainage shifting and/ or climate changes compared with the i au deposition. the absence of calcareous concretions and calcrete levels shows a different climate from that of the i au. magnetostratigraphy at ca1 the magnetostratigraphic study of the sections which were opened at ca1 allowed the palaeom..rgnetic trend across the entire ourcropping succession to be recorded. the weakly consolidated sediments exposed in the cuts were previously cleaned from more superficial fractions in order to obtain optimized surfaces. the specimens were collected in commercial plastic boxes (inner volume : 2x2x2 cm) gently pushed into the soft rock keeping the front face vertical, .rt 25 cm ìntervals (from the messinian sediments up to the top) leaving out a few levels with too much incoherent sand1. lithologies; all the c. giraudi, a. mottura, b. sala, m. s. siori k d. bormtoli specimens were oriented with bubble level and compass. the sampled section was about 16 m high and 62 specimens were taken in all. the magnetic mineralogy was investigated by isothermal remnant magnetization (irm) using a bussi pulse magnet. the resulting irm acquisition graphs were characterrzed by a quick increase up to full saturation for magnetic field values of o.z-0.+ t (fig. 3). the remnant coercive force (hrc) maintains vaiues in the range of 4050 mt. these results suggest that the magnetite is the main ferromagnetic mineral in the tested lithologies; nevertheless, since thermal tests were not possible bec:ruse of the low coherence of the rock, the presence of iron sulphides cannot be excluded. the natural remnant magnetization (nrm) was measured with ajr-5 spinner. intensity stays :rround the value of io'a/m, with the exception of the specimens collected over 13.40 rn (from the i au top upwards) whose intensity ranged between 1o'and 10'a/m. al1 the directions, across the entire sampled succession, have a variable declination, ranging between n and e, and a positive inclination. stepwise alternating field (af) dem:rgnetization on pilot specimens, up to 1oo mt peak-field, showed the presence of a viscous component which is removed after the first steps; successively the direction remains stable up ro rhe field values,of 30-50 mt (fig. 4). sometimes the situation was more complicated: the presence of different components whose cc,ercive spectra overlap made impossible to recognize the primary component. the systematic demagnetization of all the collected specimens made possible to detect the char.rcteristic remnant magnetization (chrm) direction in 50% of the cases. fisher's statistics were used to calculate the mean chrm directions for each of the sedimentary units in the ca1 succession; the resuits are reported in tab. i and fig. 5. the precision k-parameter and the half-angle confidence cone show a good grouping for the data. the mean directions were then corrected for the respective inclination of the strata: l1"-l2" for the messinian sediments; 12o for the lower complex; 1o' and 4' for the base and top, respectively, of the i au; no correction was made for the ii au which has maintained sub-horizontal strata. the inclination correction reduced the differences among all the recorded palaeomagnetic directions and so, in particular, the last three units probably assumed such corrected directions during the primary sediment deposition. al1 the corrected directions for the entire succession at ca1 correspond to a norn'ral polarity time interval since they show an n-oriented declination and inclination ol 57o, well compatible with the local co-ordinates ltab. i). the magnetic intensity values were correlated with those of the magnetic susceptibility for all the san-rpled $pecinnen: 1 'n50 ca 7a ta +j 1a jj 25 2t) 15 ,lr0 down fig. i af demagnetization dìagram (specimen from lc + i au). svmbols: full dots declination; open dots : apparenr rnclination; figures : peak field value ìn mt. specimens (fig. 6). this graph shows a good correlation among all of them and evidences the narrow grouping of data for the messinian sediments. one can observe also that the ii au shows a marked increase in the total àmount of magnetic minerals, probably because of the more intensive clastic load discharges in this unit. this fact might be correlated with a more appreciable climate variation andl or with greater drainage changes since the final i au deposition. tbe earfu pleistocene crlstagnone site the palaeontological ca1 521 record and the biochronology at the vertebrate fauna remains from the castagnone/ pontestura succession exclusively come from the two alluvial units and are best preserved in the lower one, according to the top soil weathering gradient. indeed, the ii au is almost poor in such findings because of the pedogenic demineralization. on the contrary, the i au is by far the richest in well mineralized fossiis, although often fragmented and rolled by rhe srream. the diagnostic macrofauna from the ca1 excavations is in fact mostly represented by isolated teerh, as more or less sorted within the sediments of the i au (fig. z) . diagnostic (at the genus level) non-denrary remains are only a distal fragment of the perone of a hippopot.tmus sp. and a basioccipital skul1 rvith the horns of a bison sp., coming from the base and rop, respectively, of the i au. a whole macromammal association, as has now been recognized in the lower unit, is represented by hippopotamus sp., sus sp., pseudodama sp., stephanorbinus cf.s. hundsheimensis (toula, 1l9a2), bison sp. and other proboscidae, bovini and cervidae indet. in the peratore quarry (c1), the previously reported finds (giraudi 1981) and other unpublished ones (ambrosettì pers. comm.) were from basal levels of the same i au: also dicerhorinus hemitoecbus'(recte: stephanorhinus hunrlsheimensis), megaceros cf. verticornis (recte: megaceroides cf. verticornis ) (dawkins, 1872) and [jrsus deningeri reichenau,1906. al1 these old and new findings from the i au may indicate a generic galerian (macro)mammal age (sensu gliozzi et al. 1997). an arvicolid microfauna has been samoled ar ca1 from two gravelly sandy levels of rhe i au, at the base and the top respectively (fig. 8). at the base of the unit mimomys saoini hinton, 1910, mìmomys pusillus (mehely, 1914), microtws (allopbaìomys) sp. advanced form, and ungaromys cf . u. ndnus a.re recorded. together with these, other rodents and insectivora occur: sciurus sp., apodemus sp. and talpa cf . t. fossilis pétenyi,1864. due to the presence of mimomys pusillus this microfauna is referable to the early biharian (rodent age), as defined by fejfar 8e horacek (1 990) . since -m. pusillus is accompanied also by an advanced form of microtus (allophaiomys),the possible range must be constrained ro tr late or final time of the early biharian. besides m. (allophaiomys), the (ìngaromys genus is a further biochronological marker in \le stern europe. either or both of these taxa occur rardily in rhe collecurti fauna (torre et al. nset and the vallonnet fauna (chaline 1988). by their own palaeomagnetic data both localities are calibrated in the jaramillo subchron as well as, in spite of some biogeographic differences, the northern locality of ijntermassfeld (kahlke 1992). jr (a/m) nrm n k rr,,.di chrm d,ld,lnk0,, ii'au |.19 10 148',49' 12 18 6 357",57' 357"57. 8 1 /0 1. lc+i" au 3.15 10 r j39",76' 26 5 11 6",57' 360',57" 1 1 27 9" messinian bedrock 2.69 10 , 10",5e" 2e l0 9' i 1",59' 3s3',57" 11 59 5. jl1 tab. i c. gìraudi, a. mottura, b. sala, m. s. siori &. d. bormiolt paleomagnetic results from sediments of cal successìon. jr : nrm intensitlr nrm natural remn:nt mrqnerìz:rion. chrm . chrr.rcrcri.ric r.n,,ìrn, rn:gnerization af,"rìi,r.r,ment. d, i : declination, inclination. dt, lt declìnation, inclìnation after tilt correcrion. n : number of specimens. k fisher's precision paramerer. 95' = semi-angle of confide'ce. the microfauna ar rhe rop of the i au is less represented because of the soil effects: apart from orcto/agus c[. /acosti tpomel, 1853), mimomis saoini and a .ingle m' of microtus sp. have been recorded. this latter (fig. 8, n. 6) shows poorly differentiated enamel and closely corresponds to homologous teerh of micrcttws (allophaiomys) from the same i au base (fig. 8, n.7) and from monte peglia (meulen van der 1973). this material by itself cannot permit a sufficiènt biochronologic assessment. neverthelesr. .rlong with a persi\rence ol normal mrgneric polarity up to rhe top of the unit, the cf. m. (attophaiomys) occurrence (but taking it with prudence) keeps here the correlation within the jaramillo subchron as well as, see mingly, the whole lithostratigraphic evidence (fig. 9). at ca1 the ar-vicolid elements collected from the ii aij are limited to some incisor reerh rhar do nor supporr any determination. the macrofauna there is represented only by fragmentary teerh of stepbanorhiuzs sp. and of herbivorous indet. this upper unir, because of the top soil, lacks exhaustive biochronologic markers and remains at moment unresolved in its own correlation with the possible palaeomagnetic optioht: the jaramillo subchron or alternatively the brunhes chron. the evolution of the setting and environmental inferences the above lithostratigraphic, palaeomagneric and biochronologic data allow us ro assume that at leasr the whole i au was deposited during the jaramillo subchron (1.07 0.99 m.y., according to the .orbitally tuned time" scale of cande & kent (1995)',. the lower complex was deposired beneath the i au under normal p.m. polarity but, due to the slightly unconformable contact between these units, one cannot exclude that the former deposition occurred well before the jaramillo subchron. the chronology of the ii au is by far more problematic. though this unit has a normal p.m. polarity, its unconformable contact over rhe i au misht be in favour of a somewhar younger age: ìf not in the same jaramillo subchron rhen necessarily within the brunhes chron. the age of this upper unit hence is possibly iower than 0.78 mr. in spite of the above gaps in the data, a preliminary reconsrrucrion of the geologic evolution in the area is suggested as below. after the ma1'or erosional phase supported by the top of the "messiniano 5sjimsnts (not yet deformed) due to its normal magneric polarity, at least the base of i au must fall necessarily inside the minor reversal of the jaramillo subchron (in the matuyama inverse age) and nor in the brunhes chron, since the arvicolid evolution level there precludes it. likewise, the association of m. savini, mimomys small size form, microtus (allophaiomys) advanced form, and (jngaromys cf. (j. nanus is very similar to rhat from monte peglia (meulen van der 1973), a site which has been recenrly located not lons be_ fore the jaramillo subchron (masini et al. 199g). fig. 5 equal areaprojection ofthe chrm directions (square = site mean value with alfa-95 ellipse of confidence). n--a--.. n-'-? r,qf tmcssinian bedrock lc + io ati iio au the early pleistocene castagnone site 1360 al i tgeo .tt . :î^l1fi^-' :. i i i j (ajm) 1 e+01 '1e-04 fig. 7 dental remains of large mammals îrom the i au ler.cls. 1: c, fragment oi hippopotamus sp.; 2: rjght m. ol stephanorhinus cl. huntlsheìmezsis, a) lingual viel., b) occlusal vieu'; 3: left p. of pseudodanta sp., e) lingual vìeu-, b) occlusal vicu'; 4: right nl, oí pseudodama sp., a) buccal viel'. b) occlusal vìer'; 5: rìght p,ol bíson sp., a) lingual vieq b) occlusal r.ìer.. scale bar: 3 cm. fig. 6 523 relation between magnetic intensity (j) and magnetic susceptibiìity (k). a) il'a.u. a : messinian bedrock t : i. a.u, + l,c. numbers refers to the position of the specimen in the succession. and during a subsequent time span under stable condislow colluvi.rl .rccrerion with soil co-evolurion occurred. tions, more probably before the jaramillo subchron, a this soil was characterized by the calcrete and carbonatic concreîions which heavily affect the lower complex. such pedogenic features have so far been recorded in the piedmont region only in the "viliafranchian> type succession (boano 8r forno 1996). after a further small erosional phase, the i au sedimentation took place along a water channel wirh high e nergy. in lhe -i,ljl" .,rt ^f tle i a u the energy decreased and a meandering pattern prevailed, with lateral migrations. due to the fast evaporation, along with the sand and silt aggradation, thick calcretes and concretions formed above the water table. towards rhe top ol the i au. à coarser deposition reappeared with energy renewal while the carbonrtic precipitation decreased and then stopped. the floodplain which developed during the jaramillo subchron was at the same time tectonically deformed, even though the pror-re\sive tiltinrrr" remainedb^r""'' compatibìe with the continued aggradation. this major landscape vrrirtion is hard to specify here at local detail but, on the whoie, it is certainly linked with the uplift of the north-easternmost monferrato ridge. w", @n, al w*, ót /'\ l"?ts\\ aa<\> n ffm8úb w 6789 f-\ /^\/ e) /\ u\, d a ?(,,>>r<"7\4"-r ,--k< \-1ì -..-v ==za"'z -\\ è )/t4 4 €. é= @[>è r-a>> ili \l 10 jl+ c. giraudi, a. mottura, b. sala, m. s. siori t< d. bormiolt fig. 8 an'ìcolid molar teeth from the i au levels at ca1. 1-5: m/1, microtus (allophaiomysl sp.; 6: m3/, microtus (?a/lopbaiomls); 7: m3/, microtus (allophaìontys) sp.; 8: m/2, llngaromys cf . ii. nanus;9:ml 1, mimomys pusìllus; 10: m/ 1, mimomys sa'c,itù (1,orng); 1 1 : m/ i, ùlimomys sacini (adùt) . scale bar: 1 mnt after a second erosional hiatus, the aggradation of the ii au is more recent than the offset of the major tectonic deformation event. therefore the hiatus itself contains the end of the deformation. the following desiccation of the ii au floodplain was possibly due to climatic changes or to a final but altogether different tectonic renewal. a similar pleistocene incision of the land surface is recorded from adjacent areas of central piedmont (carraro 8. valpreda 1991) in the late middle pleistocene. at this moment, such an age might reasonablv be the case also oi the reported situ.rtion. summary and conclusions the post-messinian sedimentary successron preserved in the hi11s near castagnone (pontestura) has been studied again by means of a network of suitable cuts carried out expressly for the purpose of lithostratigraphic, magnetostratigraphic and biochronologic assessment. this succession is formed by a lower complex of colluvial sediments with calcretes and carbonatic concretions of pedogenic origin, by a i alluvial unit of fluvial sediments with vertebrate macroand microfauna (most significant is the arvicolid association with two mimomys species together wrrh ungaromys cf . u. nanus and microtus (allophaiomys) sp. and by a ii alluvial unit of fluvial sediments with scarce faunal remains, in its turn covered by a thin loessic layer with embedded mousterirn rrtifacts of "vùrmiàn" age. the same succession, though broken off by erosional hiatuses, is entirely formed by sediments with normal palaeomagnetic polarity. a reliable chronological framework has been obtained for the i au which contains most of the faunal remains allowing for a direct correlation with the jaramillo p.m. subchron, between l,o70,ooo and qgo,ooo years ago. no chronological information results from the ol-ower complex" whose true age still remains uncertain. likewise, on the basis of the lithostratigraphic evidence alone, the upper and younger parts of the succession are apparently best referable to the brunhes chron. the sedimentary facies in the i au indicate a fluvial setting with energy decreasing from the bottom upwards, and a renewal of the flood energy at the top. the progressive tilting of the sedimentary beds in the lowermost succession indicates also a syn-depositional tectonic deformation, more marked during the i au deposition. such a deformation which tilted the fluvial strata southwards would not 3èerii to involve the ii au by itself. the tectonic stress over the castagnone area is seemingly related with the uplift of the north-eastern monferrato hills and, even though already noticeable in the earlier phases, it strengthened during the jaramillo time span then becoming nearly exhausted before the ii au deposition. as regards the time interval for the i a.u. mammal associations with four arvicolid species this must be limited to the late early pleistocene. the cross calibration between palaeomagnetic and palaeontological data makes it possible to refer the fossiliferous unit of the succession to the jaramillo subchron. together with the vallonet, ijntermassfeld and collecurti localities (chaline 1988; kahlke 1997:.torre et al. 1996), the castagnone site is one of the few european sites with a crucial and still poorly known early pleistocene tirne span. during the late early pleistocene, indeed, the major quaternary faunal turnover took p1ace, along with the climate cooling towards the coldest middle pleistocene glacial phases (gliozzi et al. 1997; markova & kozharinov 1998; rekovets 8r nadachowski 1995; sala et al. 1992:turner 1992). the reported succession thus represents rn improvement of the whole european biochronological time scale. the early pleistocene castagnone site fig. 9 biochronolog,v of the castagnone sitc in tbe frame of the italian pleistocene faunas (modified from masini er al. 1998) 525 acknou^ledgemen ts. \íe wish to thank p guarnero, owner of the reported quarries, for permission ro carry out the field excavatìons and support, o. girottì and d. esu ("la sapienza" university of rome) for palaeontological assessments of thc messjnian bedrock, dr. g. muttoni and an anonimous referee for the improvement of the manuscript. \ù/ork supported by the italian c.n.r (n. 96.003 /0.cto5;97 .a4fi1.cta5;98.00297 .ct05) and murst grants to the authors. refe,r birkeland p\( (1981) soils and geomorphology. oxford (uk): oxford university press. boano p & forno m. g. (1996) la successione tipo villafranchiana. litostratigrafia. in: (f. carraro ed.) revisione del villafranchiano nell'area-tipo di villafranca d'asti. 1/ quaternario, g(1): 38-62, roma. cande s.c. & kent d.v (1995) revised calibration o{ the geo*^--^':^ ^^l^-:'-time scale for the late cretaccous andl-,urdrrr) cenozoic./. geophys. res., 100: 6a%-6a95,'washington. carraro f. & \lilpreda e. (1991) the middle-upper quaternary of the asti basin. 1/ qwaternario, 4: 151-172, roma. c.g.i., 1969 carta geologica d'italia (1:100.000): foglio n'57, "verceì1i". sen'izio geologico d'italia i.g.m, firenze. ences chalinc j. (1988) les rongeurs de la grotte du vallonet (alpes-maritimes), l'environnemenr er l'age du site. lanthropologie, 92 (2), 497 -499, paris. fejfar o. 8c horacek l (1990) review of fossil arvicolids (mammalia, rodentia) of the pliocene and quarernarv of czechoslovakra. acta int. symp. e,lol. phyl. biostr. aruicolids, pp. 125-132, praha. giraudi c. (1981) presenza di depositi medio-pleistocenici in. tensamente deformati in val cerrina (monferrato settentrionale). geogr. fis. din. quat.,4: 69-71, torino. giraudi c. (1983) pontestura, loc. roletto (prov. alessandria). scavo paleontologico. quaderni della soprintendenza archeologica del piemonte 2: 1.43-1.44, torino. gliozzr e., ,\bbazzi l., argenti p., azza.roh a., caloi l., ca6 18 o ("/*) composite curve shackleton, 1995 5.0 4-5 4.0 3.5 3.0 f*,.i" 7 9 11 b 10 214+ ;_=,, <>f7 u: 20 - 1e ------3' zt,ì-=-23 )ó {> 2s-w: :+,€ 33 qs --è 35-lflpun" 4>:=1r^.4^1=__)i ?"s-;: 5b 6062o+ 6 687072 +sàq6 >__)f, €_"-,?9:-<,%" <*_y' (] 0 c a'r e c i! c e 0 c € c0 u lu 0,0 0.1 0.2 0.3 {, {, 0.5 i mammal i rodent lfaunall calibrated jaramillo i ltalian localities ages i zones i unitrl italian 'i:" l'î'' i .,i,'i.ì lirturvll ti"r!'fl' subchron i with calldrateo i microtus rf.|îî|:,f,:' i guonnaiomys) c e a f l,:,:*1',:,, uj z u o j è c .9or'. 0 i t 6 a 0 q 0.: oe (? t ro 0 c .!g cr: fo ! \ awicola i rur rtr il r -. i preffacanabnusl i'i,' f0nla na uccí f, u. isernia f, u. slivia fu, colle cu*i f.u. sauni i pirro f,u. fa rneta puaais i f u f.u, ".. 1 olivola f.u, minwmys prrocasrbs vitínia ìsotopic stage 7 torre in pìetra icnr..i. d:^è q f. ranuccio 0.458 klar blddittu etal. 1979 isernìa 0.730+/-0.04 k/ar cotort etal. 1981 vitinia f.u, 0.6 0.7 0.8 v =1.0 é ru 1.i ec r.2 : 1.3 s {e re> # ," rg'" e 1.7 1.8 l! : 1.e ! n 2,0 ll:?riíz castagnone (l a,u). m. (a.) sp. colle curti m, {a,) sp. m. peglia a m.(a.)nutiensis lvl. peglia b m,(a,)buryondíae soave c. sud m, {a,) ruffai pirro delì erba m, (a.) cf. rutroi pietrafittd m.{a)d,ntfui m. {a') chalínei pl. la n4esa m,{a)6.pliffie,1íc:/6 torre etal. 1996 le vallonet chaline 1988 u ntermassfeld kah ke 1997 ul z r.t i u f-l è u) u j 526 c. giraudi. a. mottura, b passo barbato l., di stefano g., esu d., ficcarelli g., girotti o., kotsakis t., masini f.,mazza p, mezzabotta c., palombo m.r., petronio c., rook l., sala b., sardella r.,zanalda e., ec torre d. (1997) biochronology of selected mammals, molluscs and ostracods from the middle pliocene to the late pleistocene on italy. the status .of the art. ritl. ital. paleont. strat., 1.03:369-388, milano. kahlke r.d. (1997) das pleistozàn von untermassfeld bei meiningen (thuringen). monogr. ròmiscb-germanisches zentralmuseum forschwngsinstitut fùr oorund frùhgeschichte,40/1.418 pp, bonn: rudolf habelt gmbh. masini f., abbazi l., lippi p, sala b. & torre d. (1998) review and new finds of microtws (allophaiomys) (rodenti. a ^.i.^ril".\ {"om the early pleistocene of the italian peninsula. paludicola, 2(1):75-9a, new york markova a. & kozharinov a. (i998) allophaiomys oî the southern russian plain. paludicola, 2(1): 62-69. new york. meulen a.j., van der, (1.973) middle pleistocene smaller mamsala, m. s. siori k d. bormioli mals from the monte peglia (orvieto, italy), with special reference to the phylogeny of microtus (arvicolidae, rodentia). qwaternaria,lt: 1-144, roma. rekovets l.i. ec nadachowski a. (1995) pleistocene voles (arvicolidae) of the ukraine. paleontologia i evolucio, 28-29: 145-245. sabadell. sala b., masini f., ficcarelli g., rook l. ec torre d. (1992) mammal dispersal events in the middle and late pleistocene of italy and western europe. courier forsch. senckenberg, 153: 59-68, frankfurt a m. sitec (1984) stwdio geologico e geomorfologico dell'area po 1. enel: internal report. torre d., albianelli 4., bertini a., ficcarelli g., masini f. & napoleone g. (1996) paleomagnetic calibration of pliopleistocene mammmal localities in central ialy. acta zool. cracov., 39 (1) : 559 -570, krakow. turner a. (1992) villafranchian-galerian larger carnivores of europe: dispersion and estinctions. courier forsch. senckenberg, l53: 153-160, frankfurt am m. rivista italiana di paleontologia e stratigrafia volume 117 no. 1 2 pls. pp. 29-37 april 2011 two species of profusulinella (p. aljutovica and p. ovata), early moscovian (pennsylvanian) fusulines from southern turkey and subdivision of primitive groups of the family fusulinidae fumio kobayashi received: december 6, 2010; accepted: january 10, 2011 institute of natural and environmental sciences, university of hyogo, sanda, hyogo 669-1546, japan. e-mail: kobayasi@hitohaku.jp key words: profusulinella, early moscovian, taurides, turkey. abstract. early moscovian (pennsylvanian) fusulines, profusulinella aljutovica and profusulinella ovata, from the hadim area, southern turkey are described systematically. they are contained in the bedded limestone (algal fusuline grainstone) of the yaricak formation of the aladag unit in the tauride block. morphologic analysis of these and similar species suggests: (1) aljutovella should be synonymous with profusulinella; (2) ovatella, depratina, staffellaeformes, aljutovella (elongatella), tikhonovichiella, skelnevatella, and priscoidella proposed in 1980’s and 1990’s are also synonymous with profusulinella; and (3) the families profusulinellidae and aljutovellidae are not necessary and profusulinella is included in the subfamily fusulinellinae placed under the family fusulinidae. riassunto. viene descritta la sistematica di due fusulinidi dal moscoviano inferiore (pennsylvaniano) profusulinella aljutovica e profusulinella ovata, provenienti dall’area di hadim, nella turchia meridionale. le specie provengono da un calcare stratificato (grainstone algale con fusuline) della formazione yaricak nell’unità strutturale aladag nei tauridi. l’analisi morfologica di queste forme e di altre specie similari suggerisce che: 1) il genere aljutovella dovrebbe essere sinonimo con profusulinella; 2) ovatella, depratina, staffellaeformes, aljutovella (elongatella), tikhonovichiella, skelnevatella e priscoidella, generi proposti negli anni 1980 e 1990 sono ugualmente sinonimi con profusulinella; e 3) le famiglie profusulinellidae e aljutovellidae non sono necessarie, con profusulinella inclusa nella sottofamiglia fusulinellinae, posta entro la famiglia fusulinidae. introduction fusuline faunas of the tauride block in the hadim area, southern turkey (fig. 1) have particular implications in relation to the paleogeographic location of the block on the gondwana margin facing the paleotethys sea (kobayashi & altiner 2008). devonian to triassic interbedded carbonate rocks and siliciclastic rocks referable to the aladag unit are widely distributed in the hadim area (altiner & özgül 2001). the serpukhovian, bashkirian, and moscovian limestones of the carboniferous yaricak formation of the aladag unit in the area are biostratigraphically subdivided into nine zones based on primitive fusulines such as eostaffella, pseudostaffella, profusulinella, and fusulinella (altiner & özgül 2001). profusulinella is diversified in the yaricak formation, with four species in the upper bashkirian and eight species in the lower moscovian reported by altiner & özgül (2001). in addition to them, four species of aljutovella were reported from the lower moscovian. these fusulines from the yaricak formation, however, have not been systematically described or illustrated. profusulinella and related genera are important in the early evolution of the family fusulinidae (e.g., thompson 1948; rauzer-chernousova et al. 1951). these forms also present serious taxonomic problems (ross 1999; villa et al. 2001; groves et al. 2007) mainly because solovieva in rauzer-chernousova et al. (1996) erected many new genera and subgenera that were placed under two new families (profusulinellidae and aljutovellidae). taxonomic opinions regarding these primitive fusulines vary widely among specialists. this paper systematically describes two species of profusulinella, p. aljutovica and p. ovata from lower 30 kobayashi f. moscovian (vereian) limestones of the yaricak formation in the hadim area, southern turkey. the taxonomy of profusulinella and similar taxa are discussed in conjunction with systematic description of these two species. the limestone sample used in this paper was collected from the hadim area on the occasion of the field excursion immediately after the conference on paleozoic benthic foraminifera (paleoforam 2001) held in ankara 20-24 august 2001. all the specimens herein described are registered with prefix d2 and stored in the museum of nature and human activities, hyogo, japan (fumio kobayashi collection, mnham). fig. 1 map showing the distribution of the tauride block and the hadim area, southern turkey. material and foraminiferal fauna in the hadim area, the serpukhovian is subdivided into three fusuline zones, the bashkirian into four, and the moscovian into two based on the first occurrences of zonal species (altiner & özgül 2001). profusulinella first appears in the fourth zone of the bashkirian (profusulinella zone) and ranges into the lower zone of the moscovian (eostaffella mutabilis profusulinella prisca eofusulina (paraeofusulina) zone). pseudostaffella antiqua, p. antiqua grandis, p. compressa, p. proozawai, profusulinella bona, p. parva, p. staffellaeformis, and p. rhomboides are reported from the bashkirian profusulinella zone. in addition to three zonal indicators, pseudostaffella praegorski, two species of neostaffella, eight species of profusulinella including p. primitiva and p. subovata, four species of aljutovella, and eofusulina triangula are characteristic of the lower part of the moscovian (altiner & özgül 2001). profusulinella and aljutovella were not reported from the upper part of the moscovian (fusulinella ex gr. bocki beedeina zone) according to altiner & özgül (2001). sample treated herein was collected from moscovian limestone of the yaricak formation exposed at the point (36º 54’ 44’’ n and 32º 23’ 24’’e), about 10 km sw of the town of hadim, southern turkey (fig. 1). it is highly fossiliferous and consists of algal fusuline grainstone with dominant foraminifers, subordinate red algae (ungdarella and komia) and problematic algae, and accessory brachiopods and crinoids. fusulines are mostly assignable to profusulinella aljutovica and profusulinella ovata. other genera are staffella, nankinella, eostaffella, and eoschubertella. the moscovian, consisting of the vereian, kashirian, podolskian, and myachkovian substages in ascending order, is biostratigraphically subdivided into 11 fusuline zones in the stratotype sections of the moscow syneclise (isakova 2002). eofusulina and evolved forms of neostaffella that first appeared in the kashirian in the type sections (isakova 2002) are not contained in the present material. non-fusuline foraminifers are assigned to bradyina, endothyra, planoendothyra, globivalvulina, biseriella, palaeotextularia, and spireitlina. based on the foraminiferal assemblage, the present sample is thought to be early moscovian (vereian) assignable to the lower part of the eostaffella mutabilis profusulinella prisca eofusulina (paraeofusulina) zone of altiner & özgül (2001). subdivision of primitive groups of the family fusulinidae rauzer-chernousova in rauzer-chernousova et al. (1951) proposed the genus aljutovella and assigned to it seven species and varieties formerly assigned to profusulinella, fusulinella, or fusulina, as well as 25 new species. aljutovella was distinguished from profusulinella by having characteristic “ячейки” referable to cells, meshes, alveoli, or other meanings in tangential sections that resemble the structure of the septa of the genus fusulina and the partial “поры” referable to pores in the wall of the outermost whorl. the former probably corresponds not to alveolar wall but to small cells or chamberlets formed by septal folding. the latter is found in kashirian forms (rauzer-chernosova et al., 1951, p. 21, fig. 8b). a porous wall under the tectum is also recognized in some species of profusulinella. thus, aljutovella in the original description is not clearly distinguished from profusulinella based both on slight differences of their wall structure and of an intensity and mode of septal folding in axial and polar regions in generic rank, though it might be possible in species rank. aljutovella has been widely accepted by russian workers (e.g., rauser-chernousova et al. 1951; bensh 1969; rozovskaya 1975; leven & davydov in leven et al. 2005) and by others outside russia (e.g., sheng 1958; van ginkel 1965; villa 1995). in contrast, it has been questioned by some workers (e.g., thompson 1964; loeblich & tappan 1988). ross (1999) showed that the porous wall of aljutovella is diagenetic feature commonly found in other poorly preserved fusulines in weathered zones. disagreement concerning the generic composition and classification of the family fusulinidae increased pursuant to the creation of many new genera and subgenera under the new families profusulinellidae and aljutovellidae by solovieva in rauzer-chernousova et al. (1996). the family profusulinellidae was erected to accommodate six genera, profusulinella, taitzehoella sheng, 1951, ovatella solovieva in rauzer-chernousova et al., 1996, depratina solovieva in rauzerchernousova et al., 1996, staffellaeformes solovieva, 1986, and moellerites solovieva, 1986. as indicated by early moscovian fusulines from southern turkey 31 villa et al. (2001) and groves et al. (2007), three species groups within profusulinella were reorganized by solovieva in rauzer-chernousova (1996) into ovatella, depratina, and staffellaeformes. groves et al. (2007) thought that moellerites was erected for the transitional forms from profusulinella to fusulinella. the family aljutovellidae consists of aljutovella (aljutovella), aljutovella (elongatella), tikhonovichiella, skelnevatella, and priscoidella according to solovieva in rauzer-chernousova et al (1996). these genera and subgenera were proposed by the reorganization of known species groups of aljutovella. profusulinella aljutovica elongata rauzer-chernousova, 1938 aljutovella tikhonovichi rauzer-chernousova in rauzer-chernousova et al., 1951 profusulinella skelnevatica putrya in putrya & leontovich, 1948 and profusulinella priscoidea rauzer-chernousova, 1938 were designated as the type species of aljutovella (elongatella), tikhonovichiella, skelnevatella, and priscoidella, respectively. three-layered wall structure (tectum and lower and upper tectoria) is clearly expressed in the original description of these type species. a diaphanotheca is partly developed in the terminal whorl of profusulinella priscoidea according to rauzerchernousova (1938). although shape and massiveness of chomata were added to the diagnostic features of the family aljutovellidae in rauser-chernousova et al. (1996), aljutovella and related forms are not easily distinguished from profusulinella. in my opinion, differences in the development of chomata, shape of the test and intensity of septal folding are expressed within and among populations, and these differences are insufficient to warrant the recognition of multiple genera and subgenera. the recognized generic composition of aljutovellidae by recent workers (e.g., isakova 2002; leven 2009) follows that by solovieva in rauzerchernousova et al. (1996). profusulinella aljutovica from the hadim area, shown in pl. 1, has a more elongate fusiform test and stronger septal folding than profusulinella ovata. the wall consists of thin distinct tectum and lower thicker protheca comparable to the lower tectorium of previous authors. a thin layer comparable to the upper tectorium is not always present. presence or absence, and thickness of the upper tectorium largely depend upon the state of preservation of specimens. broad morphologic variations are recognized in every test character as well as in those of profusulinella ovata illustrated in pl. 2. for example, the specimens shown in pl. 1, figs. 3 and 10 look like a form of “skelnevatella” in their inflated fusiform tests with pointed poles and massive chomata. furthermore, based on similar test characters, the specimens in pl. 1, figs. 8 and 13 appear to be a form of “tikhonovichiella”; and that in pl. 1, fig. 7 appears to be a form of “priscoidella”. obviously, these characters are highly variable and change continuously from specimen to specimen. differences among the 28 specimens illustrated are considered to only represent the intraspecific variation of profusulinella aljutovica. in conclusion, primitive fusulines in the present material are assigned to profusulinella and recognized as two species, p. ovata and p. aljutovica. aljutovella, ovatella, depratina, staffellaeformes, aljutovella (elongatella), tikhonovichiella, skelnevatella, and priscoidella are unnecessary names erected for what amount to species groups. all of them are thought to be junior synonyms of profusulinella. given this, the families profusulinidae and aljutovellidae are redundant, too. systematic paleontology suborder fusulinina wedekind, 1937 superfamily fusulinoidea von möller, 1878 family fusulinidae von möller, 1878 subfamily fusulinellinae staff and wedekind, 1910 fusulinellinae staff and wedekind, 1910, p. 112. profusulinellidae solovieva in rauzer-chernousova et al., 1996, p. 92. aljutovellidae solovieva in rauzer-chernousova et al., 1996, p. 95. genus profusulinella rauzer-chernousova and belyaev, in rauzer-chernousova et al., 1936 type species: profusulinella pararhomboides rauzer-chernousova and belyaev in rauzer-chernousova et al., 1936, p. 175. ovatella solovieva in rauzer-chernousova et al., 1996, p. 93 (type, profusulinella ovata rauzer-chernousova, 1938). depratina solovieva in rauzer-chernousova et al., 1996, p. 93, 94 (type, schwagerina prisca deprat, 1912). staffellaeformes solovieva, 1986, p. 20 (type, profusulinella staffellaeformis kireeva in rauzer-chernousova et al., 1951). aljutovella rauzer-chernousova in rauzer-chernousova et al., 1951, p. 182 (type, profusulinella aljutovica rauzer-chernousova, 1938). aljutovella (aljutovella) rauzer-chernousova; solovieva in rauzer-chernousova et al., 1996, p. 96. aljutovella (elongatella) solovieva in rauzer-chernousova et al., 1996, p. 96 (type, profusulinella aljutovica elongata rauzerchernousova, 1938). tikhonovichiella solovieva in rauzer-chernousova, 1996, p. 96 (type, aljutovella tikhonovichi rauzer-chernousova in rauzerchernousova et al., 1951). skelnevatella solovieva in rauzer-chernousova et al., 1996, p. 96 (type, profusulinella skelnevatica putrya in putrya and leontovich, 1948). priscoidella solovieva in rauzer-chernousova et al., 1996, p. 97 (type, profusulinella priscoidea rauzer-chernousova, 1938). 32 kobayashi f. discussion. all genera and subgenera listed above except for staffellaeformes were proposed for the typical forms of species groups of profusulinella or aljutovella in the systematic classification in rauzerchernousova et al. (1951). profusulinella staffellaeformis, type species of staffellaeformes was proposed, was included in the profusulinella parva group in rauzerchernousova et al. (1951). in my opinion none of these listed nominal taxa differs significantly from profusulinella so that all can be regarded as junior synonyms of profusulinella, as discussed above. the family aljutovellidae is accordingly unnecessary. moellerites proposed by solovieva (1986) with m. lopasniensis solovieva, 1986 as the type species might be synonymous with either profusulinella or fusulinella. the genus taitzehoella sheng, 1951 is distinct from profusulinella and is placed with profusulinella in the subfamily fusulinellinae of the family fusulinidae. therefore, the family profusulinellidae erected by solovieva in rauzer-chernousova et al. (1996) is also unnecessary. profusulinella aljutovica rauzer-chernousova, 1938 pl. 1, figs 1-28 1938 profusulinella aljutovica rauzer-chernousova, p. 97, 98, pl. 1, figs 10-12. 1951 aljutovella aljutovica (rauzer-chernousova); safonova and rauzer-chernousova in rauzer-chernousova et al., p. 193, 194, pl. 22, figs 1, 2. 1951 aljutovella conspecta leontovich in rauzer-chernousova et al., p. 195, 196, pl. 23, fig. 1. 1951 aljutovella arrisionis leontovich in rauzer-chernousova et al., p.196, 197, pl. 23, fig. 2. 1996 aljutovella (aljutovella) aljutovica (rauzer-chernousova); solovieva in rauzer-chernousova et al., p. 96, pl. 24, fig. 1. material: fifteen axial and thirteen sagittal sections illustrated, and others. description. test fusiform to inflated fusiform with arched to broadly arched periphery, almost straight lateral slopes, rounded to bluntly pointed poles. axis of coiling straight in general, but crossing at a large angle between inner lenticular and outer fusiform whorls in specimens. mature specimens with 4.5 tab. 1 measurement of profusulinella aljutovica rauzer-chernousova. plate 1 figs 1-28 profusulinella aljutovica rauzer-chernousova, 1, 3, 5, 7, 9, 17: ×50, others: × 25. 1: d2-025575, 2: d2-025601, 3: d2-025591, 4: d2025609, 5: d2-025623, 6: d2-025620, 7: d2-025553, 8: d2-025566, 9: d2-025620, 10: d2-025562, 11: d2025617, 12: d2-025607, 13: d2-025611, 14: d2-025562, 15: d2-025578, 16: d2-025545, 17: d2-025544, 18: d2025523, 19: d2-025537, 20: d2-025557, 21: d2-025574, 22: d2-025616, 23: d2-025584; 24: d2-025538, 25: d2025524, 26: d2-025584, 27: d2-025528, 28: d2-025530. 1 2 3 4 5 1 2 3 4 5 1 2 3 4 5 pl. 1, fig. 1 5.5 2.24 1.08 2.07 0.11 0.23 0.49 0.93 1.47 1.99 0.14 0.26 0.43 0.64 0.93 pl. 1, fig. 3 5 2.08 1.16 1.79 0.11 0.28 0.56 0.94 1.66 2.08 0.22 0.34 0.54 0.81 1.16 pl. 1, fig. 5 5.5 2.13 1.04 2.05 0.06 0.07 0.25 0.56 1.24 1.69 0.12 0.18 0.31 0.51 0.81 pl. 1, fig. 7 5.5 1.88 1.08 1.74 0.08 0.13 0.34 0.69 1.19 1.72 0.14 0.24 0.38 0.6 0.9 pl. 1, fig. 8 4.5 1.25 0.67 1.87 0.06 0.11 0.36 0.77 1.09 0.13 0.2 0.34 0.55 pl. 1, fig. 10 5 1.58 0.93 1.7 0.1 0.14 0.37 0.68 1.06 1.58 0.14 0.24 0.4 0.62 0.92 pl. 1, fig. 11 4.5 1.7 1.13 1.5 0.13 0.35 0.72 1.32 1.55 0.22 0.39 0.64 0.94 pl. 1, fig. 12 4.5 1.98 1.02 1.94 0.12 0.3 0.67 1.12 1.65 0.18 0.34 0.58 0.86 pl. 1, fig. 13 4 1.58 0.77 2.05 0.13 0.26 0.54 1.01 1.58 0.2 0.32 0.53 0.77 pl. 1, fig. 14 5.5 2.19 1.08 2.03 0.06 0.14 0.29 0.68 1.28 1.86 0.12 0.22 0.34 0.57 0.9 pl. 1, fig. 15 4.5 1.45 0.75 1.93 0.06 0.14 0.39 0.86 1.27 0.12 0.22 0.37 0.58 pl. 1, fig. 25 4.5 0.99 0.14 0.31 0.63 1.15 1.59 0.21 0.34 0.56 0.84 pl. 1, fig. 26 4.5 0.85 0.1 0.27 0.56 1.15 1.65 0.16 0.26 0.44 0.7 pl. 1, fig. 27 5.5 0.07 0.1 0.31 0.73 0.12 0.2 0.45 0.71 pl. 1, fig. 28 5 1.74 0.94 1.85 0.08 0.11 0.38 0.74 1.31 1.74 0.15 0.24 0.41 0.66 0.94 pl. 1, fig. 2 4.8 1.07 0.06 0.20 0.33 0.52 0.79 6 9 12 14 17> pl. 1, fig. 4 4.5 0.9 0.08 0.19 0.34 0.56 0.78 5 11 13 17 11> pl. 1, fig. 6 4.8 0.85 0.05 0.09 0.12 0.15 0.39 0.61 8 11 15 16> pl. 1, fig. 9 4.7 1.33 0.11 0.22 0.39 0.62 0.97 8 10 14 20 20> pl. 1, fig. 16 3.7 0.92 0.14 0.26 0.43 0.68 7 10 14 13> pl. 1, fig. 17 5.2 1.1 0.07 0.12 0.24 0.4 0.7 1.04 6 8 10 15 21 pl. 1. fig. 18 4.4 1.03 0.11 0.24 0.38 0.58 0.85 8 12 16 23 13> pl. 1, fig. 19 4.6 1 0.1 0.2 0.34 0.56 0.81 5 11 13 18 14> pl. 1, fig. 20 3.9 0.9 0.12 0.26 0.45 0.68 7 12 16 17> pl. 1, fig. 21 4.7 0.98 0.1 0.19 0.33 0.5 0.79 5 8 11 15 14> pl. 1, fig. 22 4.8 0.74 0.04 0.05 0.1 0.15 0.28 0.48 6 8 10 13> pl. 1, fig. 23 4 0.93 0.11 0.21 0.39 0.61 0.93 6 12 14 17 pl. 1, fig. 24 4.9 1.27 0.14 0.26 0.42 0.63 0.93 7 12 16 22 20> width of whorl number of septalength of whorlfig. in pl. no. whorl length width form ratio proloculus early moscovian fusulines from southern turkey 33 34 kobayashi f. to 5.5 whorls, about 1.5 to 2.2 mm in length, about 0.9 to 1.2 mm in width, with approximate length/width ratio from 1.5 to 2.1 (tab. 1). proloculus spherical to subspherical and 0.04 to 0.14 mm in its outside diameter. inner one or two whorls fusiform to eostaffelloid, and their length and width vary depending on the size of proloculus and an orientation of thin sections. beyond the second, whorls become fusiform with variable rate of expansion and form ratio, and shape of poles. length and width in corresponding whorls largely variable depended upon the size of a proloculus. length from the first to fifth whorls 0.05 to 0.35, 0.25 to 0.72, 0.56 to 1.32, 1.06 to 1.66, and 1.58 to 2.08 mm in 17 specimens. width from the first to fifth whorls 0.10 to 0.26, 0.15 to 0.45, 0.28 to 0.68, 0.48 to 0.97, and 0.90 to 1.16 mm in 28 specimens (tab. 1). wall thin, less than 0.05 mm in the thickest part of outer whorls, consisting of almost single layer in the eostaffelloid whorl, and distinct tectum, and lower thicker and upper thinner layers. upper thin layer corresponding to upper tectorium not continuous in most specimens. septa closely spaced and weakly fluted in polar regions of outer whorls. septal counts from the first to fifth whorls 5 to 8, 6 to 12, 8 to 16, 10 to 23, and 21 in 13 specimens (tab. 1). they attain 26 to 28 in their maximum in the fifth whorl. chomata present in almost all whorls and also on the proloculus in specimens with a large proloculus. their shape, size, and degree of symmetry through tunnel is variable. axial fillings absent. tunnel high and in general one-thirds to one-half as high as chambers. its path wider in outer test than in inner test in general. discussion. differences in size, shape, and expansion of the test, proloculus size, the number of septa, and development of chomata vary from specimen to specimen. therefore, they are thought to represent the intraspecific variation of this species originally described from the wells in samara bend by rauzerchernousova (1938). 1 2 3 4 5 6 1 2 3 4 5 6 1 2 3 4 5 6 pl. 2, fig. 1 4 1.57 1.04 1.51 0.17 0.42 0.77 1.24 1.57 0.26 0.48 0.72 1.04 pl. 2, fig. 2 4.5 1.06 0.78 1.36 0.12 0.15 0.35 0,61 0.88 0.18 0.26 0.43 0.67 pl. 2, fig. 3 5 1.55 1.1 1.41 0.09 0.18 0.41 0.77 1.18 1.55 0.18 0.29 0.49 0.77 1.1 pl. 2, fig. 5 6 1.53 1.18 1.3 0.05 0.08 0.44 0.78 1.13 1.53 0.14 0.2 0.32 0.48 0.75 1.18 8 pl. 2, fig. 7 5.5 1.3 1 1.3 0.09 0.1 0.25 0.5 0.81 1.19 0.14 0.24 0.36 0.56 0.83 pl. 2, fig. 8 5 1.31 0.89 1.47 0.08 0.26 0.62 0.94 1.31 0.15 0.24 0.37 0.61 0.89 7 pl. 2, fig. 12 4.5 1.44 1.03 1.4 0.12 0.28 0.58 1.02 1.31 0.23 0.37 0.6 0.86 pl. 2, fig. 13 4.5 1.45 1.01 1.44 0.1 0.24 0.44 0.81 1.23 0.19 0.32 0.55 0.85 pl. 2, fig. 14 4.5 1.44 0.97 1.48 0.11 0.2 0.38 0.82 1.22 0.19 0.31 0.51 0.8 pl. 2, fig. 15 5 1.4 0.99 1.41 0.1 0.18 0.37 0.68 1.08 1.4 0.18 0.28 0.46 0.68 0.99 pl. 2, fig. 16 4 1.29 0.91 1.42 0.13 0.3 0.6 0.93 1.29 0.21 0.38 0.6 0.91 pl. 2, fig. 17 5 1.35 1.06 1.27 0.09 0.14 0.29 0.61 0.99 1.35 0.19 0.32 0.38 0.75 1.06 pl. 2, fig. 18 5 1.3 0.93 1.4 0.09 0.13 0.29 0.58 0.98 1.3 0.16 0.25 0.4 0.64 0.93 pl. 2, fig. 19 4 1.22 0.94 1.3 0.15 0.25 0.52 0.89 1.22 0.24 0.39 0.61 0.94 pl. 2, fig. 20 4 1.35 0.95 1.42 0.16 0.32 0.61 0.98 1.35 0.24 0.39 0.64 0.95 pl. 2, fig. 21 5 1.24 0.91 1.36 0.09 0.14 0.3 0.54 1 1.24 0.16 0.25 0.41 0.64 0.91 pl. 2, fig. 22 4 1.17 0.88 1.33 0.13 0.22 0.52 0.91 1.17 0.22 0.36 0.6 0.88 pl. 2, fig. 23 6 1.43 0.97 1.47 0.05 0.06 0.38 0.71 1.05 1.43 0.1 0.18 0.28 0.46 0.69 0.97 8 pl. 2, fig. 24 4.5 1.15 0.8 1.44 0.07 0.3 0.61 0.96 0.15 0.25 0.43 0.66 7 pl. 2, fig. 31 5 1.5 1.06 1.42 0.08 0.15 0.35 0.79 1.19 1.5 0.16 0.27 0.48 0.74 1.06 pl. 2, fig. 32 5 0.99 0.08 0.63 0.94 0.16 0.27 0.44 0.67 0.99 6 5> pl. 2, fig. 33 6 0.88 0.05 0.07 0.44 0.76 1.05 0.12 0.14 0.26 0.42 0.64 0.88 pl. 2, fig. 34 5.5 1.29 0.89 1.45 0.36 0.69 1.01 0.29 0.47 0.72 pl. 2, fig. 35 5 1.31 0.84 1.56 0.06 0.31 0.58 0.92 1.3 0.15 0.23 0.36 0.57 0.84 6 pl. 2, fig. 4 4.1 0.9 0.09 0.22 0.39 0.62 0.89 7 11 12 18 3> pl. 2, fig. 6 4.4 0.76 0.07 0.16 0.29 0.45 0.64 5 8 10 14 6> pl. 2, fig. 9 4 1.07 0.14 0.28 0.46 0.74 1.07 6 12 16 21 pl. 2, fig. 10 5.1 1.19 0.06 0.16 0.29 0.48 0.77 1.19 4 7 12 18 23 3> pl. 2, fig. 11 4.8 1.2 0.1 0.2 0.33 0.55 0.87 7 12 14 18 16> pl. 2, fig. 25 4.7 1.15 0.1 0.22 0.38 0.6 0.89 6 12 13 19 17> pl. 2, fig. 26 3.9 0.79 0.09 0.18 0.32 0.56 7 11 13 18> pl. 2, fig. 27 4.1 0.93 0.14 0.22 0.4 0.6 0.88 7 11 13 19 3> pl. 2, fig. 28 4.4 0.73 0.05 0.12 0.22 0.38 0.6 5 8 10 14 7> pl. 2, fig. 29 5.1 1.01 0.08 0.11 0.16 0.27 0.44 0.7 1 8 12 14 17 3> pl. 2, fig. 30 4.2 0.86 0.09 0.19 0.33 0.55 0.84 6 10 11 17 4> number of septalength of whorl width of whorlfig. in pl. no. whorl length width form ratio proloculus tab. 2 measurement of profusulinella ovata rauzer-chernousova. plate 2 figs 135 profusulinella ovata rauzer-chernousova, 1, 3, 5, 7, 10, 11: ×50, others: × 25. 1: d2-025552, 2: d2-025614, 3: d2-025568,, 4: d2025584, 5: d2-025558, 6: d2-025619, 7: d2-025541, 8: d2-025615, 9: d2-025533, 10: d2-025599, 11: d2025604, 12: d2-025595, 13: d2-025560, 14: d2-025534, 15: d2-025555, 16: d2-025532, 17: d2-025619, 18: d2025600, 19: d2-025548, 20: d2-025549, 21: d2-025582, 22: d2-025535, 23: d2-025528, 24: d2-025557, 25: d2025566, 26: d2-025619, 27: d2-025621, 28: d2-025544, 29: d2-025592, 30: d2-025582, 31: d2-025603, 32: d2025572, 33: d2-025610, 34: d2-025539, 35: d2-025540. early moscovian fusulines from southern turkey 35 36 kobayashi f. considering wide morphologic variation of the present material, at least the following two species are probably conspecific with this species: aljutovella conspecta leontovich in rauzer-chernousova et al., 1951 and a. arrisionis leontovich in rauzer-chernousova et al., 1951 both of which were included in the aljutovella aljutovica group by leontovich in rauzerchernousova et al. (1951). aljutovella skelnevatica (putrya in putrya and leontovich, 1948), a. cybaea leontovich in rauzer-chernousova et al., 1951, and a. artificialis leontovich in rauzer-chernousova et al., 1951 were included in the aljutovella skelnevatica group by leontovich in rauzer-chernousova et al. (1951). they might or might not be conspecific with a. aljutovica. profusulinella ovata rauzer-chernousova, 1938 pl. 2, figs 1-35 1938 profusulinella ovata rauzer-chernousova, p. 101, pl. 1, figs. 14-16. 1951 profusulinella subovata safonova in rauzer-chernousova et al., p. 164, pl. 14, figs 5, 6. 1996 ovatella ovata (rauzer-chernousova); solovieva in rauzer-chernousova et al., p. 93, pl. 23, fig. 3. material: twenty-three axial, eleven sagittal, and one tangential sections illustrated, and others. description. test inflated fusiform to oval with arched to broadly arched periphery, rounded poles. axis of coiling straight in general, but crossing at a large angle between inner lenticular and outer fusiform whorls in specimens. mature specimens with 4 to 5.5 whorls, rarely 6. length about 1.1 to 1.6 mm and width about 0.8 to 1.2 mm in width giving approximate length/width ratio from 1.3 to 1.6 (tab. 2). proloculus spherical and 0.05 to 0.17 mm in its outside diameter. inner one to two whorls vary from eostaffelloid to inflated fusiform, and their length and width vary depending on the size of proloculus. outer whorls inflated fusiform to oval with variable rate of expansion and form ratio. length and width in corresponding whorls largely variable depending upon the size of proloculus. length from the first to sixth whorls 0.06 to 0.42, 0.25 to 0.77, 0.36 to 1.24, 0.69 to 1.57, 1.01 to 1.55, and 1.43 to 1.53 mm in 24 specimens. width from the first to sixth whorls 0.10 to 0.26, 0.14 to 0.48, 0.26 to 0.74, 0.42 to 1.07, 0.64 to 1.19, and 0.88 to 1.18 mm in 35 specimens (tab. 2). wall thin, less than 0.04 mm in the thickest part of outer whorls, appears to be thicker due to secondary coating of dark layer. it is almost structureless in the eostaffelloid one or two whorls. in later whorls it exhibits a distinct tectum, and lower thicker and upper thinner layers. upper thinner layer is discontinuous and indistinct in most specimens. septa closely spaced and almost plane to very weakly fluted in polar regions of outer whorls. septal counts from the first to fifth whorls 4 to 7, 7 to 12, 10 to 16, 14 to 21, and 17 to 23 in 17 specimens (tab. 2). chomata massive, roughly symmetrical through tunnel, and well developed in inner fusiform whorls. they are present on the proloculus and eostaffeloid whorls, but tend to be indistinct or absent in outer fusiform whorls. their shape and size are variable. axial fillings absent. tunnel high and probably one-thirds to one-half as high as chambers. its path becomes wider outwards in general. discussion. this species is discriminated from profusulinella aljutovica by their smaller and more inflated fusiform test in general. proloculus size, the number of whorl, height and width of inner whorls, and development of chomata vary from specimen to specimen, showing wide morphologic variation in the hadim specimens. specimens having relatively small proloculus resemble rauzer-chernousova’s (1938) original material from the upper part of the vereian (lower moscovian) of samara bend. those with larger proloculi (e.g., pl. 2, figs 1, 14, 15) are more similar to profusulinella subovata safonova in rauzer-chernousova et al., 1951. they are probably conspecific each other. those having subspherical test and tightly coiled inner whorls (e.g., pl. 2, figs. 5, 10) appear to be more like profusulinella prisca sphaeroidea rauser-chernousova in rauzer-chernousova et al. (1951) and profusulinella prisca timanica kireeva in rauzer-chernousova et al. (1951) than to profusulinella ovata. although the range of morphologic variation in these two subspecies is unclear, they should be included into the profusulinella prisca group, as done by rauzer-chernosova et al. (1951). acknowlegements. i am much indebted to demir altiner for his generous and efficient field guidance in the hadim area during the post-conference field excursion of paleoforams 2001, to maurizio gaetani for his careful editing the manuscript, and to atsuko ujimaru for her help drawing figures. many thanks are due to two reviewers, elisa villa and john r. groves for their helpful comments and suggestions, from which this paper is greatly improved. early moscovian fusulines from southern turkey 37 altiner d. & özgül n. (2001) carboniferous and permian of the allochthonous terranes of the central tauride belt. paleoforam 2001: intern. confer. paleozoic benthic foraminifera, guide book, 35 pp., ankara. bensh f. r. (1969) stratigraphy and foraminifers from the carboniferous in the southern gissar mountains. inst. geol. geofiz., izd. fan uzbeskoi ssr, 1-174 (in russian). deprat j. (1912) 1912: étude géologique du yun-nan oriental. étude des fusulidés de chine et d’indochine et classification des calcaires à fusulines. mém. serv. géol. l’indo-chine, 1: 1-76. ginkel a. c. van (1965) carboniferous fusulinids from the cantabrian mountains (spain). leidse geol. med., 34: 1-225. groves j. r., kulagina e. i. & villa e. (2007) diachronous appearances of the pennsylvanian fusulinid profusulinella in eurasia and north america. j. paleont., 81: 227-237. isakova t. n. (2002) moscovian foraminiferal biostratigraphy in the type region (russia). in: hills l. v., henderson c. m. & bamber e. w. (eds) carboniferous and permian of the world, mem. canadian soc. petrol. geologists, 19: 448-460. kobayashi f. & altiner d. (2008) late carboniferous and early permian fusulinoideans in the central taurides, turkey: biostratigraphy, faunal composition and comparison. j. foram. res., 38: 59-73. leven e. ja. (2009) the upper carboniferous (pennsylvanian) and permian of the western tethys: fusulinids, stratigraphy, biostratigraphy. trudy geol. inst., 238 pp. (in russian). leven e. ja., davydov v. i. & gorgij m. n. (2005) pennsylvanian stratigraphy and fusulinids of central and eastern iran. paleont. electronica, 9: 1-34, http://palaeo-electronica.org. loeblich a. r. jr. & tappan h. (1988) foraminiferal genera and their classification. van nostrand, 2 vol., 970 pp. plus 212 pp. and 847 pls. new york. möller v. von (1878) die spiral-gewunden foraminiferen des russischen kohlenkalkes. mém. l’acad. imp. sci. st. pétersbourg, sér. 7, 25: 1-147. putrya f. s. & leontovich g. e. (1948) toward the study of middle carboniferous fusulinids in the volga region of saratov. byull. mosk. obsh. ispyt. pri. otd. geol., 23: 11-45 (in russian). rauzer-chernousova d. m. (1938) upper paleozoic foraminifera of the samara bend and the trans-volga region. trudy geol. inst., akad. nauk sssr, 7: 69-160 (in russian). rauzer-chernousova d. m., belyaev g. m. & reitlinger e. a. (1936) upper paleozoic foraminifera from the pechora territory. trudy polyarnoy komissii, akad. nauk sssr, 28: 159-232 (in russian). rauzer-chernousova d. m., kireeva g. d., leontovich g. e., gryzlova g. d., safonova t. p. & chernova e. i. (1951) middle carboniferous fusulinids of the russian platform and adjacent regions. akad. nauk sssr, inst. geol. nauk, 371 pp., moskva (in russian). rauzer-chernousova d. m., bensh f. r., vdovenko m. v., gibshman n. b., leven e. ja., lipina o. a., reitlinger e. a., solovieva m. n. & cheduya i. o. (1996) reference-book on the systematics of paleozoic foraminifera (endothyroidea, fusulinoidea). moskva nauka, 207 pp., moskva (in russian). ross c. a. (1999) classification of the upper paleozoic superorders endothyroida and fusulinoida as part of the class foraminifera. j. foram. res., 29: 291-305. rozovskaya s. e. (1975) composition, phylogeny and system of the order fusukinida. trudy paleont. inst. akad. nauk sssr, 149: 1-267 (in russian). sheng j. c. (1951) taitzehoella, a new genus of fusulinids. bull. geol. soc. china, 31: 79-85 (in chinese and english). sheng j. c. (1958) fusulinids from the penchi series of the taitzeho valley, liaoning. pal. sinica, 143: 1-119 (in chinese and english). solovieva m. n. (1986) fusulinid zonal scale of the moscovian stage from the restudy of stratotype materials of intrastage subdivisions. voprosy mikropal., 28: 3-23, izdatelustvo nauka (in russian). staff h. von & wedekind r. (1910) der oberkarbon foraminiferensapropelit spitzbergens. bull. geol. inst. univ. uppsala, 10: 81-123. thompson m. l. (1948) studies of american fusulinids, protozoa article 1. univ. kansas pal. contr., 184 pp. thompson m. l. (1964) fusulinacea. in: moore, r. c. (ed) treatise on invertebrate paleontology, part c, protista 2: c358–c436, geol. soc. am., boulder & univ. kansas press, lawrence. villa e. (1995) fusulinaceos carboniferos del este de asturias (n de espana). univ. claude bernard-lyon, biostrat. paléozoique, 13: 1-261. villa e., sánchez de posada l. c., fernández l. p., martínezchacón m. l. & stavros c (2001) foraminifera and biostratigraphy of the valdeteja formation stratotype (carboniferous, cantabrian zone, nw spain). facies, 45: 59-86. wedekind p. r. (1937) einführung in die grundlagen der historischen geologie. band 2. mikrobiostratigraphie die korallenund foraminiferenzeit. ferdinand enke, 136 pp., stuttgart. r e f e r e n c e s rivista italiana di paleontologia e stratigrafia volume lut numero 3 pagrne jb/--./4 dicembre 1996 comparison bet\teen in larger cladistic and phenetic methods foraminifera analysis fabrizio maia key-uords: miogypsinidae, phenetic analysis, cladistic analysis. riassunto. oggetto di questo lavoro è l'analisi numerica dei miogypsinidi, macroforaminiferi presenti nel1e facies terrigene e bioclastiche alternate a facies pelitiche della collina di torino e del monferrato, nell'intervallo oligocene sup. miocene inf. in accordo con la letteratura, 1o studio dei miogypsinidi si basa sulla valutazione dei parametri biometrici relativi all'apparato embrionale e nepionico. i dati ottenuti dalle misurazioni vengono trartati statisticamenre e, successivamente. viene e{fettuato un con{ronto con i dati relativi alle unità tassonomiche riconosciute in letteratura. alcune attribuzioni specifiche ottenute in questo modo appaiono poco affidabili: ciò significa che, in diversi casi, l'elaborazione statistica tradizionale risulta inefficace. al fine di superare tale problema e rrovare nuovi metodi non soggettivi per assegnare nuove popolazioni alle rispettive specie, vengono esaminate alcune tecniche fenetiche e cladistiche. nel presente lavoro vengono quindi confronrari vanraggi e difficoltà nell'applicare 1'analisi delle componenri principali, l'analisi dei clusters, i'analisi discriminante e l'analisi filogenetica col sistema della parsimonia. lanalisi discriminante sembra fornire i risultati più interessanti. abstract. the analysis is {ocused on miogypsinidae, larger foraminifera characterizing the terrigenous and bioclastic facies that alternate with pelitic facies in the turin hill and in the monferrato area (l'{lv ltaly) from upper oligocene to early miocene. according to the literature, the study of miogypsinidae is based on biometry of embryonic and nepionic apparatus characters. measurements are processed statistically and comparison is also made with the values for taxonomic units recognized in literature. cenain specific determinations so obtained result ambiguous: this means that, in many cases, the traditional statistical elaboration appears to be inefficient. to overcome this problem and to find a new method to àssrbn new populations to their species in a non-sub.jective way, an artempt is made to use either phenetic or cladistic systems. in this work there is a comparison between the advantages and the difficulties in using techniques like principal components analysis, cluster analysis, discriminant analysis and phylogenetic analysis using parsimony. discriminant analysis seems to provide the best results. lntroduction. miogypsinidae are benthic polythalamic non-sessiles larger foraminifera, belonging to the super-family of orbitoidacea. they originate in late oligocene and disappear in burdigalian. each specimen is sectioned and studied in equatorial plane, then the data of the specimens of a sample are collected to obtain an average of the whole population, according to drooger (1952) and subsequent papers. attribution of a population to a certain species is based on biometry of embryonic apparatus parameters pertinent to the juoenarium (protoconch and deuteroconch) and to the nepionic protoconchal spirals (fig. 1). these parameters are the main spiral ienght (x), the y angle between the medio-embryonic line and the frontal-apical line, the symmerry of the nepiont (v), the protoconchal diameter (d1), the deuteroconchal diameter (d2), the ratio d2/dl, the distance between the center of the protoconch and the apical margin (e), the ratio t/dt. ffi=t y = 200a/9 fig. 1 schematic drawing showing the measures of the internal features in embryonic-nepionic stages of miogypsina. meaning of the symbols: fa : frontal-apical line; me : medio-embryonic line; cx, : angle made by the shortest spiral around the protoconch; b : angle made by both spirals around the protoconch; y : angle between me and fa lines; dl : diameter of the protoconch; e : distance bets/een the cenrer of the protoconch and the apical margin; x : number of chambers of the principal spiral around the protoconch; v : degree of simmetry of the nepiont. dipanimento di scienze della terra via accademia delle scienze, 5 10123 torino 368 f. maia species samples x 8 d1 € €.|d1 m. gunteri m. gunteri m. gunteri m. gunteri m. gunteri m. gunteri-tani m. gunteri-tani m. gunteri-tani m. guntèri-tani m. gunteri-tani m. guntefi-tani m. tani m. tani m. tani m. tani m. tani m. tani m. tani m. tani m. tani m. globulina m. globulina m. globulina m. globulina m. globulina m. globulina-intermedia m. globulina-intermedia m. globulina-intermedia m. globulina-intermedia m. globulina-intermedia m. globulina-intermedia m. globulina-intermedia m. socini m. socini m. socini m. socini m. socini m. socini-burdigalensis m. socini-burdigalensis m. burdigalensis m. burdigalensis m. burdigalensis m. negrii m. negrii m. negrii superga 1 civera 7 bis monferrato mc18 monferrato mc12 monferrato mc2 superga 33 bis baldissero 28 baldissero 34 monlerrato mu1 monferrato pc2e monferrato pc2c monferrato ro monferrato vm monferrato sm3 monferrato sm2 monferrato sm1 monferrato sb1 monferrato fb2 monferrato fa2 monferrato fal superga 93 civera 10 civera 13 civera 16 monferrato cal b superga 84 civera 29 civera 36 civera 961 baldissero 68 monferrato m10 monferrato mb superga 43 bric palouch 3a rivodora 3f rivodora 4 monferraio vdmu superga 52 baldissero 48 baldissero 41b vergnana 1 brlc palouch 3b superga 72 monferrato cal a monterrato rs t 10.44 '10.43 9.40 9.90 9.23 9.16 9.63 9.32 8.52 9.18 8.50 ó.oy 8.07 8.50 7.oo 7.97 7.91 8.40 6.95 6.17 6.42 o.ou 5.88 5.40 5.78 a.47 8.92 ó.oj 8.43 10.00 9.21 7.09 5.16 -101 .06 -109.00 -49.00 -78.00 43.00 -46.79 43.34 -49. 1 6 -24.oo -27.oo -34.00 -19.00 { 8.00 -13.00 -20.00 17.00 -30.o0 -15.00 44.oo 38.oo 't 6.48 27.o0 19.17 15.30 18.45 29.69 34.50 30.50 25.50 27.80 35.71 -1 0.1 0 42.90 -17.63 -38.00 -63.00 -52.20 4.83 4.69 1.e9 4.76 7.22 5.04 6.90 3-70 8.60 8.40 f1.00 12.10 6.50 5.60 '10.60 35.47 27.22 24.5'l 23.79 24.20 44.13 48.19 40.28 52.'14 44.43 42.50 40.60 19.87 14.43 'i 8.23 17.30 28.18 '13.64 30.99 29.83 38.61 68.10 75.90 55.30 182 151 '155 160 181 189 172 155 161 177 178 163 190 171 | /o '168 163 137 150 182 toz 141 164 185 189 184 172 '168 172 '141 157 too 174 188 183 3'12 zc9 255 254 257 308 241 251 267 253 257 269 268 261 261 270 351 213 232 241 262 273 250 292 311 312 309 3'12 309 320 301 1.74 1 .75 1.67 1.70 1.60 1.64 1.61 1.81 1.48 1.57 1.58 1 .50 1.57 1 .52 1.62 1 .50 1 .56 '1.68 2.10 '1.58 1.70 1.55 1.49 1.71 '1.75 lao i oo 1.70 '1.58 1.68 1.72 1.81 1.88 2.09 1 .71 1.91 2.06 1.70 during the evolution of this group, parameter changes permit to reconstruct an hypothetical phylogenesis. up to now; the determination of one population of miogypsinidae was based on the comparison of its parameters with the values for taxonomic units recognised in literature. student's "t" test was used to compare the specific determinations thus obtained with all the populations assigned to these forms by other workers and hence confirm our species attribution. the aim of this work is to find identification methods more reliable to assign a new population to a predetermined taxon, using more objectivity as possible and taking advantage of all the measurement collected. such methods have been identified in the realm of phenetic analysis (barrai, 1984; camussi et al., 1986; dunn & everitt, 1982; elliott, 1'977) and cladistic analysis (fofey ef al., t992). the input data for the analysis correspond to the mean values of the populations studied in piedmont (tab. 1) and those of the populations reported in literature (tab. 2). on the base of available measurements, for each analysis we used the one or the other data set. the species considered are those from m. gunteri to m. glotab. tr mean values of counts and measuremenìs on internal characteristics of miogypsina populations studied in piedmont. bulina-intermedia along the main branch of the phyletic tree, and those from m. socini to m. negrii along the lateral branch. \fle have chosen to use the mean values rather than individual values, because the objects of our study are the populations rather than the individuals that compose the populations. the mean values have been standardized before exploiting phenetic analysis: standardization consists in subtracting the mean of each variable from each value and dividing the difference by the standard deviation. a true incentive to undertake this work has come from the development of computer packages suitable for many types of numerical anaiysis. among them, we have selected ntsys-pc (a software for phenetic analysis), statgraphics (statistics) and paup (cladistic analysis). about the phenetic analysis, in this work we will describe the principal component analysis, the cluster analysis and the discriminant analysis. about cladistic analysis, we will discuss the method of maximum parsimony, used to infer the phylogenetic ree of miogypsinidae from their characters. first the principle of simiiarity will be introduced. species references samples m. gunteri m. gunteri m. gunteri-tani m. gunteri-tani m. tani m. tani m. tani m. tani m. tani-globulina m. tani-globulina m. tani-globulina m. tani-globulina m. globulina m. globulina m. globulina m. globulina m. globulina m. globulina m. globulina m. globulina m. globulina m. globulina m. globulina m. globulina-intermedia m. globulina-intermedia m. globulina-intermedia m. globulina-intermedia m. globulina-intermedia m. globulina-intermedia m. intermedia m. intermedia m. intermedia m. intermedia m. intermedia m. intermedia m. socini m. socini m. socini m. socini m. socini m. socini m. socini-burdigalensis m. burdigalensis m. burdigalensis m. burdigalensis-negrii mneorii drooger,1954(a) ferreìo, 1 965 drooger, 1 952 drooger et al., 'l 955 drooger, 1 952 drooger et al., 1 955 delicati & schiavinotto, 1985 vilizzi, 1991 drooger, 1 952 raju,1974 de mulder, 1975 fermont & troelstra, 1983 drooger, 1 952 drooger,1954(a) drooger, 1 954 (b) drooger et al., 1955 ujiié & oshima, 1969 matsumaru,1971 raju, 1 974 de mulder, 1975 schúttenhelm, 1976 schiavinotto, 1979 delicati & schiavinotto, 1985 drooger, 1 952 drooger, 1 954(a) drooger et al., 1955 de mulder, 1975 schúttenhelm, 1 976 wildemborg, 1991 drooger et al., 1955 de mulder, 1975 schúttenhelm, 1976 schiavinotto, 1985(a) schiavinotto, 1 985 (b) wildemborg, 1991 drooger, 1 954 (a) vervloet, 1 966 schùttenhelm, 1976 de bock, 1977 schiavinotto, 1979 delicati & schiavinotto, 1985 schúttenhelm, 1976 schùttenhelm, 1976 schiavinotto, 1979 schiavinotto, 1979 schiavinotto, 1979 1l 1 11a 22a',34t mor222 12 2b; 1 3-1 5; 1 5a; 1 6-1 8; m1 -6; 26" pmt16; pmtt ot2; ot4 18 g1 437 a179; dm363 80pc02 19-22 5;6;10;18 240b 1 ; 2: 5; 2oa; 24-25; 35-36; 4344; m7 shuk.a ch; ho kr36-b; g1401-b; g1401 ab; g1406-8; g1 40648; g1 421 -b; jag.w.b dm114; dm116; dm117 sm-281 87 181 1268 11 57 al 1 1 o | 1 31 | 1 35 i 1 66 | 251 i 248 i 46e i 1 40 i 1 se | 347 tls76 pmt-6/80 24;25 7', 8;9 22a';29i 3 4194; dm106; dm608 sm-2214811258 jts1 24; jt5063 2'l dm684; dm107; dm140 sm-2o51237144o ac5 ca82 jî -7980151 1 7 151 1 2 151 07 17 97 8 17977 1797 61 7 97 5 ls1 02 | s09 8 l5o9 4 17 97 4 17 1 7 7 i 5077 12 13 326-3 sm482 | 483 1234 i 267 i 447 /266 | 1 1 6 | 1 24 | 287 i 444 1283 11 881 1 87 l't 84 12261227 1298 m13 tlsl 07 pmt3; pmt2 sm-233/373 sm4781479 tls-42/39 tls-1 0s/3s tls1 1 o cladistic and phenetic metbods 369 fr ll t i similarity. the principle of similarity is essential to deal with some problems related to the principal component analysis and the cluster analysis. similarity is the resemblance or affinity among the taxonomic units, based on their characters; in other words, it's their phenetic relationship. the complement of the similarity of taxonomic units is their dissimilarity or phenetic distance, measured by means of processes that satisfy mathematical properties which make them particuiarly suitable for phenetic analysis. among the available measures of dissimilarity, in this work we used the average taxonomic distance. its expression is similar to that of euclidean distance, resulting from the pythagora's theorem. tab.2 populations oí miogypsina re.^r"j i. lit"r.r'qrs and considered in this work. see ferrero et al. (1992, 1994) íor references that are not cited at the end of this work. principal component analysis. the principal component analysis (pca) is useful to find new variables that are linear combinations of the original measures and describe the sample without the abundance of information rising from correlation among the original measures. the new variables have to be uncorrelated, so that it is possible to choose those showing the greatest variance. the first few measures that account for most of the variation of the sample correspond to the principal components. the results of this transformation can be better explained by calculating the correlation among the original measures and the new variables, so that a loading matrix is obtained in which the absolute value and the sign of the correlations allow to understand the connections among measures and principal components. a geometrical interprepri component eigenvalue percentage of variance cumulative percent 1 2 4 2.71035 1.08278 0.1 551 3 0.0s173 67.76 27.07 3.88 1.29 67.76 94.83 98.71 100.00 370 f. maia matrix parameter comp.1 comp. 2 comp.3 x 8 e ld1 o.974 -0.962 -0.914 o.027 -0.115 0.1 10 -o,267 -0.993 -0,080 0.210 -0.303 0.1 14 tab.3 results of principal components analysis. the eigenvalues (atent roots) are proportional to the variance accounted for by each of the first four components. the component loadings (atent vectors) for the first three principal components are shown in the loading matrix, in which it appears that the first component loads heavily on x, 1 and v, while the second loads heavily on e/di. tation of the pca is possible if we imagine to find the axis (or dimension) that express the variation of the characters, that is the axis which maximizes the variance of the proìections of the values onto itself. this axis is given by the line that minimizes the sum of squares of the distances between the values and itself. if there are "p" characters, the first principal component is the bestfitting straight line in the p-dimensional space. a very useful visualization of the results of pca is given by a scatterplot of the first component score for each taxonomic units against the second. how well this scatterplot describes the configuration in the original p-dimensional space may be measured by the proportion of the variance in the data acccounted for by the first two principal components. the analysis has been carrred out either on the data of the populations reported by literature or on the data of the populations studied in piedmont. in both cases the first two principal components account for more than 90olo (cumulative) of the total variance. only the results concerning the populations studied in piedmont are shown (tab. 3). the loading matrix shows the respective significance of each of the originai measures (x, y, v and e/d1) in making the components. in the scatterplot based on the first two principal components (fig. 2), the groups of the popuiations belonging to each species appear well distinct. to verify how weli this two-dimensional mapping preserves the original distances among populations (measured with the aver^ge taxonomic distance, previously discussed), we can use the minimum spanning componènt 2 1|i i i l'4 i og component 1 tr m. qunteri o m. elobulinalntermedia u m. gunteri-tani o m, socini i m. tani o msocinl-burdigalensis o m. globulina a m. burdigalensls fig.2 plot of rhe first two principal componenr scores for populations studied in piedmont. tree (lt4st) of the distance matrix. the spanning tree is a set of straight-line segments joining pairs of points such that no closed ioops occur, each point is touched by at least one line and the tree has continuous link between any pair of points. if a weight is assigned to each segment, than the lenght of the tree is defined to be the sum of these weights. the mst is defined as the spanning tree of minimum lenght. this tool helps to detect local distorsions in the diagram resulting from the ordination technique, like the case of pairs of populations which look close together in the two-dimensional representation, but actually are far aparf íf other dimensions are taken into account. for example, in the scatterplot of the pca applied to the data of piedmont (fig. 2), populations sup52 and bal48, belonging to m. socini-burdigalensis, appear to be well separated from nm-gunteri om.globulina-intermedia ta m. gunteri-tani q m. socini i m. tani o m, socini-burdigalensis o m. slobulina a m. burdigalensis fig. 3 minimum spanning tree superimposed on scatterplot of the first two principal cornponent scores for populations studied in piedmont. eo ooo o o component 2 t cladistic and pbenetic methods 371 1.8 1.5 1.2 0.9 0.6 0.3 0.00 m. ounterl m. lunterl m. soclnl-burdlgalensls m. ounterl-tanl m. soclnl m. soclnl-burdlgalensls m. soclnl m. gunterl-tanl m. gunterl-tanl m. gunterl-lanl m. ounterl-tanl m. fanl m. tani m. soclnl m. soclnl m. tanl m. tanl m. tanl m. tanl m. tanl m. olobullna-lntermedla m. 6lobullna-lntermedla m. globullnalntermedla m. globullnalntermedla m. globullnalntermedla m. globullna-lntermedla m. globulinalntermedla 0.96 0.80 0.64 0.48 0.32 0.16 0.00 m. ounterl m. !unterl m. gunterl-tanl m. gunterl-tanl m. gunterl-tanl m. ounterl-tanl m. ianl m. tanl m. soclnl m. soclnl m. tanl m. tanl m. tanl m. tanl m. tanl m. soclnl m. burdlgalensls m. ounterl-tanl m. doclnl m. globullna-lntermedia m. globullna-lntermedia m. globullna-lntermedla m. globullnalnlermcdla m. globullna-lnlermedla m. olobullna-lntermedla m. élobullna-lntermedla m. globullna m. globullna m. globullna m. globullna m. soclnl-burdlgalensls m. soclnl m. soclnl-burdlgalensls m. globulina m. qunterl m. !unîerl m. soclnl-burdlgalensls m. soclnl-burdlgalensls m. soclnl m. gunlerl-tanl m. gunterl-tanl m. gunlerl-tanl m. gunlerl-tanl m. tanl m. tanl m. tanl m. tanl m, lanl m. ianl m. soclnl m. soclnl m. tanl m. gunterl-ianl m. soclnl m. burdlgalensls m. soclni m, olobulina-lntermedla m. 6lobullna-lntermedla m. globullna-lntermedla m. globullna-lntermedla m. globu.llna-lntermèdla m. globullna-lntermedla m. globullna m. globullna m. globullna m. globullna m. globullna m. globullna-lntermedla m. globullna m. globullna m. globullna m. globullna m. burdlgalensls m. soclnl f ! i * m, globullna 1.8 1.5 1.2 0.9 0.6 0.3 0.00 f;4 dendrogram showing the results of group-average clustering applied to the populations studied in piedmont. cophenetic correlation coefficient : 0.75. the other groups. but if the mst is superimposed on the plot (fig. 3), we see that the two populations are "closer" to r[v3f (belonging to m. socini) than to each other. this means that the projection of the populations onto the first two principal components axes has not preserved, in the case of m. socini-bwrdigalensis, the original structure of the phenetic distances. cluster analysis. a cluster can be defined like a maximally connected set. for the analysis of miogypsinidae we have selected agglomerative hierarchical clustering techniques, that proceed by a series of successive fusions of the taxonomic units into groups. the two populations showing the smaller distance between them (measured with the average taxonomic distance, discussed previously) are grouped together, then distances between this two-member cluster and each of the remaining populations are calculated. the process continues with the number of groups being reduced by one at each stage, until all the populations are grouped into a single cluster. a useful means to display the results is a diagram called "dendrogram". the methods available differ in the algorithm they use to calculate the distance between two clusters. in group-average clustering (fig. 4), the distance between t'wo clusters is the averase of the distances between ail 0.96 0.80 0.64 0.48 0.32 0.16 0.00 fio 5 dendrogram showing the results of singlelinkage clustering applied to the populations studied in piedmont. cophenetic correlation coefficient : 0.64. 3.6 3.0 2.4 1.8 1.2 0.6 0.00 dendrogram showing the results of completejinkage clustering applied to the populations studied in piedmont. cophenetic correlation coeíficient : q.77. 3.6 3.0 2.4 1.8 1.2 0.6 0.00 fig. 6 discrim. function eigenvalue percentage of variance cumulative percent 'l z 12.812 1.654 0.337 86.55 11.17 2.28 86.5s 97.72 100.00 372 f. maia functions derived wilks lambda chi-square degree of freedom signific. level 1 é 0.020 o.282 o.748 686.94. 223.53 51.29 .5,5 20 i 0 0 0 tab.4 canonical discriminant functions. the eigenvalues are proponional to the variance accounted for by each function. the significance level of the three functions is tested with the \filks a statistics and the x square statistics, that show a high degree of significance (probability : 0). pairs of populations that are made up of one population from each group. in singlejinkage clustering (or nearest neighbour method) (fig. 5), the distance between two clusters is that of their most similar pair of populations. in completejinkage clustering (or furthest neighbour method) fig. 6), the distance between two clusters is that of their least similar (or most dissimilar) pair of populations. to evaluate if the original relationships among the populations (described by their measured distances) fit well with the hierarchical structure imposed on data by clustering, we can use the cophenetic correlation coefficient. it corresponds to the correlation of the original distance matrix with the cophenetic matrix, which consists of the set of similarities produced by clustering. a value above 0.8 is sufficient to give evidence of a good fit between dendrogram and distance matrix. the cluster analysis of the populations of miogypsinidae studied in piedmont, applying the complete linkage method, produced the most meaningful grouping and the highest cophenetic correlation coefficient (fig. 6). the resulting classification of populations is not the same as that produced applying only taxonomic criteria, but it provides important informations. for example, the dendrogram shows that m. globulina and m. globuli function i o lil. gunterl . m, intermedia c m. gunteri-tani o m. socini . m,tani o m,socini-burdigalènsis e m, tani{lobulina . m. burdigaleílsis o m. globulina a m. burd--negrii; m. negrii o m. globul.intermedia fig.7 plot of population means relative to the first two canonical discriminant functions. data from literature and from piedmont. na-intermedia are well distinct as regards to all the other species: this suggests an evolutionary trend that isolates these two taxonomic units from all the others. populations belonging to the phyletic branch m. socini m. burdigalensis show characters so different that it appears very difficult to group them in a single cluster. finally, m. gunteri results well distinct from the other species. discriminant analysis. the discriminant analysis technique assumes that significative differences would exist among the mean vectors of the species to which populations belong. in certain respects, it is similar to pca, but pca seeks transformed axes that account for most of the global variation of the data, while in discriminant analysis the transformed axes permit to separate the mean vectors of groups. in the case of more than two groups to be discriminated, the method is called canonical variate analysis and consists in seeking one or more new variables that would be linear functions of the original variatffi,; tab.5 classification results for species in discriminant analysis, the actual groups are cì.assified correctly when rhey correspond to predrcted groups with a high pèrqentage. 100 0 0 0 0 0 0 0 0 0 0 0 o 78 0 0 0 0 0 11 11 0 0 0 o489700000000 o20060000200000 0 0 0 i 65 16 0 0 0 4 5 0 00000740000422 000001770000013 o1730000ss17700 o2s000005025000 00002000008000 0 0 0 0 0 0 0 0 0 0 100 0 0 0 0 0 0 0 0 0 0 0 0 100 m. gunteri m. gunteri-tani m. tani m. tani-globulina m. globulìna m. globulina-intermedia m. intermedia m. socini m. socini-burdigalensis m. burdìgalensis m. burdigalensis-negrii cladistic and phenetic rnethods 373 x coeft. m. gunteri m. gunteri-tani m. tani m. tani-globulina m. globulina m. globulina-inlermedia m. intermedia m. socini m. socini-burdigalensis m. burdigalensis m. burdigalensis-negrii m. neorii 39.283 38.447 3s.41 i 33.942 éz-1j i 31 .097 30,982 39.398 40.1 01 32.102 28.125 29.489 0.1 34 0.490 0.574 0.683 0.765 0.730 0.702 u.3yj 0.545 0.511 0.564 0.691 0.682 0.571 0.456 0.509 0,970 1.336 1,738 0.793 0.934 1.182 1.326 1.485 200.51 165.37 123.27 124.73 129.54 148.71 172.82 183.46 127.20 108.66 126.52 tab.6 coefficients obtained by discriminant analysis, useful for classifying new populations. the hst column contains a constant in each function. bles. the coefficients of these functions (discriminant functions) are calculated in such a .way to maximize the between-groups variance and covariance matrix on the base of within-groups variance and covariance matrix. the first canonical variate axis is required to be in the direction of greatest variability between the means of the different species, the second axis is chosen to be orthogonal to the first and inclined in the direction of the next greatest variability, and so on. discriminant analysis was performed on the data (parameters x, y and v) of the populations of miogypsinidae studied in piedmont and those reported by literature, and three significant discriminant functions were obtained (tab. a). the scatterplot built on the base of the first two functions (which summarize the most part of discrimination) is useful for displaying the distinction between groups of populations (fig. z). in the table of results of classification for species (tab. 5), the actual and predicted groups are shown: it appears that in many cases the populations are not classified correctly. but the most important result is the set of coefficients for use in classifying new populations (tab. 6). a new population is classified by evaluating one function for each character and each species and assigning the population to the species corresponding to the highest function value. we have put these coefficients into a spreadsheet to automatically assign a new population to one species with the highest probability. cladistic analysis. the aim of the ciadistic analysis is to construct phylogenies by studying the phylogenetic relationship between species, that is to construct evolutionary trees by considering the transformations of morphological characters during evolution. to apply this approach to the study of miogypsinidae, first of all we have had to deal with the problem of coding the measured parameters. in fact, mathematical algorithms for the analysis of phylogenetic data require alphanumeric codes that represent character states, but the populations of larger foraminifera are studied by the measurement of quantitative, continuous parameters. to not derive discrete codes from quantitative data in an arbitrary fashion, we have assigned character states to the parameters of different taxa using statistically significative differences between species (or homogeneous groups of species) resulting from the analysis of variance (anova). because of the results of the anova applied to the populations of miogypsinidae suggest that the variations of parameters between species are highly significant, we used a multiple range test based on the confidence intervals to separate these species in homogeneous groups for each parameter (tab.7). then we applied a maximum parsimony method, that is a technique to search a phyletic tree that minimizes the amount of evolution needed to explain the avalaible data. as this method requires a prespecified set of constraints upon permissible character changes, we have established to consider parameters x, y and v like ordered characters (in a progressive series of character states, the transformation of one state to another that requires to skip an intermediate state is not allowed), and parameters d7, d2/d1,, e and e/dl like unordered characters (any state of the character is capable of transforming directly to any other state). the cladogram resulting by applying the analysis to the populations of miogypsinidae studied in piedmont (fig. s) is better comparable to a pattern of the similarities between species rather than to a hierarchical statement regarding genealogical relationships. this branching diagram, however, shows interesting affinities with the hypothetic phylogenesis of miogypsinidae proposed by the literature, but shows aiso unexpected deviations along the m. socini m. burdigalensis branch. tab.7 multiple range test applied ^.l-^^,,r.,;^rs studied in piedmont. the averages o{ the measurements and the character states (groups) are ,.-^é.t t^" .""1--r--.-.. --. ---.'t parameter' m. socini-burdigalensis and m. negrii are excluded because of missing data. x d1 d2ld1 sdecies group m. gunterl m. gunt.-tani m. lani m. globulina m. glob.-int. m. socini i,l. burdioal. 9.90 9.17 8.1 6 o.5u 5.84 8.71 7.39 0 1 ó 5 6 -77.8 -39,0 20.2 27.5 -jj.j -3.u 0 1 ó ó 1 2 2.94 5.43 9.16 27.83 45.o4 16.90 32.83 0 0 0 1 ó 60.3 72.8 70.5 57.3 72.1 69.4 77.3 1 e 3 0 j 2 4 t. tc 1.11 1.14 1.22 1.'16 1.11 0 1 0 j 0 1 267.8 264.o 266.4 289.8 312.8 301 .1 'ì 1 0 1 2 3 1.69 '1 .63 lj/ 1.71 1.71 1.87 1.70 ,] 1 0 3 ó 2 -lntermedia m. bufdigalensis m, socini 374 f. maia m, gunteri -{ani m. globullna fig. 8 cladogram of species studied in piedmont. numbers in square: characters that change unambiguously on branch. treelenght (number of character changes) : 31. discussion. from the working use point of view, if we compare the results of the different phenetic methods applied to the study of miogypsinidae, we can see that the most effective tool consists in discriminant analysis. in fact, this technique provides the statistics to assign a new popuiation to a certain species in a non-arbitrary manner. pca and cluster analysis cannot be seen as tools for the production of a formal classification, but only for data expioration. concerning cladistc analysis, the preliminary results suggest to probe the research, especially in improving the characters coding and the selection of constraints upon permissible character changes. a characteristic shared by cluster analysis and cladistic analysis consists in the big number of algorithms available. sometimes it appears difficult to choose the best method of studying larger foraminifera. on the other hand, the possibility of preparing a variety of classification using different techniques stimulates a global exploration of the various algorithms available, once the data are collected and coded for numerical use. this appears more and more valid as computer packages for numerical manipulation of the daí.a are increasingly available. if we consider that the data base used for this research has been intentionaliy restricted to a predeterminated number of species of miogypsinidae, it appears evident that it will be possible to reach more significant results by including other species in the analysis. moreover, it would be interesting to extend the research to other larger foraminifera than miogypsinidae. a study on the lepidocyclinidae of the piedmont basin is in progress: we hope this research will help to verify if the problems of determination are linked to the type of organism or to the palaeontological classification criteria. acknouledgements. the research was supponed by the m.u.r.s.t. grants 60% assigned to p. clari and e. ferrero. the author wishes to thank e. ferrero {or his assistance throughout the study. thanks are due also to m. delpero for his advice and helpful discussion about cladistic analvsis. references barrai i. (1984) metodi di regressione e classificazione in biometria. y. of. 170 pp., edagricole, bologna. camussi a., móller f., ottaviano e. & sari gorla m. (1986) metodi statistici per la sperimentazrone bioiogica. v. oi 500 pp., zanichell| bologna. drooger c.\f. (1952) study of american miogypsinidae. y. of 80 pp., acad. thesis, univ. of utrecht. dunn g. & everitt b.s. (1982) an introduction to mathematical taxonomy. y. of 152 pp., cambridge university press, cambridge. elliott j.m. (1977) statistical analysis of samples of benthic invertebrates. y. of $7 pp., freshwater biological ass., scientific publication, 25, ambleside. fermont \lj.j. 6. troelstra s.r. (1983) early miocene larger foraminifera from cruiser-hyeres sea mount complex @astern north atlantic). proc. kon. ned. akad.. ivetensch,, ser.b, v. 86 (3), pp. 243-253, amsterdam. ferrero e., maia f. 8r tonon m. (1992) the evolutionary patterr ol the miogypsinidae rn eastern monferrato (n\l italy). paleontologia i eoolució, *24-25, pp. 209-217, barcelona. ferrero e., maia f. ec tonon m. (1994) le miogypsine del monferrato: aspetti morfologici e tassonornici. boll. soc. paleont. ital.,v.33 (3), pp. 345-368, modena. forey p.l., humphries c.j., kitching i.j., scotland r.lw'., siebert d.j. er slilliams d.m. (1992) cladistics. v. of 191 pp., oxford university press, oxford. schiavinotto f. (1985) different evolutionary stages in the miogypsinidae from sardinia. boll. soc. paleont. ital., v. zl q), pp.38l-394, modena. ytlizzi l. (1991) studio biometrico e biostratigrafico sui miogypsinidi e sui lepidocyclinidi deile facies carbonatiche mioceniche del monferrato orientale. tesi inedita. univ. torino. receioed april 12, 1996; accepted october 3, 1996 the turolian hipparions from cioburciu site (republic of moldova): systematics and paleodiet bogdan g. răţoi1, bogdan s. haiduc1*, gina m. semprebon2, paul ţibuleac1 & raymond l. bernor3 1alexandru ioan cuza university, department of geology, carol i bd., no. 22, 700505, iaşi, romania. e-mail: bog21rat@gmail.com; haiduc.bogdan91@gmail.com; paul.tibuleac@uaic.ro 2department of biology, bay path university, longmeadow, ma 01106, usa. e-mail: gsempreb@baypath.edu 3college of medicine, department of anatomy, laboratory of evolutionary biology, howard university, washington d.c. 20059. e-mail: rbernor@howard.edu *corresponding author. associate editor: lorenzo rook. to cite this article: răţoi b.g., haiduc b.s., semprebon g.m., ţibuleac p. & bernor r.l. (2022) the turolian hipparions from cioburciu site (republic of moldova): systematics and paleodiet. riv. it. paleontol. strat., 128(2): 453-467. rivista italiana di paleontologia e stratigrafia (research in paleontology and stratigraphy) vol. 128(2): 453-467. july 2022 abstract. the cioburciu hipparions, republic of moldova, are included in a turolian assemblage, approximately dated between 9 and 7 million years. we assess herein their taxonomic position, systematics, biogeography and paleodietary habits. we have undertaken standard equid measurements as well as accessing the vera eisenmann website for measurements and images and analysed craniodental and postcranial elements. this assemblage has been determined to be of a medium-sized hipparion with an elongated muzzle, well developed preorbital fossa that is dorsoventrally extensive and placed close to the orbit, lacking a caninus fossa and having a prominent and deep buccinator fossa. as such, this assemblage is referable to cremohipparion moldavicum gromova, 1952 common to the western ukraine, balkans, romania, republic of georgia, turkey and iran. we have employed a combination of gross cheek tooth wear morphology utilizing the mesowear method and a microscopic analysis of occlusal enamel scars utilizing the light microscope microwear technique. these complementary paleodietary methods indicate that these hipparions engaged in a mixed feeding dietary behavior and that the cioburciu sample of c. moldavicum likely alternated its diet between browsing and grazing seasonally and/or regionally. a hierarchical cluster analysis based on average scratch and pit numbers positions this taxon among extant mixed feeding ungulates. large pitting and gouging assessed through the microwear technique indicates occasional consumption of relatively coarser foods than typical mixed feeders or grazers or grit-laden food just prior to death while mesowear indicates that this was not a lifetime habit. received: june 18, 2021; accepted: december 22, 2022 keywords: cremohipparion moldavicum; systematics; mesowear; microwear. introduction we review here the turolian hipparions from the cioburciu site of the republic of moldova. the geologic deposits of the cioburciu site are included in the balta formation (hubca 1969; fig. 1). according to mathosko et al. (2016), these deposits are part of an upper miocene fluvio-deltaic system in the eastern carpathians foreland. the paleontological collections for this site can be found at the paleontological museums of the iasi, odessa, and moscow universities. the hipparion materials studied in this paper are from the upper bed of the cioburciu site. the hipparion sample studied herein was collected by macarovici in 1929 during the fieldwork for his ph.d. dissertation (macarovici 1936; macarovici 1940). later, macarovici (1967) reviewed all the hipparion fossil material from the meotian of răţoi b.g., haiduc b.s., semprebon g.m., ţibuleac p. & bernor r.l. 454 romania and the republic of moldova and concluded that for this age, only four species should be considered: hipparion moldavicum, hipparion verae (previously known as, hipparion gromovae), hipparion tudorovense (small-sized) and hipparion platygenys. lungu et al. (1993) interpreted that for the meotian deposits from the republic of moldova only these species should be considered: h. moldavicum, h. verae, h. pregiganteum, h. platygenys and h.matthewi. krakhmalnaya (1996) reviewed all the neogene hipparions of the north of black sea and concluded that for the turolian only the following species occurred: h. moldavicum, h. giganteum, h. aff. praegiganteum, h. tudorovense, h. platygenys, hipparion sp. i, hipparion sp. iii and related forms. the following mammalian species, have been found at the cioburciu site (lungu & rzebik-kowalska 2011): orycteropus gaudryi, ictitherium viverinum, adcrocuta exima, machairodus schloseri, machairodus cultridens, mustela palaeatica, simocyon primigenius, choerolophodon pentelici, deinotherium gigantissimum, hipparion verae, hipparion moldavicum, aceratherium incissivum, dicerorhinus schleiermacheri, cervavitus varibilis, palaeotragus rouenyi, helladotherium duvernoyi, tragoportax frolovi, palaeoryx studzeri, and gazella deperdita. the microvertebrate fauna following the same authors is represented by: mioprotheus cf. caucasicus, rana sp., testudo bessarabica, chelidropsis sp., melanochelys sp. , sakya sp., lacerta sp., ophizaurus sp., natrix sp., vipera sp., struthio sp., anas sp., amphechinus sp., shizogalerix cf. aticus, desmana sp., crusafontina cf. kormosi, paenelimnoecus cf. repenningi, prochotona sp. nov., alilepus lascarevi, spermophilinus cf. bredai, vasseuromys theni, hansdebruijnia neutrum, castromys sp., kowalskia lavocati. a systematic evaluation of the cioburciu site hipparions entailed measurement in millimeters rounded to (0.1 mm) cranial, dental and postcranial elements: premolars and molar teeth as well as metacarpals, metatarsals, astragali, and calcanea. herein we describe skull, maxillary and mandibular morphology and analyze the third metacarpal (henceforth mciii) and third metatarsal (mtiii) by log10 ratio diagram comparisons. we employed two different tooth wear proxies (mesowear and microwear) to reconstruct the probable diet of the hipparion(s) from cioburciu. mesowear analysis (fortelius & solounias 2000) has been used extensively to interpret paleodiet (e.g., kaiser et al. 2000; kaiser & fortelius 2003; kaiser & solounias 2003; franz-odendaal & kaiser 2003; semprebon et al. 2004a; mihlbachler & solounias 2006; rivals & semprebon 2006; semprebon & rivals 2007; mihlbachler et al. 2011, ackermans et al. 2020). two variables are assessed using the mesowear technique: molar cusp shape and occlusal relief (the relative difference in height between cusp apices and intercusp valleys) in lateral (buccal) view. the technique examines attritional (tooth-on-tooth) versus abrasive (food-on-tooth) wear on tooth surfaces. differences in the shape of cusps and in the level of occlusal relief are due to the relative levels of attritive wear (e.g., sharp cusps with high relief) and abrasive wear (more rounded or blunted cusps often with lower relief). mesowear represents cumulative wear imposed on molars throughout the lifetime of an individual animal. microwear has also been used extensively to examine paleodiet (e.g., rensberger 1978; walker et al. 1978; scott et al. 2005, 2006; ungar et al. 2008, 2010; semprebon 2002; solounias & semprebon 2002; merceron et al. 2004, 2005; semprebon et al. 2004). microwear consists of examining microscopic wear that is etched into dental enamel (and sometimes dentine) via the last meals that were consumed by animals (grine 1986). microwear turns over more rapidly than mesowear, and thus it is sensitive to local, seasonal, and individual dietary variations. microwear allows for changes in dietary behavior to be discovered before changes in gross craniodental morphology (including mesowear) can be seen. the purpose of this study is to determine the systematic affinities of the assemblage and to reconstruct the dietary behavior of the turolian equids from the cioburciu site of the moldavian republic. material and methods measurements measurements are all given in millimeters and rounded to 0.1 mm. measurement numbers (m1, m2, m3, etc.) refer to those published by eisenmann et al. (1988) and bernor et al. (1997) for cranial and postcranial elements. tooth measurement numbers refer to those published by bernor et al. (1997) and bernor and harris (2003). m1m38 refers to cranial measurements as described by eisenmann et al. (1988) and bernor et al. (1997). the turolian hipparions from cioburciu site (republic of moldova) 455 element abbreviations: pof = preorbital fossa; pob = preorbital bar; iof = infraorbital foramen i3 = third upper incisor; p2 = second upper premolar; p3 = third upper premolar; p4 = fourth upper premolar; m1 = first upper molar; m2 = second upper molar; m3 = third upper molar; p2 = second lower premolar; m3 = third lower molar, mcii = metacarpal ii; mciii = metacarpal iii; mciv = metacarpal iv; mtii = metatarsal ii; mtiii = metatarsal iii; mtiv = metatarsal iv; 3phiii = 3rd phalanx of the third (central) digit; calc = calcaneum; ast = astragalus. we sampled the following museum and institutional vertebrate collections for this study: aicupm – alexandru ioan cuza university paleontological museum lpb (fggub) – laboratory of paleontology of the faculty of geology and geophysics, university of bucharest onhm – odessa national history museum mgri – moscow vernadsky geological state museum. we have used herein measurements and images of the ciobruciu hipparions from the vera eisenmann website (https://vera-eisenmann.com/ chobruchi-data-and-photos) for our morphological observations and systematic determinations. we follow bernor et al. (1997, 2016) in describing and diagnosing the cranial and dental measurements herein. in various studies, eisenmann, (see eisenmann 1995 for a comprehensive summary), has used log10 ratio diagrams to evaluate differences in hipparion metapodial proportions as a basis for recognizing taxa and their evolutionary relationships. recently, bernor et al. (2016) used log10 ratio diagrams, to evaluate and resolve the alpha systematics of maragheh hipparionine horses which includes cremohipparion cf. moldavicum. we incorporate these previously used methodologies in this work. our statistical analysis uses the skeletal population from höwenegg (hegau, southern germany, 10.3 ma; bernor et al. 1997) as a mean log standard for calculating metacarpal iii and metatarsal iii log10 ratio diagrams herein (mciii and mtiii respectively). tooth wear sample samples were acquired from the alexandru ioan cuza university paleontological museum. all original specimens are housed in the collections of this institution. the specimens were screened for potential taphonomic alteration of enamel surfaces (after king et al. 1999) under a stereomicroscope by gms (gina m. semprebon). eleven specimens were determined to be suitable for analysis and subjected to microwear analysis. microwear technique eleven tooth surfaces were cleansed, molded, and casted (after solounias & semprebon 2002). all molar casts were examined at 35 times magnification using a zeiss stemi-2000c stereomicroscope. microwear was scored by gms to exclude potential interobserver error. microwear scars were visualized on the second enamel band of the paracone of the upper m2 or the protoconid of the m2 of adult individuals (young and old adults were discarded) using external oblique illumination (an m1-150 high intensity dolan-jenner fiber optic light source) directed across the surface of casts at a 45-degree angle to the occlusal surface. the average number of pits (rounded features) and the average number of scratches (elongated features) were assessed within a 0.4 mm square area (0.16 mm2) using an ocular reticle. also, it was also noted if more than four large pits were present or absent per within the 0.4 mm square area and whether gouges were present or absent (after solounias & semprebon 2002 and semprebon et al. 2004). results were then compared to an extant ungulate average scratch and pit database (data from semprebon 2002) to determine the dietary categories of browser versus grazer. scratch textures were also scored as being either fine (i.e., narrow with low refractivity and dull in appearance), coarse (i.e., wider with a higher refractivity and bright white in appearance) and hypercoarse (i.e., very wide and/or deep and non-refractile or dark in appearance), a mixture of fine and coarse, or a mixture of coarse and hypercoarse scratch types per tooth surface. a scratch width score (sws) was ascribed to a sample as follows: a score of 0 was given to teeth with predominantly fine scratches; a score of 1 was given to those with a mixture of fine and coarse types of textures; a score of 2 was given to those with predominantly coarse scratches; and a score of 3 was given to those with a mixture of coarse and hypercoarse types of scratches. an average of individual scores for a sample produced an average scratch width score for that taxon. lastly, because extant seasonal or regional mixed feeders alternately feed on both browse and grass and may overlap the browsing or grazing average scratch/ pit ecospaces, raw scratch distributions per taxon were constructed. grazers have unimodal, high scratch distributions, browsers have unimodal, răţoi b.g., haiduc b.s., semprebon g.m., ţibuleac p. & bernor r.l. 456 low raw scratch distributions, and mixed feeders show a bimodal split of both high scratch individuals and low scratch individuals. we also calculated the percentage of raw scratches in each taxon that fell into the low scratch range (0-17 scratches) and compared them to an extant ungulate database (data from semprebon 2002). mesowear technique mesowear was scored by a single investigator (gms) to exclude potential interobserver error. we used only those specimens in which the last molar was in occlusion and the first molar retained an occlusal shape similar to that of the second to minimize potential age effects although rivals et al. (2007) demonstrated that the mesowear signal is known to be relatively stable throughout much of the adult lifespan of hypsodont taxa such as the fossil hipparion analyzed here. mesowear was assessed macroscopically from the buccal side of the paracone of the second upper molars two different ways. first, molars were scored as in fortelius and solounias (2000) whereby cusp shape in buccal view was scored as sharp, round, or blunt and occlusal relief was scored as high or low. the percentages of sharp, round, and blunt cusps as well as the percentages of high and low relief were calculated. secondly, we employed the more standardized mesowear “ruler” method (mihlbachler et al. 2011) whereby samples are compared to a standard template with seven qualitative wear stages (0-6). the ruler is comprised of extant horse cusps which represent a gradient of cusp wear from high relief and sharp cusp shape (assigned a score of 0) to completely blunt cusp shape with no relief (assigned a score of 6). the ruler is used as a reference for cusp shape and occlusal relief which are combined into one score as others have pointed out (e.g., mihlbachler et al. 2011) that the variables used in the mesowear method are not strictly independent. we slightly modified this methodology by also recognizing intermediate scores in 0.5 increments assigned if the observed cusp shape could not be definitively assigned to one of the seven original wear stages but fell in between two steps (i.e., 0.5, 1.5, 2.5, 3.5, 4.5, 5.5). an average mesowear score was calculated. finally, we employed a hierarchical cluster analysis (using past 3.25 – hammer et al. 2001) to compare the fossil hipparion mesowear scores to those of extant ungulates with known diets. geological setting the material of cremohipparion moldavicum originates from the upper miocene locality of cioburciu (ciobruciu, chiobruciu), republic of moldova (fig. 1). lungu and rzebik-kowalska (2011) have presented the detailed overview on cioburciu geology, taphonomy, and chronology. cioburciu village is situated in the eastern part of dragos –voda district, approximately 100 km southeast of khisinev town. the upper miocene section of cioburciu is exposed in the lower part yellowish – greenish clay rich in shells of khersonian mollusks (mactra caspia). in the upper part of the geological section, there are two fossiliferous layers: the first one is composed of white-yellowish sands with a large concentration of well-preserved remains of fossil mammals. the second part is composed of gray-greenish sandy clays. vangengeim and tesakov (2013) considered that the lower bed of cioburciu 1 should be assigned to mn10 and the upper bed of cioburciu 2 to mn12. this assignment, however, does not agree with data from vasiliev et al. (2011), who provided a radioisotopic age suggesting that the khersonian (an equivalent of middle tortonian) most likely correlates to the younger chron c4an and later part of chron c4ar, and therefore, the kherosnian–meotian boundary is probably drawn within chron c4ar which is 8.5 ma. this means that the “vallesian” localities in moldova are significantly younger than 11.2 ma and correlative with the lower part of the turolian (mn11). the radioisotopic data by vasiliev et al. (2011) for the khersonian–meotian boundary indicates a correlation of the normal polarity interval of the khersonian to the younger chron c4an (8.6 – 8.2 ma). the reversed intervals with the bessarabian–khersonian boundary may consequently correspond to c4ar. thus, the hipparion localities in moldova may be significantly younger than suggested (8.0 ma) by vangengeim et al. (2006) and vangengeim and tesakov (2013), probably 7.5 – 7.0 ma (mn12). the turolian hipparions from cioburciu site (republic of moldova) 457 systematic paleontology order perissodactyla owen, 1848 family equidae gray, 1821 subfamily equinae equinae gray, 1821 tribe hipparionini quinn, 1955 genus cremohipparion qiu, huang & guo, 1988 cremohippparion moldavicum gromova, 1952 holotype of cremohipparion moldavicum (gromova, 1952): p.i.n. 1256-3639, academy of sciences paleontological institute of the u.s.s.r., figured by gromova, 1952, figures 1-3 type locality: taraklia, republic of moldova, district of benderi. age: meotian (= medial turolian, mn 12) geographic range: western ukraine, romania, balkans, republic of georgia, republic of moldova, turkey and iran. diagnosis (translated from gromova 1952: 154-155): medium size hipparion, basal length of cranium = 271-273 mm.; length of cheek teeth (p2-m3) = 121-141 mm.; muzzle elongate; index of orbital-facial length = 67; index of anatomical axes = 214.6. frontals narrow, width index = about 38.2; frontal-basal index = 261.4 mm. dental series short; index of length to basal width = 33.5 mm; to the premolars, 41.2-46.8 mm; diastemato-dentary index = 66.1-28.9 mm. upper molars large relative to premolars; molar-premolar index = 82.4-91 mm. a single pof, very long and elevated, index of position relative to orbit = 26.6-37.8 mm; relative to the facial crest, 16.7-64.3 mm. nasal notch moderately deep, its posterior border is at the level of, or slightly anterior to the anterior border of p2. the diastema is well developed; its index is 60.477 mm. protocone is short and wide, length index of p3-m2 in little or moderately worn individuals is 20.7-37.5 mm, in very worn teeth, 25-43.3 mm; index of form in the same conditions 42.8-78.3 and 57.1-92.3 mm. enamel plication is moderate in upper cheek teeth, on the posterior wall of the prefossette and anterior wall of the postfossette; when p3, p4, m1 and m2 are very worn or moderately worn, they have 3.5-6.5 to 9.5 plis. cheek teeth have a height-length index on p3, p4 of 156-195.5 mm; on m1-m3, 204.3 – 232.5 mm; on p3, p4, m1 and m2, 159-200 mm. a double knot (metaconid-metastylid) of hipparion type (rounded with intervening v-shaped linguaflexid). external depression in lowfig. 1 geographical and geological map showing the deposits of the cioburciu fossil site within the balta formation of the republic of moldova. răţoi b.g., haiduc b.s., semprebon g.m., ţibuleac p. & bernor r.l. 458 er cheek teeth deep, complementary elements little developed. the islette occupies the anterior portion of the i3 crown. the postcranials are gracile and elongate; index of width in the lower articulation relative to the width of mciii, 14.5-16.3 mm; to the length of mt iii, 12.4-14.2 mm. metapodial length relative to width: mt/t = 74.4 mm. lateral digits are moderately developed; moderate indices of lower extremities diameters mc ii and iv to mc iii, 75.8 mm – mt ii and iv to mt iii, 66.8 mm; moderate indices of length of the first phalanges of the lateral digits to those of the anterior median digits, 58.2 mm; posterior 53.8 mm. extremities recurved to the level of articulations; moderate index of the pisiform bone 114.3 mm; anterior 3phiii narrow; length-width index of posterior 3phiii 71.4-86.6 mm. emended diagnosis (after bernor et al. 2016): a medium-sized hipparionine with an elongate snout. the pof is single, subtriangular shaped, anteroposteriorly oriented and elongate, dorsoventrally and medially deep, with slight posterior pocketing, a distinct anterior rim, and strongly expressed peripheral outline. lacking a caninus (= intermediate) facial fossa. the pob is short with the lacrimal bone invading the posterior aspect of the pof. the nasal notch is incised at a level either just above the mesial border of p2, or slightly mesial to it. middle wear adult cheek teeth have moderately complex plications of the preand postfossettes; protocones are round to oval and show lingual flattening in some individuals. the p2 anterostyle is usually elongate but can be short and rounded in some individuals. mciiis and mtiiis are elongate and slender. p2-m3 length ranges from 120.0-145.2 mm. description the most complete material includes 4 crania from cioburciu, odessa 3801, odessa 2064 odessa 2067 (or alternatively odessa 2078). of these the best preserved is odessa 3801 (supplementary fig. 1; https://vera-eisenmann.com/ chobruchi-data-and-photos). there is also a nearly complete cranium, lpb(fggub) 522 lacking the snout with a mandible in occlusion (fig. 2). there are a number of maxillary cheek tooth series that are diagnostic of the hipparion species (fig. 3 a-d): aicupm4003 (p2-m3), aicupm4004 (p2-m3), aicupm4006 (p2-m3), aicupm4010 (p2-m3). eisenmann (https://vera-eisenmann.com/ chobruchi-data-and-photos) has further provided excellent images of 5 left maxillary cheek tooth series (odessa 2064, 2067or2061, 2085, 3078-1, 3078-2 and 3801) which augment the basis for our amended diagnosis of the species (supplementary fig. 2). odessa 3801 (supplementary material – supplementary fig. 1) is a complete, well preserved cranium characterized by the following features: a very large, subtriangular shaped preorbital fossa (pof) placed close to the orbit (with a short pob) that is dorsoventrally and medially deep and distinctly invaded by the lacrimal bone; caninus fossa is lacking; buccinator fossa is large and deep; infraorbital foramen (iof) is placed near the anterior-most limit of fig. 2 the nearly complete cranium lpb(fggub) 522 within the original plaster collection jacket with the mandible associated. scale bar: 15 cm. the turolian hipparions from cioburciu site (republic of moldova) 459 the pof; there is no intermediate (= caninus) fossa; buccinator fossa is distinct and deeply excavated; nasal notch is shallowly incised extending anterior to p2. the canine is large indicative of a male individual; cheek teeth have m3 in full occlusion and p2 is very worn indicative of a fully adult individual; cheek teeth have oval protocones; preand postfossettes are moderately complex; pli caballins appear single or double; hypoglyphs are moderately deeply incised. p2-m3 length is 136 mm in length. odessa 2064 is a moderately crushed skull and lacks the posterior cranium and snout. pof is as in odessa 3801. odessa 2067 (or 2061) is dorsoventrally crushed and preserves little anatomical detail. odessa 2078 is a mandible viewed labially with no extraordinary features. (length p2-m3 – 149 mm). both odessa 3801 and bpu clearly have a large preorbital fossa, subtriangular shaped, medially very deep, with a distinct anterior rim and placed very close to the orbit with a short preorbital bar. there is no apparent intermediate (= caninus fossa) diagnostic of cremohipparion moldavicum and not the closely related species c. mediterraneum. while odessa 2064 (supplementary fig. 1) is more fragmentary it displays all the facial characters of the other two crania. lpb(fggub) 522 (fig. 2) is a skull with mandible in occlusion retained within the original plaster collection jacket. the skull lacks the snout and the mandible is lacking the symphysis. in occlusion one can see the same essential salient features of the face: pob is short; pof is subtriangular shaped, medially deep and has a distinctly well-developed peripheral rim with iof at the anterior limit of the pof. occlusal details of either the skull or mandibular cheek teeth are not visible. p2-m3 is 125 mm in length and p2-m3 is 130 mm in length. figure 3 (a-d) includes occlusal views of aicupm4003 (fig. 3a), 4004 (fig. 3b), 4006 (fig. 3c), and 4010 (fig. 3d) p2-m3. aicupm4003 (fig. 3a) has protocones rounded to oval; p2 has a short anterostyle; pli caballin is double to complex on p2m2 on all cheek teeth; fossette are complex on all cheek teeth. aicupm4004 (fig. 3b) is similar to aicupm4003 but not as in advanced wear. p2 also has a short anterostyle; protocones are oval; pli caballin is single to double on the cheek teeth; p2 has preand postfossettes linked which is a primitive feature for old world hipparions. aicupm4006 (fig. 3c) is a young adult with p4 and m3 just coming into occlusion. p2 has a very short anterostyle; protocone is small and round in p2; p3 – m2 protocones are lingually flattened and otherwise short-oval shape due to their early wear; preand postfossettes of m1 and m2 are in wear and are very complexly plicated; all cheek teeth have poorly developed pli caballin due to relatively early wear and the protocones are lingually flattened on p3m2. p2-m3 length = 141 mm. aicupm4010 (fig. 3d) is another young adult p2-m3. the cheek teeth are in too early stage of wear to characterize (p2m3 137 mm). eisenmann (https://vera-eisenmann.com/ chobruchi-data-and-photos) figured 6 maxillary cheek tooth series from odessa (2064, 2067 or 2061, 2085, 3078-1. 3078-2 and 3801, with associated skull, supplementary fig. 1 and supplementary fig. 2). these series have the following salient features: protocones short, oval to rounded; p2 anterostyles mostly short, but some extended; pli caballins vary from double to single; preand postfossettes fig. 3 occlusal views of: a) aicupm4003 (p2-m3), b) aicupm4004 (p2-m3), c) aicupm4006 (p2-m3), and d) aicupm4010 (p2-m3). scale bar : 6 cm. răţoi b.g., haiduc b.s., semprebon g.m., ţibuleac p. & bernor r.l. 460 are complexly plicated; linking of p2 preand postfossettes occurs on odessa 3078-2 and 3801 (a primitive feature; bernor et al. 2017). eisenmann (https://vera-eisenmann.com/ chobruchi-data-and-photos) has figure 7 mandibular cheek tooth series (odessa 3077, 3078-2, 3078-3, 3078-4, 3078-5, no number and 2078) (supplementary material fig. 3). the most salient features of these cheek teeth are that metaconid and metastylid are mostly rounded, pli caballinids and protostylids are lacking and, primitively, ectoflexid of p2 and p4 can sometimes extend in between metaconid and metastylid (also a primitive feature; bernor et al. 2017). figure 4a is a log10 ratio analysis of mciii comparing sinap cormohipparion sinapensis (csin mean; bernor et al. 2003), pikermi cremohipparion mediterraneum (cmedpik mean; koufos 1987a, b; bernor et al. 2003), maragheh cremohipparion cf. moldavicum (mnhn mean; bernor et al. 2016) and cioburciu cremohipparion moldavicum (mgr mean) to the hoewenegg hippotherium primigenium mean log10 standard (bernor et al. 1997). the results show that the total mciii sample has strikingly narrow mid-shaft dimensions (m3) contrasting with relatively deeper midshaft depths (m4); bernor et al. (2003) termed this the “esme-acakoy” effect and heralded this as an adaptation to open country running. of the entire sample, the maragheh form, ca. 8 ma (bernor et al. 2016), has a markedly longer maximum length (m1). all mciiis illustrated in fig. 4a have similar log10 trajectories and in most dimensions are more slightly built than the hoewenegg population of hippotherium primigenium. figure 4b shows very much the same results as fig. 4a. maragheh maximum length (m1) is greater than the rest of the sample. cormohipparion sinapensis again exhibits the “esme acakoy” effect whereby midshaft width (m3) is much more slender than the hoewenegg standard but midshaft depth (m4) is about the same. the contrast in m3 and m4 measurements is further accentuated in maragheh, pikermi and the cioburciu hipparions and the latter two, pikermi and cioburciu are virtually identical. remarks the genus cremohipparion was originally recognized at the subgenus rank (hipparion (cremohipparion) by qiu et al. (1987) for the chinese species hipparion (cremohipparion) forstenae and licenti. bernor and tobien (1989) recognized cremohipparion as being a distinct lineage warranting its generic rank for the small samos hipparion c. nikosi. bernor et al. (1996; 2021) later recognized a number of species a b fig. 4 a) depiction of the log10 ratio analysis of mciii comparing sinap cormohipparion sinapensis (csin mean; bernor et al. 2003), pikermi cremohipparion mediterraneum (cmedpik mean; koufos 1987a, b; bernor et al. 2003), maragheh cremohipparion cf. moldavicum (mnhn mean; bernor et al. 2016) and cioburciu cremohipparion moldavicum (mgr mean) to the hoewenegg hippotherium mean log10 standard (bernor et al. 1997). b) moldova, maragheh and pikermi cremohipparion mtiii compared to sinap cormohipparion sinapensis log 10 ratio, ho std. the turolian hipparions from cioburciu site (republic of moldova) 461 of cremohipparion and currently these include: c. mediterraneum, c. proboscideum, c. forstenae, c. licenti, c. moldavicum, c. macedonicum, c. matthewi, c. nikosi, c. periafricanum and c. antelopinum. according to zouhri and bensalmia (2005), the spanish species “h.” concudense and “h.” gromovae are also referable to cremohipparion. all cremohipparion taxa are characterized by having a short pob and the more primitive forms have dorsoventrally and medially deep pof that are also subtriangular shaped. cremohipparion mediterraneum, c. proboscideum and c. licenti all have an intermediate (= caninus) fossa and chinese c. licenti, the youngest cremohipparion (= 4 ma, china) had an additional malar fossa and posteriorly deeply pocketed buccinator fossa. cremohipparion licenti has an enhanced development of its facial fossae, accompanied by a strong, deep nasal notch incision to p4, were likely in support of a highly mobile snout for feeding. cremohipparion species never achieved a maximum crown height greater than 55-60 mm. the cioburciu sample of cremohipparion moldavicum exhibits primitive characters including: skull, with very large, medially deep and dorsoventrally deep preorbital fossa placed close to the orbit; nasal notch short, incised anterior to p2; cranium lacking caninus fossa; maxillary cheek teeth with p2 having a short anterostyle and rounded protocone, pli caballins varying from single to complex; pre and postfossettes complexly plicated with opposing borders that occasionally link together; mandibular cheek teeth mostly with rounded metaconids and metastylids; mciiis and mtiiis that are elongate and slender and exhibit sharply contrasting midshaft width (m3) and depth (m4) dimensions. these characters support the hypothesis that cremohipparion was most likely derived from a form like sinap cormohipparion sinapensis, but without excluding the possibility that cremohipparion was derived from a more primitive member of the cormohipparion clade (sensu woodburne 2007). microwear and mesowear microwear. raw fossil hipparion microwear is shown in tab. 1 and summary statistics are summarized in tab. 2. figure 5a shows the average scratch and pit results for the fossil hipparion plotted in specimen # p s lp sws st g cusp shape relief mws aicupm4000 28 20.5 7 1 mixed absent blunt low 6 aicupm4001 24 21.5 5 2 coarse present blunt low 6 aicupm4002 19 25.5 5 2 coarse absent sharp high 0 aicupm4003 20.5 21.5 4.5 2 coarse absent round high 1.5 aicupm4004 26 16.5 6.5 1 mixed absent round low 1.5 aicupm4005 30.5 20 10.5 1 mixed present aicupm4006 blunt low 6.0 aicupm4007 35.5 21.5 4 2 coarse present sharp high 1.0 aicupm4008 19.5 21.5 3.5 2 coarse absent round high 2.0 aicupm4009 25 11 9.5 2 coarse absent round high 1.5 aicupm4010 30.5 20 10.5 1 mixed present round high 1.0 aicupm4011 round high 2.0 aicupm4012 36 17 13.5 2 coarse present round high 2.0 tab. 1 microwear and mesowear results for fossil horses. abbreviations: p = average number of pits; s = average number of scratches; sws = average scratch width score; lp, average number of large pits; st = scratch texture; g = presence or absence of gouges; mws = mesowear score. taxon p sp s ss lp f m c sws g sc rc bc h mws ciuburciu hippparion 26.8 6.0 19.6 3.8 7.1 0 36.4 63.6 1.6 45.6 16.7 58.3 25.0 66.7 2.5 tab. 2 mesowear and microwear summary statistics for ciuburciu hipparion. p = average pits, sp = standard deviation of pits, s = average scratches, ss = standard deviation scratches, lp = average number large pits, f = percentage of finely textured scratches, m = percentage of mixed finely and coarsely textured scratches, c = percentage of coarsely textured scratches, sws = scratch width score, g = percentage of gouges, sc = percentage of teeth with sharp cusp shapes, rc = percentage of teeth with rounded cusp shapes, bc = percentage of teeth with blunt cusp shapes, h = percentage of teeth with high occlusal relief, mws = average mesowear score. răţoi b.g., haiduc b.s., semprebon g.m., ţibuleac p. & bernor r.l. 462 reference to gaussian confidence ellipses (p=0.95) on the centroid for extant browser (b) and grazer (g) data adjusted by sample size (from semprebon 2002 and solounias and semprebon 2002). figure 5a shows that the fossil hipparion has an average scratch and pit count that falls in the gap between the extant browsing and extant grazing ecospace. figure 5b displays raw scratch and pit results for the fossil hipparion plotted in reference to gaussian confidence ellipses (p=0.95) on the centroid for extant browser and grazer data adjusted by sample size (from semprebon 2002 and solounias and semprebon 2002). figure 5b shows that the fossil hipparion has scratch and pit results that overlaps the browsing and grazing ecospaces but most samples fall in the gap between them. figure 6 shows the percentages of finding more than 4 large pits and the percentages of gouges found within the optical reticle imposed on the microscopic field of view for both extant ungulates with different known diets (data from semprebon 2002) and the fossil hipparion examined in this study. what is clearly shown, is that the fossil hipparion has more large pits and gouging than is typical of the extant ungulates studied via microwear. interestingly, fig. 7a shows that the fossil hipparion has a percentage of low scratches that falls within the extant mixed feeder range even though its scratch width score (fig. 7b differences in the range of scratch textures) is more similar to that of extant grazing ungulates (data from semprebon 2002). figure 8 shows the percentages of sharp, round, and blunt cusps as well as the percentages of teeth that showed high and low occlusal relief for the fossil hipparion. it is clear that the fossil hipparion has relatively few sharp cusps (17%) typical of most extant browsers. the majority of cusps are rounded (58%) followed next by some blunt cusps (25%). figure 9 shows the results of the hierarchical cluster analysis. the cioburciu hipparion clusters with a group of extant mixed feeding ungulates (i.e., those that alternate regionally or seasonally between browsing and grazing) and away from a cluster of extant browsers and grazers. this designation matches the scratch/pit results obtained through microwear. fig. 6 bar graph showing the percentages of finding more than 4 large pits and the percentages of gouges found within the optical reticle imposed on the microscopic field of view for both extant ungulates with different known diets (data from semprebon 2002) and the cioburciu fossil hipparion examined in this study. fig. 5 scratch and pit results for the fossil hipparion from the cioburciu deposits plotted in reference to gaussian confidence ellipses (p=0.95) on the centroid for extant browser (b) and grazer (g) data adjusted by sample size (semprebon, 2002; solounias & semprebon 2002). a) the placement of the cioburciu hipparion in the gap between the extant browsing and extant grazing ecospace (average scratch and pit data). b) raw scratch and pit results for the cioburciu hipparion plotted in reference to gaussian confidence ellipses (p=0.95) on the centroid for extant browser and grazer data adjusted by sample size (semprebon 2002; solounias & semprebon 2002). a b the turolian hipparions from cioburciu site (republic of moldova) 463 discussion cremohipparion moldavicum first appeared in the ukraine and balkans during mn 10 (gromova 1952; bernor et al. 1996), while the somewhat smaller species c. macedonicum appeared at this time in greece (koufos 2016). cremohipparion moldavicum occurred in the ukraine, balkans, romania, georgia and iran during mn11 to 13. the dwarf form cremohipparion matthewi overlapped its stratigraphic range with c. moldavicum in the maragheh section (bernor et al. 2016). cremohipparion mediterraneum and c. proboscideum occurred in greece also in mn12 and 13, c. antelopinum in the siwaliks in mn10-13, c. forstenae in china in mn13 and 14, c. nikosi in greece in mn13 and c. periafricanum in southern europe and north africa in mn13 and 14. cremohipparion licenti is the latest occurring member of the genus and was restricted to china in mn 15 (ca. 4.0 ma). the genus cremohipparion was one of the chronologically and geographically longest old world hipparion lineages extending its chronologic range from mn10mn15 (9.7-4.0 ma), from spain to china and the indian subcontinent and north africa (bernor et al. 2021). most species were medium sized to very small. cremohipparion proboscideum was the largest member of the clade and distinguished by having very retracted nasals (bernor et al. 2018). the evolutionary origins of cremohipparion are a matter for debate. bernor et al. (1996) suggested that cremohipparion was derived from hippotherium primigenium. another suggestion is that the turkish early vallesian species cormohipparion sinapensis could be a likely ancestral stock for cremohipparion. for this to be supported, we would have to accept the hypothesis that the preorbital bar reduced secondarily in length after increasing in length for both hippotherium primigenium and cormohipparion sinapensis which in turn was derived from a member of the cormohipparion occidentale complex of woodburne (2007). another possibility is that the somewhat more primitive north american cormohipparion, c. quinni with a shorter pob and large pof, could be the genetic source for cremohipparion. cremohipparion moldavicum would appear to be the most primitive member of the cremohipparion clade and the size, great medial depth, distinct peripheral rim and subtriangular shape is shared with hippotherium primigenium and hippotherium weihoense (sun et al. 2019). crea b fig. 7 a) box plots of the percentage of scratches per taxon that fall within the low scratch range (i.e., between 0 and 17) for the cioburciu fossil hipparion compared to results for extant ungulates (extant data from semprebon 2002). b) box plots of the scratch width score of the cioburciu fossil hipparion compared to results for extant ungulates (extant data from semprebon 2002). 33 figure 8. bar chart of the percentages of sharp, round, and blunt cusps as well as the percentages 822 of teeth that showed high and low occlusal relief for the cioburciu fossil hipparion. 823 824 825 826 827 828 829 830 831 832 833 834 835 fig. 8 bar chart of the percentages of sharp, round, and blunt cusps as well as the percentages of teeth that showed high and low occlusal relief for the cioburciu fossil hipparion. răţoi b.g., haiduc b.s., semprebon g.m., ţibuleac p. & bernor r.l. 464 mohipparion moldavicum has mciii and mtiii log10 ratio trajectories that are elongate and slender, albeit more elongate than cormohipparion sinapensis. bernor et al. (2020) noted that both cremohipparion and african eurygnathohippus feibeli has elongate-slender metapodial iiis which were similar but longer, and potentially derived from, sinap cormohipparion sinapensis. there is no current radioisotopic data, or for that matter well corroborated magnetostratigraphic data, that supports the occurrence of cremohipparion moldavicum, or for that matter any member of the cremohipparion clade occurring as early as the cormohipparion datum in central europe at 11.4-11.0 (bernor et al. 2017), or the first occurrence of cormohipparion sinapensis in turkey, 10.8 ma., or cormohipparion sp. in the potwar plateau, pakistan at 10.8 ma. (bernor et al. 2021). nevertheless, cremohipparion has primitive features of the pof, dentition and postcranial morphology that on morphological bases cannot deny the possibility that it represents a primitive morphology of old world hipparion. the cioburciu hipparion is referred herein to cremohipparion moldavicum, a medium sized, primitive hipparion common to moldova, ukraine, turkey, iran and likely the democratic republic of georgia. cremohipparion moldavicum was likely adapted for cursorial locomotion in eurasian “pikermian” open county woodlands (eronen et al. 2009; kaya et al. 2018). our results show that both dietary proxies employed here (i.e., microwear and mesowear) point to a mixed feeding dietary behavior in the fossil hipparion studied. as shown in fig. 5a, average scratch and pit results place the fossil hipparion clearly in the gap between extant browsers and grazers and individual raw scratch/pit results cluster mainly in the gap between extant browsers and grazers but also overlap both browsing and grazing ecospaces (fig. 5b). these results indicate that the fossil hipparion from cioburciu most likely alternated between browsing and grazing regionally or seasonally. figures 5a and 7 corroborate this conclusion by showing that the fossil hipparion has a percentage of low scratches that falls within the extant mixed feeder range and clusters on a hierarchical cluster analysis with extant mixed feeders. however, microwear also revealed that the fossil hipparion from cioburciu had more large pits and gouges in its dental enamel than is typically seen in the extant ungulates studied by gms (semprebon 2002 & solounias and semprebon 2002) as well as scratch textures more typical of grazing ungulates than mixed feeders. since microwear captures only a snapshot of dietary behavior (i.e., short-term) just before death, these results indicate that the fossil hipparion, though typically engaging in a mixed feeding diet, consumed relatively coarser foods (i.e., coarser grasses or browse) and/or grit-laden food just prior to death but most likely not typically (i.e., mesowear results are not coarse enough to indicate a lifetime habit of consuming highly coarse or grit-laden foods). conclusions the turolian cioburciu hipparions from the republic of moldova were investigated for both their likely paleodietary behavior and systematic position. regarding systematics and alpha taxonomy, both craniodental and postcranial measurements and observations have allowed us to refer this assemblage as cremohipparion moldavicum gromova 1952 – a medium-sized, primitive hipparion with an elongated muzzle and well-developed and dorsoventrally extensive preorbital fossa which is sit0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0 d is ta n ce a e p e yc e ro s m e la m p u s g a ze ll a g ra n c a p ri co rm is s u m a tr a e n si s g a ze ll a t h o m so n i o vi b o s m o sc h a tu s c io b u rc iu h ip p a ri o n o d o co il e u s h e m io n u s tr a g e la p h u ss c e rv u s ca n a d e n si s ta u ro tr a g u s o ry x r e d u n c a re d u n ca g ir a ff a c a m e lo p a rd a li s o d o co il e u s vi rg in a n u s o k a p ia jo h n st o n i d ic e ro s b io co m is r h in o ce ro s so n d ia cu s a lc e s a lc e s d ic e ro rh in u s su m a tr e n e si s a lc e la p h u s b u se la p h u s c o n n o ch a e te s ta u ri n u s h ip p o tr a g u s n ig e r k o b u s e ll ip si p ry m n u s h ip p o tr a g u s e q u in u s b is o n b is o n c e ra to th e ri u m s im u m e q u u s b u rc h e ll i e q u u s g re v yi d a m a li sc u s lu n a tu s fig. 9 results of a hierarchical cluster analysis of extant ungulate browsers, grazers and mixed feeders and the cioburciu hipparion. the present analysis depicts the fossil hipparion clusters within a group of extant mixed feeding ungulates. the turolian hipparions from cioburciu site (republic of moldova) 465 uated close to the orbit. cremohipparion moldavicum has been shown here to be closely related to pikermi cremohipparion mediterraneum in cranial, dental and postcranial anatomy, and both of these taxa are turolian age. we concur with previous studies (bernor et al. 2003, 2017, 2020, 2021; woodburne 2007) that cremohipparion was most likely derived from a species close to cormohipparion sinapensis from the vallesian of sinap, turkey. two paleodietary proxies – microwear and mesowear indicate that the cioburciu hipparions engaged in seasonal or regional mixed feeding and thus alternated between browsing and grazing. microwear reveals the occasional consumption of relative coarser foods or foods laden with grit. the systematic and paleodiet results are consistent with previous attributions that cremohipparion moldavicum inhabited western eurasian pikermian mixed woodland-grassland biomes (bernor et al. 2021). acknowledgements: bogdan haiduc`s research was supported by pn-iii-p1-1.1-mc-2019-1758, project funded by the romanian ministry of research and innovation, cncs-uefiscdi, pncdi iii. ray bernor wishes to acknowledge research funding by nsf ear grants 8806645, 0125009, 1113175, and 1558586; dbi grant 1759882 for the futres database and support from the smithsonian human origins program and university of florence for a visiting professorship in 2017 and 2018. this is publication number 29 for the nsf sponsored futres database program, bernor co-pi. references ackermans n.l., martin l.f., codron d., hummel j., kircher p.r., richter h., kaiser t.m., clauss m. & hatt j. 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(2007) phyletic diversification of the cormohipparion occidentale complex (mammalia; perissodactyla, equidae), late miocene, north america, and the origin of the old world hippotherium datum. bulletin of the american museum of natural history, 306: 1-138. https://vera-eisenmann.com/chobruchi-data-and-photos accessed on 10th june 2021 rivista italiana di paleontologia e stratigrafia volume 104 numero 2 r*ole 1-2 pagrne 279-286 agosto 1998 noa breve spirorbid polychaetes as in the mediterranean boreal guests pleistocene rossana sanfilippo received january 1, 1988; acceted april 17, 1998 keytaords: spirorbidae, boreal guests, pleistocene, mediterranean, north atlantic. riassunto. sono stati trovati numerosi tubi di policheti spirorbidi attribuibili alle specie spirorbis spirorbis (linnaeus, 1758) e spí rorbis corallinae de silva 6c knight-jones, 1962, provenienti vari depositi del pleistocene freddo della sicilia, nonché da sedimenti wiirmiani del mar tirreno. tali spirorbidi vivono lungo le coste dell'atlantico settentrionale, in acque basse della piattaforma continentale, e sono assenti nell'attuale mediterraneo. nei sedimenti pleistocenici questi sono invece associati a faune di media piattaforma ed epibatiali, probabilmente perché spiazzati da biotopi pirì superficiali. viene attribuito a s, spirorbis e s. corallinae un significato paleobiogeografico e paleoclimatico, in base alla loro attuale distribuzione biogeografica. tali specie, entrate in mediterraneo durante una o più fasi fredde del pleistocene, assumono quindi il significato di ospiti nordici. speciale attenzione è stata, infine, rivolta alla morfologia e struttura del tubo, le quali hanno ponato all'individuazione di caratteri diagnostici a livello specifico. abstract. a first report of spirorbis spirorbis (linnaeus, 1758) tnd spirorbis corallinae (de silva & knight-jones, 1962) from early and middle pleistocene deposits in sicily and submerged late glacial sediments in the \western mediterranean is presented. today both species live on shores and very shallow bottoms in the nonh atlantic and are unknown from the recent mediterranean, such differences in their present and past biogeographic distributions suggest that these species were boreal guests (bgs) in the mediterranean pleistocene. special attention ìs paid to tube morphology and structure, which bear some diagnostic features for species identification. lntroduction. the appearance in the mediterranean of boreal shelf species, together with the progressive disappearance (since the piacenzian) of warm-temperate taxa is a distinctive element of the cold pleistocene phases (raffi, 1986; raffi & marasti, 1982). these cold-water species are currently known as "northern guests" or "boreal guests" (bgs) as they spread mainly in the north atlantic ocean, from arctic latitudes southward to the southern british isles and the bav of biscav. during cold phases the bgs funnelled inro the mediterranean, superimposing on the mediterranean autochthonous temperate faunas. their migration occurred in successive, cumulative, increasingly cold floods, which correspond in part to the chronostratigraphic repartition of the early pleistocene (raffi, 1986). bg species, early used as strarigraphic markers (n4ars 1963; ruggieri et al. t976), were employed subsequently as palaeoclimatic tools (di geronimo, 1974; malatesta & zarlenga, 1986; raffi, 1986; taviani et al., 1997). deposits rich in bgs have been studied since the last century and crop out extensively in southern kaly. records of boreal species are known from the colder periods of both the mediterranean and the atlantic ocean (taviani et al., 1991). nevertheless, bgs are quire poorly documented within the east atlantic pleistocene sediments (submerged thanatocoenoses) and are recorded mainly from the mediterranean quaternary. most bgs are molluscs, such as the bivalve arctica islandica, the best known north atlantic species in rhe mediterranean, which is now regarded as a marker for the plio-pleistocene boundary @uggieri er al., 1984). other taxonomic benthic groups, nohbly serpulids (zibrowius, 1979; zlbrowius & ten hove, 1987) have been considered sporadically as palaeoclimatic-oceanographic tools. however, these plio-pieistocene taxa, are more widely distributed, belong to the deep-sea mediterranean benthos and have an atlantic or more generally oceanic affinity. in the present work, the first mediterranean fossil records of two north atlantic spirorbid species are discussed, together with an account of their present-day geographic distributions and ecology. material and methods. the material examined (fig. 1) originates from early to middle pleistocene sandy deposits, cropping istituto policattedra di oceanologia e paleoecologia, università di catania, corso ltalia 55,i-95l29, catania, italy, e-mail: rosso@mbox.unict.it 280 fio i out in .western (valle del belice) and eastern (augusta and salice) sicily, and from submerged late glacial sediments in the tyrrhenian sea (canyon "del toro"). the valle del belice section crops out along the right side of the belice river, in south'western sicily (fig. 1). it consists of a biogenic sandy succession, ca.3.5 m thick, within a clayey sequence of uppermost eariy pleistocene age. the sands are highly fossiliferous, with corallinaceous algae, rhodoliths and molluscs. the spirorbid tubes were found in a bed with arctica islandica. palaeoecologic indications inferred from the benthic macrofaunistic assemblages, suggest an upper-circalittorai soft bottom (ca. 35-40 m depth) (di geronimo et al., 1994). several species of north atlantic origin are present. s. spirorbis (10 specimens) and s. corallinae (21 specimens) are associated with other serpulids, i.e. hydroir. sanfi.lippo fossil (closed symbol$ and recent (open symbols) records of spirorbis spirorbis (triangle$ and, spirorbis corallinae (circle$. 1) augusta; 2) valle del belice; 3) salice; 4) canyon del "toro". des noroegicus, spirobrancbus po/ytrema, serpula concharum and pomatoceros triqueter. most s. corallinae tubes show a concave attachment surface, indicating grorà/th on articulate coralìine stems (p1. 1, fig. z, 8), while some s. spirorbis tubes have flat attachment surfaces, suggesting that they may have been attached to kelp fronds. the augusta outcrop is located along the left side of the marcellino river (se sicily), ca.7 km s\l of augusta (fig. 1). it consists of an organogenic sequence of muddy-gravelly sands of uppermost early pleistocene age (di geronimo et al., in press), over 1 m thick, contained in bluish sandy muds. palaeoecologic observations from the sandy levels indicate a mid-shelf area (upper circalittoral) with palaeocommunities characterized by north atlantic immigrant species, among which ,4. plate 1 scale bar : 400 pm fig. 1 spirorbis spirorbis (linnaeus, 1758). augusta, early pleistocene. fig. 2 spirorbis spirorbis. valle del belice, early pleistocene. fig.3 spirorbis corallinae de silva ec knight-jones (1962), forming irregular ascending coils. salice, early pleisrocene fig. 4 spirorbis corallinae, with regularly flat coìls. valle del belice, early pleistocene. fig. 5 tube of sptrorbis corallinae on a corallina stem. valle del belice, early pleistocene. fig. 6 tube of spirorbis corallinae, attached on corallina stem. belice, early pleistocene. frg.7 bioimmuration of corallina stem lry spirorbis corallinae. augusta, early pleistocene. fig. 8 detail of frg. 7 showing casts of sporangiae. pleistocene soirorbid pohchaetes in the mediterranean 281 282 r. sanfilippo islandica is dominant (di geronimo et al., in press). some layers of the sequence supplied tubes of s. spirorals (6 specimens) and s. corallinae (14 specimen$. they are mostly detached from the original substrata, probably coming from neighbouring shallower biotopes. among the associated serpulids, hydroides noraegicus is the more abundant species. moreover, several tubes of an undetermined circeis species occur. the salice section crops out near the town of salice, ca. 15 km w of messina (fig. 1). the middle pleistocene outcrop (di stefano & lentini, 1996) is an organogenic sandy sequence, 13 m thick, overlying in unconformity limestones with bathyal corals. the sands are often cross-bedded and increase in muddy fraction upwards. the same two spirorbid species (13 s. spirorbis and 25 s. corallinae specimens) were obtained within some layers of the sands, becoming more abundant in their upper part. the associated benthic assemblages indicate an epibathyal palaeoenvironment swept by currents, with infraand circaiittoral elements transported down from shallower biotopes located on top of the steep sides of the palaeobasin. faunas are characterised by species with a markedly atlantic affinity (such as neouermilia falcigera among serpulids). placostegus tridentatus and serpula lobiancoi are the dominant associated serpulids. some tubes of circeis sp. are also present. s. corallinae tubes are usually detached from the substrate, showing a concave area of attachment, suggesting an originai settlement on cylindrical substrata \ke corallina. síems, s. spirorbis and s. corallinae were found also in a sample dredged from "del toro" canyon, off southwestern sardinia (stn. dg03, 38o43.28'n, 8o20.59'8, i95 m). the sample consists of muds rich in oyster shells and scleractinians, indicating a late glacial age. s. spirorbis and s. corallinae (both with 6 specimens) occur together with deep circaiittoral and bathyal serprlids (meuverrnilia rnultioistau, semioermilia agglutinaa, hyalopomatus oariorugosus and filogranula stellau). as in the previous sample, spirorbids seem to have been displaced from the infralittoral environment, down to epibathyal depths. some specimens of spirorbids are free in the mud, showing limited areas of attachment, as though they had been attached to comparatively thin srems of algae, hydroids or bryozoans. a stock of living specimens of s. spirorbls from iceland, collected by drs. p. and e.\l knight-jones, was used for observations and comparison of tube structures. they are densely settled on a blade of. fwcus serratus. for sem observations, before coating with gold palladium, fossil and recent tubes were cleaned with fizoz. some tubes were broken to investigate the wall slructure. tube morphology and structure. differences between the tubes of s. spirorbi.s and s. corallinae are seemingly slight, both tubes being sinistrally coiled, smooth and similar in shape. nevertheless, careful observations of tube morphology allow a distinction at the species level to be made, even when only fossil material is available. size is also similar, the outer coil diameter being ca. 2-3 mm in s. spirorbrs and 2-2.5 mm in s. corallinae. the opening is simple, without peristomes, in both species (pl. 1, figs. 1,,2,4). a weak sculpture occurs oniy in s. corallinae, consisting of wide-spaced rounded gros/th ridges, indicating the previous openings (p1. 1, fig. a). in s. spirorbls the outer edge of the tube spreads out to form a more or less evident peripheral flange (pl. 1, figs. 1,2). this feature is distinctive for s. spirorbis (see rzhavsky, 7994), but lacking in s. corallinae, the cross section of which is regularly circular (pl. 1, figs. 3,4,5). when attached to the small and convex fronds of corallina, a tube of s. corallinae grows often as irregular ascending coils (p1. 1, figs. 3, 5, 6). on fronds of larger algae it is regularly flat and planispirally coiled. the tube diameter of the external coii does not increase roward the opening (pl. 1, fig. a). the outer coil of s. spirorbis, however, lies partly over the inner ones and the tube diameter increases more rapidiy with growth (p1. 1, figs. 1, 2). on the attachment surface of s. corallinae, on casts of corallina stems and sporangia are often visible (pl. 1, fig. 7, 8). plate 2 scale bar : 400 pm (fig. 1),20 pm (figs. 2-7) fig. 1 spirorbis spirorbis on fucus serratus. iceland, recent, f;, t."-."-...-^'l^'1 of tube wall of spirorbis spirorbis sho*ing an inner chitinous layer, a mid opaque and an outer glossy calcareous layer. iceland, recent. fig. 3 microstructure of the two calcareous layers. opaque layer with crystals arranged in a criss-cross pattern. outer glossy with isorientated crystals. iceland, recent. fig. 4 detaii of outer glossy layer. needle-like crystals, perpendicular to the outer surface. fig. 5 detail of mid opaque layer. crystals of prismatic shape and criss-cross arrangement. fig. 6 transverse wall section of fossil spirorbis spirorbis, with homogeneous structure without layers. augusta, early pleistocene. fig.7 detail of fig. 6. crystals have lost the original shape structure and have crowded together. pl.2 pleistocene spirorbid polychaetes in the mediterranean 284 r. sanfilippo the tube wall of living s. spirorbis specimens has a glossy surface and is composed of two calcareous layers (p1. 2, fig. 2,3). the inner layer is opaque, ca. 50 pm thick, and with crystals arranged in a criss-cross pattern (pl. 2, fíg. 5); the outer iayer is porcellaneous or almost vitreous, ca. 20 1tm thick, with larger crystals, showing a common axis orientation. at higher magnification, crystals appear needle-like and perpendicular to the outer tube surface (pl. 2, fig. 4). this kind of structure results in transparency (zlbrowius & ten llove, 1,987). a third chitinous layer rarely occurs around the lumen of the tube (pl. 2, fig. 2). several burrows, probably by boring algae, are visible near the outer surface (p1. 2, fig. 2). fossii tubes of both species are thick and rather porcelaneous, with a glossy surface. they are always recrystallized. sem micrographs show a unique homogeneous layer, 70 ptm thick, in which crystals are closely packed without a regular orientation (p1. 2, fig. 6, 7). the original wall structure is lost. biogeography and ecology. the present geographic distribution of the two species investigated is atlantic, north of latitude 40on. s. spirorbis is a boreal-arctic species (fig. 1), ranging from the arctic sea to the northeast coast of the u.s. (knight-jones e/ a1.,1.99t).it is frequent in the celtic and north seas (knight-jones & knight-jones, 1,977; knight-jones, pers. comm.). it was found also in the \x/hite and barents seas (rzhavsky, 1,992). s. corallinae is boreal-arctic too, but less vzidely distributed, ranging from the barents sea (rzhavsky, l992) to southern brittany (fig. t). it is particularly common in the celtic sea, from ireland to the scilly isles (knight-jones & knight-jones, 7977). in the southern area of distribution of both species (english channel) the mean bottom water temperature is 6-10o c in winter and does not exceed 16" c in summer (holmes, 1966). both species are mesolittoral to upper infralittoral, occurring from rock pools to shallow depths (knight-jones et al., 1975; knight-jones & knight-jones, 7977; rzhavsky, 1992). they are mainly epiphytic on fronds of various algae. s. spirorbis typically settles on brown fucoids (especially fucus serratus) but may occur also on kelps (larninaria, sacchorbiza) (knight-jones et ^1., i975)" s. corallinae settles almost exclusively on corallina officinalis, and rarely on other red algae. exceptionally it was found on the soft red alga chondrus, in higher pools (knight-jones er knight-jones, l977). gregariousness is seen in both species and the larvae swim for no more than a few hours (knight-jones, 1951). the boreal distribution of s. spirorbis and s. corallinae ís a character common to many species belonging to the genus spirorbls daudin. s. rupestris, s. inornatus and s. tridentatus are exclusively boreal (knight-jones et al., 1.991; knight-jones ec knight-jones, 1995). s. strigatus, as well as the recently described s. (wlorbis) gesae, live in the atlantic at midlatitudes (off west africa). only three species live in the mediterranean: s. infundi" bulum, which is probably endemic, s. cuneatus which lives also in ne atlantic, and the circumtropical s. (spi rorbella) marioni, whose recent introduction into the mediterranean was by ship transporr (zibrowius & bianchi, 1981). the last species, and three other easrern pacific spirorbids (5. spatulatus, s. bifurcatus, s. rothlisbergi) are dextraily spiraled. oniy two other spirorbis are tropicai, s. placophora and s. bidentatus, the former known only from the galapagos and the latter having a circumtropical distribution (knight-jones et al., 1991). in the fossil assemblage the associatio n of s. spirorbis and s. corallinae with a deep water fauna can be explained easily by post-mortem down slope displacement from shallower bottoms. it is very unlikely that these shallow water spirorbids were fossilized within their original environment, exposed to dispersing.wave action. discussion. the present boreal and arctic distribution of s. spirorbis and s. corallinae, here reported as pleistocene fossils, suggests significantly colder conditions than today. their immigration into the mediterranean pleistocene probably took place by a gradual southward expansion of their biogeographic range, in step with the progressive cooling. the movement of their populations into the mediterranean would have been helped by their short planktonic larval phase and gregariousness during settlement. s. spirorbis and s. corallinae probably made a relatively "late" appearance in the mediterranean during the cold pleistocene. they are recorded from sediments not older than the uppermost early pleistocene (sicilian age), corresponding to the third molluscan migration flood (raffi, 1986). as already srressed by buccheri (1985) this age couid be correlated to the first cold pleistocene peak recognised by shackleton & opdyke (lze) at abour 0.8 ma. a similar southward movement of spirorbis was invoked to explain the recent finding of a species which may be endemic to madeira, s. (wlorbis) gesae kníghtjones & knight-jones (1,995). this is quite like s. spirorbis, and the authors suggest that a common ancestor of both species may have reached madeira during an ice age, and that s, (wlorbts) gesae may be its modified descendent. as mentioned above, tubes of circeis and of the serpuiid h. noruegicui occur together with the fossil splrorbis snder study. circeis is a circumboreal spirorbid genus, quite common in northeastern atlantic infralittoral waters, but not represented in the present-day mediterranean (knight-jones & knight-jones, 1977). it may thus have the same palaeoclimatic significance as the two other spirorbids. h. noraegicus, still common in the recent mediterranean (zibrowius, l972), appears more abundant in pleistocene mid-shelf deposits. this could also suggest lower pleistocene temperatures. the diagnostic value of shape and other morphologic features of spirorbid tubes is demonstrated here. identification at the species level is possible only when empty or fossil tubes are available. sem observations of the tube wail of s" spirorbis show a particular microstructure, not observed in other spirorbids (two distinct pleistocene spirorbid polychaetes in tbe mediterranean 285 layer$. this suggests that at least in recent materiai, not affected by diagenetic changes, tube microstructure can also be a diagnostic character at the species level. achnouledgements. i am greatly indebted to p and e. w. knìght-jones (depanment of zoology, swansea) for constructive comments and loaned material. i also thank h. zibrowius (stat. mar. endoume, marseille) for critical review of the manuscript, s. di geronimo (istituto policattedra dì oceanologia e paleoecologia, catania) for useful suggestions and o" torrisi (c.n.r., catania) for assrstance rn sem observations. this paper was supported financially by 40/o (di geronimo) m.u.r.s.t. erants. references buccheri g. (1985) osservazioni paleoclimatiche al limite pleistocene inferiore-pleistocene medio della foce del belice (sicilia sud-occidentale) mediante l'uso degli pteropodi. boll. soc. geol. it., v. 104, pp. 1l5-1.22,f.orr'a. di geronimo l (1974) significato paleoecologico di cylichna alba nuovo "ospite nordico" del pleistocene deila sicilia. mem. mus. cia. st. nat. wrona, v. 2a, pp. 569572, yerora. di geronimo i., costa 8., la perna r., randazzo g., rosso a. 8c sanfilippo r. (199a) the pleistocene "case catarinicchia" section (belice, s\l sicily). studies on ecology and paleoecology of benthic communities. r. matteucci et al. (eds.), boll. soc. paleont.1r., vol. spec. 2, pp. 93-l15, modena" di geronimo l, di geronimo r., la perna r., rosso a. ee sanfilippo r. (in pres$ cooling evidence from pleistocene shelf assemblages in se sicily. proc. european paleontological congress, lt;jy 1997, lù/ten. di stefano a. & lentini f. (1996) ricostruzione stratigrafica e significato paleotettonico dei depositi pleistocenici del margine tirrenico tra villafranca tirrena e faro (sicilia nord-orientale) . studi geologici camerti, vol. spec. n. 2, pp. 21.9-237, camerino. flolmes n. (1966) the bottom fauna of the engiish channel j. mar. biol. ass. u.k., v. 41, pp.397'461, [,ondon. knight-jones e.\f. (1951) gregariousness and some other aspects o{ the settling behaviour of spirorbis. j. mar. biol. ass. u.k, v. 3, pp. 201,-222, i-ondon. knight-jones p., knight-jones e.'w. & al-ogily s.m. (1975) ecological isolation in the spirorbtdae. proc. 9th europ. mar. biol. sy*p., pp. 539-561,, aberdeen. knight-jones p. & knight-jones e.\l (1977) taxonomy and ecology of british spirorbidae (polychaety'. j. rnar. biol. ass. ij.k., v. 57, pp. 453-499, london. knight-jones p., knight-jones e.\l & buzhinskaya g. (1991) distribution and interrelationships of northern spirorbid genera. bull mar. science, v. 48, n. 2, pp. 189-197. knight-jones p. 8c knight-jones e.'sl (1995) spirorbidae (polychaeta) from madeira including a new species and subgenus of spirorbis. mitt. hamb. zool. mus. inst^, v" 92, -^ eq_1n1 l-l"-h,,,^ ^*^.^*rg. malatesta a.. k zarlenga f. (1986) northern guests in the pleistocene mediterranean sea. geol. romana, v. 25, pp. 9l-154, roma. mars p. (1963) les faunes et la stratigraphie du quaternaire méditerranéen. réc. tia<.,. stat. mar. endoumo v 2rnn 61-97, marseille. raffi s. (1986) the significance of marine boreal molluscs in the early pleistocene far,rnas of the mediterranean area. palaeogeogr. palaeoclim., palaeoec., v. 52, pp. 267289. amsterdam. raffi s. ec marasti r. (1982) the mediterranean bioprovince from the pliocene to the recent: observations and hypotheses based on the evolution of the taxonomic diversity of moiluscs. in: paleontology, essential of historical geology, proc. int. meet. venice 1981, e. montanaro galitelli (ed.), pp. 151-177, mucchi, modena. ruggieri g., buccheri g., greco a. & sprovieri, r. (1,976) un affioramento di siciliano nel quadro della revisione della stratigrafia del pleistocene inferiore. boll. soc. geol. it., v. 94, pp. 889-914, roma. ruggieri g., rio d. & sprovieri r. (198a) remarks on the chronostratigraphic classification of lower pleistocene. boll. soc. geol. ll., v. 103. pp. 251-259, roma. rzhavsky a.v. (l992) review of circeinae and spirorbinae (polychaeta, spirorbidae) from seas of ussr with a description of a new species, circeis gurjanotae. zool. zhurn., v.7 l n.7. pp. 5-13. moscow. rzhavsky a.v. (1994) on the morphoecology of spirorbid trbes. ophelia, v. 39, n. 3, pp. i77-182, fielsrngor. shackleton n.j. e. opdyke n.d. (1976) oxygen-isotope and paiaeomagnetic stratigraphy of pacific core y28-239 late pliocene to latest pleistocene. geol. soc. am. mem. v. 145, pp. 449-484. 286 taviani m., bouchet p., metivier 8., fontugne m. er delibrias g. (1991) intermediate steps of southq/ards faunai shifts testi{ied by last glacial submerged thanatocoenoses in the atlantic ocean. palaeogeogr., palaeoclimat., palaeoecol., v. 86, pp. 331,-338, amsterdam. zibrowius h. (1,972) hydroides noroegica gunnerus, hydroides azorica n.sp. et hydroides capensis n. sp. (polychaeta serpulidae), espèces vicariantes dans l'atlantiqre. bull. mu* hist. nat., paris, s. 3, v. 39 (33), pp. 433-446,paris. zibrowius h. (1979) vitreotubus digeronimoi n. g., n. sp. (polychaeta serpulidae) du pléistocène inférieure de la sicile et de l'étage bathyal des agores et de l'océan indien. 7étlrys , v. 9, n. 2, pp. 183-190, marseille. zibrowius h. ec bianchi c.n. (1981) spirorbis rnarioni er pileolaria berkeleyana, spirorbidae exotiques dans les ports de la médlterranée nord-occidentale. ropp. comm. int. mer mèdìt., v. 27, pp. 763-t64, monaco. zibrowius h. & ten hove h. (\987) neoztermilia falcigera (roule, 1898) a deepand coldwater serpulid polychaete common in the mediterranean plio-pleistocene. bull. biol. soc. wash., v.7, pp. 259-271, \tashington. r. san/ìlippo rivista italiana di paleontologia e stratigrafia yolume luj numero 2 tavole 1-2 pagrne 173-1,82 settembre 1997 conodont biostratigraphy of the late triassic sequence of monte cocuzzo (catena costiera, calabria, italy) adelaide mastandrea*, fabio ietto*+, claudio neri*** & franco russo**** key-uords: catena costiera calabrese, basin deposits, conodonts. norian. rhaetian. riassunto. nel presente lavoro sono riponati i risultati preliminari di uno studio biostratigrafico, basato sui conodonti, dei terreni carbonatici triassici della catena costiera calabrese, affioranti nella finestra tettonica di monte coctzzo. la successione studiata (colle del crapio) si è depositata in contesti di piede scarpata/bacrno ed ha fornito faune a conodonti ricche e ben presenate. sono riconoscibili due biozone: la prirna, nella pane inferiore-media dell'unità, è marca'ta dt epigondolella slwahensis e può essere riferita al norico superiore (sevatiano); la seconda è caratterizzata da misikella bernsteini, in associazione con m. postbernsteini, ed ha un inquadramento cronostratigrafico pir\ controverso. secondo krystyn (1990) e golebiowshi (1990) potrebbe essere riferita al sevatiano sommitale (norico superiore), mentre per kozur 6c mock (1991) ia comparsa di m posthcrnsrcini rndicherebbe il passaggio al rerico. abstact. preliminary results are reponed from rn investigation of the conodont associations found in the late triassic carbonate succession of the so-called "catena costieru calabrese" that crops out in the tectonic window of monte cocrzzo. the succession of colle del crapio consists of alternating carbonate mud, breccia and calciturbidites deposited in a toe-of-slope to basin setting and contains rich and well-preserved conodont faunas penaining to two biozones. the iower zone is characterised by the occurrence o{ epigondolella sloz,ahensls and may be referred to the late norian (sevatian). the upper zone is characterised by misikella bernsteini associated wrth m. postbernsteinì. the chronostratigraphic setting of the latter zone is more controversial, as it may be regarded as latest sevatian (upper norian) according to krystyn (1990) and golebiowski (1990), while according to the zonation of kozur 6c mock (1991) the first occurrence of m, postbernsteini marks the beginning of the rhaetian stage. introduction. this paper is the first contribution on conodont stratigraphy of the late triassic succession cropping our in the so-called monte cocuzzo tectonic uindotu auct., pertaining ro the "catena costiera calabrese" and loca' ted west of cosenza. the succession, mainly consisting of strongly dolomitized carbonate rocks, has been interpreted by amodio morelli et ai. (1976) as a rracr of the african margin of apulia, cropping our as a recronic window below the thrust sheets perraining ro rhe "calabrian (calabridi) units" (ogniben, 1969). the stratigraphic section studied .was measured and sampled near colle del crapio (fig. 1). the section is 73 m thick, including the upper part of rhe "intermediate complex or monte cocvzzo complex" and the lower part of the "upper complex or colle del crapiovalle delle pernici complex" of ietto et al. (1995) (fig. 2). observations on the same stratigraphic interval were also made in a quarry on the southern slope of monte cocuzzn (fig. 1). the little data on the age of this succession available to date were based on benthic fossils, such as the dasycladacean green alga griphoporella cuntata (giimbel, 1972) pia, l915, occurring in the upper complex of colle del crapio (ietto et ai., 1993) and referred to the late triassic (norian-rhaetian) (barattolo et al., 1993). the finding of a rich conodont fauna in the section studied allows more accurate biostratigraphic dating. two distinct conodont biozones were recognized within the relatively thin interval sampled. the stratigraphic section lithostratigraphy and sedimentology. the stratigraphic section studied (fig. z) inciudes, from a lithostratigraphic view point, the upper part of the "intermediate complex" and the lower part of the "upper complex" of ietro et al. (1993; 1995). 'f istiruto di paleontologia, università di modena, via università,4, i-41100 modena. 'f" dipanimento di scienze della terra, università federico ii, largo s. marcellno, 10, i-80138 napoli. ',"" dipanimento di scienze geologiche e paleontologiche, (jnrversità di ferrara, c.so ercole i d'este,32, i-44100 ferrara o'r+* dipartimento di scienze della terra, universirà della calabria, i-82030, arcavacata di rende (cosenza). 174 a. mastandrea, f. ietto, c. neri & f. russo intermediate complex. the intermediate complex (52 m, excluding a not precisely determinable lower tract of the complex) consists mainly of dark grey to blackish dolostone with minor marly interlayers. the main iithofacies are as follows. a black dolomicrite with minor thin (mmto cm-thick) fine-grained, graded dolomite intercalations, interpreted as fine, distal or starved, turbidites embedded into basin lime mudstone. b coarse grained dolostone, deriving from the dolomitization of a calcarenite and a fine breccia, forming individual beds 3-4 ro 30-40 cm thick, with erosional base, normai grading, even lamination and, occasionally, current ripples. these beds are interpreted as turbidites (usually t"6); they are either amalgamated or separared by centimetre-thick dolomicritic interlayers. the grains are usually too deepiy recrystallized to be recognizable; however, in some rare case, bioclasts such as dasy' clads (gryphoporella sp.) or fragments of microbialitic boundstone can be observed. calabrian (calabridi) crystalline-metamorphtc units carbonate complex border of lhe lectonic window sampled outcrops : a. colle del crapio seclron b. quarry south of m. cocuzzo fig. 1 a, location map of the area studied; b, structural units and location of the ourcrops sampled. c doiomitic breccia, mainly with flat clasts, forming beds ranging in thickness from 50-60 cm to 2-3 m; clasts are mainly represented by tabular fragments of beds, 1-5 cm thick and 20-50 cm wide. they consist both of basinal dark dolomicrite (lithofacies a) and of calciturbidites (ithofacies b); platform-derived clasts are more rare, usually smali in size (4-5 cm) and consist of microbialite boundstone (stromatolitelike laminite with associated skeletal cyanobacteria and encrusting foraminifers). the breccia bodies show quite different degrees oí organization, ranging from chaotic mud-supported deposits (mud flows) to normal-graded, clast-supported beds capped by laminated doioarenite (turbidites). latera1 and vertical transition between siumped beds and flar pebble breccia is frequent. these lithofacies are frequently organized into coarsening-up sequences, mainly made of facies a and b, or, more rareln into fining-up sequences. the intervals dominated by flat clast breccia do not show any defined organisation. moreover, the tract of the intermediate complex studied does not show a clear vertical evolution, although an upward increase in the amount of breccia occurs. the frequent occurrence of slumped ho[] [l *-:::1 r v :fiurnefreddo :=:= î------------j** gririard x ! a) u x j u f o f z conodont biostratigraplry of monte cocuzzo : 'z175 .cc l0 .cc9 .cc 8 . cc 5 .cc 48 .cc 4a occ 4 2_ o cc 1 rizons and chaotic deposits consisting of tabular slabs derived from slope deposits in the upper part of the unit may indicate a pronounced slope instability, probably due to synsedimentary tectonics. upper complex. only the lowermost paft of the upper complex (20 m) was investigated. this unit is very poorly exposed, due to the abundance of marls intercalated in the succession. outcrops consist of dm-thick blackish dolomite, represented by both dolomitized mudstone and calciturbidites, and of yellow to grey dolomitic marls. poor exposure makes facies analysis and paleoenvironmental interpretation difficult, but according to the available data the overall depositional setting indicates a bakey fig. 2 colle del crapio stratigraphic sectton. key: 1) dolostone; 2 and 3) marly dolostone and dolomitic limestone; 4 and 5) fine-grained doloarenites; and dolomitic calcarenites; 6 and 7) coarse-grained dolostone and dolomitic limestone; 8 and 13) dolomitic breccia with platy laminated clasts; 9 and 18) dolomitic breccia with marly clasts; 1o) alrernating lithologies; 11) marls; 12) slumped deposits; 14) thin turbidites; 15) coarse amalgamated turbidites; 16) carbonate mud (possibly dolomitized); 17) marlylpelitic mud; 19) slump; 20) erosive surface; 21) normal grading; 22) even lamination; 23) small scale crosslamination; 24) covered interual; 25) sample. sinal environment. the unit records a marked reduction in the carbonate gravitational sedimenration and an increase in pelite input. biostratigraphy. the conodont fauna. nine samples, with a minimum weight o{ 7 kg, were collected in the coile del crapio section. six were obtained from the intermediate complex and three from the upper complex (fig z). two additional samples collected from ^ qu rry south of mr. cocuzzo, ar the boundary between the intermediate and upper complex, were consistent with the data from colle del 25 24 23 22 21 20 19 18 17 16 15 -) a lo l/ o a. mastandrea, f, ietto, c. neri g f. russo rarniform elements of conodonts in the colle crapio. four samples taken from the underlying lower complex (serra mezzana) yielded large mono-specific collections of epigondolella slwakensis. all the samples from colle del crapio were collected from black dolomicrites, quasi-stoichiometric and very rich in organic matter (russo f. et ai. in preparation), with thin graded intercalations interpreted as fine turbidites. the samples yielded a very rich conodont fauna, well preserved as single elements or less frequently as clusters. conodonts showed a colour alteration index (cai) of 5 (epstein et al. 1.977). conodont occurrences and vertical ranges are given in fig. 3. discussion. fauna 1 samples cc1 to cc5 from colle del crapio (fig. 3) yielded a rich monospecific conodont assemblage exclusively containing specimens of epigondolella slwakensis (kozur) emend. budai & kovacs, 1986 (fig. 3). this species is very abundant (approximately 530 platform elements and some clusters) and well preserved. mostly broken specimens of ramiform elements (i..., prioniodiniform, enantiognathiform, hideodelliform, and hibbardelliform) of the apparatus were also found. budai er kovacs (1986) and kovacs & nagy (1989) reported the presence of epigondolella slwakensls from the rezi dolomite formation of keszthely mts. and fekete-hegy limestone formation ("avicuia limestone") of pilis mts. in the balaton highland (hungary) where it ranges from upper alaunian to sevatian. the e. slwa' kensis population described by these authors includes elements with great morphological variability, similar to those observed in the colle del crapio fauna. orchard (1,983, 1,991) recognised a significant morphological variety in the epigondolella population of the norian of british columbia and therefore defined four conodont biozones in the columbianus zone (ammonite standard scale of canada, tozer,1994), corresponding to the middle norian (alaunian 2 and 3 sensw krystyn in zapfe,1983). colle delcrapio section è f bè feffè=eee * è è e è 9è e x; èé é fi s é fio i rcc 10 .cc 9 .cc 8 .cc 6 .ll 5 rcc 48 occ 4a .cc 4 .ll i occurrence and venical rrnge del crapio section. plate 1 upper norian conodonts from colle del crapio (monte cocuzzo) section (cc). eq. f-l epigondolella sloaakensis (kozt4 1972) emend. budai & kovacs, 1986. 1) upper view, ipum 25715, cc1, x 80. 2) upper view, ipum 25716, cc4, x 80. 3) lateral view, ipum 25717, ccl, x 80. 4) oblique lateral view, ipum 25718, cc4, x 1oo. 5a) upper view; 5b) lower view, ipijm 25719, cc4, x 80. 6) upper view, ipum 25720, ccl, x 80. z) upper view, ipum 25721, cc4, x 80. 8) lateral view, ipum 25722, cc6, x 80' 9) lower view, ipum 25723' cci' x 80' fig. 10 enantiognathiform elements, ipum 25724' cc4, x 100. fig. 11, 12hibbardelliformelernent. i1) ipum 25725,cc10,x200. 12)ipum25726, cc4a,x 150 fig. 13 ' grodelliform element, ipuìi{ 25727 ' cc10' x 100' ril.r+,rsprioniodiniformelements. 14) ipum25728,cc4,x1oo. 15) ipum.?5729,cc4b,x70. r;g.to,tzhindeodelliformelemenrs. 16)ipum25730,cc9,x100. 17) ipum2573l,cc4b,x10o. fig, 18 panicuiar o{ the pit of epigondolella slwabmsis, fig' 9, x 800' x 'j at u rr'. rrl x f-.1 (^) u f! f f z conodont biostratigraphl of monte cocuzzo krystyn (in zapfe, 1983) kryslyn, t980; 1988 kozur, l9e9; kozur s mock, 1991 z e i (f marshi posthernsteini delrei rhaetica ultima stuezenbaumi paucdentata poslhemsteini steinbergensis z é. z z ua '6 f q reliculalus hemsteini hemsteini andrusovi ouinquedunclalus bidentata bidentata z z f [iacer posíerae n.sp. d hogani postera oo ó amn/onoid zone€ conodont zones 178 a. mastandrea, f. ietto, c. neri g f. russo fig.4 biostratigraphic zonations based on ammonoids and conodonts of the upper norian and rhaetian stages. roghi et al. (1995) noted, in the e. slwakensis population collected in the dolomia di forni (carnia), an evolutionary trend similar to the one described by orchard (1983; 1991) tn two differenr species, e. postera and e serrulata. on the basis of this observation they referred the dolomia di forni to the alaunian (middle norian). according ro kozur (1972; 1989) and kozur er mock (1991) epigondolella slwahensis is a typical sevatian (upper norian) species. its main occurrence marks the upper part of the e. bidenwa zone and its disappearance corresponds to the lower boundary of the misikella hernsteini-paruigondolella andruswl assemblage zone (sensu kozur & mock, l99i) of the uppermosr sevarian (fig. a), this species has been reported also from the calcari selctferi of the late sevatian age in the lagonegro basin, southern italy (amodeo et al., 1993). fauna2. in sample cc6 specimens of epigondolella slouahensls (kozur) occur together with misikella postbernsteini kontr & mock. the occurrence o{ m. posthernsteini earlier rhan m. hernsteini (which firsr appears in sample cc8) is problematic, since m. posthernsteinl is thought to have evolved from the latter. samples cc8, cc9 and cc10, from the upper complex, yielded a conodonr fauna composed exclusively of the genus mísikella, represented by m. hernsteini and m. posthernsteinl associated with ramiform elemenrs. the first species is quanritatively dominant over rhe se_ cond. transitional forms between m. lternsteiní and m. posthernsteinl occur in sample cc8 (fig. 3). misikella bernsteini (mostler) has been considered typical of the suessi zone (upper norian)(mostler er al., 1,978; gazdztcki et ai., 1979; kovacs & kozur, 19go), as well as the stuerzenbaumi subzone (lower rhaetian) in the zlambach marls and koessen beds of the northern calcareous alps (krystyn, 1980; golebiowski, l986; 1ee0). misikella postbernsteinl was originally described from the i-ower rhaetian in the zlambach beds of maly mlynsky vrch (slovakia) by kozur & mock (192a). since then it has been reported many times from stratigraphic successions attributed ro rhe upper norian and/or rhaetian of europe and japan (mostler et a1., 1978; gazdzícki, 1978; gazdzicki et al., 1979; kovacs & kozur, 1980; isozaki 8c matsuda, 1982; krystyn, 1980, 1988; golebiowski, 1,986, 199a; gulio, 1996). recently, m. postbernsteini was collected from the uppermost triassic (crickmayí zone) of canada (orchard, 1991). m. posthernstetni rs generally considered a species characteristic of the rhaetian. fiowever, according ro krystyn (1990) and golebiowski (1990) it appears from the uppermost sevatian (suessi zone); and the predominance of m. hernsteini on m. posthernsteini is diagnostic of the uppermost noriar age. kozur & mock (1991) suggesr that the first occurrence of m. postbernsteíni can be used as a marker ro define the norian,/rhaetian boundary, an event easily recognisable in the tethyan successions. in this case, most of rhe cochloceras suessi ammonoid zone would belong to the rhaetian. on the other hand, krystyn (1988, 1990) defined the base of the rhaetian in the hallstatt region as the first occurrence of " cboristoceras" baueri (stuerzenbaumr zone). this corresponds, in terms of conodont biostratigraphy, to the last occurrence datum (lod) of plate 2 upper norian/early rhaetian conodonts from colle del crapio (monte cocuzzo) section (cc). fig. 1-6 misikella hernsteini (mostler, 1967). l) oblique lateral view, ipum 25732, cc9, x 100. 2a) upper view; 2b) lateral view, ipum 25733, cc9, x 1zo. 3) lower view, ipum 25734;4) upper view, ipum 25735; cc8, x 170. 5) later,rl view, ipum 25736, cc9, x 100. fig.6a-b misikellabernsteini-misikellaposthernsteini6a) obliquelateralview;6b)lowerview, ipum25737,cc9,xi7a. fig.7-10 misihella postbemsteini kozur 6c mock, 1974. 7a) upper view; zb) lateral view; 7c) oblique lateral view; ipum 25738, cc6, x 170. 8a) lower view; 8b) oblique lateral view; ipum 25739, cc6, x fia. 9) upper view, ipum 25240, cc6, x 170. 10) oblique lateral view, ipum 25741, cc6, x 170. fig. 11 cluster of mkikella hernsteini.ipum 25742, cc10, x 150. fig. 12, 13 clusters of hideodelliform elements. ipum 25743, cc8, x 150. fig. 14 cluster of grodella delicatula.ipum 25744, cc8, x 100. fig. 15 cluster of epigond.olella slmakmsis. ipum 25745, cc4, x 150. conodont biostratigraplry of monte cacuzzo 180 epigondolella bidenaa, first occurrence datum (fod) of misikella rhaetica and m. koessenensis, all species not documented in the catena costiera calabrese. following the scheme of kozur & mock (l991), the conodont fauna from samples cc6 to cc10 could be referred to the early rhaetian (m. hernsteini m. postbernsteinl subzone, m. postbernsteini zone), whereas according to krystyn (1990) it should be ascribed to the latest norian (ì4. hernsteini zone) (fig a) conclusions. 1. the lower part of the colle del crapio sequence (samples ccl to cc5) is characterised by the occurrence of epigondolella slwakensis, a taxon referred to the aulanian-sevatian (middle-upper norian) by many authors (kozur, 1972, 1.989; budai 6c kovacs, 1986; kovacs & nagy, 7989; kozur & mock, 1991; roghi et a1., rees). 2. the chronostratigraphic attribution of the upper part of the succession (from samples cc6 to cc10) is more controversial, as it involves the unsolved definition of the norian-rhaetian boundary. samples cc8 to cc10 record the occurrence of the genus misikella, represented by m. hernsteini and m. posthernsteint. 3. according to many authors, m. hernsteìnì and m. posthernsteini indicate. an ljpper norian-rhaetian a. mastandrea, f. ietto, c. neri & f. russo references age, even if a larye part of the m. posthernsteini range is typically rhaetian. the presence and abundance of m. hemsteini still at the top of the section, and the absence of diagnostic species such as m. rhaetica and m. koessenensis, may indicate that the upper parr of the the colle del crapio section (upper complex) belongs ro rhe uppermost norian (krystyn, 1990; golebiowski, 1990). on the contíary, according to the scheme of kozur er mock (1991), the upper part of this section would be regarded as early rhaetian in age. 4. since this preliminary study is based on a single stratigraphic section, we are not able to address the chronostratigraphic problems connected with the norian,/rhaetian boundary. f{owever, we provide a description of the verticai distribution of conodonts within the stratigraphic succession of the "catena costiera". acknooledgements. $7e ere grrteful to a. nicora (milano) and m.j.orchard (vancouver) for revisions of the original manuscript. field and printing expenses were supported by a grant of murst 40o/o and cnr 96.00364 (f russo, cosenza). english revision by e. fois erickson (cleveland). illustrated specimens are housed in the collections of the institute of paleontology, university of modena ipum 257 15-257 45) . amodeo f., molisso f., kozur h., marsella e. & d'argenio b. 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(1983) das forschungsprojekt "triassic of the tethys realm' (igcp proj.4). abschlussbericht. ósterr. akad. ly'iss. schrrftenr. erduisl komm., v. 5, pp. 7-l6, 'wien. received december 19, 1996; accepted may 5, 1997 brandoni 111..124 �� ��� �������� � � ��� �������� �� ������� �� ���������� �������� ���� � ��� ���� � ����� � ������������ ������ �� ��� �� � ��������� ��� ��� ��� ��� ����� �������� ������� � ������� � ��������� �� � �� ����� ��������� ������ � ���� ��� �� ��� ���������� ������������� ���� �������� ��������� ������ �� �������� � ���� �!��"����! ���� ��� #�� $������� �% ��� !&$%�"�'� �������������� '���� �� (��� '���� �����!����' !'���! ����%��) $� ����� � ��� �&(���� �� ������ ���� �*���)! %��" ��� "�))'� ������� �% +�������� �� ��� '��� +'��!��� ��������'� ,�'����� ������������! ������) ����� �����!� )�(��!��� �� ��� '�-�� '�(�'! ./00����'�"���)� �!12%���33� ���� �������4 �% ��� ��&� 5�����2 ���"����� �� ����� �12�! +��(����� �������!���� ��������� �"��� ��� %�&� "�����������! ���� ���&� �� ���! &���� ��� ������ � � ����� � ��� �"������� !������ � ��� 6���'��(��� ��) �"���� ��� � ��� 6���'��(��� �����!��� ��'���(�'� !"�'' �� "�)�&"�!�5�) ��*� ,��� -� )�!���$� ��"������' �!!����) �� �� �������� � ��� ��� '����!� �% ��� %�&� ������ %��" ��� !�"� $�) ��) ��"����$'� �� !�5� �� !�"� 7&�������� !�����! �% ����� � ��� #���� (�'�) !�����! �% �� �������� � ��� ��(� $��� �������5�)� �'' )�!���$&��) "���'� �� �����-�!���� ���������� %��" ��� ���� ������� �� ����"���� +��(���� .��) ���$�$'� �'!� �� #&�&"�2� +��(����4� ��) ��� '��� +'������ �� �� ���8� +��(���� #�� ���!���� �% ���! ���&! �� �������!���� ��������� �*���)! ��! 9��-� ��'��$�������������' )�!���$&���� ��������� �' $�������� )�''� !����%�"��'�� �������������� ���� )����)� )� )�"��!���� ����� "�'�� ����)� �!�"�'�%����� )� ����� � ��� �&(���� !� �!���)� �� ��������� )�' ������� ��)�� )�''� +�������� !��� �' ���)� +'��!��������'����� $�!�'� � "����������� �����&�!��� '� '��� )�(��!���: ��&: ����&������ ��� '�(�''� ��%������ ./00����'�"���)� �!12%���33� ���)� �������4 )�''� ���"�5���� ��&5�����2 ��''� +��(����� )� ����� �12�!� ��' ���)�!� )�''3��������� #�� '� ;&����� %��"� )� "����������� ��� !� ���(��� �� ;&�!�� &����:� � ������ � ������ � ��� �"������� !������ � ��� 6���'��(��� � �"������� � ��� 6���'��(��� ������!��� ���� ��*� )� ���'�� )� ��'���(�"���� �����'� � "�)�� � )�!���(� ;&� �&�(� "������'� �����$&��� � �� �������� � ��� '� %��"� )� "������� )�"��!���� ��� � ;&����� ������ )�''� !��!!� !������ )� ���'�� ���%������ $�'� ��� �'�&�� !����� ;&��������� )� ����� � ��� ��� !���� ������� !��&�� ��"� (�'�)� ��� !����� )� �� �������� � ���� �&��� ���!���� !�� �����&��� ��' ���)�(�!� )�''3���������� )�' ���)� ������� )�''� +��(��� ��� )� ����"���� .� ���$�$�'"���� ����� ��''� +��(����� )� #&�&"�2�4� � ��' ���)� +'������ )�''� +��(����� )� �� ���8� �� )��&"������ ���!��� 5� )� ;&�!�� ������ ��' ���)�!� )�''3��������� �� �!���)� '� )�!���$&� 5���� ��'��$��������%��� !����� ���� ������������ #�� <�������� ��� � ���&� �% "�""�'! -��� �� �'"�!� �*�'&!�(�'� ����������' )�!���$&���� #��� ��� ����������5�) $� � !9�'���' ���������&�� )�!����� %��" �'' ����� �'������' "�""�'! #��� ��� %&����� )�!���� �&�!��) $� "�'��&'�� �(�)���� !&�������� ����� ��!����� �! ��� �% ��� %�&� "�8�� �'�)�! �% "�""�'! .��'!&� �� �' �== � �==�> ��)!�� �� �' �== > �&���� �� �' �== 4 <���������! ���'&)� ����� �'�)�!� 4 ��� ����&'���� ��(� ���) -��� � $��� )��"�' !��''> �4 ��� ?��"�'���&�� ����'� !�����'�5�) %�� "��"��������> ��) @4 ��� #��)�� ���)� �� +��''������� ��� ��$����' ��) �����!����' !'���! �&�!�)� ����&'���� ��'� � %��*����� %��"! �% #��)�� ���)� ��!!�!!�) � ���&"��� -��� !&$�&�����&! �!���� )��"! ���" ��� ��"� �% ��� ���'��!� �&$'�!��) �����)! �% %�!!�' #��)����)� .!�� ���'��� �� �' aa=�� $4� ��� )�(��� !��� �% ���! ���&� ��! $��� ����'� (����$'� �� "�""�' �!!�"$'���! ��! )�(��!��� �� ��� '�-�� '�(�'! .���� ���� ����4 �% ��� ��&5�����2 ���"����� .���� ��������+'��� ����4 �� ����� �12�! +��(���� �! 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'6% 0-&' �h >3.! 0-$-%()$",-'") ",0$")-'"(*&� c" ; ��" /���� ��! h��+h��� �6")6%&'%.� �-*)6%''��� �(*-#(**-� ����� �"-,0-$"*"� ��� �-$$")>� �� �� �%(*%� ��� �-.)($"*"� �� � �")6%$/))"� �� 7�gg�9 + �*'.-'3..6%*"-* )(($"*5 %2%*' #%#/)%# 13 *(*+,-."*% ,($$/&) -&&%,1$-5% -' �"$$�� �"(.5"( 7�"2(.*(� �'-$39� =���" /��" #��" (��� ��! ��f+��d� �(#%*-� �-*)6%''��� �%(*% ��� �-$$")> �� � � �(*-#(**���� 7�gg�9 + �835%* "&('(0% )(,0(&"'"(* (: $"2"*5 $-*# &*-"$ &6%$$&! #-':.(, �'-$3� #�����-��"� #��������$ �����"� #���������"� ���! �g+��� �,&'%.#-,� rivista italiana di paleontologia e stratigrafia volume 104 numero 3 15 dicembre 1998 noa brevry . short note ctenognathichthys bellottii (de alessandri, 1910): nomenclatural problems and stratigraphical importance of this middle triassic actinopterygian fish andrea tintori* receioed february 6, 1998; accepted september 2, 1998 key-uords: middle triassic, paleopterygians, nomenclature, holotype, neotype, stratigraphy. riassunto. durante la revisione della collezione paleontologìca depositata presso il liceo gìnnasio 'a.volta' di como, è stato rinvenuto l'esemplare figurato da de alessandri (1910) come esemplare tipico della sua nuova specre heterolepidotus (?) bellottii. ciò solleva una questione nomenclaturale, poìchè, considerando perso i1 materiaìe originario, è srato recentemente istituito un neotipo in seguito all'attribuzione di questa specie al nuovo genere ctenognathichtlrys (bùrgin, 1992). seguendo il codice internazionale di nomenclatura zoologica, la questione è stata sottoposta alla commissione internazionale di nomenclatura zoologica. poichè alcuni requisiti necessari per i'erezione di un neotipo non sono stati soddisfatti, l'olotipo deve essere considerato a tutti gli effetti come l'esemplare portatore del nome. viene inoltre messa in evidenza i'importanza stratigrafica di questa specie presente sia nella formazione di perledo-varenna (materiale originario) che nella formazione di besano (scisti ittiolitici di besano o grenzbitumenzone auct.). questo conferma che la fauna di perledo, generalmente considerata un tutt'uno, è in realtà composta da almeno tre diverse associazioni di fossili avendo alcune specie (probabilmente 3) in comune con la fauna della formazione di besano e del1a parte inferiore del calcare di meride (ladinico inferiore) e aimeno cinque specie in comune con la fauna della kalkschìeferzone (membro sommitale del calcare di meride, ladinico sommitale) rinvenuta à ca' del frate piggiù, varese, italia) e nei pressi di meride (tr cf . abstract. the existence of one of the origìnal specimens of ctenognatbicbtlrys bellottii (de alessandri 1910), figured and labelled as 'type specimen' by de alessandri himself, raises a nomenclaturaì question as there is also a neotype designated by bùrgin (1992). the international commission of zoological nomenclature has been asked to rule on the validity of the neotype. as some of the requirements for the designation of a neotype were not met, the holotype should be treated as the name-bearing type. the discovery of this species in both the besano formation (grenzbitumenzone auct ) and the perledo-varenna formation is very importànt lrom a stratrgraphic point of view, confirming that the so called perledo fish-fauna is composed of different assemblages. in fact, the perledo fauna has most species in common with the ca' de1 frate (viggiù, varese, italy) fauna from the kalkschieferzone (uppermost meride limestone), which is latest ladinian in age, while a few of the perledo species have been found also in the grenzbitumenzone and/or in the low-er meride lìmestone o{ lower ladinian age. lntroduction. during the revision of the paleontological coliection stored in the liceo ginnasio 'a.volta' school in como (italy), i was charged by the archeological survey of iombardy with the triassic vertebrates. this small collection consists of twenty specimens, mainly found in the quarries around perledo (lecco, northern italy). this material proved to be very important because, like almost all the other specimens known from perledo, it s/as collected in the first half of the last century and was used by de alessandri (1910) for his monograph on the triassic fishes from lombardy. since then, very fe'w specimens have been collected and, in any case) nobody has ever described them. furthermore, the museo civico of milano, which housed the main collection, was destroyed during the second world war, so that few of the described specimens have survived, the ones stored in the senckenberg museum in frankfurt and in the italian geological survey in rome, other than those in como. the reason why almost all perledo vertebrates have been found during the last century is still unknown. quarry.work, in fact, was carried on until 40 years ago, but new paleontological finds were limited to a few remains of sawricbthys and lariosaurus, whlle some other specimens have been collected by the author during field surveys. as for many other old collections, the exact stratigraphic position of the fossils inside the periedo-varenna formation is not known; this is particularly regrettable since the unit is about 600 m thick (gaetani et al., 1992). nonetheless, comparison with the other middle triassic faunas recovered not far from perledo (the kalkschieferzone from ca' del frate (viggiù, va, italy) and meride (ti, ch) and the besano-monte s.giorgio 'r tlinarìmenrn ,l; qcienze della terra, università degli studi di milano, via mangiagalli 34,lz)fi3 milano (italia). e-mail: andrea.tintori@unimi.it a. tintori fauna) helps us in the srratigraphic definition of the perledo fossils" the lack of a complere modern revision of the perledo fishes can be easily ascribed to the shortage of new material, giving even grearer importance to historical collections. bùrgin (1992) erected the new genus ctenognatbichthys, to which he assigned a few grenzbitumenzone specimens of the species ctenognathichtlrys bellottii, oríginally described on perledo marerial as heterolepidorus (?) bellottii by de alessandri (1910). bùrgin (1992, p.78) designated also a neotype for this species, assuming the original specimens to be lost. actually, the type specimen still exists in the collection of the liceo ginnasio 'a.volta' in como, where it has been housed for at least 60 years (fig. t). the type specimen ot ctenognathichthys bellotti (de alessandri, 191 0). bùrgin (1992, p. z8), describing the new genus ctenognathichthys, wrofe "the holotype .was said to be located at the museo civico di como, italy (de alessandri: 111). however, an intensive search failed to locate this specimen (g. achermann, pers. comm.). thus, it is necessary to designate a neotype. the neotype has been chosen from the tessin collection and represents an almost complete specimen (pimuz t.4349) lrom point 902 (layer 165), preserved in lateral aspect (figs. 83 & 84)". actually, i was also asked to help in locating the type specimen. at the time i did not know rhe exacr composition of the como school collection, so i gave bùrgin (phone comm.) the address and phone number of the liceo ginnasio 'a.volta'. on the other hand, g. achermann, a swiss reporrer living in italy, apparently never asked the director of the museo civico di como about the perledo marerial (l castelletti, wrirten comm.). thus, neither was the director of the museo civico di como directly inquired about the perledo mafig. 1 ctenognathichtlrys bellottii (de alessandri 1910): the holotype from perledo, stored et the liceo ginnasro statale 'a.volta', como, italy (catalogue n. 96"3890.10623). specimen dusted with ammonium chloride. scale bar 20 mm^ terial, norwas my suggestion foilowed. therefore, bùrgin's reasons for believing the holotype to be lost are very weak (iczn 1985, art. 75 d(3)). furthermore, the designation of a neotype is not compulsory (iczn 1.985, art" 75a); rn this case it was not necessary because no "exceptional circumstances" (iczn 1985, art. zsb(il) exrsted, ctenognathichtbys bellotti being the only species ascribed to the genus, with no possibility of being confused with similar species. in addition, other rules were not fulfilled by bùrgin (1992) in erecting the neotype, especially regarding its locality and supposed age. it comes, in facr, from the grenzbitumenzone of monre s.giorgio, whose stratigraphic and geographic positions are remarkably different from those of the type specimen (iczn 1985, art. zsd(s)). thus, in my opinion, even assuming the type specimen to be 1ost, there was no need to designate a neotype. f{owever, as rhe iczn (1985, art. z5h) states that similar cases must be referred ro rhe commission, i submitted the question ro the international commission on zoological nomenclature. finally, as some of the requirements for the designation of a neotype were not met, the holotype should be treated as the name-bearing type without any further commission pronouncement (p. tubbs, written comm.). by the way, in that same paper (búrgin, 1992), erecting the new species peltoplewrws nothocephalus, búrgin (1992, p. 123) gave the correcr etymology of the specific name, bur then used peltopleurus notbosomoides throughout the description as well as in the caprions (but see bùrgin, 1992, pag. 5 and 155). that the two names referred to the same species is proved by the holotype (pimuz 7.2902) which is figured as peltopleurus nothosomoides (bùrgin, 1992, fíg. 148) after having been cited as pehopleurus notbocephalu.s. as bùrgin himself agrees (written comm.), i consider pebopleurus notbocephalus as the valid specific name, for which the etymology has been given, under the iczn (1985, art. 24). the specific name peltopleurus nothosomoides must rhen be considered a junior synonym. the original marerial of ctenognathichtl4ts bellottii stored in como consists of a complete specimen, though the body and the posterior half of the skull are preserved as counterparts. glued to the specimen was a hand-written label (fig. 2), most probably by de alessandri himself, as all the specimens he cited in his work (de alessandri, 1910) bear similar labels. being on the underside of the specimen, the label is somewhat worn, but still legible: 'esempl(are) tipi(co)': type specimen. also the locality 'perledo' is clear, even if de alessandri wrote (1910) that the specimen was without previous 1abel, but being together with the other perledo specimens and in rock of the same lithology it could well have come from the perledo area. i agree with his interpretation because its preservation is that peculiar to the perledo fishes. in de alessanciri's (1910) picture this specimen appeared incomplete (see also bùrgin, 1992) because the head remained unprepared and covered by a thin laver of matrix. i have prepared the hidden region, but, as in most of the perledo material, the scarce bone preservation makes the boundaries of the individual bones very uncertain. the large tooth-bearing bones are detectable and the peculilr teeth. which gave lhe name to this genus, are also distinctly visible. the broad frontals and the other dermal bones of the sku1l roof cannot be traced in detail, but they seem very close to the restoration made by biirgin (1992). the opercular is more rectanguiar th.rn in bùrgin's restoration, and its size is comparable to that of the subopercular. ijnfortunately, the preopercular region is disturbed and its peculiar shape cannot be detected. squamation is composed of +l-++ transverse rows, each with 18 scales (respectively about 45 and 20 in the grenzbitumenzofle specimens). trunk scales are rectangular, and higher than long except in the six most ventral elements of each row, where scales become suddenly 419 fig. 2 the original label. most probably handwritten by de alessandri, which was glued on the lower surface of the slab of the holotype of ctenognatbícbthys bellottù (de alessanclri 1910). .r"r,, l^*, r h" l".oth hcino t qriee rhcir hciohr tnwrrd rhe cludrl region, scales become gradually rhomhic. losinq the denticularion o[ the posterior edge thar is quite strong in the trunk region. also. the ornlmenretion of r he scrles v,lries wir h position on rhe body. along the j^-^.1 ._ .-^::.. f-^rt oi theuvrùdr rrrdréllrr rll rrvl dorsal fin, dorsalmost scales are heavily ornamented with longitudinal ridges. thinner ridges rre present .ill along the trunk region, being somewhat stronger in the ventrai part, iust above the low ridges" the small dorsai fin originates at the 31st scale-row, while the anal begins four rows in advance; both fins are very small. the caudal fin shows 6-7 epaxial rays. more than 30 segmented lepidotrichia are preserved and the central ones probably have been destroyed in the past during preliminary preparation. the ventral lobe of the caudal fin, as well as the ana1, is preserved as impression: this kind of preservation, with the extremities of the fish remaining on the counterpart, is comrnon jmong the few known specimens from perledo. after the preparation of and the new observations on the como specimen, described and frgured by de alessanclri (1910), its conspecificity with the ne.w material described by btirgin (1,992) and its status as the original type specimen of the type species of ctenognatbtchth^t,s is bevond anv doubt. stratigraphic significance oî ctenognathichthys beliottii. vhile very little interest has been payed recently to the perledo vertebrate fauna. another famous middle triassic paleontologicai area, the besano-monte s. giorgio, h.rs seen r renell,rl of sttrdies, especiallv concerning fishes. the two localities are about 35 km apart from each other and both fossiliferous sequences span almost the whole ladinian. they formed in the same sedimentary basin, but under different environmental conditions (bernasconi, 1991; tintori, 1.992; bernascont ta rrva, 1993; i-ombardo, 1,997, tintori & lombardo, in oress). c. bellottii holotype rediscoaery + 1\) monte san giorgio a. tintori p. porroi, p. altolepis, a. nothosomoides, a. macroptera, f. trottii peltople u rus sp.n.a .--r.._.-'t-ll:-#l t:-ft-f:ff--t-i-rl r-r---t_---t-1 e l-r-----7r-t// // | @ t"ry t-v-;;71 r-r=-r=--] e;eitffil+ffi w=r-/--7---7c o o, .e j fig. 3 stratigraphy of the besano-monte s.grorgio and the perledo-varenna area with correlation based on fishes present in both the sequences ksz, kalkschieferzonel' pporroi, prohalecites porroi; paltolepis, perleidus altolepis; a.notbosomoides, àllolrpirtot6 nothosomoi'd.es; a,macroptera, aneurolepis macroptera: etrottii, furo trottii; c.bellottii, ctenognathicht/rys bellottii; s.cortasquornàsus, sauricbtlrys costdsqu+înosrts. lithology: 1, sandy limestones; 2, marls; 3, thin bedded limestones, sometime larninated; 4, organic matter rich shales; 5, massive dolostones; 6, volcanic ashes; 7, biocalcarenitic limestones; 8, intrabasinal breccias; 9, thin bedded dolostones. saurichthys macrocepnalus upper salvatore dolomite grenzbitu menzone -, c. bellottii , ;'' habroíchthys sp. s. cosfasquarnosus in the besano-m.te s.giorgio-viggiù area, the site of ca' del frate (viggiù-va) has been exploited since the beginning of the 80s (tintori et al., 1985; tintori & renesto, 1990; tintori, 1990a,b). it is situated in the kalkschieferzone, the upper member of the meride limestone (senn, 1924). this member yields vertebrates from several levels (bùrgin, 1995; furrer, 1,995; yirz, 1945; pers.obs.) across its 100 m of thickness; fish species, though, seem to have an uneven distribution, some of them being very common throughout the unit, and others being represented by single or raîe specimens. lombardo (1997) descrlbes 15 species from ca' del frate, while in different levels in rhe area of meride only a few species have been found, some so far unknown from ca' dei frate (tintori et a1., in press). after the 1996-97 field seasons in ca' del frate and meride, the recovered fish specimens number more than 3,000, of which at ieast 95o/o belong to the small species probalecites porroi and fewer than 5o/o have been collected around meride. a few fish species are common to perledo and ca' del frate: prohalecites porroi, perleidus altolepis, allolepidotus bellotti, peltopleurus sp. n. a, aneurolepis macroptera aîd fwro trottii. other species found in the perledo area are unknown in the kalkschieferzone, but have been found in the older vertebrate levels of the monte s.giorgio area (fig. 3). for instance, aparf from a detached mandible (tintori et. al, in press) and a iarge skull recently found perledo -varenna area lierna formation perledo member esino formation in the lower kalkschieferzone near meride, saurtchtlrys seems to be absent in the middle kalkschieferzone fossiliferous level of ca' del frate. this fact led tintori (1990b) to assume) for the first time, that the perledo fauna, where saurichthys is well represented, embraces different fossil assemblages. actually, rieppel (1985), in his revision of the middle triassic saurichtltys, observed that saurichthys macrocephalus is presenr in both the perledo and the lower meride lm. faunas and that saurichthys costasqua?nosus has been found in the grenzbitumenzone and the perledo faunas, without discussing the stratigraphical implicar ions. ascribing grenzbitumenzone specimens ro rhe species ctenognathíchtlrys bellottií, which is not found above this level, bùrgin (1992) fixther srrengrhened the simiiarity between the grenzbitumenzone and part of the perledo faunas, to which belongs the original specimen described by de alessandri (1910). the fact that saurichtlrys costasquamosus and ctenognathcbtlrys bellottii are found in the perledo-varenna formation, and are yielded also by the earliest ladinian beds of the grenzbitumenzone, seems ro contradict the age of the base of the former unit, which is considered upper early ladinian on rhe basis of conodonts (gaetani et al., 1992). however, the actual age difference between the two units yielding saurichtlrys costasquamosus and ctenognathichtlrys bellottii may be very small (a. nicora, pers. comm). the fact that these two species are found also in the late ear|y ladinian suggesrs their dit i n u) yo o q) c j .t o) = c. bellottii bolotype rediscovery references 421 stribution could cover a wider time-span than we previously knevr. actually, two species of cnenognatbichthys, one very close to cnenognathichtlrys bellottii, have been recently found in the middle ladinian prosanto formation of eastern switzerland (bùrgin, pers. comm.). the same can be hypothesized for other fishes of the grenzbitumenzone: some factors, such as a change in the depositional environment in the besano-monte san giorgio area, may have prevented their local preservation later in the ladinian. we suggest that during the lower ladinian the required conditions for vertebrate fossilization ended in the area of besano-monte san giorgio, while, with slightly different characteristics, they persisted in the perledo-vareflna area. only in the latest ladinian were both areas again suitable for preserving a significant vertebrate record: ar. fhaf time the kalkschieferzone and the perledo member yield a very similar fauna. the stratigraphic distribution of fish species or genera will be more precisely known when all the species contained in the three major vertebrate levels of the lower meride limestone have been individuated (furrer, 1995). thus, it seems that much evidence supports the hypothesis that the perledo fauna consists of three fossil assemblages, corresponding to the grenzbitumenzone (owermost ladinian), the lower meride limestone (lower late ladinian) and the kalkschieferzone (uppermost ladinian). acknouledgenents. thank are due to m. $filson (edmonton) and t. bùrgin (st. gallen) for the revision of the menuscript. i am grateful to dr. j. lorenzi of the soprintendenza archeologica della lombardia who gàve me the possibility to study the perledo material stored in the liceo ginnasio 'a.volta' in como. dr. l. castelletti (como museum) helped in undestanding the history of thìs collection. p tubbs (iczn, london) very efficiently helped in settling nomenclatural questions. photos are by g. chiodi; drawing by g. landra and m. r. torazzi. bernasconi s.m. (1991) geochemical and microbial controls on dolomite formation and organic matter production/preservation in anoxic environments: a case study from the middle triassic grenzbitumenzone, southern alps (ticino, switzerland). diss. e.t.h.-zúrich, n. 9432, l98 pp. bernasconi s.m. e{ riva a. (1993) organic geochemistry and depositional environment of a hydrocarbon source rock: the middle triassic grenzbitumenzone formation, southern alps, italylswitzerland. in: spencer a.m. (ed.) generation, accumulation and production of europe's hydrocarbons ili, spec. publ. europ. assoc. petrol. geoscientists, n. 3, pp. 179-190, springer verlag, heidelberg. bùrgin t. (1992) basal ray-finned fishes (osteichthyes; actinopterygii) from the middle triassic o{ monte san giorgio (canton tessin, switzerland). schueiz. palàont. abh., v. 114, pp. 1-164, basel. bùrgin t. (1995) actinopterygian fishes (osteichthyes; actinopterygii) from the klalkschieferzone (ijppermost ladinian) near meride (canton ticino, souther switzerland). eclogae geol. helv., v. 88, n. 3, pp. 803826, basel. de alessandri g. (1910) studii sui pesci triassici della i-ombardia. mem..!oc. 1t .lc. nat., v.3, n. 1, pp. 1-145, milano. furrer h. (1995) the kalkschre{etzone (upper meride limestone; ladinian) near meride (canton ticino, southern switzerland) and the evolution of a middle triassic intraplatform basín. eclogae geol. helv., v. 88, n. 3, pp. 827-852,basel. gaetani m., gnaccolini m., poliani g., grignani d., gorza m. ec martellini l. (1,992) an anoxic intraplatform basin in the middle triassic of lombardy (southern alps, italy): anatomy of a hydrocarbon source. riv. it. paleont. strat., v. 97, n. 3-4 (1991), pp. 329-354, milano. lombardo c. (1997) ittiofauna della kalkschieferzone (calcare di meride, ladinico superiore) di ca' del frate (viggiù, varese). tesi di dottorato, università degli studi di milano, 198 pp., milano. international commission of zoological nomenclature (19s5) international code of zoologicai nomenclature. 338 pp. international trust for zoological nomenclature, i-ondon. rieppel o. (1985) die gattung saurichthys (pisces, actinopterygii) aus der mittieren trias des monte san giorgio, kanton tessin. schaeiz. palàont. abh., v. 108, pp. 1-103, basel. senn a. (1924) beitràge zur geologie des alpensùdrandes zwischen mendrisio und varese. eclogae geol. helv., v. 18, n. 4, pp.552-632,basel. tintori a. (1990a) ca' del frate (varese). in pesci fossili italiani: scoperte e riscoperte, pp.25-35, milano. tintori a. (1990b) the vertebral column of the triassic fish saurichthys (actinopterygii) and its stratigraphical signi ficance. ri,u. it. paleont. strat., v.96, pp. %-ia2, mi-lano. tintori l. (1992) fish taphonomy and triassic anoxic basins from the alps: a case history. ria. it. paleont. strat., v. 97, n. 3-4 (1991), pp. 393-408, milano. tintori a. & lombardo c. (in press) late ladinian fish faunas from lombardy (njtaly): behaviour and paleoenvironment. .in: arratia g. er schultze h.p. (eds.) 422 a. tintori proceedings of second international meeting on mesozoic fishes "systematics and the fossil record", pp. ,buckow. tintori a., lombardo c., danini g.l., felber m.,marazzib. & vendico m. (in press) scavi paleontoiogici nella kalkschieferzone di meride (canton ticino, svizzera): risultati preliminari della campagna 1997. geol. insubrica, lrgano. tintori a., muscio g. 8r nardon s. (19s5) the triassic fossil fishes localities in ltaly. rio. it. paleont. strat., v.97, î. 2, pp. 197-21,0, milano. tintori a. & renesto s. (1990) a new lariosaurus from the kalkschieferzone (uppermost ladinia) of valceresio (varese, n. italy). boll. soc. paleont. it., v.29, n. i, pp. 309-319, modena. \lirz a. (1945) beitràge zur kenntnis des ladinikums im . gebiete des monte san giorgio. scbweiz. palàont. abh., v. 65, pp. 1-84, basel. rivista italiana di paleonrologia e stratigrafia dicembre 1999 nota breve short note amphiblestrum (aviculamphiblestrum) ruggeroi sp n., subgen. n. (bryozoa) from the \testern mediterranean sea a. rosso recehed april 4, 1999; accepted september 6, 1999 ,^n"^n.u, ruords: bryozoà' new taxa' deep-sea, recent, mediterriassunto. amphible stwm (aviculamphible strwm) ruggeroi sp.n. viene descritta e per essa viene creato un nuovo sottogenere all'interno del genere,4mphiblestrum. la specie proviene da fondi cìrcalitorali profondi ed epibariali del canale di sicilia e da località del mediterraneo nord-occidentale. 11 nuovo sortogenere è creato per distinguere, all'interno del genere amphiblestrwm, specie che hanno oltre ai tipici aviculari awentizi, anche grandi aviculari interzoeciali, entrambi sviluppati a partire dt dietellae. abstract. a new species and a new subgenus amphibles*um (aoiculampbiblestrum) rwggerol sp.n. are described from deep circalittoral-epibathval bottoms from the sicily strait and the north-western mediterranean. the new subgenus is created to distinguish, within amphiblestrum, species u.ith both gymnocystal adventitious and large interzooidal avicularia, both originating frorn basal pore chambers. introduction. during the study of some fossil and recent species belonging to rhe genus amphiblestrwm gray, i848, the examinarion of specimens from the mediterranean lead to the discovery of a new species. it roughly resembles amphiblestrurn auritum (hincks) from atlantic-mediterranean shallow water environmenrs (see rosso, in press). fiowever, the new species, besides gymnocystal adventitious avicularia, shows also large interzooidal avicularia, a feature not recorded, until nos/, for species referred to this genus. the generic attribution of the presenr new species and rhe introduction of a new subgenus aoiculamphiblestum within amphiblestrwm gray, 1848 is discussed. systematic paleontology class gymnoiaemata allman, 1856 order cheilostomatida busk, 1852 suborder anascina levinsen, 1909 family calloporidae norman, l9o3 genus amphiblestrum gray, 1848 subgenus aoiculamphiblestrurn subgen. n. etymology. from "avicularivn" tnd amphiblestum refcrrins to the presence of addirional interzooidal avicularia. fig. 1 di'rributron of an[ltiblc strum (av i c u latnp b tb le s t um ) ,uueroi rp. n., subgen. n. encircled star : studied .etnples; r rirngle : or hcr localities. drpartimento di scienze geologiche, sezione di oceanologia e paleoecologia, catania university, corso itala, 55, i-951,29, ross o @ mbox. unict. it. a. rosso fig.2e-banphiblestum (ariculatnphìbles*um) ruggeroi sp. n., subgen. n. holot,vpe. periancesrrular zooid (a) s'itir six margin;il spines and the proximal avicularium and ,r mature zooid (b) with four oral spines persisting in prcsence of the ,.r-icell. note the peculiar position of thc avicul:rrium. sicily strait, sample cr-90-8. scale bar = 2oo fm. diagnosis. as the genus, but with the addition of interzoordal avicularia, besìdes the gymnocystal adventitious ones, both originating fron'r basal pore-chambers. remarks. as for the species: see above. amphiblestrum (aviculamphiblestrum) ruggeroi sp. n. (fig. 2-a) 1988 amphiblestrunt minax zabah & maluquer, pl. 3b. 1988 nonamphiblestum rninar za'bilt 8r maluquer, p. 82, fig. 93. 1993 amphiblestrum sp. di geronimo et al., tab. 3. 1993 amphiblestwn fletningi, zrbah et al., p. 68, fig. 2. material. specimens come from post-\wùrmiin thanatocoenoscs (fig. t), collected from 201 (sample cr-90-15) to 212 m (semple cr-90-8), on and near the graham bank, in the sicily straits (di geronirno et al., 1993). etymology. frorn the author son's name: ruggero. types. the seìected holotype is a colony encrusting a valve of neopycnodonte cochlear (poli) on a scleractinian calvx of dendrophyllia cornigera (lamoroux). p:rratypes: one large and five young srnall colonies from the same sample. they all are dead and originate from st. cr-90-8, 212m,37" 14.38'n; 12" 42.31.'e. types are housed in the palaeontological museum of catania university (pmc) and are ìabelled pmc. 89. 26-2-1998. description. colony encrusting, flat. zooids oval ro pyriform (figs. 2a, b),425-566 (512, n = lo) gm long and 288-517 (334, n : 10) lrm 1arge, contiguous but well separated by grooves. area oval ro pyrifornl, occupying 4/5 ol the total length, marked by. a rhin crenellate raised rim. opesia oval ro subquadrangular, rarely with almost midway consrricrions. six to four distal thin spines (see fig. 2), four persisting on ovicellare zooids with the distal pair close to the orifice of the ovicelì. cryptocyst finely gr,rnuì.rr. nrrrorv rrnd sreeply rising from the distal edge of the opesia; proximally large and grading into the rim. gymnocyst smoorh, sreeply dipping away from the rim, only proximally produced, partly obscured by the avicularium ir supporrs. a singie gymnocystal columnar avicularium proximally located, occupying one third of the zooidal width (figs. 2a, b); the sharply triangular rosrrum sianted transversely or distal-laterally upwards; condyles stour; 'cross-bar absent; opesium elliptical. zoojds placed distally to an ovicell with one, often two, gymnocysral avicularia, laterai or disto-1ateral, symmetricaliy and transversely placed to the ovicell; the rostra pointing towards the middle forming a "roof ". interzooidal avicularia more ofren near the colony boundary (figs. 3a, b), polygonal c.a. one half rhe length of the autozooids. often as wide as an aurozooid. rosarnphíblestrum (aaiculamphiblestrum) from ruestern mediterranean sea fig. 3e-b ampbibles*urn (ariculanphiblestum) rugeroisp. n., subgen. n. holotypc. interzooidal avicularium at the colony edge and (b) an orher sector showing several interzooidal avicularia and an ovicell ovcrarched by two conver{ing ldventitions avicularìa. sicriy streit, sample cr-90-8. scale bar = 200 urn. trum horizontally 1ying, long, straight with thin, raised walls; the apex curving downwards. mandibular pivot blunt. oral shelf wide, opesium e1liptical, proximally bordered by a finely granular, concave rim. oviceli globular, prominent, seemingly not closed by the operculum, with a frontal, finely granular crescentic area separated by a raised suture from the marginal smooth ectoecium (figs. 2b, 3b) . ancestrula (figs. 4a, b) smaller than other zooids (340 ptm long and 250 prm wide) but similar in shape, with nine spines, budding a first distal zooid and tvro secondary disto-lateral ones, each having six spines, the more proximal pair at the beginning of the opesrum. new zooids are budded contemporaneously in distal and latero-proximal directions allowing the colony to achieve a rounded outline. the three first budded zooids lack the gymnocystal avicularium. remarks. colonies are very similar to species of the genus ampbib/estrum gray, 1848 for most of their characters, nevertheless, they have interzooidal avicuiaria, a very important diagnostic feature (cf. gordon, 1984). such avicularia actually originate from basal pore chambers (see fig. 3a). this condition is shared with smaller, f rontal adventitious avicularia of the same species and also with those described in other species referred to amphyblestrum such as a. flemtngl (busk) from rhe ne arlanric. the new subgenus aaiculamphi blestrum is, thus, introduced, within amphiblestrum, to stress rhe nresence ^{ ltto" i.t"t"^^ì.{ .l .rvicullria,.'6" besides the sma11 gyn-rnostal ones, both having a conrmon origin and different position and size. this also happens in species of chaperiopsis uttley, 1949, such as c. multifida (busk) (gordon, pers. com.) and callopora gray, 1848, such as c. smttti (kluge). virhin the genus amphiblestrum, this peculiar feature is presently known only for a. ruggeroi. large interzooidal avicularia develop from lateral or disto-lateral basal pore chambers, attain large sizes, almost sin-rilar to rhose of zooids, and show a random location among t1-re zooids, although rhear rr" mnre enmmnn ncîr rl" "^l^.t"".,../ nr,ìrglns. mandibles were not observed but thelare presumably straight and pointed rs, in rhe family calloporidae, mandibles are usually quite close in shape to their skeletal rostra (vinston, pers. com.). the specimen figured loy zaba\a & maluquer (1988: fig. 38) and zabala et al. (tll:) have been synonymised with,,1. ruggeroi as it seems to share with the present species all diagnostic features. zabala et al. (1993), indeed, doubtfully referred their material to a. flemingi (busk) stressing differences in the number of 168 a. rosso !ig. 'la-bamphiblestutn (aticulamphiblestrum) rugerol sp. n., subgen. n. a paratype. ancestrula (a) and periancestrular arer (b) showing firsi astogenetic budding pattern. note the interzooidal avicularium laterally budded from the rncestrula. sicill' g152i1, sanple cr-90-8. )cîtc dîr luu urn. spines (4, rarely 6), the absence of the proximal gianr spine, the shape of the opesium and that of the ovicell frontal area. all these characters are obvidus from photos and they are shared with ,4. rugeroi. moreover, the figured ovicells show distal single or paired avicularia transversely placed to form rhe "roof " peculiar to ,4. ruggeroi and not described for any other species in the genus. the specimen from rhe zabala collection figured by lópez de la cuadra & garcía gomez (199a) is different and actually belongs to a. flemingl (busk). distribution. the species s/as found in two samples from sicily straits, between 201 (sample 15) and 212 m (sample 8) (di geronimo et al., tl13;. specimens were all dead, when sampled. community is rich and diversified in the former and extremely scant in the latter. thanatocoenoses (i. e. skeletal remains present in the bottom sediment) are more rich and diversified. both testify the mixing of outer shelf (deep circalittoral) rocky bottom organisms or skeletons (belonging to the rocky offshore assemblage) with faunas peculiar to upper slope (epibathyal) muds (bathyal mud assemblage). samples contain sma1l branches of the gorgonacean corallium rwbrum (linnaeus), large fragments of the scleractinian dendrophyllya cornigera (lamoroux) and large valves of the mollusc neopycnodonte cocltlear (poli). living bryozoans are scant but diversified comprising both rigid erect species (annectocyma tubwlosa, entalopboroecia deflexa, e, gracilis) and unilaminar species among which puellina (cribrilaria) innomrnata, p. (glabrilaria) ped.uncwlata, escharina hyndmanni, e. au lgari s and cras sim ar gi n a t e i i a s o i i d u i a encrus rin g lar ge molluscan and scleractinian skeletons. dead specirnens mainly belong to erecr species: terr.,ta irregularis, hornera frondiculata, h. iichenotdes, palmicellaria c[. elegans, sertella couchi biarticwlata, s. septentrionalis, tessaratloma borea/e and scrupoce/larta delili. e,ncrusting taxa are also common pue/lina (cribrtlaria) lnnomtnata, p. (c.) radiata, crassimarginate/la solidula and escbarina oulgaris. a. ruggeroi is also present in the blanes canyon from 180-350m where colonies encrusr a par-rly alive colony of the deep-sea scleractinian madrepora oculata (ztbala et al., tllt;. the species is, therefore, known only from deepsea (lower circalittoral to epibathyal) sites from the western mediterranean. acknouied,gements, i arn grrteful to: l)r. c. m. lópez de la cuadra (dep. fisiologia y biologia animal, sevilla), dr. j. vinston (american museum of natural history new york) for useful inforrnation; dr. d. p gordon (new zealand oceanographic institute, .wellington), dr. i.-g. harmelin (station marine d'endoume, marseille) for discussions and suggestions. i would also thank mr. o. torrisi (isrituto internazionale vulcanologia, cnr, catania) for sem assisrrnce. paper financially supported by min. ris. agr. al. for (prof. di geronimo) and murst lrosso). di geronimo i., rosso a., sanfilippo r. (1993) -the corallium rwbrurn fossiliferous banks off sciacca (strait of sicily). in: cicogna f. & cattaneo-vietti r. (eds): red coral in the mediterranean sea: art, hystory and science. min. ris. agr. al. for., pp. 75-107, roma. gordon d. p (1984) the marine fauna of new zealand: bryozoa: gymnolaemata from the kermadec ridge. nezo zealand oceanographic institute memoir 91, pp. 1-198, \x/ellington. rosso a. (in press) amphiblestrum gray, 1848 (bryozoa cheilostomatida) in the atlantic-mediterranean area, with description of a new species. journ. nat. hist., london. 469 lópez de la cuadra c.m. & garcía-gómezl.c. (99a) bryozoa cheilostomara: the genus amphiblcstrum in the \flestern mediterranean and the f vst sessibugula of atlantic waters. journ. nat. hist., v. 28, pp. 683-693, london. zabalam. & maluquer p (1988) illustrated keys for the classifications of mediterranean bryozoa. treballs del museu de zoologia, v. 4, pp. 1-294,barcelona. zabalam., maluquer p & harmelin j.-g. (1993) epybiotic bryozoans on deep-water scleractinian corals from the catalonia slope (western mediterranean, spain, france). scientia marina, v. 57, pp. 65-78, barcelona. amphiblestrwm (aoiculamphiblestrum) from zaestern meditenanean sea refe,rences lucas 323..326 ����� ���� � ���� �� � ����� ������ �� ����� �� ��� � �� � ����� � ���� ��� ����� � ����� �� ������� � ����� �� ��� ����������� ���� � � � ��� ����� ����� ��������� �� ��� �� ���� ��������� ����� ��� ��� ��� ������ ������� � �!"#$ �$� %%�&� � '!(!�)� �*+� ( �+�%� �+!,#(� � ��������� ��# -�$%. $#&!$/ ! -!%%�+ ������ ( �( �+!,#(� �% -$ 0�#(. ! $��� 1. " % &!++#&.#/ �( .�# �!"#$ �$� %%�& !.�# �*+� ( �+�%� . .�# �.*/!$%2� �$#, + $# � ��# & $�!( .# %*&&#%%�!( !.�# �!"#$ �$� %%�& . �.*/!$%2� �$#, + �% &� $ &.#$�3#/ �) +.#$( .�!(% !/!+!��.#� +��#%.!(#� � $+) +��#%.!(# (/ � $+%.!(# "�.� � $.�&*+ $ �#/% !!!+�.�& +��#%.!(#� ��# � &$!*( -!*(/ . .�# %�.# �(/�& .#% ( �+#(#2� ( 0#� ��# �!(# -$ 0�#(. �% $!/�+�2# #+#�#(. 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(foraminiferida) from the middle triassic (anisian) of northern dolomites, italy roberto rettori*. baba senowbari.daryan's* & rainer zuhlke*** key-utords: paleontologl foraminifers, triassic, anisian, alps, nonhern dolomites, italy. riassunto. nel presente lavoro viene descritto un nuovo gener€ di foraminifero sessile a parete agglutinante: flatscbkofelia (specie-tipo: flatschkofelia anisica gen et sp. nov.). questo foraminifero è stato rinvenuto nelle brecce di talus recifale di età triassico medio (anisico) de1la formazione del serla superiore (dolomiti di valdora, italia). flatscbhofelia, gen. nov., è riferito alla famiglia placopsilinidae rhumb1er. abstract. the new genus of sessile and agglutinated foraminifer, flaxschhofelia (type-spectes: flatschhofelia anisica gen. et sp. nov.) from the middle triassic (anisian) reef talus of the olang dolomites (ljpper serla formation, dolomiti di valdora, italy) is introduced. flaxcbkofelia, gen. nov., is referred to the family placopsilinidae rhumbler. lntroduction. after the end-permian mass extinction, reefs and reefal carbonate buildups did not exist during the late permian to early triassic. first occurrence of reefs has been reported from the early-middle anisian (middle and late pelsonian) of the dolomites, italy and from southern china (see senowbari-daryan er al., 1993). in the northern part of the western tethys additional anisian reefs are known from the carpathians and from the karawanken mountains. from the southern paft of the western tethys no anisian reefs are known. most of late anisian reef carbonates in the olang dolomites (southern alps, italy) are represented by allochthonous talus reef blocks of one or several cubicmeter size. autochthonous reef buildups are extremely rare. both, the allochthonous reef blocks and autochthonous buildups occur within the middle to late pelsonian recoaro formation. the fossil content as well as the microfacies types of these reef carbonates are described by bechstàdt 6c brandner (1.970), fois & gaetani (198a) and by senowbari-daryan et al. (1993) in detail. olangocoelia ozr bechstadt & brandner, a problematic fossil (sponge or alga?), is the most abundant organism in the anisian reef carbonates, followed by different types of sphinctozoid and inozoid sponges, bryozoans, solenoporacean algae, corals, encrusting epibionts (porostromate algae, cyanophycean crusts, microproblematica, worms and foraminifers) (see for detaii senowbari-daryan et a1., 1,993). foraminifers are relatively rare and mostly represented by meandrospira dinarica kochansky-devidè & pantic, pilammina densa pantic, endoteba sp., endotebanella sp., endouiada sp., endotriadella sp., glomospira sp., glomospirella sp. duostominidae etc. sessile foraminifers are also not abundant and represented by the questionable gena,s bullopora? andby an agglutinated type, described here as flatschhofelia anislca gen. et sp. nov. systematic description foraminiferida eichwald, 1830 suborder text u i a ri i n a delage & hérouard, 1896 superfamily l i t u o i a c ea de blainvitle, 1827 family placopsilinidae rhumbler, 1913 subfamily placopsilininae rhumbler, 191.3 genus flatschkofelia gen. nov. type-species: flatschkofelia anisica gen. et sp. nov. derivation of name. after the locality flatschkofel, olang dolomites, italy (fig. 1), where the holotype was found. diagnosis. test simple, agglutinated and solid structured, attached on the early coiled stage, free and uncoiled on the final stage. chambers are biserially ar'r dipanimento di scienze della terra, università degli studi di perugia, piazza università 4, 06100 perugìa, italy. 'r'r institut fùr palaontologie, universitàt erlangen-nùrnberg, loewenichstrabe 28, d-91054 erlangen, germany. ooo geologisch-palàontologisches institut, universitàt heidelberg, im neuenheimerfeld234,d-69120 heidelberg, germany. /'"\:--' )--r-)-^ ì-v;fir*.--'; / a n a b@en mamolada o.-l--30* furkelpm piz da per6 dreifrngerspitze locality ,é maurerkopf ^^flatschkofel o12|lrn r. rettori, b. senoubari.dartan €r r. zùblke fig. 1 location map of the dolomites and flatschkofel, where flatscbkofelia anisica gen. et sp. nov. was found. ranged in the initial stage, biand uniserial in later stage. aperture simple and terminal located in the first biserial stage, moving to the margin in the second uniserial stage. composition of the genus. at the moment monotypic, only represented by the type-species. stratigraphic and geographic distribution. middle triassic (anisian), northern dolomites, italy. remarks. flatscbkofelia gen. nov. is herein included in the family placopsilinidae rhumbler, 1913 deilned by i-oeblich & tappan (1987, p. 80) as follows: "test attached, early stage coiled, arcuate, or biserial, later uncoiled; wall agglutinated, solid, aperture terminal, single or multiple. l. jurassic to flolocene" tho f"-il-' i..l'161s5 two subfamilies: placopsilininae rhumblen i9i3 and adhaerentiinae loeblich ec tappan, l986. flatscbkofelia gen. nov., is referred to placopsilininae because having a coiled early stage even if all the genera belonging to the subfamily lacks a biserial paft. the biserial arrangement of the chambers is a morphoiogical feature of adhaerentiinae, but the unique genus (adhaerentia plummer, 1938, type-species adhaerentia midaayensis plummer, 1938) included in the subfamily lacks an early coiled stage. the presence of a coiled early stage, a biserial second part and a final uniserial part, which is common both in placopsilininae and adhaerentiinae, rogerher with aperturai features, could justify the introduction of a new subfamily for flatschkofelia, gen. nov. flowever, the material from the northern dolomites is not enough to introduce a new subfamily. at the momenr, flatscbkofelia is referred to placopsiiinidae also because the lower paleocene genus adhaerentia shows, in the final srage, aperrure multiple with two to many separate openings and for stratigraphic reasons too. according to loeblich & tappan (1987) the stratigraphic range of placopsilinidae spans from i-ower jurassic to holocene, but we can affirm the range of the family starts from triassic, nor only on rhe basis of the existence of the new genus flatschkofelia, bur also because of previous record of the genus placopsilina in triassic sediments (trifonova 1967, 1992; brónnimann & zanínettt. 1972\. flatschkofelia anisica gen. et sp. nov. pl 1., íig. 1-12 1993 sessile agglutinated {oraminifer (gen. et sp. indet.) senowbaridaryan, ztihlke, bechstid & flùgel, p. 230, pl. 64, fig. 2,4,5. derivation of name. because rhe occurrence of the soecies in anisirn limestones. holotype. longitudinal section illustrated in pl. 1, fig. 1 (thin section: 89lv4,/18). paratypes. all specirnens illustrated in pl. 1, fig. 2-12 from different thin sections (see explanation of plate 1). type locality. flatschkofel, olang dolomites, italy (see fig. 1). type level. upper serla formation, middle triassic (anisian, pelsonran). material. several specimens in different planes from thin sections: 89/y7/49, 89/vr/85, 89/v4/t8 (holotype), 89/vr, 86/2, 89/y7/32, deposited at the institute fùr pa1àontologie, universitv of erlangen-nùrnberg. diagnosis. see diagnosis of the genus. italy; middle triassic (anisian, 8, ia) 8e/y4/1.8r 7) 89/y1t e) plate 1 fig. 1-12 flatscbhofelia anisica gen. et sp. nov. from flatschkofel n, upper serla formation, olang dolomites, pelsonian). fig. 1: holotype, thin section 89/v4/1,8. fìg. 2-72: paratypes, thin sections: 2) 89/v7/32i 3, 4, 5, 6, 86/vr/85r 1r, 12) 89/v1/8s. 415pl. 1 flatschkofelia anisica gen. et sp. no'u. .,,ts o o.smm r. rettori, b. senoubari-daryan & r. zilhlke description. the test is attached, at least for the initial part, then, probably, may grow free of the attachment. it is irregulariy coiled in the reduced early stage, later uncoiled and rectilinear, slightly contorted above aii in the final part. the coiled early stage is followed by a short stage with 2-3 series of subangular chambers biserially arranged. the final part is uniserial and the chambers (5-7 ín the more developed specimens) are irregular in shape, increasing more rapidly in breadth than height. the aperture is simple, terminally located in the biserial stage, moving to the margin in the uniserial part, at the end of a short neck in any chamber offset toward the opposite margin in respect with the previous chamber. the chamber wall is solid, agglutinated and simple in structure. dimensions of the test (in mm): lensnt: u.)-1.+u maximum diameter: 0.4 height of the chmbers in the uniserial stage: 0.06-0.08 breath of the chambers in the uniserial stage: 0.16-0.25 thickness of the *all: 0.05-0.06 stratigraphic and geographic distribution. middle triassic (anisian, pelsonian), northern dolomites, italy. association. flaxcbkofelia anisica, gen. et sp. nov., was found within upper ramp and lagoonal facies of the prograding upper serla formation. the underlaying recoaro formation and the upper serla formation contain allochthonous reef facies (reef talus biocks, reworked bioclastic respectively). for details on anisian reef organisms of olang dolomites see senowbary-daryan et ai. (1993). the species is associated with rare foraminifers mainly represented by endoteba sp., endotebanella sp., endotriada sp. and endotriadella sp. in the reef facies; meandrospira dinarica kochansky-devidè & pantic and pilammina densa pantíc in the lagoonal facies. acknouled.gments. the investigations were forschungsgemeinschaft (pro;eu priority program "global and sedimentation". supported by the deutsche be 641/17, anis-rifq in the regionel controls on biogenic references bechstàdt t. & brandner r. (1970) das anis zwischen st. vigil und dem hóhensteintal (pragserund olanger dolomiten, sùdtirol). festband geol. inst., 300-jahr-feier, univ. innsbruck, pp. 9-103. brónnimann p. & zaninetti l. (1,972) foraminifera from the basal upper muschelkalk at hyères, western basse-provence, southern france. rio. it. paleont. strat., v.78, n. l, pp. 43-44, milano. fois e. 8r gaetani m. (1984) the recovery of reef-building communities and the role of cnidarians in carbonate sequences of the middle triassic (anisian) in the italian dolomites. paleontograpbica amer., v. 54, pp. l9l2a0, ithaca/new york. loebiich a.r. e tappan e. (1987) foraminiferal genera and their classification. van nostrand reinhold comoanv inc., vol. oî 970 pp., new york. senowbari-daryan b., zùhlke r., bechstàd t. 6c flùgei e. (1,993) anisian (middle triassic) buildups of the northern dolomites (italy): the recovery of reef cornmunities after the permian,/triassic crisis. facies. v. 28, pp. 181-256, erlangen. trifonova e. (1964 some new triassic foraminifera in bulgarla. ann. unio. sofia, geol., geogr., fac. 60 (1 geol.) pp. 5-6, sofia. trifonova e. (1992) taxonomy of bulgarian triassic foraminifera. i. families psammosphaeridae to nodosinellidae. geol. balc., v. 22, n. 7, pp. 3-50, sofia. receioed june 20, 1996; accepted october 3, 1996 rivista italiana di paleontologia e stratigrafia rrrorne 4ì9-444 drcembre 1999 nota breve short note a ne\t multidiscus ? species (foraminifera) from a fusulinacean-rich succession encompassing the carboniferous-permian boundary in the hadim nappe (central taurus, turkey) cengiz okuyucu received nooember 1 1, 1998; accepted june 3a, 1999 key-uords: m uhidiscus ?, fusulinacean, carboniferous-permian boundary, biostratigraphy, hadim nappe, turkey. riassunto. in questo articolo vengono studiati gli stratì ricchi in fusulinidi intorno al limite carbonifero-permiano, provenienti dalla sezione di qatalkatran tepe, falda di hadim, nei taundi centrali. la comparsa di sphaeroschwagerina sp. vtene vrllizzar^ per definire il limite carbonifero/permiano in questa sezione. la presenza dt psewdofusulina ? bwzuluh.ensis dobrokhotova, pseudofusulina paruiflucta zhou, ps eudofus ulina hot unensls davydov, o ccidentoscbzr;agerìna (7) konaensis echlakov e di rugosofusulina stabilis rauzer-chernousova conferma l'età asseliana della oarte superiore della sezionc. abs*act. the hadim nappe carboniferous-permian boundary and its fusulinacean-rich strata from the qatalkatran tepe stratigraphic section were studied. the appearance of spbaeroschuagerina sp. determines the carboniferous-permian boundary in this section. the presence of pseudofusulina ? buzulwhensis dobrokhotova, pseudofusulina paruij'lucta zhoq pseudofusulina horunensls darydov, occidentoschuagerina (?) kosvaensìs echlakov and rwgosofusulìna stabilis rauzer-chernousova confirm the asselian age. a new multidiscas ? species, mubidiscws 7 tauridiana n. sp. was discovered from the early permian (asselian) of turkey (hadim nappe, central taurus). lntroduction. the study area is located in the southwestern part of sariveliler (karaman) in the central taurus region (fig. t). this area belongs to the hadim nappe that was described as an allochthonous tectonic unit to the south of hadim by blumenthal (1944,1951). gùvene (1965, 1969, 1974) studied different sections extensively and described a rich microfauna and microflora of visean, bashkirian, moscovian, kasimovian, gzhelian, and early and late permian deposits. subsequently, ózgul (1976, 1984) studied the structural geology of the hadim nappe. the last biostratigraphic studies of this region were carried out by góktepe (1996), okuyucu (1997), and okuyucu and gùven e 0997). the carbonif erous-permian boundary (c-p boundary) of the hadim nappe is represented mainly by quartzitic sandstones with iron oxide concrerions and a very shallow marine limestone with very special facies known a"s gin)anella limestone (gúveng, 1991). the pseud.oschwagerina zone of the asselian is represented by the girz,,anella lime stone (gùvene, 1.965, l977 b), composed of algae, fusulinids and fragmenrs of fossils (brachiopods, echinoids, ostracods erc.). the c-p boundary beds that are rich in fusulinaceans s/ere investigated and a new muhidiscus ? species, mwhidiscus 7 tauridiana n. sp. was described from early permian deposits of the qatalkatran tepe stratigraphic section. stratigraphy. tl^ rr^l:'nt^^pe contains carbonate and clastic deposits from the late devonian to lare creraceous. the late carboniferous and early permian srrata include a well-known giroanella limestone facies (fig. 1) . gúveng (1977 b,1980) examined the different sections of the hadim nappe and divided the late carboniferous and early permian strata into two formations, gavuralani and dikmentepe, respectively (fig. 1). only the lower levels of the gavurala formation were investigated. the levels studied begin with red, more or less sandy limestones. the upper levels are rich in iron oxide and characterized by girz,anella limestone. the upper carboniferous age is indicated by the exisrence of gin,anella sp., globioalpwlina bamensis reitlinger, maden tetkik ve arama genel mùdúrlùgù, jeoloji etútler dairesi 06520 sogiitcizti, ankara, turkey. e-maii: okuyucr:@mta.govtr c. okuyucu location and geological map of the strldv rre,r.rnd (.,rrlk.rt rrn tcpe strrtigrrphic .ccr10 n. globiotah,ulina sp. a, and ammoaertella sp. (platr 2). biostratigraphy. ruzhentsev ( 1950) developed a scheme for the c-p boundary interval utrlizing ammonoid evolutionary stages and established four successrve stratigraphic units. later, ruzhentsev (1952) correlated these stages with fusulinid data from rauzer-cl-rernousova and rozovskaya. in 1954, ruzhentsev placed the lower boundary of the permian sysrem at the base of the asselian stage and accepted the sphaeroschwagerina z, u lgaris sp h aero s chraagerina fusifòrmis zone as the base of the asselian (bogoslovskaya et al., lq95). the recently àccepred definition of the c-p boundary at aidaralash creek (d"rrydov er rl.. 1995) ;pproxim:rtes the horizon proposed by ruzhentser' (19 52). chuvashov et al., (1986, 199a,1993) also accepted ruzh-entsev's definition (the brrse of asselian begins with the first appearance ol sphaeroschzaagcrtna genus). in this study, the aurhor accepts ruzhentsev's (195a) asselian definrtion (fig. z). gùveng (1965) detertetrataxis minwta morozova, lrleotuberitina maljavkini mined the kasimovian stage of the upper carbonifermikhailov, quasifusulina sp., ozawainella sp., schuous based on rhe presence of quasifwsulinoides sp., bertella obscura (lee & chen), and schubertella obscura schubertella sp., fusiella sp., pseudofusulina sp., and the procera rauzer-chernousova (plate 2) . gzhelian stage by the presence of daixina sp., pseudothe lower levels of the dikmentepe formation fusulina sp. and schubertella sp. gúvene (1965) suggesrcharacterized by the girvanella limestone and a red ed that the conglomerate and sandstone beds represent limestone were studied. the asselian age is demonstratthe upper part of the gzhelian, and the asselian may be ed by the presence of sphaeroscbzaagerina sp., occidenmarked by an unconformity. later, gùveng (1965,1977 toschwagerina (?) bosaaensls echlakov, psewd.ofusulina ? b) defined the girvanella lirnestone as asselian or equivbuzwlukensis dobrokhotova, pseudofuswlina hoounensis alent to the pseudoschraagerina horizon in the góksu davydov, rwgosofusulina stabilis rauzer-chernousova, valleysection. rugosofusulina stabilis longa rauzer-chernousova, tritin this study, fossils of gzhelian age are repreicites aff. pan)us chen, hemigordius sp., mwltidiscus ? sented by giroanella sp., epimastopora sp., garooodia tauridiana n. sp., tetrataxis parviconica lee & chen, sp., eotuberitina reitlingerae miklukho-macklay, tetratat-r:iîl ei=t{ti cìaysy ginànàlla limestone rtlttl fi , --i quartzitic sanosrone ezi ,------l covered interval ttr] ffi limestone fr-tî5tl ginanella limestone tt| l-l-t-.1rtclavey llmesrone tr à f a a rt f a a (, a àp z tì 'a a foramini l-e fusulinacean argae s 'ì s rlri-.3= l-: ì; è.è iì p^\ sèè \è s<-\\ fe è è d\ : \pì r !r $* i ; =ùè.èè :i= 5: sisé = i *esè"i 3éèéièèès <*ìfì -;èi=èsùíyi liiii i3i èèèèièèèì;èèèèèèèèè sss;èssàèèìièiéeèèi 3€i o.xx úè 3f i *è ì èùèèè f.èf-èè:è: !:sìtt :!ù::.:è cú,ì'ùèù z è, 130 129 t28 i2'1 126 t25 124 r23 \22 121 12c 119 118 117 116 i 15 tl4 113 ,,;4, :]:::: ,:l:,:oo ,ooi:; l 'o':iaaoo i : : ooa : : r : aao aa l o:o o.o r oo ,a. a: :o ool t,' o:o t o o a o u) 3 z (, n tii tll t1( t0f t0t ró, 10( 10i l0r l0: oo: .4,,4 oaaa or ì 'oo l :l ,l :, .'l ':l :'o, io: o a:o .f '.,'. ,f 'o to: a.: , xis minuta morozova, tetrataxis hemisphaerica elongata morozova, tetrataxis plana morozova, tetrataxls sp. a and b, bouhonia zuillsi lee, quasifusulina sp. a, and qwasrfusulina sp. b. asselian fossils are represented by sphaeroschwagerina sp., occidentoschruagerina (?) kos'uaensis echlakov, triticites schuageriniformis kauzerclrernousova, triticites afî. paruus chen. triticires sp., pseudofwsulina i bwzulukensis dobrokhotova, pseud,ofw s u lin a p ail en s i s fergan en s ls dutkiev rt ch, p s e u d ofus w lin a hoztunensis davydov, pseudofuswlina. ex gr. ellipsoides grozdilova, pseudofusulina mikhai/ovi leven, pseudofusulina saratoztensis. chernova, psewdofusulina sp. a, rwgosofusulina stabilis rauzer-chernousova, rugosofusulina stabilis longa rauzer-chernousova, hemigordiws sp., multtdiscws ? tawridiana n. sp.,ammodiscus sp., palaeonubecularia sp., and glomospira sp. (fig.2). in the qatalkatran tepe section, sphaeroschwagertna, boultonia, schubertella, psewdofusulina and quasifusultna are found together. the c-p boundar,v is char441 fusulinrcern, .rlgre, end 'mrll [orrrninr[er di.rribution of qatalkatran tepe 'rrrrigrephrc secrion iriter okuyucu .rnd gúvenc. 1997. modified). -^*^-:-^l i-,..l^ ^-^^arance ofurl lrr! &$ �� ��� ������� ������������� ��� �!� ���������� '��" ���� ���� � ������� ���,���+� ��� ���� ��� ��� �� �"� ����$ ���" � !������ "�� '��� ��!����� ���� !�� �� ���������� %������� 666&$ �� �"� '���� �! �"��� �������� ������������ %��� ����� 11<��'� 666= (��'��� ����� 66 & �! ����� ��-��� �� ����� ����������� ����� ���� �� '��� � ���� ��'�� �� ����#��*� �������� ����������� ��� ����$ ?� � �����;������ @����� ��,�� % 66 & ������!��� � �"��� ���������� ����������� �"������ #����� !�� �"� !� �� �"��� ����" $�� ��% ��� a������ ���� ��� ��������� ������� 7����� % 66>& ��� 7����� ������� % 66b& ����#��*�� �������� ����������� ���-���� -����'��� � ��# �"� ������������� -����'���� ���� '��#�� �! 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(2019) on the validity of terebratula sinuosa (brocchi). riv. it. paleontol. strat., 125(3): 631-636. rivista italiana di paleontologia e stratigrafia (research in paleontology and stratigraphy) vol. 125(3): 631-636. november 2019 abstract. here we aim to fix some nomenclatural problems relating to the definition of terebratula sinuosa. in 1616 fabio colonna first described two different brachiopod specimens from italy which were later attributed to the genus terebratula. in 1758 linnaeus erected anomia terebratula in reference to the drawings of colonna. he described the heavily sulciplicate specimen figured on the upper left (specimen number 4) but addressed the specimen as if it was the number 1 in the figure (upper right). several authors later inadvertently followed the error of linnaeus. the neotype for t. terebratula, indicated in 1998 by lee & brunton, refers to the specimen number 1 in colonna’s figure (the one to the upper right). the two specimens in colonna were originally considered synonyms. however, the sulciplicate specimen number 4, originally figured by colonna, refers to a distinctive miocene terebratula species, which has been often referred to as terebratula sinuosa. we review evidence in favour of such a designation and provide stratigraphic and morphological evidence that t. sinuosa deserves the full rank of species. the name t. sinuosa should be maintained given the long tradition of the name in the literature, and the definition of t. terebratula should therefore be amended. received: april 10, 2019; accepted: june 04, 2019 keywords: brachiopoda; miocene; nomenclature problems; fabio colonna. introduction in 1616 fabio colonna first described two brachiopod specimens (fig. 1) as “concha anomia vertice rostrato”. on page 22 of his book he figured the specimens and referred to the individual on the upper right as n. 1. this specimen comes from the “calcareniti di gravina” formation (near andria, apulia, southern italy) and is pliocene-pleistocene in age. the individual on the upper left, indicated as n. 4, refers to a specimen with sulciplicate commissure coming from the private collection of ferrante imperato, a famous xvith century neapolitan natural scientist. the provenance of this particular individual is uncertain, although it was possibly found in miocene outcrops of apulia (southern italy). unfortunately, both specimens were lost. linnaeus (1758) applied his formal nomenclature scheme to the drawings of colonna erecting the species anomia terebratula. unfortunately, he misinterpreted the captions of the original illustration. while the description of linnaeus clearly refers to colonna’s specimen n. 4, he reported it as if it was the specimen originally labelled as n.1. consequently, the reference of linnaeus to the formal name anomia terebratula did not match the original numbering scheme. in 1814, brocchi erected the species anomia sinuosa and anomia ampulla. he referred to the former describing a brachiopod possessing “testa oblonga, valva superiore uniplicata, altera biplicata, margine infero sinuoso, apice perforato. column., de purp., pag. 22, fig.1 (fossilis)” addressing the same figure as linnaeus’ a. terebratula (which is indeed specimen number 4 in colonna). then, he described and figured a. ampulla as having “testa inflata, valva inferiore basim versus obscure biplicata, altera rotundata, laevi, apice prominente taddei ruggiero e., serio c. & raia p.632 pertuso” referring to a specimen figured in agostino scilla’s vana speculazione (1670, tab. 14, figs. 1, 2). from this moment onward, most authors have followed linnaeus’s wrong reference to colonna’s illustration, or used brocchi’s a. sinuosa in reference to a heavily sulciplicate, late neogene terebratula from the mediterranean area. this enduring attitude is obviously wrong because the definition of either terebratula terebratula or terebratula sinuosa was clearly incomplete. to fix this problem, in 1998 lee & brunton designated the specimen bm(nh) bg152 from the pliocene of andria (southern italy) as the neotype of t. terebratula. lee et al. (2001) later revised the species t. terebratula. they grouped under t. terebratula exclusively sulciplicate miocene specimens and uniplicate to sulciplicate pliocene forms, thus considering the two individuals of colonna as conspecifics. here, we aim to fix the nomenclatural problems regarding the genus terebratula. in particular, we find it timely to propose an amendment of the current definition of t. terebratula recognizing t. sinuosa as a separate valid species. several studies have highlighted the profound morphological distance between typical t. terebratula and specimens usually referred to as t. sinuosa. we revise these studies and the stratigraphic distribution of the two species. previous reference to terebratula sinuosa in the scientific literature after brocchi (1814), the name t. sinuosa was used by davidson (1864) to indicate fossil specimens from malta. davidson himself later (1870) described several tertiary brachiopods from italy including terebratula bisinuata, t. sinuosa, terebratula rovasendiana, terebratula grandis, terebratula pedemontana, terebratula ampulla and terebratula regnolii. the name t. sinuosa was used by seguenza (1865) in reference to specimens found near messina, and later re-used by the same author in reviewing the collection of costa (seguenza 1870). subsequently, seguenza (1871) reviewed tertiary brachiopods from southern italy. in his focus on terebratula, seguenza described t. sinuosa, t. ampulla, t. pedemontana, t. philippi, t. regnolii, t. romboidea, and t. siracusana. in the same paper he erected the species terebratula calabra, terebratula costae, and terebratula scillae. it is worth noticing that seguenza erected t. costae from the miocene of monteleone (now vibo valentia, calabria) and malta, including the taxon t. biplicata previously recognized by costa within the list of synonyms. further reference to t. sinuosa was made by bertrand & kilian (1889) for spanish individuals coming from near granada. sacco (1902) cited t. sinuosa in a study on tertiary brachiopods from piedmont and liguria (northern italy). in 1933 boni proposed the name terebratula maugerii for a miocene specimen of monte vallassa. studying a large number of well-preserved specimens from the same locality, boni (1934) recognized the similarities between t. maugerii and t. fig. 1 page 22 of purpura (colonna 1616). the small numbers close to the specimens in the upper part of the figure are respectively number 4 and 1, from the left to the right. the specimen on the upper left (number 4) probably came from the miocene of the pietra leccese formation (apulia, southern italy). the specimen on the upper right came from the pliocene calcarenites of andria (southern italy). it represents a topotype of terebratula terebratula. on the validity of terebratula sinuosa (brocchi) 633 sinuosa, but, despite this, he divided the latter into a number of subspecies and ‘varieties’. in 1966 sirna reviewed the stratigraphic distribution and systematics of t. sinuosa. he further collected this species in serravallian (miocene) deposits from scanno and monte maiella (abruzzo, central italy). these levels could be dated up to the messinian according to cornacchia et al. (2017). sirna considered t. sinuosa pedemontana, boni’s (1934) subspecies, and t. costae, described by nelli (1910) in reference to specimens found close to l’aquila (central italy), as synonyms of t. sinuosa. marasti (1973) described a number of beautifully preserved specimens from the miocene deposits of the stirone river (emilia, northern italy) as t. sinuosa. cooper (1983) described t. ampulla from pliocene sediments at monte mario (roma, central italy), illustrating individuals morphologically similar to t. terebratula, and t. sinuosa from malta. in the same paper he established the new genus maltaia which externally resembles terebratula but yields a different brachidium. cooper further erected the new species maltaia maltensis. he pointed out that “the loop of maltaia is similar to that of terebratula in its great width, lack of inner hinge plates, and well-defined terminal points. the loop of terebratula, however, has the transverse band flattened medianly and not extended ventrally to form a protuberant tongue” (cooper 1983, p. 232). gaetani & saccà (1983), studying tertiary deposits from calabria and sicily (southern italy), described t. sinuosa from cessaniti (calabria) and considered t. pedemontana, t. costae, m. maltensis and all of boni’s (1934) t. sinuosa subspecies to be synonyms. they further accepted as valid t. calabra (seguenza, 1871), for a number of specimens described from the upper pliocene of santa tecla (sicily) and terreti-monte gonì (calabria). they considered t. biplicata described by costa (1851), t. sinuosa by seguenza (1865, 1870, 1871) and taddeiruggiero (1983), and t. terebratula by pajaud (1976) as synonyms of t. calabra. taddei ruggiero (1994) identified t. sinuosa, t. calabra, t. siracusana and t. scillae in neogene sediments of salento (apulia, southern italy). lee & brunton (1998) selected a neotype for terebratula terebratula from the “calcareniti di gravina” formation near andria (apulia), since the specimen originally figured by colonna (1616) was lost. lee et al. (2001) discussed the considerable taxonomic confusion surrounding the taxon t. terebratula. they considered t. sinuosa as a synonym of t. terebratula. they further put t. calabra and t. costae into the synonymy of t. terebratula, and therefore accepted that the latter species extends back to the miocene. borghi (2001) described neogene brachiopods from emilia (northern italy), reporting that both t. sinuosa and t. ampulla were present. in 2006, garcía-ramos provided an extensive revision of european tertiary brachiopods. he recognized t. sinuosa and t. maugerii as conspecifics and took the latter specific name as valid. he further acknowledged the validity of t. calabra and of the genus maltaia, including the species m. maltensis, m. costae and the new species m. pajaudi. garcía-ramos considered the specimens referred to as t. sinuosa by sirna (1966) as maltaia costae. he further listed a number of iberian fossil sites where t. sinuosa is actually present. according to garcíaramos (2006) further possible conspecific (or at least closely related) forms are terebratula carryensis from the lower miocene of france and terebratula hoernesi found in the miocene of hungary. bertolaso et al. (2009) figured a specimen of t. ampulla from the pliocene of castell’arquato (piacenza, emilia) and two tortonian specimens of t. maugerii, which are remarkably similar to each other in spite of their different origins. one of the specimens of t. maugerii comes from scipione ponte (parma, emilia, northern italy) and the other from los brianes, corvera (murcia, spain). the validity of terebratula sinuosa keys to determining whether t. sinuosa deserves the status of a full species include the stratigraphic distribution and the peculiar morphology. typical t. sinuosa shells are not just sulciplicate. the most distinctive external character is a long, massive and well-defined fold running longitudinally along the ventral valve from the umbonal zone (fig. 2). specimens with these characters are known in, at least, miocene deposits of italy, spain, and possibly malta (see garcía-ramos 2006 for an extensive review). many researchers have used the specific name terebratula sinuosa in reference to such heavily sulciplicate terebratula brachiopods. we have taddei ruggiero e., serio c. & raia p.634 statistically tested the morphological uniqueness of t. sinuosa in reference to other species belonging to this genus. we performed a three-dimensional geometric morphometric analysis of 142 undeformed brachiopod specimens belonging to four nominal species (t. ampulla, t. sinuosa, t. terebratula, and t. scillae; taddei ruggiero et al. 2008a). the individuals tentatively referred to t. ampulla in taddei ruggiero et al. (2008a) come from valle botto (asti, northern italy), but they could actually belong to a different species. nevertheless, we found that, from a morphological point of view, t. sinuosa is the most derivate terebratula species (see fig. 5 in taddei ruggiero et al. 2008a). all the specimens analysed by taddei ruggiero et al. (2008a) were adult, meaning that the morphological distance between t. sinuosa and other terebratula species could depend on a late ontogenetic variation within an otherwise highly variable terebratula lineage. to deal with this issue, we studied the developmental basis (ontogeny and ontogenetic allometry) of the same four species, still using three-dimensional geometric morphometrics. we demonstrated that the typical, heavily sulciplicate anterior commissure in t. sinuosa is already present at juvenile stages and did not depend on size differences between the morphotypes (taddei ruggiero et al. 2008b, fig. 2). this indicates that the difference between t. sinuosa and other terebratula is profound, justifying its recognition as a distinct species. in addition, t. sinuosa specimens are miocene in age, whereas larger terebratula, and most importantly its presumed synonym t. terebratula, are pliocene to pleistocene in age. hence, the ontogeny, morphology and stratigraphy all point to the same evidence, that is t. sinuosa is a different species, which deserves its own taxonomic species status. other authors have advanced similar evidence in support of our conclusions, among the others gaetani & saccà (1983), garcìa-ramos (2006) and bertolaso et al. (2009). it should be noted that sulciplicate specimens, not as strongly folded as t. sinuosa, are known in the miocene and in the pliocene as well. fig. 2 terebratula sinuosa. row a) juvenile specimen from cessaniti (calabria, southern italy). rows b-c-d) specimens coming from the stirone river section (b juvenile; c-d adults, specimen d is housed in the collections of the castell’arquato geological museum). from the left: dorsal view (first column), ventral view (second column); side view (third column), frontal view (fourth column). on the validity of terebratula sinuosa (brocchi) 635 such individuals were recognized as t. sinuosa in seguenza (1865, 1870, 1871) and taddei ruggiero (1983), and as t. calabra in gaetani & saccà (1983) and garcía-ramos (2006). the presence of such a sulciplicate anterior commissure was considered part of the morphological variability of t. terebratula by lee et al. (2001). however, it is important to consider that these forms lack the long and prominent fold on the ventral valve which is present in t. sinuosa. we deem it is probably incorrect to keep considering these specimens as synonyms of t. terebratula, yet they clearly do not belong to t. sinuosa and therefore deserve further study. the same holds true for terebratula costae. the extensive collections for these forms including the specimens studied by costa (held at the museum of paleontology in naples), the collection of seguenza (held at the “gemmellaro” museum in palermo), and the possibly conspecific sample from águilas (murcia, spain, garcía-ramos 2006) deserve further analysis. several authors have used the taxonomic name t. maugerii as a junior but valid synonym of t. sinuosa. however, the species epithet maugerii was erected on a single deformed specimen by boni (1933) and is only valid under the hypothesis that t. sinuosa is a synonym of t. terebratula (garcìa-ramos 2006), which we proved to be incorrect. in addition, given the obvious reference of the ‘sinuosa’ morphotypes to the heavily sulciplicate specimen of colonna (e.g. sacco 1902; marasti 1973; gaetani & saccà 1983), we propose to protect the taxon t. sinuosa as a ‘conserved name’, in keeping with the rules of the international code of zoological nomenclature (iczn, art. 23). therefore, t. maugerii falls within the valid name t. sinuosa. the provenance of the specimen n. 4 figured by colonna (1616) is unknown, although in his description colonna refers to it as “replete erat concretione candida terrea” (filled with pale sediment, most probably a calcarenite) which is consistent with the pietra leccese formation. we found t. sinuosa specimens from silts of stirone river (emilia, northern italy, fig. 3), from sandstone of cessaniti (calabria, southern italy), and from calcarenites within the pietra leccese formation (salento, apulia, southern italy). in erecting anomia sinuosa brocchi explicitly referred to colonna’s sulciplicate specimen and further indicated that fossil deposits near piacenza (emilia, northern italy) yield the species. it is thus plausible that colonna’s specimen belongs to the pietra leccese calcarenites. hence, we deem that the neotype and type locality for t. sinuosa must be found and eventually defined (icnz, art. 76.3). the miocene outcrops in emilia indicated by brocchi could be used for a suitable neotype investigated for the same aim (icnz, art. 76.2). fig. 3 the specimen illustrated in figure n.4 by colonna (1616) together with a typical miocene t. sinuosa specimen coming from the stirone river section. taddei ruggiero e., serio c. & raia p.636 conclusions we remark that miocene, sulciplicate terebratula having a massive, well-defined and distinctive fold running longitudinally along the ventral valve from the umbonal zone should be attributed to terebratula sinuosa which we therefore consider a valid taxon of full species rank. this species occurs in miocene deposits of italy, spain, and possibly malta. a neotype and the relative type locality should be selected for the taxon terebratula sinuosa. terebratula maugerii should be considered a junior synonym of t. sinuosa. the definition of t. sinuosa as a valid species requires the amendment of the stratigraphic distribution of t. terebratula, which is consequently limited to pliocene-pleistocene deposits. acknowledgements: we are grateful to carlo francou, head of the castell’arquato geological museum for the beautifully preserved t. sinuosa specimens he provided for this study. the staff of the “gruppo paleontologico tropeano” took one of us (ert) to cessaniti, tropea and several other sites to look at the fossil material of terebratula sinuosa in situ. antonella taddei kindly took the pictures figured here. daphne lee and diego garcía-ramos kindly provided us unvaluable comments on an earlier version of the manuscript. references bertolaso l., borghi e. & garcía-ramos d. (2009) brachiopodi neogenici e pleistocenici dell ‘emilia (parte ii). parva naturalia, 8: 3-42. bertrand m. & kilian w. (1889) études sur les terrains secondaires et tertiaires dans les provinces de grenade et de malaga. missión d’andalousie memoires academique sciences paris, 30: 378-582. boni a. (1933) fossili miocenici del monte vallassa. boll. soc. geol. it., 52: 73-156. boni a. (1934) studi statistici sulle popolazioni fossili: chlamys scabrella lam. e terebratula sinuosa brocchi. riv. it. paleontol. strat., 40: 1-275. brocchi g.b. (1814) conchiologia fossile subappennina, 2 vols. stamperia reale, milano, 712 pp. borghi e. (2001) osservazioni sui brachiopodi neogenici e pleistocenici dell’emilia. parva naturalia, 45-81 colonna f. (1616) purpura. hoc est de purpura ab animali testaceo fusa, de hoc ipso animali, aliisquibus rarioribus testaceis quibusdam. j. mascardo. roma, 42 pp. cooper g.a. (1983) the terebratulacea (brachiopoda), triassic-recent: a study of the brachidia (loops). smithsonian contributions to paleobiology, washington, 290 pp. cornacchia i., andersson p., agostini s., brandano m. & di bella l. (2017) strontium stratigraphy of the upper miocene lithothamnion limestone in the majella mountain, central italy, and its palaeoenvironmental implications. lethaia, 50: 561-575. costa o.g. (1851) fauna del regno di napoli animali molli classe v brachiopodi. napoli 60 pp. davidson t. (1864) description of the brachiopoda of the maltese islands. in: adams l. (ed.) outline of the geology of the maltese islands 2: 7. annals and magazine of natural history, london. davidson t. (1870) on italian tertiary brachiopoda iii. geological magazine, 7: 359-370. gaetani m. & saccà d. (1983) brachiopodi neogenici e pleistocenici della provincia di messina e della calabria meridionale. geologica romana, 22: 1-43. garcía-ramos d. (2006) nota sobre terebratulinae del terciario de europa y su relación con los representantes neógenos del sureste español. bol. as. cultural paleontol. murciana, 5: 23-83. lee d.e. & brunton c.h.c. (1998) terebratula müller, 1776 (brachiopoda): proposed designation of anomia terebratula linnaeus, 1758 as the type species. bull. zool. nomen., 55: 220-223. lee d.e., brunton c.h.c., ruggiero e.t., caldara m. & simone o. (2001) the cenozoic brachiopod terebratula: its type species, neotype, and other included species. bull. nat. hist. mus. geol. ser., 57: 83-93. linnaeus c. (1758) systema naturae, sive regna tria naturae systematicae proposita per classes, ordines, genera et species, 10th edition, tom 1 regnum animale. holmiae, stockholm, 824 pp. marasti r. (1973) la fauna tortoniana del t. stirone: limite parmense-piacentino. boll. malacol. it., 12: 76-120. nelli b. (1910) fossili miocenici del modenese. boll. soc. geol. it., 28, 489-523. pajaud d. (1976) les brachiopodes du pliocène i de la sierra de santa pola (sud d’alicante, espagne): terebratula terebratula (linné, 1758) et phapsirhynchia sanctapaulensis nov. gen., nov. sp. ann. soc. géol. nord, 96: 99-106. sacco f. (1902) i brachiopodi dei terreni terziarii del piemonte e della liguria. carlo clausen, torino, 50 pp. scilla a. (1670) la vana speculazione disingannata dal senso. a. colicchia, napoli, 168 pp. seguenza g. (1865) breve cenno di ricerche geognostiche ed organigrafiche intorno ai brachiopodi terziari delle rocce messinesi. annali dell’accademia degli aspiranti naturalisti napoli, 5: 24. seguenza g. (1870) dei brachiopodi viventi e terziarii di o. g. costa. boll. malacol. it., 3: 145-160. seguenza g. (1871) studi paleontologici sui brachiopodi terziarii dell’italia meridionale. boll. malacol. it., 4: 1-79. sirna g. (1966) brachiopodi miocenici dei dintorni di scanno e della maiella. boll. soc. paleontol. it., 2: 184-196. taddei ruggiero e. (1983) strutura del guscio dei generi gryphus e terebratula (terebratulidae, brachiopoda). societa dei naturalisti in napoli, bollettino, 90: 177-201. taddei ruggiero e. (1994) neogene salento brachiopod palaeocommunities. boll. soc. paleontol. it., 33: 197-213. taddei ruggiero e., raia p. & buono g. (2008a) geometric, morphometrics species discrimination within the genus terebratula from the late cenozoic of italy. fossils and strata, 54: 209-217. taddei ruggiero e. & raia p. (2008b) ontogeny in terebratula species. functional convergence and allometry but not heterochrony. proc. royal soc. victoria, 120: 320-331. rivista italiana di paleontologia e stratigrafia volume 104 numero 2 a:oìne )9.7-)9l agosto 1998 noa brevry remarks on the stratigraphy and biochronology of the late pleistocene deposit of ingarano (apulia, southern italy) carmelo petronio* e. raffaele sardella*,i receiaed decenrber 10, 1997; accepted march 19, 1998 key-oorcls: biochronologn stratigraphy, late pleistocene, vertebrates, southern italy. riassunto. ulteriori considerazioni sulla stratigrafia del deposito di ingarano (foggia) consentono di rpotrzzare la presenza di tre associazioni faunistiche riferibili ad un intervallo ben definito del pleistocene superiore. la associazione pirì antica (ingarano a) può essere riferita allo stadio 4 delle paleotemperature, quelle più recenti rispettivamente allo stadio 3 (ingarano b) e allo stadio 2 (ingarano c). i dati faunistici si accordano con i'alternanza delle diverse condizioni climatiche nel corso del pleistocene superiore. abstract, new field data on the late pleistocene deposit of ingarano (foggia) allow us to hypothesise the occurrence of three faunal assemblages (ingarano a, b, c) respectively referable to isotopic stages 4, 3 and 2 of the palaeotemperature scale. the palaeontological data match with the alternating palaeoclimatical conditions during late pleistocene times. introduction. the late pleistocene karst cave deposit of ingarano is exposed at 270 m a.s.l. along the railway of gargano) near apricena (foggia, southern ltaiy). the first studies started in the second half of '8oties, but only sínce 1992 the complete stratigraphical succession has been exposed by quarry works. these activities and the previous quarry ril/orks partially disturbed or hid the stratigraphy of the cave succession, which appears chaotic. from this succession a rich vertebrate fauna and some lithic tools have been recorded and described (capasso barbato er. ai., 1992; petronio et ai., 1996). the study o[ the fossils allow us to point out rwo possible biochronological hypothesis: a single or some distinct faunal assemblages. in order to test these two hypothesis and to point out a more detailed stratigraphy of the cave succession new geological field surveys have been carried out in the area. the vertebrate fauna has been considered following the updated biochronological framework for the late aurelian mammal age (glíozzr et ai., 1997). these new researches allow us to point out a more detailed stratigraphy and biochronology for the locality of ingarano, with the definition of three distinct faunal assemblages and the general lines of the paleoecological evolution of the area. stratigraphical framework. two fossiliferous deposits can be identified (fig. 1): 1) a sandy-clays deposit (b), includingvery big calcareous blocks, which is exposed in external position to the cave deposit;2) a successiofl oí 12 m, which rcstifies fig. 1 stratigraphical sketch of the late pleistocene fossiliferous deposit of ingarano (foggia, southern italy): a) calcareous bedrock (calcari di sannicandro fm.); b) silty-clays with calcareous blocks; c) alabastrine and encrusted phosphatic layer; d) conglomerate with round and flattened calcareous pebbles of smrll size (less than one cm) with silty reddish matrix; e) conglomerate with calcarenitic large sized pebbles (1 m length), cemented by a calcareous matrix; f) conglomerate with calcareous pebbles scarcely rounded (sized more than 10 crn), cemented, with a prevalently calcareous -"r,;*. ol i."^h".".î ";lr."..1;-"., sapienza" ptazza aldo moro 7 , roma, italy. e-mail: petronio@axrma.uniroma.it università di roma "la sapienza". 'f dip. scienze della terra, università di roma "la 'r'r borsa post-dottorato dio. scienze della terra. 288 c. petronio & r. sardella different phases of the filling of an ancient cave, today destroyed by the quarry works. the sandy-clays sediments (b) are in unclear stratigraphical relationship with the cave succession and contain remains of several fossil mammals: stephanorhinus hernitoechus, coelodonta antiquíatis, hippopotamus ampbibius, elepbas a.ntiquus, pantbera spelaea, vulpes oulpes, apodemus syhtaticus. the calcare di sannicandro (a) constitutes the bed rock of the cave, in which an intense karst activity formed stalagmites and a thick alabastrine layer. these layers are encrusted with phosphatic material (c). a geof;o 2 l.'..,'^ /f^..;' s^"rl.'.^ ,b. " italy). layer b: 1) stephanorhinus bemitoecbus: fragmentary maxillary bones with teeth (occlusal view); 2 and 3) coelodonta antiquitatls: [ngmentary humerus (posterior and anterior view). all the figures ''" ...'^"i-"r.ì., a(or^ .'r"ral size. .l^-: .-r .-r...: . with thellìcrlrrldr 4rr4iyjrù rrrth/2r4u merhod on differenr samples of the speleothem gave an age of 40.000 +2.000 for the phospatic concretion due to the guano of the birds which lived in the area, while the speleothem stopped its growing àt about 58.000 +2.000 (petronio et al. 19961 from lrwer r come the vertebrate'bones represenîing several taxa: aves aquila cbrysaetos, columba livia, pyrrhocorax graculus; mammalia cervus elaphus, dama dama dama, lynx lynx, canis lupus, canis ex gr. arnensis-mosbachensis, vulpes oulpes, martes sp., mustela ni,ualis. above layer c a conglomerate with round and flattened calcareous pebbles of small size (less than one cm), with silty reddish matrix, poorly cemented is exposed (d). this conglomerate gradually evolves upv/ards into another conglomerate with calcareous scarcely rounded pebbles o[ larger size (more than 10 cm), more cemented, with a prevalently calcareous matrix (e). a grear number of fossil bones (represented often by some isolated remains) comes from these two layers: equus ì.rydruntinus, bos primigeniws, cerous elapbus, capreolus capreolus, rupicapra sp., felk silaestris, lynx lynx, panthera pardus, crocuta crocuta, canis lupus, canis ex gr. arnensis-mosbachensis, vulpes oulpes, ursus arctos. at the top of the succession, with a thickness of almost 3 m, a .o.rg1o-..rte with calcarenitic large sized boulders (1 m length), cemented by a calcareous matrix (f), is exposed, testifying the phase of closing of the cave, when the vauit fell down. the passage between the layers e and f is characterised by a different inclination of the deposits; twelve lithic implements of levaliois technique comes from the stratigraplty and biochronology of tbe late pleistocene of ingarano 289 ingarano (foggia, southern kaly). lryer c: l) lynx lynx skull (at the top) (palatal view); emimandible (at the bottom) (labial view). al1 the figures are approximately 70o/o natural size. top of layer e and their state of preservation allow to hypothesise a very low degree of transportation (petronio er ai., 1.996). some smail galleries, probably connected together, exist into the levels d and e. they are partially filled with a silty incoherent sediment (g), with severai fossil bones of micromammals and birds (petronio et al., 1996). some bones of mammals and birds are partially covered by phosphatic encrustation and for this reason are clearly coming from layer c. the taxa surely coming from iayer g are: aves aquila chrysaetos, falco peregrinws, falco tinnunculus, alectoris graeca, circus nov. sp., perdix perdix, columba lioia, nyctea scandiaca, pyrrbocorax graculus, pynhocorax sp., corvus corax, corvus rnonedula; mammalia erina' ceus europaeus, myothis blythi, oryctolagus cuniculus, kpus europdeus, arvicolidae indet., microtus (fenicola) sp., microtus ex gr. aroalis-dgrestis, apodemus syhtaticus, eliorrrys quercinus. fauna. layer b. the fauna assemblage coming from the sandyclays sediments is characterised by the presence of the "warm" species stepbanorbinws hemitoechus and the "cold" coelodon ta an l iqu ita lis. stephanorhinus hemitoecbzs is represented by skull fragments with a complete series of teeth fig. 2.1-, a complete tibia and other parts of limb bones, probably belonging to the same juvenile specimen. even if it is a juvenile specimen some dental features allow a quite sure taxonomical determination. premolars and molars show the peculiar undulate profile of the ectoloph and the tendency to develop quite ipsodont teeth. these characters are typícally ol stephanorhinus bemitoechus and are shared with the "etruscoid" forms. coelodonta dntiquitatis is represented by a fragmentary upper tooth (capasso barbato et al. t992), a paftial humerus (fig 2.2, 2.3), carpal bones and some metacarpal in anatomical connection, radius and tibia). the fauna is completed by hippopotamus amphibius (a fragmentary molar and a proximal epiphysis of a femur); elephas antiquus (a fragment of a molar), panthera spelaea (part of a very large sized tibia), vulpes oulpes (some limb bones) and apodernws syloaticus (a complete emimandible). cave succession. layer c. bird remains are represented by several limb bones, with pyrrhocorax graculus the most common species. among mammals, the boreal lynx is the best represented taxon with skull fragments, mandibles and limb bones referable to almost five specimen (fig 3). a study still in progress is evidencing some peculiarities of these lynxes. the skulls are comparable in size to the actual scandinavian qnx lynx, but show proportionally less developed teeth, intermediate in size between the northern lynxes and the pieistocene samples of lynx pardina spelaea (\ferdelin 198 1). canids and mustelids are quite common in this layer. the occurrence of canis lupus ts testified by several limb bones, while also a small wolflike dog occurs. the presence of a middle sized canid in late pleistocene deposits from southern italy have been considered as a persistence of a taxon of galerian origin (di stefano et al., 1992; capasso barbato &. glíozzí, 1996). the systematic position of this dog is still unclear. the mediterranean galerian deposits are characterised by the occurrence of canís aff. arnensis (rook & torre 1996), while the small wolf canis mosbachensis is a peculiar element 294 c. petronio g r. sardella fìg.4 ingarrno (foggia, southern italy). layer c: vulpes vulpes 1) emimandible (labial tiew); 2) fragmentary skull (palatal view); 3) skull (sagittal vrew); canis lupus 4) emimandible (labìal view); 5) skull (palatal view). all the figures are approximately 607o natural size. of the coeval european localities. until new studies will clarify the relationships between these middle sized canids and their dispersal during pleistocene times, we refer the fossils to canis ex gr. "arnensis-mosbachensis". vulpes vulpes (fig. 4.1, 4.2, 4.3) is a very common element and the skulls, the mandibles and the limb bones testifies the occurrence of at least six individuals. martes sp. is represented only by an os penis, whtle mustela niaalis occurrence is testified by four almost complete skulls. among cervids an adult specimen of ceraus elaphus have been recorded (mandible, vertebrae and limb bones of), while dama dama dama is represented only by few fragmentary remains (fig. 4.4, a.5). the modern subspecies of the fallow deer is an important biochronological marker. the modern fallow deer appears in itaiy at the beginning of the aurelian mammal age in correspondence of the isotopic sfage 7 with the subspecies dama dama tiberina (di stefano 8c petronio 1997). this cervid evoives in dama dama dama in isotopic stage 5 and it characterises the late pleistocene cieposits. despite the fragmentary remains the evolved features in premolars and molars allow to refer the fossils recorded to the modern fallow deer. layers d and e: these layers are characîerised by the abundance of the carnivores remains (in particular almost complete skulls or cranial fragments). ursus arctos is represented by a complere skull with mandible fig. 5 -, partially crushed, atlas and epistropheus; among fehds lynx lynx is the most common taxon (several skulls, mandibles and limb bones), whiie the occurrence of panthera pardus and felis sih,estris is recorded only by fragmentary remains. canis lupus and vulpes vw/pes are quite common (at least three specimens for every taxon), while seems there is no evidence of the presence of the middle sized canid; crocuta crocuta, usually very common in cave deposits, is really poorly represented (a fragmentary cubitus and a metapodial). herbivores remarns are less frequent; equus lrydruntinus is represented by a complete mandible (capasso barbato et ai., 1992) and some jugal isolated teeth, while the occurrence of cerous elaphws, bos primigenius, capreolus capreolus, rupicapra sp. is testified only by fragments of mandibles and isolated jugal teeth. layer g: among the birds, to which belong a very high percentage of the fossil bones of this layer, the most paleoecologically significant taxon is nyctea scandiaca. the avifauna is characterised most of al1 by the abundance of ràptors and corvids. several micromammals come from this layer. the best represented species rs microtus ex gr. agrestis, while apodemus sylaaticus, eliomys quercinus and \èrricola sp. are quite rare. probably micromammals may be considered as prey of the raptors which inhabit the area. discussion. in the first study on this deposit (capasso barbato et al., 1992), the presence of two distinct faunal assembiages, respectively referred to the stage 4 of the isotopic scale (the fauna with "pachyderms') and to the stage 2 stratigraplry and biochronolog of the late pleistocene of ingarano (in particular the avifauna, with the "cold" taxon nyctea scandiaca), have been hypothesised" in a further work, (petronio et a1., 1.996) the hypothesis of a single faunal assemblage, referred to the isotopic stage 3, was considered as more probable for the coexistence of "cold" and "warm" taxa. new field data allow us to test these different hypothesis showing a more articulated stratigraphical framework outlined in this paper. at the present time, there is no evidence of direct stratigraphical relationships with the cave succession for deposit b, with the "pachyderm" remains. no fragments of "pachyderms" come from the other layers. the layer b may probably be older than the cave succession. the coexistence of two species of rhinos, ecologically distinct, of the hippo and of the forest elephant may be explained, if we consider the faunal assemblage coming from the layer b as homogeneous, referring it to the isotopic stage 4 (ingarano a, fig. 6). in particular the chronology of this fauna is defined by the occurrence of coelodonta antiquitarls, which was widespread in italy in late pleistocene since the isotopic stage 4, while the other iarge mammals are frequent in the isotopic stage 5 and survive until the isotopic stage 3 (late aurelian mammal age, gliozzi et a1., 1997). however the coexistence oí coelodonta and ste' phanorhinus, even if rare, has been recorded in some european localities as achenheim (alsace, france) and balauzière (france) (in guerin, 1980, with bibliography). the occurrence of the woollv rhino, in association with i-ig.5 irrgrrrno (foggir, southern italy). layer d: ursus arctos skuil and mandible (approxìrnately 45% naturel size). r ho ^r ho, ^,.l-,,1.,*-.l;^1"'-rlc ulrllr pdllr)uqr lll)r wllllll show.r wider ecological srgnific:ìnce. may restify rhe begrnnrng of the climaric detenoration referred to the isotopic stage 4. considering the cave succession, the geochemical analysis wirh the 2lath/234ij method of rhe phosphatic encrustation g,rve :rn.rhsolr:re,rpe of 40.000 +2.ooo (petronio et a1., 1996). in consequence this is the age of the fossil bones coming from level c. for this reason it is possible to refer the faunal assemblage to the isotopic stage 3 (see gliozzi et al., 1992). also rhe vertebrate fauna coming from the layers d and e can be referred to the isotonic sr.rpe 3. hrrr ir resti[ies the progressive change toward more temperate-cold climatic conditions (reiative abundance of cervus elaphus, occurrence of equus hydruntìnus and rupicapra sp.)the medium sized dog, even if rare, coexists with the wolf in layer c, while doesn't occur in layers d and e, where there is the predominance of cants lupus. a similar tendency seems to be present in the grotta romanelli succession with the canis ex gr. "arnensis-mosbachensis" more frequent than wolf in "terre rosse" and very rare in the younger "terre brune" with the predominance of canis lupus qagliacozzo, personal comm.). as a matter of fact, the maintenance of delicate anatomical structures as nasal coanes (vulpu aulpes, canis lupus, felis silaestris from layer d and e) and encepha1ic casts (lynx lynx from layer c), besides some examples of fossil bones in anatomical connection (ursus arctos from layer d), suggest a 1ow degree of transportation for the fossils and a quite high degree of the sedimentation ratio. also the study of the lithic implements coming from the top of leve1 e match with both the deposition setting than with the chronological considerations (the levailois technique is not used after approximately 3537.000 b. p.) (petronio et a1., 1996). the fossils coming from layers c, d and e mat be included in the faunal assemblage ingarano b (fig 0)" the fauna coming from g (ingarano c) is younger than the other faunal assemblages, probably referable to the isotopic srage 2, with cold climatic conditions testified by the predominance of the snow-owl. lhronostratigraphy absolute ^ge nfagnetostratigraphy mammal aqes isothopic scale sclcctcd localities from southcrn ltaly holocene 0.05 0,1 0.1 5 0,2 i c o lr l (t) l'r ì o rlt ; :z f-l t\/. frl f ,l z j l'rl crotta romanelli ("terre brune") grotta b di spagnoli (upper levels) grotta di cardamone i n g a r a t o ingarano ingarano grotta del cavallo, grotta del sarcofago grotta b di spagnoli (lower levels), grotta romanelli (k-g), grotte delle striare, melpignano, s. sidero, grotta uluzzo c rtì z frl u f (t) rrì j rrì f j f r'ì .t trì j grotta lina cerveteri, sedia del diavolo, torre in pietra(upper levels), vitinia 292 fio a palaeoecological notes. from the fossiliferous deposit of ingarano have been identified three faunal assembiages (a, b, c) referable to the late aurelian mammal age. the study of the assemblages allow us to stress some considerations about the palaeoenvironmental evolution of the area. the fauna wíth coelodonta aîd stepbanorhinus (ingarano a) is probabiy to refer to the beginning of the climatic deterioration occurred in isotopic stage 4. the presence of these two genera of rhinos testifies the existence of plains and open areas. the fauna ingarano b is characterised by the occurrence of the modern subspecies of the fallow deer in layer c, which may be referred to warm-temperate climatic conditions. the fauna of the layers d and e testify instead the changing of the climate (occurrence of equus lrydrwntinus and rwpicapra sp.), with the development of more temperate-cold conditions and the reduction of the forested areas. the coldest phase, related to the last pleniglacial (isotopic stage 2), is testified by the fauna ingarano c, characterised by the abundance of the snow-owl nyctea scandiaca, which today is widespread in northern latitudes and is specialised in preying lemmings. c. petronio & r. sardella revised chronological framework of ingarano faunal assemblages in comparison with some late middle and late pleistocene selected iocalities from southern italy (from petronio et a1.7996, modified). conclusions. the analysis of faunal assemblages ingarano a, b and c allow to implement the knowledge on the late aurelian mammal faunas (gliozzi et al., 1997). the late aurelian, is referable to a time span included between the eemian and the end of the last glaciation. no faunal units have been defined because the late aurelian faunas are characterised by the disappearance of large and medium mammals. in the corresponding time span some climatic events occur. these events, in particular in the adriatic coast, have a great influence in the faunal composition and in the causes of their disappearances. flowever, another important element to consider is the rule of homo neandertalensls and homo sapiens. fínally, the great amount of data referable to late aurelian faunas if on the one hand gives the possibility to define a paleoecological framework of the microclimatic and environmentai conditions for the different areas of the peninsula, on the other hand do not allow to choose a single fauna to define a faunal unit. for this reason the ingarano faunal assemblages testify the disappearance of the large "pachyderms" in an earlier moment than 35.000 b.p. (ingarano a), a quite progressive climatic deterioration (ingarano b) corresponding to the isotopic stage 3, toward the cold phase of the isotopic stage 2 (recent wurm) with the disappearance of several taxa of mammals and the abundance of cold taxa as nyctea scandíaca. 293 acknotaledgernmts. 'sfe wish to thank prof . m. gaetani (lvlilano) rnd t-o anonymous reviewers for the useful suggestions, l. codebò for the field observations, g. di stefano, p ferraro and e. squazzini for suggestions and technical support. photographs by l. spinozzi e g. d'arpino. this work was supported by murst 40% grants. stratigraplry and biochronology of the late pleistocene of ingarano references capasso barbato l., cassoli p.f., minieri m.r., petronio c., sardella r. & scarano m. (1992) note preliminari sulla fauna pleistocenica di ingarano (apricena, foggia). boll. soc. paleont. 1t.,v.31, n. 3, pp" 325-334, modena. capasso barbato l. & gliozzi e. (1,996) biostratigraphical and palaeogeographical implications of the late middle pleistocene well balanced fauna from quisisana-certosa (capri, southern italy). boll. soc. paleont. it., v. 34, n. 2, pp. 235-261., modena. di stefano g., petronio c., sardella r., savelloni v. & squazziti e. (1992) nuove segnalazioni di brecce ossifere nella costa fra castro marina e otranto (lecce). il qu* ternario, v. 5, n. 1, pp. 3-10, roma. di stefano g. & petronio c. (1997) origin and evolution of the european fallow deer (dama, pleistocene). n. /&. geol. palaont. abh. v.203, n. 1, pp. 57-75, stuttgart. g|iozzie., abbazzi l., argenti p., azzarolí a., caloi l., capasso barbato l., di stefano g., ficcarelli g., kotsakis t., masini f., mazzap., mezzabotta c., palombo m.r., petronio c., rook l., sala b., sardella r., zanalda e. & torre d. (1997) biochronology of selected mammals, molluscs and ostracods from the middle pliocene to the late pleistocene in ltaly. the state of the art. rlz. it. pal strat., v. 103, n. 3, pp" 369-388, milano. guerin c. (1980) les rhinocéros (mammalia, perissodactyia) du miocéne terminal au pleistocène superiéur en europe occidentale comparaison avec les espèces actuelles. docum. lab. géol. lyon, v.79, n. 3, pp. 785-1185, lyon. petronio c., billia e., capasso barbato l., di stefano g., mussi m., parry s.j., sardella r. & voltaggio m. (1996) the late pleistocene far.rna of ingarano (gargano, italy): biochronological, paleoecological, paleoethnological end geochronological implications. boll. soc. paleont. it., v. 34, n. 3 (1995), pp. 333-339, modena. rook l. & torre d. (1996) the latest villafranchian-early galerian small wolves in the mediterranean area. acta zool. cracoo., v. 39, pp. 1-7, cracovia. verdelin l. 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*����& �$����)���%%� ����� �!� (��% �% �% ��% )*� �����+� ���, + �� �$� �������� �+ �$� -����� -�������� ?> rivista italiana di paleontologìa e stratigrafia volume ruj numero 2 tavole 1 pagine 183-192 settembre 1992 ofrheupperhlfsijè3'.8îr'''f, è.""lfi |+"?uy"s-"s%t;'.navalle' (udine, friuli-venezia giulia, ne italy) \tith reptile track\tays guido roghi* & fabio m. dalla vecchia'i key-uords: biostratigraphy, upper triassic, carnian, tuvalian, *ln*tt, southern alps, palynology, vertebrate tracks, palaeoenviriassunto. vengono riponati nuovi dati e considerazioni sulla biostratigrafia e il paleoambiente di una sezione stratigrafica costituita da alternanze dolomitico-marnose del triassico superiore affioranti nella val dogna (udine, friuli, ne italy). in panicolare è stato investigato un intervallo con uno strato contenente piste di rettili arcosauromorfi terrestri. nel livello sopràstànte, quello che diretramente ricopre le impronte e' stata rinvenuta una abbondante associazione palinologica tuvalica (carnico superiore) con enzonalasporites aigens; vallasporites ignacii, patinasporites d,ensus, zonalasporites cinctus, pseu. doenzonalasp orites summn s, samaropo/lenites speciosus, camerosporites secdtus; partitisporites spp. evidenze floristiche e sedimentologiche indicano un clima arido e un ambiente di deposizione costiero soggerro . ri-ot"to "-...;^-; abstrdct. new data and considerations about the biostratigraphy and the palaeoenvironment of a section in the late triassic dolomitic-marly sequence which crops out in the dogna valley (jdine, friuli, ne italy) are reported. in panicular a unit wìth a surface bearing tracks of archosauromorph terrestrial reptiles has been investigated. in the layer immediately overlaying the track-beanng one, a rich palynological assemblage with enzonalasp orites vigens; vallasp orí tes ignacii, patinasporites densus, zonalasporites cinctus, pseudomzona" lasporites surnnlus, satnaropollenites speciosus, camerosporites secatus and partitisporiter spp. was found, indicating a tuvalian age (late carnian). microfloral and sedimentological evidence indicate a dry climate and a coastal depositional environment subject to repeated emersions. lntroduction. the canale di dogna (dogna channel), usually reported as "val dogna" (dogna valley), is an e-w oriented incision in the julian alps and is run through by the dogna creek, near the village of dogna, udine province (fig. 1) ladinian dolostones (dolomia de1lo sciliar), some poorly known units of carnian age, and the dolomia principale (hauptdolomlt of german authors) outcrop in the dogna valley (selli, 1963; jadoú et al., 1995). the discovery of reptile tracks in the dolomitic-marly sequence cropping out in the lower dogna valley (dalla vecchia, i996a, b) creared new inreresr in this lithostratigraphic unit. the present nore concerns the stratigraphy, in particular the biostratigraphy, and the palaeoenvironment of the section containing the footprints. g. roghi contributed the palynological work and f.m. dalla vecchia the palaeoichnological study; other parts of the account were written jointly. stratigraphy and sedimentotogy. the well bedded alternations of black marls and grey and dark grey dolostones cropping out along the lower dogna valley were attributed to the monticello formation by jadoul et al. (1995). they are considered lateraily equivalent or even coinciding wirh the carnizza formation (fadoul et al., 1995, fig. 2) which crops out at the nearby valbruna and cave del predil (formerly raibl) localities. these alternating beds appear to lie below the dolomia principale, but this is difficult to confirm because of the steep morphology of the valley. no biostratigraphical and paiaeoenvironmental data have been published about this formation in the dogna valley. the monticello formation is well exposed along the dogna valley downstream from the aqueduct intake situated to the east of the small viilage of chiout di puppe. near the second waterfall downstream from the intake the foliowing succession is exposed (from the top downwards; fig. 2a): 3) well bedded grey dolostones alternating with black marls in tabular, centimetric-decimetric layers (about 70 m); in the lower part of this unit the pelitic intercalations are more frequent and there is often evidence of mud-cracking on the upper and lower surfaces dipartimento di geologia, paleontologia e geofìsica dell'università degli studi di padova, via giotto 7,i-35737 padova. kà dogna mt. lof gi dogna chiout ,./1962 _" ---1/-î\,//v !1 cortina à/or?heas?grlv italy trevìso fl f o 'lo 184 of the dolomitic layers; in the study area in the dogna valley the sequence ends at a prominent ledge which may be related to a change in lithology; 2) grey dolostones in banks with faint and irregular joints possibiy representing the "orizzonte di mestri" (giannolla, pers. comm.; see jadoul et al., î995) (about 20-25 m); in the upper part in a passage to the overhanging unit (3) there is a 1-3 cm-thick greenish dolostone; 1) some metres of well bedded, tabular, centimetric-decimetric alternations of grey dolostones and black marls, the lower limit of which we have not verified because unimportant for the scope of this paper. the reptile tracks occur in the middle part of unit 3 (fig. 2a), in the valley floor on the north side of dogna vailey, upstream from the intake of the aqueduct. the beds dip about 3o.ssw. the section containing the tracks is shown in fig. 2p. at the base of the section there is a breccia layer (dga98) with millimetric clasts, followed by two layers of marly'dolomitic mudstone (dga99-1oo). the track-bearing layer (dga1oo) is a disturbed mudstone with spots of wackestone at the top and deep mud-crack fissures; microfossils are completely absent. the track-bearing layer is overlain by black silty marls (dga101) which were sampied for palynomorphs. above this level there is a mainly marly unit with thin intercalations of sterile, black dolomitic mudstones (dgai02-1,03). the upper pan of the section g, rogbi & f. m. dalla vecchia location of the site comprises beds of fine grained dolostone, sometimes separated by very thin interwals of black silty marls; structures indicating subaerial exposure (fenestrae with vadose silt) are common at this level. ostracods are the most frequent microfossils (dga104, 107, 108 and 110) foraminifers (aulotortws?, nodosariidae) are less common (dga104, 105 and 110). the depositional environment could be a restricted lagoon, represented by beds with only ostracods, sometimes more open (dga110), which passed to a tidal flat represented by beds with fenestrae, fine grain dolomitization due to strong evaporation, perhaps stromatolitic lamination (dga104 ) and with long lasting supratidal exposure represented by vadose silts. this environment was involved in the repeated arrival of pelitic sediment which was probably deposited in a reducing environment, indicated by a black coiour and the presence of well preserved palynomorphs. the lack of marine fossils in the carbonate layers of the lower part of the section, where black marls are prevalent, could suggest a brackish or even fresh water environment. the frequent subaerial exposures, even in the iayer with a more marked marine influence (dga110) are noteworthy. they rcstify to the osciliating emersion of the zone and are in agreement with the presence of terrestrial reptiles. the monticello formation is considered to represent deposition on the inner platform (iadoul et al., l995) and the existence of an emergent area to the s and sw of the palynology and reptile trackuays in dogna valley c : mdst.pkst 5 & .9 wkst-pkst c : & m-w-p 185 dgal 07 dgal 06 dga105 dgal 04 dga103 dgal 02 dga1o1 dgaloo dga99 o =@ o oe oa,îf ^nyóro pd'e;.' ; c ! {@ (y'èq): --;s.^ = de--e a3>e ó,.€'-ò tvos*,sàp?ef h;;=e =o-> èó!t -:oè=+ = i q à ì -! j;e:€= è?e èse6e_3èèfiù;è !éèi.?ph--j3èe e.ièsé 8,:3:eeè eè:sg l'èhè*fe*è*esèí ósèosatyùíhèpèèpònfèdó;ì::f* rbsèès&8sèiei'=9rò p c o.e ò x è è f o * è bèg 3è b*rrrs sgeèff*óèéés*è èìefi&$ss:fn&è @ o palynological assei\,lblage sedimentologjcal structures and fossils f'.) a\s.h"-.ti."t'lliglapfii6sectionoftheobservedpanofthemonticelloformationalongdognacreek. 1,2,3:interualnumberl,_'b'2, 3 (see text). b) the section containing the layer with the reptile tracks. legend: 1) dark grey breccia with millimetric clasts, 2) dark grey to black marly dolostone, 3) black silty marls, 4) grey to dark grey dolostone, 5) light grey dolostone with undulating or mdst : mudstone, \ikst : wackestone, pkst : packstone, m-\fl-p : bands of mudstone, wackestone andrfféorrlrf r^rnt.. packsrone. fenestrae with vadose silt v desstcatton cracks (rvud cracks) 5 bioturbation. (disturbed sédiment) v reptile prints dogna valley during the deposition of the carnizzaformation was suggested by the same workers (ibidem, p. 87). vertebrate paleoichnology. the reptile trackways were described by dalla vecchia (1996a, b). there are at least four trackways and two of them are 2.40 m and 2.80 m long. they were produced by a quadrupedal plantigrade reptile, with a pentadactyl pes 17-20 cm long aîd aterr^dactyl or, most probably pentadactyl manus at least 50o/o smaller than the pes (fig. 3a,b; see also dalla vecchia, 1.996b, fig. 3-5). the free portions of the toes are rather short, narrow and sharply pointed. digit v is short, thin, placed rather posteriorly and is not laterally oriented but is curved forward; digits i-iv have similar lengths but digit iii is the iongest and i the shortest (fig. 3a,b; see also dalla \y'ecchia, 1996b, fig. 3-5). in the best preserved trackway the pace rs 43-47 cm, the stride 67-72 cm and the width about 30 cm; the pace angulation of pedes is osrracods e 1 17-t fo oesrnopoos a7 ' & fofat\,4'n,fers i . bt'alvta [-] 4 6è pollens a 5 about 100o. tracks not strongly outwardly directed, no evidence of feet dragging and absence of tracks of the tail, of the chest and of the belly indicate a relatively erect posture and a non-sprawling gait. therefore the trackmaker was well adapted to locomotion on land. the relation between tracks and mud cracks (dalla vecchia, 1996b) shows that these reptiles walked across a surface on which mud was drying or had just dried (see thulborn, 1990, fig. 5.19). another bed surface with mud-cracks and probable reptile tracks occurs in the creek bed, between the intake of the aqueduct and the first waterfall downstream (r. azzoia, pers. comm.). unfortunately this bed, which is at a stratigraphically lower level, is at present covered by debris. track morphology and, to some extent, the parameters of the trackways are reminescent of those of the crocodiles (see dalla vecchia, 1996b, fíg.8) and suggest a phytosaur or primitive crocodylomorph affinity (daila vecchia. 1996b). r mdst-pkst ] tocm 1,86 g. roghi & f. m. dalla vecchia b;= the tracks do not correspond exactly to any described ichnotaxon (haubold, 1971., 1984, 1986; olsen & padian, 1986). their formal description and the institution of a new ichnotaxon is beyond the scope of this paper and will be made elsewhere (dalla vecchia, in progress). crocodylomorphs are known from upper carnian deposits but are relatively smali, delicate reptiles represented by poor remains (long & murry, 1995). primitive crocodylomorphs of the late triassic and early jurassic were small, cursorial animals. it is unlikely that the earliest crocodylomorphs were larger than the succeeding late triassic and early jurassic forms and had a body shape and posture more similar to that of present crocodiles. the pes of primitive crocodylomorphs is functionally tetradactyl, with digit v reduced to a metatarsal spur (see dalla vecchia, 1996b, fig. z). the posi fig. 3 manus-oes set s from t he two main trackways. a) right set from trackway a (t' dalla vecchia, 1996b, frg. 1), light from rhe lefr iower corner; b) right set (5th) from trackway b (ibident),light from the left side. pictures taken from sili con rubber casts of the origi nal tracks. scale bar: 10 crn. tion of digit v in the pes tracks from dogna could be in agreement with its position in the primirive crocodylomorphs but this digit is more strongly deveioped in those tracks. phytosaurs were typical late triassic reptiles, with a distribution ranging from the middle carnian in north carolina to the rhaetian in switzerland, germany, new jersey and connecticut; maximum diversity was reached during the late carnian (tuvaiian) (benton, 1995). even if the general shape of the pes print is similar to the reconstruction of a phytosaur footprint (parrish, 1986, fig. 4.9a), the pedal skeletons of parasuchus lyddeker, 1885 and rutiodon tenuis camp, 1930 (see parrish, 1986, fig. 4.4) and pseudopalatus pristinus mehl, t928 (long & murry, 1995, îigs 53, 55) do not correspond exactly to the footprints from dogna valley, mainly in respect of the position and shape of digit v (fig. 3; cf. dalla vecchia, 1996b, fig. 3). therefore, comparison should be made with a phytosaur with a digit v characteristically placed very posteriorly and such a phytosaur is not known amongst the available skeietal remains. a third possible trackmaker considered by dalla vecchia (1996b) is an aetosaur. aetosaurs were typical terrestrial quadrupedal plant-eaters of carnian to norian times. the oldest remains are from the ?middle carnian of north america, the youngest are from the "rhaetian" of argentina and england, with a higher diversity during late carnian-early norian (benton, 1995, long & murry, 1995; lucas & fleckert, 1996). large aetosaurs are usualiy reconstructed as plantigrade and have a short and rather posteriorly placed digit v of the pes, with a curved metatarsal y. stagonolepls agassiz, 1844 and paratypotordx long & baiiew, 1,985 are late carnian aetosaurs with a size comparable to that of the trackmaker of dogna tracks (i-ong ee murry 1995). the rauisuchians are excluded as the potential trackmaker because of the different parameters of their trackways. the possibility remains that the trackmaker was an unknown taxon. trackmaker length on the base of the body proportions of the well-known phytosaur parasucbus, can be roughly estimated as slightly iess than 3 m. on the basis of the reconstruction of the aetosaur stagonolepis the length was about 2.5 m. the morphological peculiarity of the tracks may make them useful for the biostratigraphic correlation of non-marine units (haubold, 1986) and for correlation of the monticello formation with non-marine formations. palynology. the black silty marls (dga 101; fig. 28) just above the track-bearing surface were sampled for palynological investigation. a rich and very well preserved miospore association was found. seventeen taxa have been identified and a quantitative analysis on more than 2oo isolated grains was made. in the sample, the miospore are organized in single grains (85%) and in tetrads (15%). treatment of the monosaccate group follows that adopted by scheuring (1970;1978) and brugman (1983) and the specimens found are in agreement with the descriptions of these authors. some problems exist in the taxonomy of the bisaccate group. the descriptions of van der eem (1983) and brugman (i983) have been used for the identification of the circumpolies; taxonomy is problematic when the grains are organized in tetrads. a detailed taxonomic treatment will be given in an account of the palynological study of the whole section. palynology and reptile trackzaays in dogna valley t87 the results of the quantitative analysis are as follows: circumpollen 560/o unidentifiedbisaccatepollens 27o/o monosaccate and vesicate pollens 8vo alete bisaccate pollen 7o/o rok taeniate bisaccate pollen 0.5o/o trilete spores 0.5o/o the following taxa have been recognised (the recognised taxa are listed in fig. 1 and the most significative miospore are illustrated in plate 1): monosaccate: enzonalasporites oigens leschik, 1956 (pl. 1, fig. 1) vallasporites ignaciileschik, 1956 (pl. 1, íig. 2-3) patinasporites d.ensus (leschik, 1956) scheuring, 1970 (pi.1, fig. 6) zonalasporites cinctus leschlk, 1956 pseudoenzonalasporites summus scheuring, 1970 (pl.1, fig. a-5) bisaccate: sarnaropollenites speciosus goubin, 1965 (p1. 1, fig. 9) ovalipollis pseudoalatus (thiergart, 1949) schuurman, 1976 (pl. 1, fts.7) chordasporites spp. lueckisporites spp. lunatisporites acutus leschlk, 1956 klausipollenites sp. a i i sp orit es / fa i c ispori t es spp. circumpolles: camerosporites secd.tus leschik, 1956 (pl. 1, fig. 12) partitisporites maljaukinae (k.laus, 1960) van der eem, i983 (pl. 1, {is. i 1) partitisporites nnimundanus leschik, 1956 partitisporites quadruplicis (scheuring, l97a) yan der eem, 1983 corollina aff. sp. a (sensz schuurman, 1.977) (p1. 1, fig. 10) the combined occurrences o[ enzonalasporites aigens, vallasporites ignacii, patinasporites densus, pseudo enzonalasporites slirnrnus, samaropollenites speciosus and carnerosporites secatus are indicative of tuvalian age (late carnian). this palynoflora is nearly identical to that described by visscher & krystyn (1978) ín the carnian-norian transition sequence of monte triona, sicily, corresponding to the boundary between the ammonoid tropites subbullatus and anatropites bereichi zones (krystyî, 1974, tuvalian 2-3 boundary). a microfloral association containing all the taxa found in sample dga 1,01 was described from a carnian evaporite succession in albania (cirilli & montanart, 1994) at a stratigraphic level attributed to the tuvalian. the same raxa are found from litwin & ash (1993) in the chatham group, deep river basin (north carolina, usa), and are, with the exceptio n of samaropollenites speciosus (dunay & fisher, 1924), present in the chinie formation and dockum group (s\f usa). fisher & dunay (1984) reported the presence oí samaropollenites concin' nus, a form similar to s. speciosus, in a microflora from the blue mesa of arízona, which contains all the taxa found in sample dga 101. 188 g. rogbi g f. m. dalla wcchia dunay & fischer (1978) reported a microflora with camerosporites secdtus, enzonalasporites oigens, patinasporites densus, vallasporites ignacii and brodispora striata (pseudoenzonalasporites sumtnus and sarnaropollenites speciosu.r are not present) in levels of the opp.nitzer-kalk (austria) of tuvaiian age (tropites subbullatus zone). following these authors, brodispora striata has a particular biostratigraphic significance since it was found with the above mentioned association only in the late carnian arden sandstone of england (clarke, 1965; fisher, 1972; warrington, 1970) and in some localities of the chinle formation and dockum group of s\f usa (dunay & fisher, 1,974; 1979; fisher and dunay, 1984). fiowever, brodispora striata was not found in the late carnian microfloral assemblages of the southern alps (this work), sicily (visscher ec krysryn, 1.978) and albania (cirilli & montanart,1994). therefore the presence of brodispora s*iata could be attributed to palaeogeographical causes. in sample dga101 a form very similar to corollina sp. a, sensu schuurman 1977 and van f.we 1977 (: círculina sp. a) has also been recorded as cf. corollina aff. sp. a. the specimens are or3 nized in tetrads, have a subequatorial circular furrow, a smooth sexine and faint equatorial infrastriature; a distal pseudopore is absent, or is ill-defined (pl. 1, fig. 10). palaeogeography and palaeoenvironment. the palynological assemblage reported here shows a noteworthy likeliness with the onslow microflora (dolby & balme, 1976). the latter is a mixed association with elements belonging to the gondwana and to the laurasia continents and is different to the high latitude boreal and austral microfloral associations. all the taxa found in sample dga101 are present in the onslow microflora but in different proportions. bisaccate pollen such as sulcatisporites are prevalent in the onslow microflora, while circumpolles are most abundant in sample dga101. visscher & van der zwan (1981) identified three late triassic floristic zones: a northern one characterized by camerosporites and ooalipollis, a middle one wíth camerosporites, ooalipollis and samaropollenites, and a southern one with camerosporites and samaropollenites. since the dga101 microflora includes camerosporites, ovalipollis and samaropollenites it belongs to the middle belt, known mainly from localities in north africa. the dga101 microflora consist aimost entirely of circumpolles, bisaccate and monosaccate pollens, which belong to the typical triassic conifers; only one spore was observed. the monosaccate forms have been found associated with the conifer brachypbyllum (yan konijnenburgvan cittert, 1971). the circumpolles show affinities with cheirolepidaceae; in fact corollina was found in association with the conifer hirmeriella (francis, 1983; van konijnenburg-van cíttert, 1.97 1). the pollen, particulary the circumpolles, are rypical xerophytic elements, which indicate an arid climate (cirilli & montanarr,1994; visscher et a1.,1994). aridíty is also indicated by the total lack of miospore produced by hygrophytic plants (ferns or equisetales). this is in agreement with the position of the mediterranean region near the palaeotropic, where the cli, mate was arid and warm during the late triassic (robinson, 1973). the presence of gypsum in the upper carnian of carnia (pisa, 1,971) and of aupa valley (bianchin et a1., 1980) is additional evidence of aridity (hallam, 1984). the microflora is represenrative of the vegetation living at or near the site where the reptiles walked leaving their tracks. this is indicated by the relatively high number of tetrads, as these usually disarticulate during prolonged transport and generally form a very low proportion of a palynomorph association. acknouledgements. '!le thank sandro venturini for the description of the thin sections and the help in the interpretation of the depositional environment. 'we are grateful to maurizio tentor and to the gruppo speleologico a.d.f. of monfalcone (gorizia) for the preparation of the thin sections, to stefano castelli for the pictures, to nicola michelon for the drawings and to andrea vorlicek for technical advices. thanks to piero giannolla for the useful discussion concerning the stratigraphy of dogna valley. the writers are grateful to prof. g. 'sfarrington and to prof. m. prosperi evans who revised the english text. this work was sponsored by a m.u.r.s.t. grant (ex 40% v de zanche;600lo mietto). plate i the slide collection. coordinates of the figured specimens were taken with the england finder using leitz 'wetzlar no. 5345 with attached camera. all the slides are housed in the dipanimento di geologia, paleontologia e geofisica of the university of padova. 1) enzanalasporttes vigens i-eschrk, l956, (45 pm); slide dognal ii, g 3l/4;2-3) vallasporites ignaciileschlk, 1956 (48 pm); slide dogna 1 i1,v 4a/),;4-5) pseudoen. zonalasporites sunmt{l scheuring,1970 (3 8 pm); (4, single grain, slide dogna 1 y, l 46/ 1,; 5, tetrad, slide dogna 1 v n 46); 6) patinasporites densus (leschik, 1956) scheuring,1970, (4a pm), slide dognal iii, k39/i;7) ooalipollis pseudoaldlus (thiergan, 1949) schuurman, 1976 (ength 81 pm), slide dognal, lii,m 42/1.;8) lueckisporites sp. (48 pm), slide dognal, s 48/3i9) samaropollenrtes speciosus goubin,1965, (ength ez pm), slide dognali,534/3.i0)cf.corollina aff.sp.a(sensuschuurman,tstt), (diameterof singlegrainintetradis33pm),slidedognalil,ka1/\1.1) partitisporites maljauhinae (klaus, 1960) van der eem, 1983, (diameter in single grain in tetrads is 40 pm), slide dognal v,d 41,14; 1.2) camero" sporites secatus leschik, 1956, (45 pm), slide dognal iv, g 3i/4. palynology and reptile tracktoays in dogna valley 1ì 189 -*"-,. .., i$i* 3l'':wp 190 g. roghi & f. m. dalla vecchia references benton m.j. (1995) late triassic to middle jurassic extinctions among continental tetrapods: testing the pattern. in n.c. fraser and h.-d. sues (eds) in the shadow of the dinosaurs early mesozoic tetrapods, pp. 366-397, cambridge. bianchin g., carulli g.b.,frizzo p., longo salvador f., mantovani f., masé g., mezzacasa g. tr semenza e. 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(1994) rejection of a carnian (late triassic) "pluvial event" in europe. reo. palaeobot. palynol., v. 83, pp. ./.r/-226, amsteroam. visscher h. & van der zwan'c.1., (1980 palynology of the circum-mediterranean triassic: phytogeographical and palaeociimatological implicatíons. geol. rundsch., v. 7 0, no. 625-67 4. stuîtsart.rr'"-" warrington g., (1970) the stratigraphy and palaeontology of the "keuper" series of the central midlands of england. ql geol. soc. london., v. 26, pp. 183-216, london. received july 31, 1996; accepted may 26, 1997. acosta 23..34 ��� �������� � �� ���� � � � ������ ����� ����� �� �� ��� ���� � ��� � ������ �� ��� ���� � �� � �������� �������� ������ �� ������ � � ������ �� ����� � ��������� ��� �� � � ����� ��������� �� ����� ��� ���� ��� � ���� ������� ����������� ������ ���� ���� � �������� !��" �#����$� %�&��� !��#�����' � ������' �#� ������� ����� ����% �(������ )� **"+"�,"� ��!�� ���! �#� ���� ��&��� �� �� )����� -��!����� .��$!� � �����%� ����������/ �� �#� �� �#0������ ����� �� �#� ������ ' �#��� ��!���� 0��� �� �% �� �#� � 1!����� ����!�!1��� 0��#�� �#� ����� � ���� � ������� � �� 2��3���� %���% �� 45'� )� ���� ������� ! 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accepted april 17,2aa0 key-uords: tanystropheus, carnian, middle-late triassic, carnie aìp.. northeartcln itrlt. riassunto. vengono descritti i printi resti ossei rinvenuti ne11'italja norclorientale scnza dubbio appartenenti al grande rcttile prolacertiformc tdqtstropheils. si tratt;r di una vertebra caud:rle prossimale dcl triassico medio della val aupa (udine, friuli) e di una vertebra cervicale del carnico di fusea (uiline). la vertebr:r cervicalc r:ìppresenta 1a prima segnalazione clel gencre nel carnico e la sua testimonianza più re cente, se si csclude t. jossai tlel norico. dur;rnte il triassrco superiore thnystropheu5 1,i1,g1'1 lungo le coste della tetide nordoccidentale mentre era scompàrso nell'europa centrale, dominata da ambienti continentali. abstrdct. thc first diagnostic remains of the large prolacertiform tanystrophezs arc rcported fron northeastcm ltall'. thc;include a proximal caudal vcrtebra from the mjddlc tiassic of aupa valley (udinc, friuli) and a cervical vcrtebra fron the carnian of fusea (udine). the cervical vertebra rcpresents thc first record of tanystropheus in the carnian arrd is the geologically 1-oungest occurrence other tlran the norian z fossai. tanystropheus hved along the coasts of the northwestern tethl-s during thc late trjassic r.hile it disappeared in central europe u'herc continental cnvironments t-ere prevaìling. lntroduction. during the last ten vears the amount of reptile remaihs found in the mesozoic formations of northeastern italy increased dramatically (sirna et al., 1994; dalla vecchia, in press; rieppel & dalla vecchia, in press, and references therein; marco ar-anzini, pers. comm.) with the discovery of most of the main reptilian clades, such as chelonia, placodontia, sauropterygia, ichthyopterygia, prolacertiformes, crocodylia, pterosauria, and dinosauria. some remains of coastal reptiles from the middle to upper triassic of northern friuli n'ere described by dalla vecchia (1994) and most of the sauropter\-gians, the placodonts and the truly marine reptiles (ichthyopterygians) from the middle-upper triassic of ne italy are described by rieppel ec dalla vecchia (in press). new specimens were found in two different localities and triassic horizons of the northern part of the friuli region (fig. 1). they belong to the prolacertiform tanystropheu.t v. me)rer, 1852, and enrich the list of the mesozoic reptiles found in this eastern area of southern alps. abbreviations: mfsn, museo friulano di storia naturale, udine. systematic palaeontology class reptilia t,aurenti, 1768 subclass diapsida osborn, 1903 superdivision neodiapsida benton, 1985 division archosauromorpha von huene, 1946 order prolacertiformes camp, 1945 genus tanystropbeus h.v. meyer, 1852 lvpe species: t. conspicuws h.r meyer' 1852 tanystropheus sp. specimen mfsn25761 description of tbe specimen dnd comparisons. the specimen was completely freed from the matrix. it is a complete, isolated proxirnai caudal vertebra 37 mm long and 52 mm tall of a prolacertiform reptile (fig.2). its size and morphology closely corresponds to that of the proximai caudal vertebrae of tanystropheus figured by wild (1973, figs. 57-61, pls.7-9;1980, pl. 5). the specimen is slightly deformed due to compression. the right prezygapophysis, right postzygapophysis and right pleurapophysis are clearly more developed that the left ones. this cannot have been caused by compression museo paieontolosico cittadino, via \ralentinis 1i.{, 14074 monfalcone (goriz-ia), italr,: e-mail: fabdalla(rr'tin.it a ustria )-'-'-'-'-.-^--.-'' 1f -t=.-=-. .t ) ,a'j 't* a' (r! \ u\*.-. friuli ) .r' slo,:.( \ udine t-ì \í o o j 136 f. m. dalla vecchia because compression can deform a body but cannot reduce its size. the skeletal element was probably original11' 6i5rh"pen because of a rrauma which affected the base of the tail, a part of the body subject to the highest stresses. despite the fact that the specimen is presen-ed within a conglomerate, representing a relatively highenergy environment of deposition, it did not suffered a prolonged transport. for example, it does not show any darnage to the long and thin pleurapophyses. only the tip of the left pleurapophysis was posteriorly crushed by a clast and appears spoon-like. the amphicoelous centrum (24 mm long) is lareraliy constricted just beiow the pleurapophysis. the ventral profile is also concave (arched) in lateral view and the cenrrum is spool-shaped. the articular surfaces are elliptical because of a slight deformation. there is a narrow longitudinal groove along the ventral side, like in the caudal 6 or 7 o{ t. conspicuus figured by \(/ild (1973, fig.6oe). the elongated and'narrow pleurapophyses (the right one is 3z mm long) are directed laterally, posteriorly and slightly downward. they attach to rhe cenrrum between the basis of the pedicel of the neural arch and the dorsal margin of the centrum. the pleurapophyses are fiattened dorsoventrally but the distal tip was probably swollen and rounded. the neural arch is fused to the centrum but a suture is possibly present at the base of the right pedicel. the prezygapophyses are very short and do not project beyond the cenrrum. their articular surface is eliiptical and faces forward, dorsally and only slightly outward. they are connecred to rhe anterior neural spine by short supraprezygapophysial laminae. another channel-like srrucrure is developed between the prezygapophyses and just above the small neural channel. its floor is a short intraprezygapophysial lamina. the postzygapophyses project posterolaterally beyond -rlg. i location of the fossiliferous localities in friuli, ne italy. 1) dell'andri creek, aupa valley;2) fusea. the centrum and are larger than the prezygapophyses. they have elliptical elongated articular surfaces, facing downward and slightly posteriorly and laterally; however, the orientation is somewhat distorted by deformation. supraposrzlrgapophysial and intrapostzygapophysial laminae are presenr. the intrapostzygapophysial laminae converge medially and ventrally inside the neural arch, just above the neurai channel. there is a deep infra-postzygapophysial cavity. the neural spine is quadrangular, wide and slightly inclined backward. irs crosssection in anteroposterior view is a tall, upside-down triangle, and its top is thickened and flat. the surface of the neural spìne i. rugose, rhe anreroventral parr just above the prezygapophyses is blade-like and becomes a prespinal larnina. the centrum is comparatively less eiongated than the centra of the caudai vertebrae o{ t. iongobardicus (bassani, 1886) and t. conspicuus. the shape of the pleurapophyses is similar ro thar of the pleurapophyses of the anterior caudal vertebrae of t. /ongobardicws (wild, 1973, p|s. 7-9; 19sa, pl. 5). the shape of the pleurapophyses differs from that of the anterior caudal vertebrae of t. conspicuus which are much broader (\7i1d, 1973, {igs. 57-61). there are other characters of mfsn25761 which differ from those of the caudal vertebrae of t. conspicwus: the posterior inclination of the neural spine of mfsn25761 is higher, the prezygapophyses are shorter and have articular facet facing more forward (i... the facet is subvertical), the intraprezygapophysiai laminae are shorrer and thinner, the neural channel is smaller, and the lateral sides of the centrum are probably more excavated. the size indicates that the vertebra belongs to a rather large individual which, however, was smaller than t. conspicuus. tanystropheus remains frctm friult 137 proximal portion and it enllrges fan-wise distally. although this lrone ir not actually diagnostic. it clorely resemble. the posterior thoracic ribs of t. longobardicus (\íild, 1973, fig.35 left). since it was found in the same block with the caudal vertebra and it belongs to a large reptile of comparable size, it could plausibly belong to the same taxon of the caudal vertebra and possibly to the same individual. other very fragmentary bones (mfsn25z63, mfsn 25764, mfsn25765, mfsn 25766) found in the same block are not cornpletely freed from the rock matrix and cannot be identified with certaintl-. geological setting. the specimens were found by dr. corrado rosenfeld in a single isolated block near the confluence of the dell'andri creek fig.2 -mfsn25'61, anterior caudar..ertebra t:\1:.1"1:::t"-111":t^::i' a'rrrr.rior rrcn. b) po\rerior rieu. cr f ne drlohe ol tne roao de\-ordorsal r.icn,, d) r.entral vìer. e) left latchians-studena alta (aupa valeral vien'. the centrum is 2/ mm long ley, moggio ljdinese, udine and the wholc vertebra is 52 nm hìgh. province; fig. 1). the rock conlegenda: pr:prez,vgapophvsis; pz: i"irrir.,g the specimens is con_po'tzvq:pophr''ì' glornerate with.r silty-carbonate matrix and \\ ith d;rrk gray. the proximal half of a thoracic rib, without the millimetric carbonate clasts, bivalves, and echinoderm articular head (mfsn 25762; fig. 3), was found in the elements. only the carbonate upper serla formation same block with mfsn2576|. the proximal portion of (1ate anisian) and the terrigenous "torbiditi d'aupa" mfsn 25762 is anteroposteriorly flatteneci with an (1ate anisian-early ladinian) outcrop in the catchment ellipgical cross-section and presents an anterodorsal and basin of the creek (jadoul & nicora, 1929). the limea posterodorsal longitudinal ridge. the distal portion is stones and dolomitic limestones of the upper serla forelliptical in cross-section, it is less flattened than the mation are decidedly different from the lithology of the l\lfsn25762. p:rrial po.r, rior rhoracic rib. scrle bar : i cm. trio l 138 f. m. dalla vecchta bone-bearing block, whrch shows a significativc relriqenous content. the "torbiditi d'aupa" is a n-rar1-sandstone basinal, deep-rvater unit. i{or-er.'er, manv other lithostratigraphic units rrnqing fron.r brs:rl triassic to norian ourclop .rlong rhe r:rlley and. lor crrmple. 'inrilar conglon.rerates with crrbonrre-terrigenous components could be present in other middle trirssìc units (e.g. the "terrigeno ladinico"; jadoul er nicora, 1929). the block could come frorn the morainal deposits all around the creek and could har.e been transported frorn nearby valleys bv glaciers during pleistocene timcs. an1'n'ay, a stratigraphic frarnework similar to that of the aupa valley is found along these vallc1's. specrmen mfsn25760 description of the specìmen and comparisons. the specin'ren is the posterior half of a single isolated cenical vertebra, the anterior half of which was weathered away (fig. a) . the prescrr-ed pert of rhe vertebra, bone .-'l -"','.'l -^l.l i, | )r -lorrrnr'l rhn .""."rrr",-ldilu li!tluldl lllulu! lj r-u rilrrr rvrròt artu portion of the centrum is 123 mm long. the part of the centrum preserved as bone is 68 mm long. the middle to anterior part of the centrum, lacking the anterior end carrying the prezygapophyses, is presen'ed only rs natural moid. the tube-shaped centrum is very loq about 1o mm in the middle, and collapsed because ir is hollow inside. the posterior articular end of the centrum, l4 mm high. i. enl.rrged.rnd slightly deflected ventraìly. the processus spinosus of the neural arch projecrs shortly above the postzygapophyses, is triangular and similar to the lin of rn airplane t:ril. ending cranialll'.rgain,r rhe fig. ,+ x,ifsn25z60, cenicel r-ertebra, right latcral r,ien-. a) photograph, b; tlrari.inr.'1'he scale blr is ccntirnctric in a and 2 cm in b. legend.r: cì centrum; p$ : processus spinosusi pz : ìrost7)eapophvsis. dorsal mrrgin of the ccntrum. it is similar to the processus spinosus of the cerr.ical vertebra 7 and 1(wild, i973, pl. 9), z-8 (\fild, 1923, pl. 11) or 5-z (wild,1973, pl. 16) of t. longobardicus. the neural:rrch is exrremelv lo$t:.r feature of the cervical vertebrae oí tanystropheus. in f;ct, rccording to pever & kuhn-schnvder (1955, p.591, translated from frcnch) in the cervical vertebrae of thnystropheur "..the neural arch is not rised above the r.ertebral centrum, and the neurai channel seems ro pàss inside the latter". this character can be observed in the cervicals of t. conspìcuus figured in wìld (1973, see the r.'ertebral cross sections in figs. 43, 11, 51). thc rìght postzvgapophysis is relativeiv srrong and long. project;-,. *.^ll h.-,^-l ,l^ ^osterior end of the centrum. the specimen mfsn25760, with its elonsated tubular aspect, and very lon neural arch, resemble s most closeh' the middle to posterior cen.ical r.errebrae of tanystro, pheus longobarc{icus (\lild, 1923" pls.9, 11, 13, 16). geoìogica/ setting. thu specimen n-rs found in the fusea site (tolmezzo, udine; fig. 1). i collected the bone ar rhe top surface of the laver e (fig. 5) of this site n'hich is particularly rich in fossii ve rtebrates. abundant disarticulated elemenrs ol notbctsrturus sp., ìt/. cf. giganteus múnster, 1834 and of a cyamodontoid placodont (dalla vecchia, 1994; rieppel & dalla vecchia, in press), together with acyodus-like teeth and other fish teeth, and plants were found on the ex, posed upper part of layer e. scattered bones and armour elements of cyamodontid placodonts, bones and teeth of l"lothosaurus sp. and fish teeth are also found in the middle part of the layer and in the layers a, d and f. in layer d, skull roof bones of a dipnoan fish were also found. tanystropheus remains from friult 139 1o cm 25760 m fsn aaa aaa 't, l a' a a a ,aa 4a a.a aa fig. 5 stratigraphìc column of the fusea site, lower carnian. legcnda: 1) black limestonc (n-achestones-packstones), 2) conglomcrate wìth prei'ailìng carbonate clasts, 3) s'hite to ' light-gral' dolomitic 1ìmcstone, poorlv bedded and with lenses of intraformational breccia. the arrowpoint to the posìtion of the vertebra n{fsn25z60. the stratigraphic sectiotr contr;ning the bonebearing layers is just at the transition between the top of a thick carbonate platform sequence and well-bedded black limestones hundreds of metres thick. the carbonate platform sequence is referred to as dolomia cassiana by carulli et al. (i995, p. 76) and as dolomia dello schlern by prsa (in braga et a\.,1971'), ^nd dated as late ladinian-middle carnian in both papers. it is the iast carbonate platform before the deposition of the dolomia principale. the thick sequence of "black limestones" is the basal portion of the so called "raibl group" (pisa in braga er al., i97i). this basal part is considered middle carnian in age by pisa (in braga et al., [_lll v{,1lo"ooì e-a;l lo oil 1971) because of the presence oî myopboria leefersteini (mùnster, 1845) and clypeina besici (pantic, 1966). the layer where mfsn25760 was found is at the very base of these "black limestones". for references on the geology of the surroundings of fusea, see dalla vecchia (1994). the comparison with the well-studied stratigraphy of dolomites suggests that the stratigraphic position of the section is most prob;ably in the upper part of the lower carnian, between car2 and car3 depositionai sequence (de zanche et al., 1,993). this dating is also supported by carulli et al. (1995, p.76). the tentative of biostratigraphical correlation of the fossiliferous section by palynomorphs w.as not successful (guido roghi, pers. comm.), and the rare foraminifers indicate only a possibly late ladinian to earll. carnian age (presence ol trocholina cf. cordeoolica, sandro venturini, pers. comm.). conclusions. most of the fossil record of tanystropbeus comes from the muschelkalk of europe (southern spain, france, germany, slesia,transylvania) (.i.e. t. antiquus v. fiuene, 1908, z conspicuus and z. sp.), the muschelkalk of israel (peyer, 1955), the upper anisian-anisian/ ladinian grenzbitumenzone (= besano formation) of tessin (southern switzerland) and lombardy (nv ital;') (z longobardicws), and the lower ladinian lower meride limestone of tessin (7. meridensis \fild, 1980)' a fragmentary cervical vertebra is reported from the middle triassic of saudi arabia (vickers-rich et al.' 1999). a partial, single tooth attributed rc t. cf. meridensis (wrld, 1980) was also found in the lowermost keuper (upper ladinian) of sv germany. a partial dorsal centrurn tentatively identified as tanystropheus 6y \flild (1980, p. 14, fig. 1o) comes from the "tufi a pachicardie", upper ladinian of alto adige/sùd t;rol (n italy) . the youngest record ol tanystropbeus is fron-r the upper norian argilliti di riva di solto of lombardy with the sn-ral1 z/ossal \flild, 1980 (\fild, 1980; benton, 1994). t. fossai is represented only by a segment of four vertebrae from the cervical vertebral column. the specimen belongs to a very small individual, the vertebrae are not longer than l5 mm. ]t is rery similar to a segrncnt oi the n-riddle tail of a basal pterosaur, and pterosaurs have been found in the same formation. hor''ever, some structures (e.g. the rised facets for the articulation of the cervical ribs) support the identification as cervical vertebrae of tanystropheus. the specimens mfsn2576a and mfsn25761 here described are the first remains unambiguously attributable rc tanystrophews fron northeastern itaiy. the caudal vertebra from aupa valley partly differs from the proximai caudal vertebrae of tanystrophezzs described in literature but the material is too incomplete to war2 ^o qgoq ipo o oc ogoc)oooo oo o oooo a oooòoooò ooooòooo soòooooo-o oo og 006 ^ o o o ó o oo "oo oo^oó oò aooo"ooo "o o o oo o o o ooooo(2oo 140 f. m. dalla veccbia rant the erection of a new species. the specimen from fusea is the first carnian evidence of tanystropbeus and, other than the norian t. fossai, is the geologically youngest occurrence of the genus. the absenc e of tanystropbeus in the central europe during the late triassic (keuper) is most probably related ro the reduction of coastal environinenrs and the predominance of continental environments not suited for the way of life of this reptile. the genus survived during the carnian and the norian along the coasrs of rhe western tethys. the tubular vertebra mfsn25z60 confirms the phylogenetic trend of the cervical vertebrae of tanystropheus toward a more and more elongation, hypothesized by \lild (1980, p. 25-26, fig. 1a). acknouleclgenents. i thank dr. carlo morandini and dr. giuseppe muscìo, respcctivell' director and curaror of the museo friulano di storia naturale of udine for thcir pernission to study the material. thanks to dr. corrado rosenfeld, rvhose collaboratìon has always bccn fundamental for the progress of the vertebrate paleontology in thc frjuli region, to dr. sandro venturini for the biostratigraphic and the nicrof:rcies analysìs and to dr. guido roghi for the research of palynomorphs in the samples frorn the fusea site. i am grateful to mr. maurizio tentor who realizcd the thin scctions of the samples at the laboratoires of the museo paleontologico cittadino of monfalcone (gorizia). thanks to dr. stefania nosottì, dr. silvio rencsto and dr. rupert wild for thc ìnformations and the discussion and to dr. anna p;rganoni for the access to the collections of the museo civico di scienze natur;rli of bersamo. fina1l1', i thank dr. stefania nosotti and dr. olivier rieppel for the criricel rer:ew ol rhe m.rnrscripr. refe,rences benton m.j. (1991) late triassic to middle jurassic exrinctlons among continental tetrapods: tesring the prttern. in fraser n. c. & sues h.-d. (eds.) 1n the shadow of the dinosaurs early mesozoic tetrapods, pp. 366-397, cambridge. braga gp., carloni g.c., colantoni p, corsi m., cremonini g., frascari f., locateìli d., monesi a., pisa g., sassi f.p, selli r., vai g.8., zirpoli g. (1971) note illusrrative della carta geoloeica d'italia alla scala 1:1oo.ooo: fogli 4c-13 monte cavallino-ampezzo. sen. geol. italia, pp. 1-108, roma. carulli g.b., longo salvador g., ponton m. (1995) lc unità ladino-c:lrniche nella carnia centro-occidentale. ann. unio. ferrara, sci. terra, v. 5 (sr.rppl.), pp. 75-81, ferrara. dalla vecchia f.m. (1994) reptile remains from the middleupper triassic of the carnic and julian alps (friuli venezia giulia, northeasrern italy). gortama atti museo friul. st. nat., v 15 (1993), pp. 19-66, udine. dalla vecchia f.m. 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(in press) marine reptiles from thc triassic of the tre venezie :lrear northeasrern kely. fielcliana, chrcago. sirna g., dalla vecchia f.m., muscio g. & piccoli g. (1994) catalogue of paleozoic and mesozoic verrebrares and vertebrate localities of the tre venezie area (north eastern italv). mem. scí. geol.,v.46, pp.255-281, padova. vickers-rich p, rich t.h., rieppel o., thulborn r.a. & mcclure h.a. (1999) a middie triassic vertebrate fauna from the jilh formation, saudi arabia. n. /ú. geol. palàont. abh., v.213 (2), pp.2a1-232, stuttgrrt. \i/ild r. (1973) die triasfauna der tcssiner kalkalpen xxiii. thnysnopheus longobardicus (bassani). sclttoeiz. palaont. abh.,v.95, pp. 1-162, basel. sfild r. 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�) )���' 4��' 4���' &��"�� �� ������' ��� ���"�<�) << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /all /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /warning /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /syntheticboldness 1.00 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments 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(http://www.color.org) /pdfxtrapped /unknown /description << /fra /enu (use these settings to create pdf documents with higher image resolution for improved printing quality. the pdf documents can be opened with acrobat and reader 5.0 and later.) /jpn /deu /ptb /dan /nld /esp /suo /ita /nor /sve /kor /chs /cht >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [595.000 842.000] >> setpagedevice rivista italiana di paleontologìa e stratigrafia volume 104 numero 3 tavole 1 15 dicembre 1998 ne\/ deep.sea nou brew . short note bryozoan species from of southern italy the pleistocene antonietta rosso* receioed september 21, 1997; accepted april 17, 1998 taxa. key-ztord.s: bryozoa, pleistocene, deep-sea, southern ltaly, new riassunto. vengono descritte due nuove specie ed una sottospecie di briozoi: 1èroia barrieri (cyclostonatida), heliodorna angusta (cheilostomatida) e crisia tenella calvet longinodata (cyclostomatida). tutte provengono da sedimenti essenziàlmente marnosi af{ioranti nell'area dello stretto di messina (italia meridionale) e depostisi in paleoambienti degli orìzzonti sommitali de1 piano batiale. si tratta di taxa endemici de11'area mediterranea, estinti, probabilmente stenotermi che potrebbero rivestire un interesse paleoclimatico e stratigrafico. abstract. two new species: térvia barrreri (cyclostomatida) rnà heliodoma angusta (cherlostomatida) together with the subspectes crisia tenella calvet longinodata (cyclostomatida) are described from pleistocene deep-sea sediments cropping out in southern ltaly. they are mediterranean palaeoendemics, closely related to recent deep-water atlantic species. lntroduction. this study is part of a program to define the composition and structure of benthic palaeocommunities from pleistocene deep-sea sediments (southern italy). this area, and above all the messina strait, is particularly suitable for the study of deep-sea palaeocommunities, as strong neogene tectonic activity caused a remarkable uplift of deep-sea plio-pleistocene sediments (barrier et al., 1986; barrier, 1987). the sediments are very fossiliferous. fossil assemblages are abundant and diverse with bryozoans, molluscs, scieractinians, isidids, serpulids, brachiopods, crinoids and barnacles. ail these systematic groups show a clear atlantic affinity as attested by numerous atlantic guests and northern or boreai guests (gaetani ec saccà., 1984, zibrowius, 1987, roux er ai., 1988, barrier et al., 1989, di geronimo & la perna, 1996, 1997a, b; rosso & di geronimo, 1998). the former are atlantic deep-sea species which spread in the mediterranean fo1lowing the general climatic cooling since the middle pliocene. the latter are bjorth atlantic shelf species whose latitudinal range shifted southwards during cold pleistocene phases, thus comprising the mediterranean. these atlanticlike palaeocommunities are characterised also by the presence of some species closely related to recent atlantic taxa from which they differ by slight morphologic characters (di geronimo et al., 1996; di geronimo & la perna 1996; 1997b). these species may be mediterranean pleistocene palaeoendemics. in this paper, new bryozoan species with paleoendemic characteristics are described. materials. the material studied originates mainly (fig. 1) from three pleistocene fossil localities, two (lazzàro: 2q samples; and archi: 12 samples) located in southernmost calabria, and one (furnari: a single sample) in ne sicily, along the tyrrhenian side of the peloritani mounts. fossil bryozoans came {rom sediment bulksamples ranging in volume from2 to 10 dmr. additional material (30 samples belonging to eocumms4-95 campaigns) comes from upper pleistocene (? wùrmian) submerged sediments (300-1,500 m depth) from the se tyrrhenian sea (gioia and cefalù basins adjacent to the eoiian islands) and from the northern part of the tyrrhenian sea (2 samples from the bs/77 and bs/78 campaigns, 500-1500m depth), off corsica and sardinia" samples were collected mainly with a box-corer; a few ones were dredged. sediments were disaggregated in water and gently washed using a 63 pm mesh to obtain even very smail bryozoan specimens and fragments. dried fractions ) 250 pm 'were routinely sorted, whereas finer materials were usualiy checked to iook for rare and/or the smallest species. 'llstituto policattedra di oceanologia e paleoecologia, corso italia, 55 95729 catania, italy. sea lyrrhen\an l éo\i?1 a \.r3nòt to" a s oul qrup at l_ r{ * 0,,u,.1"ìj" d 424 a. rosso the materiai described is deposited in the paleontological museum of the istituto policattedra di oceanoiogia e paleoecologia, caranía university (ipop museum, in the following). geological and palaeoecological setting. the messina strait area (between sicily and calabria) offers exceptional opportunities for the study of pleistocene deep-sea communities. the plio-quaternary sedimentation in this area has been strongly affected by tectonics which split the substrate into several "tectonicsedimentary entities", each evolving independently from the neighbouring sectors (barrier, 1987). yertical displacements, occasionally exceeding 1,000 m, allowed the deposition of deep-sea sediments and their subsequenr upjift to 200-400 m above the present sea ievel (barrier et ai., l986; barrier & montenat, 1987). bathyal palaeoenvironments are often recorded perfectly by fossils and contain in situ palaeocommunities belonging to both hard and soft bottoms which have been referred to the deep-sea corals (cap) and the bathyal mud (vp) palaeobiocoenoses sensu pérès & picard (tlo+) and di geronimo (1,987). the lazzà.ro section belongs to the motta s. giovanni tectonic-sedimentary ení.ify, bordered landwards by a nsí-se fault system (barrier et a1., x986). the section comprises basal tortonian sandstones overlain by five sedimentary units, outlined by submarine rruncations. except for the topmost unit, the sequence was deposited in circalittoral to epibathyal palaeoenvironmenrs fig. 1 location of studied samples. triang)es: submerged, lare pleistocene thanatocoenoses; circles: plio-pleisrocene ourcfops. indicating a sudden uplift to infra-circalittoral depths. epibathyal sediments, mainly marls, were deposited during a cold or cold-temperate phase of the eariy pleistocene (violanti, 1988) or, partly, during the middle pleistocene (aifa et al., 1987). marls containing embedded blocks colonised by deep-sea, hard-bottom species were deposited in a nearby fault palaeoescarpment, ca. 600 m deep. fossil assemblages are rich and diverse, consisting mainly of scieractinians, gorgonians, molluscs, serpulids and bryozoans. they comprise mud-dwelling taxa and species living on hard substrata (boulders) andlor coarse biogenic bottoms around the boulders (di geronimo er ai., 1996). bryozoans consist mainly of slender erect species. setoselliniform species, parricularly adapted to colonise sand grains are common, whereas encrusting species, represented mainly by runners, are scarce. dominant species are "lèssaradoma boreale (busk), gemellipora eburnea smitt, reteporella sparteli (calvet), caberea ligata juliien, euginoma vermrformis jullien, setosellina roulei caivet, hornera lichenoides (linnaeus), anguisia uerrucosa jullien and a new species belonging to the genus tèruia (described below). the section of archi belongs to the reggio recronic-sedimentary entíty, controlled by a n4oo-5oo fault system (barrier, 1986). sediments comprise rransgressive, shaliow-water deposits evolving into bathyal marls overlain by deltaic sands and gravels. bathyal beds contain boulders encrusted by deep-sea scleractinians and serpuiids. marl deposition occurred in a cold phase of the upper part of the early pleistocene-middle pleistocene as demonstrated by foraminifers and nannofaunas (di geronimo et ai., 1997). fossil assemblages indicate a strong atlantic affinity and a bathyal palaeoenvironment 500-1,000 m deep. benthic faunas consist mainly of molluscs. bryozoans are usually rare but common and well diversified in the coarsest (silty or sandy) samples. slender erect, both rigid and flexible, often rhizoid-bearing species and setoselliniform taxa prevail. encrusting species are rare. tessaradoma boreale (busk), zrvia irregulaùs meneghini, hornera frondiculata lamouroux, anguisia oerrucosa jullien, setosellina rowlei lullien and setosella oulnerata (busk), together with a ne'w species belonging to heliodorna (desurbed below) are the most common components of these assemblages. the furnari outcrop is located in the "furnari block" recognised by kezirian (1992). this fault-controlled entity is formed by a miocene substrate overlain by shelf calcarenites and silts. a small n-s directed graben is located along the northern edge of the block. deepsea faunas encrust fault escarpments and the basin is filied by grey bathyal marls deposited during the early pleistocene as indicated by foraminifers. benthic assemblages originate from a deep-sea muddy bottom community living îear a fault paleoescarpment as demonstrated by the presence of sessile species needing a hard bottom or) at least, coarse biogenic substrates interspersed in the bathyal muds. fossils consist mainly of molluscs with deep-sea mud-dwelling species and subordinate sessile bivalves (see di geronimo & la perna, !997b). bryozoans are common but poorly diversified. they consist mainly of cemented, erect-rigid morphotypes with subordinate rhizoid-bearing erect-rigid and flexible specimens. a single setoselliniform species (described below) is presenr. tèreia inegularis (lr4eneghini), tèssaradoma boreale (busk), homera lichenoides (linnaeus) and anguisia oerrucosa lullien are the dominant species. the eoclimm samples, studied incompletely by di geronimo et al. (1995) originate from bathyal (ca. 300-1,500 m deep), muddy to silty-muddy bottoms rich in planktonic gastropod and foraminiferan tests. benthic assemblages are dominated by deep-sea molluscs, mainly nuculoids and small mud-dwelling species. bryoznans are rare and quite absent from several samples. they comprise a few, mainly erect species among whìch an' guisia aerrwcosa jullien and reteporella sparteli (calvet) prevail. the bs samples come from bathyal (ca. 500-1700 m) muddy bottoms, including blocks originating from rocky scarps above. benthic assemblages consist mainly of molluscs (deep-sea nuculoids, pectinids and arcids); isidid gorgonians and brachiopods are locally abundant" bryozoans are subordinate but usually diverse. erect-rigid specimens prevail, mainly belonging to tèssaradoma boreale (busk), palmicellaria cf . elegans alder and jaculina cf . tessellata hayward. systematic accounts class s t e n o i a e m at a borg, 1926 order cyclostomatida busk, 1852 family crisiidae johnston, 1838 genus crisia lamoroux, 1812 pleístocene deep-sea bryozoans frorn sicily 425 crisia tenella calvet, 1906 longinodata ssp. nov. (pì. 1, fig. 6-10) 1995 lcrisia tmella,moissette er spleldnaes, pl. i, fig. 5-6. 7998 crisia sp.1, rosso & di geronimo, tab. 1, fig. a, i-2. 1997 crisia sp.1, di geronimo et al., tab. 3. material. lazzà,ro marls: over 100 internodes or fragments from 3 samples, collected in the basal, l-ower pleistocene pan of the marls; archi section: a single internode from the topmost sample (middle pleistocene); eocumm campaigns: few specimens from three samples ranging from 485 to 750 m depth (ll-ate pleistocene); bs 78 campaign: few specimens from 1,2931,7a2 m depth pate pleistocene). etymology. longinodata from longus : long and nodatus : with nodes, i.e. characterised by long internodes. types. holotype: a fenile internode (pl. 1, figs. 8-10) from the basal part of the lazzàro marls (boulder 1993). paratypes: 32 sterile internodes and a single fenile one from the same sample. ipop museum, ipop. 82. 26.1.t995. diagnosis. slender, long internodes comprising up tò ten autozooids, characterized by a very acute basal angle. description. very slender, poorly calcified, biserial internodes (pl" 1, figs. 6, z). basal width of internodes small; basal angle (o) very acute (10-13'); 2"d zooid long (l2 : 600-845prm; n:10). each internode shows o-2 basis rami. gonozooid sub-erect (p1. 1, figs. 8-10) consisting of an apex-down cone with a convex base, rising frontally and obliquely ar an angle of about 50o to the internode (lgon : 39opm, lgon : 260pm; n:2). ooeciostome (pl. 1, figs. 8-10) sub-terminal and dorsai; it is a short flared tube antero-distally compressed and facing upward. remarks. these fossil fragments agree rather well with c. tenella as described by harmelin (1990) but differ in some morphometric characteristics such as the smaller gonozooid dimensions and the general shortening of l2. moreover, internodes are longer and often comprise 8-9 zooids vs. 3, (range 1-7: harmeiin, 1990). these features are constant in the fossil specimens studied. therefore, the designation of a subspecific taxon is proposed for these specimens. distribution. the subspecres c. tenella longinodata is presently known only as a fossil from the mediterranean area where it lived in both western and eastern basins. c. tenella longinodata was found in sediments cropping out in the messina strait zone (southern ltaly) and in submerged sediments of the se tyrrhenian sea. few additional specimens come from the northern tyrrhenian sea, off sardinia. the specimens from the plio-pleistocene deep-water sediments of rhodes) recorded as c. tenella by moissette & spjeidnaes (1995) seem to belong to this new taxon. the stratigraphic distribution ranges from the upper part of the late pliocenelower part of the early pleistocene (moissette a spjeldnaes, t995) to late pleistocene (submerged wúrmian sediments from tîrrhenian sed. this is a 426 a. rosso deep-sea species as it 'was recorded from sediments deposited in depths ranging from ca. 4oo to 1,200 m. bathymetric and geographic distributions partially overlap those of the nominal species, known from bathyal bottoms in the ne atlantic (calvet, 1906a; harmelin & d'hondt, 1982; 1992; flarmelin, 1990), from epi-bathyal bottoms of the alboran sea (harmelin & d'hondt, 1992) and from deep-circalittoral botroms off marseille (harmelin, 1928). family t e r a i i d a e canu and bassler, 1920 genus 7èrztia i:ullien, 7882 tervia barrieri sd. nov. (p1. 1, fis. l-sj l998 teroia sp.1, rosso & di geronimo, tab. 1, fig. 4, 4. 1.997 tèroia sp.1, di geronimo et al., tab.3. material. lazzà,ro marls: over 4oo fragments from 8 samples from the basal pan of the marls (hwer pleistocene). archi section: over 40 specimens from 4 samples of the middle pan of the section (middle pleistocene) where fragments are more abundant rn coarser (silty-sandy) sediments. eocumm campaigns: a few fragments from 3 samples from 300-360 m depth (late pleistocene); bs ll campaign: over 100 specimens írom718-742 m depth (late pleistocene). etymology. name in honour of the french geologist pascal barrier who actively studied the plio-pleistocene palaeoenvironmental evolution of the messina strarr. types. holotype: from the pleistocene of lazzàro (lazzà,ro cava antica: pl. 1, figs. 2, 5). paratypes: ca. 2oo sterile and fenile branches from the same sample. ipop museum, ipop b3. 26.7.1995. diagnosis. very slender branches with 1-3 zooids per rransverse series. small, sac-shaped gonozooid with flared ooeciostome. description: colony vinculariform, dichotomous, with slender, sometimes curved branches (p1. 1, fig. 1), round in cross section. zooids arranged in alternating transverse series with peristomes facing frontally and laterally; each series with 2 tubes (range 1-3); the outermost the longest (pl. 1, fig. 1). sometimes tubes isolated in the middle line between the series. inner spines absent (p1. 1, fig. 4). gonozooid (pl. 1, fig. 2-3, 5) dorsaily placed, sac-shaped, short (length once or twice the vridth), suddenly swollen in its proximal part and reaching its maximum in the mid-distal region. gonozooidal wall with transversally growing grooves and densely spaced, round pseudopores. ooeciostome (pl. 1, fig. 2, 5) terminal, placed against the dorsal wall of the branch; it consists o{ a very short, transversally elongated, sli ghtly flared tube facing distally. remarks. a single tèraia specres is known presently from the mediterranean, t irregularis (meneghini). it lives in lower circalittoral to bathyal environmenrs (harmelin, i976;harmelin & d'hondt, 1982) and has a long (eocene to recent) and wide (.ùtestern europe from holland to spain and italy) distribution. t banieri differs from specimens of t irregwlarls showing a feeble calcification. it presents a more slender appearance due to the few number of zooids per series, a smaller and a shorter gonozooid and a flared ooeciostoma. inner spines, regarded by harmelin (1976; 1977) as diagnostic for t irregularis, are lacking. another similar species t superba, was described by jullien (1ss2) on a few sterile specimens from bathyal bottoms (896 m) in the biscay gulf (ne atlantic). harmelin (1977), although with reseryations, referred some specimens from the concepcion banc, canary island (200 m) to this specres, previously considered by harmelin (1976) as a deep morphotype of t irregularis. he separated t superba taking into account the absence of inner spines and the characters of the gonozooid described as "une vésicule renflée, assez courte, dont la partie proximale tubulaire est visible et comparable à celle d'un autozoide. looeciostome est un tube court terminal qui s'ouvre en direction distale au niveau de la zone de croissance de la branche". thus, the pleistocene specimens show affiniti es fo t superba: zoarial morphology, a few tubes per series, and absence of inner spines. nevertheless, they differ from t superba as the gonozooid is longer and pear shaped and overall, has a differenr ooeciosrome consisting of a compressed and flared tube, artached to the dorsal side of the branch. such features are consrant in the fossil specimens and allow an easy recognition of the species. plate 1 scale bars: 1o0mm for figs. 4-5 and 8-11; 5o0mm for figs. 1-3, 6-7 and 12. fig.1-5 tèrvia barrieri sp. nov., lazzàro, early-middle pleistocene. 1) sterile, slender branch: liv 1 (1993);2,5) holotype (lazzàro, ancient quarry): 2) complete view of a fenile branch with a dorsally placed gonozooid; 5) detail of the swollen gonozooid and the ooeciostome, a short, slightly flared, proxìmal-distally compressed tube, medially located at its distal end; 3) fertile branch with a very shon, distally inflated gonozooid: liv. 1 (1993); 4) longitudinal section of two peristomes to shothe ebsence of inner spines: liv. 1 (1993). fig. 6-10 crisia tenella calvet longinodata ssp. nov., lazzà"ro: boulder 1993, early-middle pleistocene. 8-10) holotyp.; s; p". of a fertrle internode wìth a suddenly inflated gonozooid and its flared ooeciostome compressed in a proximal-distal fashion; 9 and 10) lateral close-ups of the gonozooid to show its semi-erect position and the distally located ooeciostomel 6 and 7) paratypes: slender, curued sterile internodes. fig. 11-12 -heliocloma dngusta sp. nov., furnari, early pleistocene. holotype. 11) detail of two zooid-avicularium couples with an ovicell (arrowed). note the very narrow, beaded, cryptocyst, the entire, smooth mural rim and the very reduced frontal pore of the ovicell; 12) ' ^^*^l't' -"r"'e colony with twenty-one zooids: note the ancestrula and the peculiar periancestrular budding pattern givrng rise to two clockwise spirals of zooids and the ovicell (arrow). pl. i plerstocene deep-sea bryozoans from sicily 427 428 a. rosso distribution. the species is presently known only as a fossil from the central mediterranean area. it was found in lower-to-middle pleistocene sedimenrs cropping out in the messina strait (rosso & di geronimo, 1998; di geronimo et a1., 1997) and in submerged wùrmian thanatocoenoses of the tyrrhenian sea. this species seems to be limited to bathyal bottoms from ca. 300 m (se tyrrhenian sea) to ca. 1,000 m (inferred depth of deposition of the archi fossiliferous layers). ciass gymnolaemata allman, 1856 order cheilostomatida busk, 1852 family setoseiiinidae haywardand cook, lgtg genus heliodoma calvet, 1906 heliodoma angusta sp. nov. (p1. 1, fis. 11-12) 1998 heliodoma sp.1, rosso & di geronimo, tab. 1, fig.5, 1. 1,998 heliodoma sp.1, di geronimo et al., tab. 3. pi.3, fig.2. material. furnari marls: 6 colonies (early pleistocene); lazzaro marls: over 20 colonies from coarse biogenic sands around the boulders (early pleistocene); archi section: over zo colonies from 9 samples from the base to the top of the section (early-to-middle pleisrocene). etymology. from its very narrow (angustu) cryprocysr. types. holotype: a complere fertile colony (p1. 1, fig. 11-12) from furnari marls (early pleistocene). paratypes: 1 entire and 4 incomplete colonies encrusting sandy lithic and biogenic (bivalve fragments and forams) grains from the same sample. ipop museum: ipop. b 4. 26:11995. diagnosis. heliodotna well characterised by an evenly narrow cryprocysr. description. roughly elliptical colonies, 1-2 mm in diameter (pl. 1, fig. i2). zooids with well developed gymnocyst and a narrow (ca. 15 pm), depressed cryptocyst bordering lateral and proximal sides of the opesium (p1. 1, fig. 11). mural rim smooth. ancestrula budding a distal avicularium (sensu hayward &. cook, 19) and a mid-lateral narrow and a distal-lateral, normal primary zooids (pl. 1, fig. l2). each of them gives rise to a separate, not dichotomous, clockwise, spiral of zooids. new zooids are laterally budded each with a distal-lateral avicularium so thar the two zooidal spirals are separated by an avicularial spiral. periancestrular zooids show compiete, beaded ciosures, with a median longitudinal umbo, which covers the enrire opesium except for the opercular zone. ovicell large, terminal, not prominent, with a small, subcentral foramen, sometimes very reduced (pl. 1, fig. 11-l2: arrowed), presumably closed by a large operculum. the zooidal size increases rapidly from the approximately rounded ancestrula (la: l7o185pm; ia 1.60-1751tm) and the first zoojd of each spiral ftg.2 heliodoma angusta sp. nov. schetch showing the peculiar colony budding pattern giving rise to the two clockwise spirals of zooid-avicularial couples. (l21,: 234ytm; iz7: 177ytm and lzl: 234yl"m;1zi: i95p,m) in the holotype, to the fifth-sixth zooid. size of zooids budded afterwards become constanr (lz: 380-41opm; iz:240-27}p.m). remarks. heliodoma trngusta is very similar to the recent deep-sea atlantic species h. irnplicata calver, 1906b, for colony organization and periancestrular budding pattern (fig. 2), typical of the genus (see hayward & cook, 1929). neverthless, the latter differs from fi. angusta for its wider, laterally extended cryprocyst (see prenant & bobin, l966; hayward & cook, 1979) glrng the opesium an elongated key-hole shape. ancestrula and zooids from the reperirion zone are consrantly longer rn h. implicata (ancestrula: 2ao-220 fm vs. i7o-i85 pm; zooids: 400-445 pm vs. 380-41opm) (see hayward & cook, 1979). moreover, measures stabilise earlier in astogeny, from about the 4'h zooid instead of the 6,h as ín h. angusta. distribution. h. angusta seems ro be restricted to the mediterranean early-to-middle pleistocene where it has been recorded exclusively from deep-sea sediments deposited in 500-1,ooo m depths. conclusions. the three newly described taxa are known exclusively from the e,arly-to-late pleistocene of the vestern mediterranean area, excepr for crisia tenella /onginodata which was recorded also from the late pliocene of the eastern mediterranean sea. furthermore, all have deepsea counterparts in the present day atlantic ocean, whereas only one (c. tenella longinodata) has a probable deep-circalittoral to bathyal mediterranean relative: the nominal species. furthermore, the new taxa belong to palaeocommunities showing strong atlantic affinities indicating the psycrosphaeric conditions of the mediterranean during the pleistocene. c. tenella longinodata, t barrieri and h. angustd could therefore represent cold stenothermic extinct taxa endemic to the mediterranean area, and are potential indicators of palaeoclimatic and stratigraphic conditions (see rosso 8c di geronimo, 1 ee 8). pleistocene deep-sea bryozoans from sicily references 429 acknoaledgements. thanks are due to dr. j.-g. harmelin (statron marine d'endoume, marseille) who mrde rvrihble deep-sea recent specimens and for heipful suggestions; prof . di geronirno (ist. policattedra di oceanologia e paieoecologia, catania) for usefr.l dìscussions; sig. o. torrisi (cnr, catania) and g. dafnis (stazione zoologtca, a. dhorn, napoli) for sem assistance. paper financrally supported by m.u.r.s.t. brents (rosso 60o/") " aifa t., barrier p., feinberg h.,pozzr j.-p. 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"($ �3 3 ��8e8� � )) &+!"&#' !'1 &%&"!"&#' #. .&//&#'� "$!+0 ,(#+*$#'# #,"# s5!"($'!$"()*$!/3 /���� +��� ��� �� �?�:b� �&1 &',"#'3 �(/",!"( d3 3 ��8e8� � �/#"*($%! ) !"(!5 .&//&#' "$!+0 !,(/ #. *-1$!"(1 , !// /*!$1/ .$#% /& &+&+ "()*$! 4(1/3 0� �) , ����� -��� #���� 8?� ��b���:� %/"($1!%3 $������%& ��' (����! �� �)� � �� $� ������ :8 rivista italiana di paleontologia e stratigrafia volume tuj numero 2 pagíne 259-262 settembre 1997 a ne\t find of hea/tisyntrachelus (cetacea, delphinidae) from piacenzian sediments of rio stramonte (northern apennines, italy) giovanni bianucci key-uords: cetacea, delphinidae, systematics, piiocene, northern apennines, italy. riassunto. viene descritto un cranio incompleto di hemisyntachelus sp. (cetacea, delphinidae) scoperto nelle marne argillose ptacenziane di rio stramonte (provincia di piacenza). questo nuovo ritrovamento conferma l'alta frequenza, nel pliocene, di hemisyntrachel u s nell' are a periadriatica. abstract. an incomplete skull of hemisynnachelus sp. (cetacea, delphinidae) is described from the piacenzian clayey marls of rio stramonte (piacenza, italy). the new find confirms the abundance of henisyntrachelus in the peri-adr)atrc area during the pliocene. introduction. in the spring of 1.991. mr. migliorini discovered a fossil delphinid skull preserved in pliocene clayey maris that crop out near rio stramonte, a small stream in the northern apennines qracenza, italy) (fig. 1). the specimen was collected by the gruppo paleontologico "la xenophora" of castell'arquato (a village near the discovery locality) and following restoration, is now kept in the geological museum of castell'arquato. the pliocene sequence of rio stramonte has been referred to the piacenzian (monegatti ec ranieri, 1e87). the holotype of hernisyntracbelus cortesii (fischer, 1,829) and another almost complete skeleton of this species were found in the same locality in the past (cortesi, 1819; cuvier, 1823; del prato, 1.897; cígala fulgosi, 1990). more generally, several significant cetacean fossils, including two almost complete skeietons of balenopterid, have been discovered in the northern apennines of pracenza province, in the same facies known in the literature as "argille azzvrre"(see francou, 1985). another probable balenopterid was recently excavated (1986) in this area by the gruppo paleontologico "la xenophora". this specimen, retrieved from lower pliocene sediments of monte oliveto, also is preserved in the geological museum of castell'arquato. systematic description class mammalia linnaeus, 1758 order cetacea brisson, l762 suborder odontoceti fiower, 1762 family d e i p h i n i d a e gray, 7825 genus hemisyntrachelws @randt, 1873) type-species: h. cortesii (fischea 1829) hemisyntrachelus sp. (fis.2-3) the specimen consists of an incomplete skull (acking the rostrum, part of the left side of the neurocranium, the right antorbital process and the hamular processes of the pterygoid$. the auditory bones, the mandible, the teeth and all of the postcranial skeleton are not preserved. the skull size is larger than that of contemporary tursiops, but more closely resembling that of hemisyntracbelus, a genus recently re-described and referred to the family deiphinidae (bianucci, 1996, 1997). the wide exposirion of the frontals, in dorsal vieq suggests an immature animal, like the specimen described by sacco (1893). the left dorsal portion of the braincase is iost and an endocranial cast is exposed. in lateral view, the temporal fossa is relatively large and is antero-posteriorly elongated as in tursiops and hemisyntrachelws, while differing from the round-shaped fossa of the condipanimento di scienze della terra, via s. maria, 53,56126 pisa (italy) 260 g. bianucci fig. 1 geographic location of hemisyntracbelzs sp. described. temporaneous genus astadelphis bianucci, 7996. the orbit is placed relatively anteriorly, as in hemiryntrachelus, and the zygomatìc process of the squamosal is anteroposteriorly elongated. among the living genera, this process more closely resembles that of pseudorca than the short process of tursiops. the skull size, the shape of the temporal fossa, the advancement of the orbit and the elongation of the zygomatic process of the squamosal indicate that the specimen belongs to the genus hemisyntracbelus. a specific attribution is not possible since the rostrum and the mandible, defining characters of the two species h. cortesii and h. pisanus are lacking in this specimen. vidth of rostrum at base vidth of premaxillae at base of rostrum greatest oostorbital width 180 106 320 tab. 1 heznislntachelzs sp. estimated measurements (in mm). discussion. the discovery of a new specimen of. hemisyntrachelus confirms the reiative abundance of this fossil genus in the pliocene of the peri-adriatic area. in fact, !1. of 16 significant delphinid records are referred to this genus (bianucci, 1996, 1997). partícularly in the emiiia piacenzian delphinidae hemisyntrachelzs sp. rio stramonte (northern apennines, italy), fr, frontal; mx, maxilla; n, nasal; occ, occipital; pmx, premaxilla 261 piacenzian. lncomplete skull in dorsal view. ec, endocranial cast; scale : 5 cm. fio 2 f;o 1 hemisyntracbehr sp. rio stramonte (nonhern fr, frontal; mx, maxilh; occ, occipitalr pop, squamosal.scale:5cm. apennines, italy), piacenzian. neurocranium of the incomplete skull in lateral view. postorbital process; sq, squamosal; tf, temporal fossa; zp, zigomatic process of the area (provinces of piacenza, parma, and bologna) all well-preserved delphinid records (z specimens) belong to hemisyntrachelus. the abundance of this genus might be related to its feeding habits (hemisyntrachelus was probably a generalist and opportunistic predator) and the restricted width of the peri-adriatic basin. this last condition might not have favoured the diffusion of open sea delphinids such as stenella, that instead inhabited the paleo-thyrrenian sea. achnouledgements. i thank dr. p miculan for information concerning the new cetacean finds and for his assistance during my visit to the geological museum of castell'arquato. i also wish to thank the director of this museum, dr. c. francou, for granting me access to the fossil cetacean collection. i am grateful to prof. v. landini (dipartimento di scienze della terra, university of pisa) for useful discussions and suggestions. i am also indebted to dr. d. j. bohaska (\washington) for critically reading the manuscripr. the illustrations are by the author and the photos by mr. m. gini (dipanimento di scienze della terra, university of pisa) and the author. 262 g. biznucci references bianucci g. (1996) the odontoceti (mammalia, cetacea) del prato a. (1897) il thrsiops capellinii sacco del pliocene from italian pliocene. systematics and phylogenesis of piacentino. palaeont. ital.,v.3, pp. i-14, pisa. delphinidae. palaeont. ital.,písa (inpress). fischer j.b. (1829) synopsis mammalium. v. of 512 pp., bianucci g. (1.997) hemisyntrachelus cortesii (cetacea, delsumtibus j.g. cottae, stuttgart. phinidae) from pliocene sediments of campore quarry francou c. (1985) i cetacei del pliocene piacentino. v. off (salsomaggiore terme, italy). boll. soc. paleont. ital., v. 77 pp., arr'nrrn. prov., ptacenza. 36 (1,2), modena (in press). monegatti p. & ranieri g. (1987) osservazioni paleoecolocigala fulfosi f. (1990) predation (or possible scavenging) giche sulla sezione pliocenica di rio stramonte (piacenby a great white shark on an extinct species of bottleza). boll. acc. gioenia sci. nat., v. 20, pp. 2873a8,catanose dolphin in the italian pliocene. tèrt. res., v. l2, nia. pp. 1,7-36, leiden. sacco f. (1893) il delfino pliocenico di camerano casasco cortesi g. (1819) saggi geologici degli stati di parma e pia(asdgiana). mem. soc. ital sci., '.9, pp. 1-15, napoli. cenza.y. of x + 165 pp., maino, ptacenza. cuvier g. (1823) recherches sur les ossements fossiies, où i'on rétablit les caradères de plusieurs animaux dont les révolution du globe ont détruit les espèces. tome cinquième, i partie. v. of 405 pp., g. dufour & e. d'ocaqnelibr.. paris. received decetnber 11, 1996; accepted april 24, 1997. rivista italiana di paleontologia e stratigrafia i i i 0"r,""--;--;ilicembreleee l nota breve short note kogia pusilla from the middle pliocene of tuscany (italy) and a phylogenetic analysis of the family kogiidae (odontocetr, cetacea) giovanni bianucci 't e \ialter landini '!'t receired. januart 22, 1999; accepted juq 27, 1999 key-oords: cetacea, kogiidae, systematics, phylogeny, middle pliocene, tuscany, italy. riasswnto. viene preso in esrme un crrnio quesi completo di cetaceo odontoceto provenienre dai sedimenti del pliocene medio di monte voltra.io (provincia di pisa, toscana). tale esernplare, in pessato erroneamente attribuito alla famiglia ziphiidae e descntto come olotipo della specre hyperoodon pusillus, viene qui riferito al genere kogla (farniglia kogiidae). la specie kogia pusiila è ridescritta e messe a confronto con le specie atruali k. brniceps e k. simus. da un punto di vista filogenetico, viene ipotízzart una separazione antica (ahneno nel miocene inferiore) dei kogiidae e physeteridae (physeteroidea). il mancato ritrovamento di reperti attribuibili con sicurezza alla famiglia dei kogiidae in sedimenti piìr antichi del miocene superiore è probabilmente dovuto alla raritàl di quesri cetecei. lanalisi delle relazioni filetiche tra i kogiidae è stata condotta prendendo in esame supposte apornorfie relative alla morfologia e/o estensione del bacino sopracraniale che alloggia le sacche aeree e l'organo dello spermaceti. abs*act. a partial skull of an odontocete cetacean from middle pliocene sediments of monte voltraio (pisa province, tuscany, italy) is exanrined. f'his fossil, erroneously referred to the fanily zrphitdte and described in the past as holotype of the species hyperoodon pusil/as, is assigned here to the genus kogia (family kogiidae). the species kogia pusilla is redescribed and compared to the living species k. breoiceps rnd k. simus. phylogenericalìy, an old separation (at least in the lower miocene) of kogiidae and physeteridae is suggested. the lack of substantiated kogiid records until the upper miocene is probably due to the rarity of these cetaceans. phyletic analysis within the kogiidae is undertaken and supposed apomorphies in the morphology andlor the extension of the supracranial basin that houses the large air sacs and the spermaceti organ are considered. lntroduction. the kogiidae are a family of small odontocetes (cetacea) similar and closely reiated to the extant sperm whale (physeteridae). !íell-preserved records of kogi"1,-1r l nulero 3 idae are very rare: only two almost complere skulls from late miocene sediments of mexico (barnes, 1973,1984) and peru (muizon, 1988) have been described previously. those specimens are the holotl'pe oi rhe extincr genera and species praekogia cedrosensis and scaphokogia cochlearis, respectively. other presumed records of kogiids are fragmentary and consist essentially of mandibular fragments (kel1ogg, 1929), isolated teeth (matsumuto,1927) and/or auditory bones (pilleri, 1986a, 1986b, 1987,1988; pilleri et al., 1989). the skull from the pliocene of tuscany described here represents the third known significant record of this family. this specimen s/as referred erroneously to the ziphiidae in the past. capellini (1893) assigned it to placoziphiws and pilleri (1987) described it as holotype of the species hyperoodon pusillus. this fossil sku1l, already referred to the genus kogia by bianucci (1997) and bianucci et ai. (1998), is redescribed in detail in this paper. kogia pusilla represents the ancesror of kogia breviceps and k. simus, the only two living species of this family. systematic description. class mammalia linnaeus, 1258 order cetacea brisson, 1762 suborder odontoceti flower, 1867 superfamily p hys e t e r o i de a (gray,1821,) g111,1.872 family k o g i i d a e (gill, 1871) mlller, 1923 genus kogia gray, l846 'rmuseo di storia naturale e del territorio, via roma, 103, 5601 1 calci, pisa (italy). e-mail: bianucci@dst.unipi.it 'r''t dipartimento di scienze della terra, via s. maria, 53,56126 pisa (italy). e-mail: landini@dst.unipi.it 446 g. bianwcci f: w land.ini kogia pusilla (pilleri, 1982) (fig.1-aa) 1893 placoziphizs, v beneden capellini, pp. 287-288. 1987 hyperoodon pusillus pilleri, pp. 35-36, fig. 11, pl. 13. 1997 kogia pusilla (pillerí) bianucci, pp. 172-1,73, tig. 6b. 1998 kogia p wsilla (p rllerl) bianucci et al., pp. 1,24, 126, 127, íigs. 3t1,22 . emended diagnosis of species. a species of the genus kogla that differs from k simus and k. brmices in having a more elongated rostrum, a less anteriorly extended antorbital processr a smaller lacrimal, a slighrly more prounonced dorsal asymmerry and a lesser elevation of the posterior region of the neurocranium. it differs from !r"?:"r*ot in irs sn'raller size and in having a more narrow sagittal holotype. igf1540v distorted and incomplete skull, lacking part of the right side and of the ventral portion of the braincase. auditory bones, teeth, rnandible and postcranial skeleton are not presewed. the specirnen s.as donated by lawley in l877 to the museo di paleontologia of the university of florence, where it is preseroed today. type locality. la rocca locality in monte voltraio hill, near volterre, province of pisa, tuscany (italy). age. middle pliocene: globorotalia crassaformis crassaformis subzone of iaccarino el salvatorini (1982), approximately between 3 m-a and 2.6 ma (bianucci et al., 1998). description. the condylobasal iength of the only known but incomplete skull is estimated to be about 270 mnr. this skull size is similar ro rhar ol kogia simus and it is slightly smaller than that of an adult of kogia bredceps. in fact, according ro ross (1979), the condylobasal length of 24 skulls of kogia simws range between 201 mm and 323 mm whiie the condylobasal length of 22 skulls of k. breviceps range berween 237 mm and 467 mm (less than 300 mm are young animals). compared to the living species, the rostrum has a simiìar triangular shape from a dorsal view; but differs in being rnore elongate (fig. a). moreover, it has a welldeveloped dorsal concrrvity and a venrral convexiry rhar u.'e harve not observed in extant species. post morten-r deformation may have accenruared these last characters in the fossil specimen. the cerebral skull has a sernicircular outline in dorsal view. in this respecr the skull is similar to thar of k. breoiceps and k simus but differs from the anreroposteriorly elongated skulls of both praekogia and scaphokogia. nevertheless, this skull differs from those of extant kogia spp. in having a more pronounced dorsal asyrnmetry (due to the displacement of the external nares tosrard the left side and to the displacement of the sagitt:ìl crest tor,'ard the right side), and in hrving smaller elevation of the posterior region of the neurocranrum. the first of these two fearures may have been emphasized by dost mortem deformation of the skull. in dorsal view (figs. 1,a1, la2) the premaxillae, the maxillae and probably rhe vonìer (covered by marrix) extend as far as the apex of the rosrrum, as in living species. a simiiar exrension of the rosrral bones was observed also in scaphokogia by muizon (1988). muizon (1991) considered this feature an apomorphy of the kogiidae. two asymmetricai superior nares are present at the base of the rostrum as in all physeteroidea (physeteridae and kogirdae). in particular, the left naris is 14 mm wide and 10 mm long whiie the right is 6 mm wide and 10 mm long. the nasals are missing, as in the other kogiidae. the ieft premaxilla exrends posteriorly as far as the postero-dorsal margin of the skull and its medial margin joins the right maxilla ro form a very promlnenr sagittal crest, as in living species. this crest is bent left in its anterior porrion. it is delirnited by vertical walls and is very narros/ ar the verrex (width: 13.3 mm), as in k. simus. the width of the sagirtal crest is a diagnostic character for the distinction of the two living specles (handley, 1966). ross (1979) observed that in 26 skulls of k. simus the minimum wrdth of this crest ranges from 5 to 20.5 mm while in 22 skulls ol k. breaiceps this n-idth ranges from 14.5 to 49 n-rm (from 23 to 49 mm excluding five young specimens). the right premaxilla forms a small elevated basin on the neurocranium, extending from the sagirr.ll crest on the left and to its lateral margin on rhe right. a sìrrilar depression is present in the living species of kogia where the spermaceri organ (kernan & shulte, 1918; raven & gregory, 1933; schenkkan ec purves, 1973) resides. in contrasr, in praekogia, rhe lareral rìlargin of the right premaxilla is not protuberant and the r.igbr maxilla and the right prerl-raxilla forrl a single craniai fossa that probably resulted in a more developed spernaceri organ rhan n kogill. scaphokogia shows an even more dissimilar dorsal cranill architecture than kogia.in fact, the skull of scaphokogta h:rs no sagitt:r1 cresr ar all (the facial crest is displaced near rhe left m:.rrgin of the neurocraniunr) rnd con\eqlrenrly it is likely rlr.rr the spermaceti organ also invaded the left side of cerebral skull (muizon, 1988). the antorbital notcl-res penerrate the supracranial basin as in irving species o[ kogia and in praeleogia,but nor in scapbokogia and in the other physeteroidea. this character is considered by muizon (1988) to be an apomorphy of the kogiinae (kogia and praelectgia) re1:rted to the large antero-lateral development of the supracrxnial basin. kogta pusilla (pilleri, 1982) holotype; la rocca (lgf1540v). a1,a2, dorsal view; b1,b2, vcntral view sagittal crest; vo, vomer. iocaìity, monte voltraio (province of pisa, -l'uscar-ry). almosr complete skull cf, cranial fossa; fr, fronral; l;r, lacrirnal; mx, maxilla; n, naris; pn-x, prerlaxillae; sc, fig. 1 kogia pusilla from middle pliocene of tuscany 448 g. bianucci & v/. landini 10 cm the antorbital process is narrower than in living species and does not extend anteriorly on the proximal portion of the rostrum. as a consequence, the cranial f ossae (antero-laterally delimited by the antorbital processes) are less developed antero-posteriorly in this fossil species. the anterior portion of left cranial fossa is almost completely covered dorsally by the lateral and sagittal crests, both being bent towards the fossa. posteriorly, the cranial fossae are slightly more elevated than in k brear.ceps. this feature makes k. pwsilla very different from k. simws, in which the posterior part of the craniai fossae rises strongly (handley, 1966; ross, 1979). the iateral view (figs. 2ay 2a2) shows a larger antero-posterior exposure of the rostrum than in the living species. this condition is due to both the greater elongation of the rostrum and the smaller anterior extension of the rrntorbital processes (in k. brer:iceps and k. sìwws the antorbital processes cover the proximal part of the rostrum) . the anterior portion of the lacrimal is smaller than in the living species and is comma-shaped as in the physeteridae, whereas it is triangular in the other kogiidae. there is no evident lacrimal-maxilla suture. figs" 2a2 and 3 present the probabie course of this suture: the postero-dorsal process of the lacrimal is large and posterifig.2 kogta pusilla (pilieri, 1982) holotype; la rocca localits monte voltraio (province of pisa,'iuscanv). almost conplete s kull (i gf 15 40v). a1,a2, laterel view; b1,b2, posterior view. fr, frontal; la, . lacrimal; mx, naxilla; oc, occipital; sc, sagittai crest. orly exìended, to a greater extent than in the other kogiidae. muizon (1988) noted that the posrero-dorsal process of the lacrimal is not present in the physeteridae except in a young specimen of awlophyseúer, and the presence of this process in the kogiidae is considered as a plesiomorphic character. the supraorbital process is thinner than in both iiving species and praekogia. the lateral view shows also an elevation of the posterior portion of the cranium, smaller than rn living species and in praebogta. in posterior view (figs. 2bi.2b') the sagirral crest is high, narro'w and delimited laterally by vertical wal1s. in ventral view (figs. 1b1, 1b2) the skull is badly damaged and most of the ventral surface of the neurocraniun is not preserved. ln particular, only the left frontal, the left lacrimal and a small porrion of the left squamosal are preserved. the medial groove of the rostrum allows the vomer to be observed. along the right preserved margin of the rostrum is a sma1l deep furrow but without distinct alveo1i. in this respect k. pusilla is similar to the living species and differs îrom scaphokogia (with distinct but shallow alveoli). comparisons. a peculiar kogiid feature of the only skuli known of l{ogia pusilla is the lack of both nasals. the presence o[ a supracranial basin and the strong asymmetry of the skul1 are other characrers typical of all the physeterida (kogiidae and physeteridae). kogia pusilla is similar to k. breoiceps and k simus in the peculiar outline of the skull in dorsal view (the cranium is semicircular and the rostrum is triangular), and in having a vertical sagittal cresr on the cranium (fig. a). kogia pusilla from middle pliocene of tuscany 449 fig. 3 kogia .pusilla (pilleri, 1982) holotype; la rocca localitv, monte voltraio (province of prsa, tuscany). neurocranium ìn lateral view of skul.l (igf1540v). fq fronral; l.a, lacrimal; mx, rnaxilla; oc, occrpital; porp, posrorbital p roce \s i \q. : qur mo 'eì; r f. temporal fossa. moreover, as rhe extant species, k. pwsilla shows a srnall basin on the cerebral porrion of the right premaxilla that extends as far as the postero-dorsal margin of the skull. k. pusilla is more similar to k. simus than k. breuiccps in the relatively small size of the skull and in the narrow sagittal crest at the vertex. fig. 4 comparisons of the skull in dorsal view of: a, kogia pusilla (restoration of the skull based on holotype, igf 1540v); b, kogia brniceps (laboratoire de anatomie comparée de paris, 1833-483); c, kogia simus (accademia dei fisiocritici di siena). ,1 cm, 450 g. bianucci g \y landint k. pwsilla differs from both k. brersiceps and k. simws in its longer rostrum, smaller lacrimal and perhaps grear.er asymmetry of its skull in dorsal view. phylogenetic relationships. the genus kogia shows evident affinities with physeter. common apomorphies of these ts/o genera concern particularly the facial anatomy (heyning, 1989) and the concave shape of the dorsal surface of the skull (scaphidiomorphy) for housing the large air sacs and the spermaceti organ. consequently, the singie bones of the skull suff ered large modifications, in parricular, a marked asymmetry and the reduction of the right external nares. even if there is consensus on the close affinities of kogia and physeter, precise systematic relationships between these two genera are debated. some authors (caldwell & caldwell, 1989; fraser & purves, 1960; physeteridae quaternary 1.8 ma handley, 1966;heyning, 1989; simpson, 1945) consider that kogia and physeter belong to the same family physeteridae. others (barnes, 1,973; kaxrya, 1.973; miller, 1.923; muizon, 1991) think these rwo genera have such distinctive characters that their separation in two families (kogiidae and physeteridae) is justified. this iast hypothesis was strengrhened by recenr studies on cetacean mitochondrial dna (arnason et al., 1.993; milinkovitch et al.,1.994) that confirm a large divergence between physeter and kogia. we believe that the fossil and phylogeneric evidence reveal that the separation of these two groups may be relatively old. in particular, all fossil skulls of physeteridae show a marked enlargement of the posterior portion of the right premaxilla, an apomorphy absent in kogia and in fossil taxa relared to this genus. as observed by muizon (1991) this feature is relatively less emphasized in living pbyseter than in fossil physeterids. this derived character is also present in the most ancient known skull of physeterid, assigned to diaphorocetus kogia pusilla kogiidae pliocene miocene oligocene fig. 5 phylogeny of the kogiidae. dotted lines indicate no fossil or fragmentary record. kogia pusilla from mid,dle pliocene of tuscany 451 k. breviceps k. simus k. pusilla praekogia scaphokogia diaphorocetus squalodon fig. 6 maxinun-parsimony cladogram (cosistency index : 100) conputed from the data in tab. 1 using henning 86, version 1.5 (farris, 1988). powcheti (early miocene from argentine; moreno, 1892; lydekker,1894) the primitive state of kogiidae, regarding the shape of right premaxillae, suggesrs rheir origin is no younger than the beginning of rhe miocene (fig. 5). the lack of of kogiid records prior ro the late miocene may be due to their low frequency in the ancient sea. flowewer, some isolated auditory bones and reerh from the lower miocene of piedmont (north italy) and ba1tringhen (austria) and middle miocene of visiano (north italy) were referred ro this family by piileri (1986b, 1988) and pilleri et al. (1989). nevertheless, because these elements (periotics and teeth) are rhe only remains of kogiids found in the sediments, their conclusion must be viewed with caution. as in kogra, some periotics show an articular surface for the tympanic flat and parallel to the general plane of the periotic and not ventrally oriented; nevertheless, on the whole rhey differ substantially from kogia periotics and show also affinities with those of some fossil physeterids. the distinction of the two families on rhe basis of rhese structures is problematic, considering also that skulls of fossil physeterid and kogiid genera with preserved auditory bones are rare. the isolated teerh are referred to miohogia elongata (emended of miokogia elongatum), a genus and species created on rhe basis of three teeth from balringen (piileri, 1986b). in fact, all teeth referred to this presumed kogiid are similar in shape and size to those of the physeterid scaldicetws bellunensls frorn the early miocene of belluno (north italy) (dal piaz, 1977).in accordance with orher authors (barnes, 1976; muizon, 1988) we attribure a 1ow systematic value to isolated teeth and therefore consider nr.tmina dubia borh senus and species names of miokogia elongata. considering other described kogiid fossils, only two skulls were referred correctly to this family. one, from the late mìocene of mexico (barnes, 1973,1984), was named praekogia ced.rosensis, and the other, from the late miocene of peru (muizon, 1988) was named scaphokogia cochlearis. praekogia and scaphokogia share wrth kogia the loss of nasal bones, an apomorphy of the family kogiidae. nevertheless, the rwo fossil genera differ from kogta by having a different architecture of the supracranial basin. in parricular, in praekogia the basin [or the spermaceri orgrn exrend\ ro rhe entire right portion of the dorsal surface of the cerebral skull that is formed by the premaxilla and maxilla. as observed by barnes (1.973), this genus lacks a distinct right maxillary fossa that is present in kogia. the basin for the spermaceti organ in scaphokogia takes up rhe entire dorsal surface of neurocranium and only a large cranial fossa is present. because of this peculiar shape of the supracranial basin and other peculiar characrers, scapholeogia was assigned to the subfamrly scaphokogiinae by muizon (1988). apparently scaphokogia shows affinities with pbyseter in having a large cranial basin for the spermaceri organ. nevertheless, rhe cranial basìn of scapbokogta does not extend on the rostrum as in pbyseter. a cranial basin more anteriorlyextended than rn scapholeogia is present also in kogia and praekogia, both having the dorsal depression that exceeds the external nares. moretaxon character i 2 7 + 5 6 7 8 9 121110 l3 i4 siua,lodon diaphorocetus 0 0 1 0 1 2 2 2 2 2 00 ; t;ui ,t>:0 00 -00 f-00 0 00 00 , : o _f t01 r01 t01 0 0 0 i. 1 1 000 ò o-nnln oo:i ll110 111 11. 1 ;--t--" ^-^uu olo ::0 00 0 ll 0 scaphobogia 0 praekogia 1 1 1 1 k"4o p"tr\ kogia simus kogia breoiceps 1 1 1 data matrix used in the cladistic analysis. 't= missing characters. characrer states: 1) supracranial basin: o, no; 1, yes; 2) strong rsymmetry: 0, no; 1, yes; 3) lack of nasals: 0, two nasalsl 1, one nasal; 2, nasals absent; 4) vomes premaxillae and maxillae reach the apex of rostrum; 5) enlargement of posterior portion of right premaxillae: o, no; 1, yes; 6) antorbital notch penetrating rhe cranial fossae: o, no; 1, yes, 7) supracranial basin formed by only one hemispherical depression posteriorly developed and anteriorly closed (see muizon, 1988, 1.991.,for more detail on the bone modifications): 0, no; 1, yes; 8) craniurn half-circle shaped: o, no, 1, yesl 9) rostrum triangular: 0, no; 1, yes; 10) small basin on the cerebral portion of right premaxilla: o, no; 1, yes; 11) very prominent sagittal crest: o, no; 1, yes; 12) very large antorbital process and lacrimal; 0, no; 1, yes; 13) rostrum verv shorr: 0, no; 1, yes; 14) relarively broad sagittal crest: 0, no; 1, yes. tab. 1 452 g.bianucci&wlandini over, in these last two genera, the cranial basin takes up the antorbital processes and consequently the antorbital notches penetrate the cranial fossae. this last character was considered as an apomorphy of the kogiinae (kogia and praekogia) by muizon (1988). considering interrelationships among the three species of the genus kogia, apparently k. brez,iceps and k. simws are more derived than k. pusilla in having more anterioriy extended antorbitai processes (and consequently more elongated cranial fossae) and a shorter rostrum. a parsimonious interpretation would suggest that the wide sagittal crest of k. brer.ticeps is an apomorphy, considering the primitive status of the narrow crest observed in k. pusilla and k. simus. the characters discussed belovr are used to create a maximum-parsimony cladogram of the relationships of kogia pwsilla using henning 86, version 1.5 (farris, 1988) (tab. 1, fig. 6). regarding other fossils referred to the family kogiidae in the past, we agree with barnes (1973) and muizon (1988) on the attribution of kogiopsis floridanus from the miocene of florida (kellogg, 1929) as a probable physeterid and in considering the name of the species kogia prisca (miocene?, from japan; matsumoto, llt1l/ ) a nomen au0tum. pilleri (1986a, 1982) identifies the living species kogia simus and k, breoiceps in the pliocene of tuscany on the basis of isolated teeth and periotics. \fe consider these fossils and other unpublished specimens as belonging to the kogiidae and most probably to the gentts kogia. nevertheless, we do not agree with pilleri in referring these f ossils to the two living species because the periotics show different characters and the teeth are not diagnostics at this systematic level. it is possible that most of these specimens belong to the fossil species kogia pusilla. fossil kogiids were reported recently from pliocene sediments of the yorktown formation (vhitmore, 1994) and northern appennines (cigala fulgosi, 1996), but they were not described in detail. conclusions. the assignation of the specimen examined here to the kogiidae confirms the presence of this farnily in the pliocene of tuscanl', already documented by pilleri (1986a) on the basis of isolared teeth and periotics. other auditory bones from these sediments are under study by the authors. k. pwsilla is a primitive species of the genus kogla that differs significantly from living k. breaiceps and k. simus in having a smaller antorbital process and a more elongated rostrum. the phyletic relationships between the kogiidae and the physeteridae were examined and an origin of the kogiidae older (at least early miocene) than that suggested by the paleontological data (late miocene) is suggesreq. in accordance with muizon (1988, 1991), the kogiid,ae are separated into two subfamilies on the basis of different architecture of the supracranial basin: the kogiinae (kogia and praekogia) and the scaphokogiinae (scaphokogia). the morphology of rhe supracranial basin of kogia pusilla is similar to that o[ k. breaiceps and k. simws eyen if the basin of the fossil species differs in having a smaller anterior exrension. acknouledgements. 'we are very grateful to the following persons for grrnting us access to cetacean collections and for their assistance during our vrsirs at the museums: p. mazza and f. cozzini (museo di geologia e paleontologia, università di firenze); d. robineau (laboratoire de anatomie comparée de paris); f. cancelli (accademia dei fisiocririci di siena). \7e also wish to thank c. de muizon (museurn natinal d'histoire naturelle de paris) for critically reading the rnanuscnpr. the photographic plates were prepared by the euthors and m. gini (dipartimento di scienze della terra, f inir.ersità di pisa) re,ferences a.r,rror ú., grllb..g a. 6c \x/idegren b. (1993) ceraceen mitochondrial dna control region: sequences of all extapt baleen whales and two sperm r.vhales..species. mol. biol. eool.,v. 10 (5), pp. 960-97a, chicago. barnes l.g. (1973) praekogia cedrosensis, a new genus and species of fossil pygmy sperrn whales from isla cedros, baja california, mexico. contr. sci., nat. hist. mus. los angeles c., v. 247, pp. 1-20, los angeles. barnes l.g. (1976) outline of eastern noth pacific fossil cetacean assemblages. syst. zool., v. 25 (a), pp.321-343, new haven. barnes l.g. (1984) fossil odontocetes (mammalia: cetacea) from the almejas formation, isla cedros, mexico. paleobios,v. 42, pp. 1-46, berkeley. bianucci g. (1997) the odontoceti (mammalia, cetacea) from italian pliocene. the ziphiidac. palaeont. ital., v. 84, pp. 1,63-192, pisa. bianucci g., sarti g., catanzariti r. & santini u. (1998) middle pliocene cetaceans from monte voltraio (tuscany, itaìy). biostratigraphical, paieoecological and paleoclimatic observations. rio. ital. pal. strat., v 104, pp. 1.23-13a, milano. capellini g. (1893) nuovi resti di ztftoidi in calabria e in toscana. atti. r. accad. lincei, rend., v. 2 (7), pp. 283288, roma. cigala fulgosi f. 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(1960) hearing in cetaceans. evolution of the accessory air sacs and rhe structure and function of the outer and middle ear in recent cetaceans. bull. britisb mus. (nat. hist.), zool., v. 7 (1), pp. l-140. l.rndon. iaccarino s. & salvatorini, g. (1982) a framework of planktonic foraminiferal biostratigraphy for early miocene ro late pliocene mediterranean area. paleont. strat. eaol.. v. 2, pp. 115-125, roma. handley c.o. jr. (1,966) a synopsis of the genus kogla (pyg-y sperm whales). in norris k.s. (ed.) \whales, dolphins, and porpoises. univ. calif. press, pp. 62-69, berkeley and los angeles. heyning j.e. (1989) comparative facial anatomy of beaked whales (ziphiidae) and a systematic revision among the families of extant odontoceti. contr. sci., ndt. hist. mus. los angeles c., v. 405, pp. 1-64, los angeles. kasuya t. (1973) systematic considerations of recent toothed whales based on the morphology of rympanoperiotic bone. scl. rep. whal. res. inst., v 25, pp. 1-103, tokyo. kellogg r. (1929) a new fossil toothed whale from florida. amer. mws. nooit., v. 389, pp. 1-10, newyork. kernanj.d. 8r shulte h. von\il (1918) memoranda upon rhe anatomy of the respiratory tract, foregut, and thoracic viscera of a foetal kogia breviceps. bull. am. mus. nat. hist., v. 38, pp. 231,-267, new york. lydekker r. (1894) contributions to a knowledge of the fossil vertebrates of argentina. pan ii. cetacean skuìls frorn patagonia. ann. mus. la plata, v. 2, pp. 1-14, buenos aires. matsumuto h. (1927) on some fossil cetaceans of japan. scl. repx. toh. imp. unio., 2nd ser. (geol.), v. 10 (1), pp. 17-27, sendai. milinkovitch m.c., meyer a. ec powell j.r. (1994) phylogeney of all major groups of cetaceans based on dna sequences from three mitochondrial genes. mol. biol. eool., v. 1 1 (6), pp. 936-948, chicago. miller g.s. jr. (1923) the telescoping of the cetacean skull. smiths. misc. coll.,v.76 (5), pp. 1.-71, baltimore. moreno f.p (1892) lijeros apuntes sobre dos géneros de cetóceos fósiles de la república argentina. rer.t. mus. la plata, v. 3, pp. 393-400, buenos aires. muizon c. de (1988) les vertébrés de ìa formation pisco (pérou). troisième partie: les odontocètes (cetacea, mammalia) du miocène. inst. fr. etud. andine.r, mem. 78. pp. l-244. paris. muizon c. de (1991) a new ziphudae (cetacer) from the early miocene of \vashington srate (usa) and phylogenetic analysis of the major groups of odontocetes. bull. mus. ndtn. hist. nat.,v. 12 (3-4'5, pp.279-326,pa,ris. pilleri g. (1986a) pygroy sperm whales (kogia) in the italian pliocene. inuest. on cetacea, v 18, pp. 133-154, vammala pilleri g. (1986b) the taxonomic status of "shizodelphis elongatus" : miokogia elongatus (probst, 1bb6) (cetacea, physeteridae). inaest. on cetacea, v. 18, pp. 155-162, vammala. pilleri g. (1987) the cetacea of the italian pliocene with a descriptive catalogue of the species in the florence museum of paleontology. v of 1eo pp., printed by vam-.r.v:..:.-.:î1, vammala.lrrdrrrr r\tt.,dpdrrtu \ pilleri g. (1988) preand post-messinian cetacea in the mediterranean tethys and the messinian "salinity crisis". in pilleri, g. (ed.) contributions to the paleontology of sorne tethyan cetacea and sirenia, pp. 19-4a, printed by vammalan kirjapaino oy, vammala. pilleri g., m. gihr & c kraus (1989) odontoceti (mammalia: cetacea) from the lower miocene of rosignano, piedmont, north italy. inrtest. on cetacea, v. 22, pp. 1,89-291, vammala. raven h.c. & gregory \(k. (1933) the spermaceti orgen and nasal passages of the sperm wl-raie (physeter catodon) and other odontocetes. amer. mus. nc,,oit., v. 677, pp. 1-18, new york. ross g.j.b. (1979) records of pygrny and dwarf sperm whales, genus kogia, frorn southern africa, with biological notes and some comparisons. ann, cape proo. mus. (nat. hìst.),v. 11 (14), pp.259-327, grahamstown. simpson g.g. (1945) the principles of classification and a classification of mammals. bull. am. mus. nat. hisr., v. 85, pp. v-xvi, 1-150, new york. schenkkan e.j. & purves pe. (1973) the compertative anaromv of the nrsal rract and rhe [unction of rhe sner-_"_'" " '/ màceti organ in rhe physeteridae (mamrralia, odontocetr). bijdragen tot de dierkwnde, v. 43, pp. 93-112, amsterdan. \lhitmore f.c.jr. (199a) neogene climatic change and emergence of the modern whale fauna of the norrh atlantic ocean. in berta a. & deméré t.a. (eds.) contributions in marine mammals paleontology honoring frank c. \ú/hitmore jr. proc. san diego soc. nat. hist., v. 29, pp. 233-227, san diego, california. rivista italiana di paleontologia e stratigrafia paleomagnetic and palynologic investigations in the upper valdarno basin (central italy): calibration of an early villafranchian fauna andrea albianelli, augusto azzafioli, adele beriini, giovanni ficcarelli, giovanni napoleone e. danilo torre key-uords: magnetic stratigraphn palynologn middle pliocene,early villafranchian, upper valdarno, central italy. riassunto. la fauna a vertebrati rinvenuta nella pane inferiore dei depositi del 1o ciclo fluviolacustre del bacino del valdarno superiore (successione di castelnuovo dei sabbioni) è stata attribuita all'unità faunistica triversa del villafranchiano inferiore. i più antichi giacimenti dell'europa occidentale, come layna (spagna), suggeriscono un'età della base del villafranchiano non più antica di 3.6 ma. in italia i'u.f. triversa è stata datata tra 3.0 e 3.3 ma. le indagini paleomagnetiche hanno permesso il riconoscimento della porzione superiore delf intervallo a polarità normale gauss e della prima breve inversione sottostante (k.aena); in quest'ultima è contenuto il tetto delle ligniti basali e dei primi livelli argillosi sovrastanti dove furono raccolti i resti di mammiferi attribuiti all'u.f. triversa. lassociazione a mammiferi è quindi databile tra 3.10 e 3.17 ma. la base della successione conserva la testimonianza palinologica di una vegetazione indicativa di un clima caldo-umido a carattere subtropicaìe, che evolve verso condizioni più fresche con progressiva rìduzione degli elementi tropicali-subtropicali. nel loro insieme le assocìazioni vegetazionali ricostruite possono essere correlate con quelle del reuveriano dell'europa centrale. in prossimità del tetto del corpo sabbioso che chiude il ciclo di castelnuovo. in località "rena bianca", si hanno evidenze di una fase marcatamente arida seguita da condizioni più calde ed unide. quest'ultimo cambiamento porrebbe corrispondere al passaggio pretigliano-tigliano oppure a una fluttuazione stadiale-interstadiale del pretigliano. sulla base dei dati raccolti, quindi, si può indicare per l'inizio del vìllafranchiano un'età più antica di 3.1 ma. abstract. the silty clays embedding an early villafranchian mammal fauna of the triversa faunal unit (f.u.) have been paleomagnetically and palynologically studied in a continuous sequence exposed in the santa barbara quarry. the study has allowed to date the earliest occurrence in ltaly of a faunal assemblage of this unit and to define the corresponding climatic conditions. the sampled section has provided a màgnetic polarity sequence of the late gauss, where the fossiliferous layer fits the kaena reversed interval. its age of ca. 3.1 ma, during subtropical climate conditions correlatable to the reuverian of netherlands, suggests an older age for the beginning of the villafranchian, possibly assocìated to a more dramatic scenario able to trigger the faunal turnover. introduction. the beginning of the villafranchian was darcd bet.ween 3.0 ^nd 3.6 ma on the basis of radiometric and paleomagnetic analyses carried out in sites with most significant local faunas. in spain the layna local fauna (province of soria) is characterized by a well differentiated association in which the occurrence of leptobos sp. and mimomrys cappettai, alongside ruscinian species, lends the fauna a character transitional towards the villafranchian (perez and soria, 1992). the sediments in which rhe fauna was collected show normal polarity magnefiz fion most plausibly assigned to the gauss chron and therefore the deposit cannot be older than 3.6 ma, according to the magnetic time scale of baksi (1,993).in the gaudíx-baza basin, where a complete continental sequence from latest pliocene to middle pleistocene is presenr, the paleomagnetic survey has also placed the beginning of villafranchian in the gauss chron, after the 2ln.2r chron (agusti et a1.,1995). in france, the vialette fossil bearing level, characterrzed by a typical early villafranchian association (triversa f.u. : mn16a), overlies a volcanic bed and is probably older than the azanieras lava flow. according to the k/ar dates of the volcanic rocks (bander et al., 1978) the fauna is no older than 3.3 ma, but older than 2.6 ma. still in france, the well known les etouaires fossil site of early villafranchian age, was dated between 3.3 and 2.6ma by couthure 6. pasrre (1983), on minerals collected from the fossil bearing deposit. ly et al. (1983) proposed an age of 2.6-2.5 ma, on exrrapolation from the grande nappe volcanic event, in the frame of a generai geological reconstruction of the history of the area. savage and curtis (197a) reported an age of 3.4-3.5 ma from a sediment rich in volcanic elements underdipanimento di scienze della terra, università di firenze, via la pira 4,5ad1 firenze, italy ;ae rùe = 4ruvlal--_ o.r*yyr^o"r^"""f t "" :/ o i palustrine' lacustrine rf{" ,.,delt o ::w 9!-!1,9: o 6 ó :j-._qe-l!!ll fluvialw sw n.e' tt2 fio i schematic stratigraphy and environment interpretation of the valdarno basin fi1l. modified after sagri et al. (1994). lying the fossil bearing level. the age proposed by ly et al. (1983) appears anomalous in comparison with other datings and deserves deeper inquiry. in ltaly, the poggio mirteto lignite-bearing clays (near rome), with scanty fossils referred to the triversa f.u., are heteropic of marine sediments of the globoroulía crassaformis zone, in which a voicanic ash level gave a reversed polarity magnetization, in all likelihood corresponding to the mammoth subchron. in the type locality of the triversa f.u. (boano & forno, 1994), lindsay et al. (1980) made a palaeomagnetic survey in the section of rdb brick factory (san martino formation). they found two reversed intervals separated by a normal polarity zone, and interpreted the reversed intervals as kaena and mammoth subchron, respectively. the short vertical extent of the section and the low quality of magnetic response left their interpretation uncertain. the majority of datings reported above indicates a most probable age for the oldest villafranchian faunal unit (triversa f.u. : mn16a) of 3.0-3.3 ma. in order to improve our understanding of the chronology and climatic evolution, a paleomagnetic and palynologic study was carried out in the sediments of the oldest fluviolacustrine cycle of the upper valdarno (castelnuovo dei sabbioni sequence). the sequence is well exposed in the castelnuovo dei sabbioni area, on the left bank of the arno river (se of florence). the deposits consist (sagri et al., 1994) of basal aiiuvial and fluvio-deltaic sands and gravels (spedalino sands and gravels), grading upwards into well-bedded lacustrine silty clays and sands (meleto clay$. lignite seams occur in the lower part of the lacustrine deposits and have been exploited industrially in the santa barbara mine. the lacustrine deposits are conformably overlain by more coarse-grained deltaic sands and well sorrcd, light coloured fluvial sands (san donato sand$ (fig. t). the main lignite seam, and the silt beds overlying it, yielded a scanty mammalian fauna represented ficcarelli, g. napoleone & d. torre irm acquisition curves from two extreme samples, labeled in met€rs, starting from the base of the 13 m thick lignite seam. sample n. 2 is from the core, sample 148 from the upper outcrop: in the most anoxic deposits the low magnetic coercivity behavior does not differ from that fixed in less reducing environment. by [jrsws minimws, thpirus arvernensis, dicerorhinus sp., lcptobos sp. and anancus araernensis. from the same deposits, but from an unknown locality, came a molar tooth of zygolophodon borsoni. this association is typical of the triversa f.u. (torre et al., 1996). magnetic investigations sampling site. a section of the meleto clays, in the open santa barbara pit, has been chosen for the paleomagnetic analyses. along an outcrop of gently dipping layers, the siity clays, including the lignite, were sampled up to the base of san donato sands for 150 meters. a reference level was fixed at a sandy layer 0.6 m thick, 17 m above the lignite seam. the sampling extended downwards for more than 14 m, as close as possible to the lignite bed, since the mine was being filled and the section is now covered almost completely. during the filling a borehole 62 meters long was drilled alongside the sampled outcrop; it passed the lignite and penetrated the underlying sandstones; another 18 m long core penetrated the sandy ievel and reached the top of lignite. from the outcrop and the two cores, cubes of about 1.0 cc 'were prepared for the magnetic measurements. these were carried out at the eth magnetic laboratory in zirich. magnetic properties. the quaiity of the magnetic character of the rock type has been enhanced by measuring the isothermal remanent magnetization (irm), for several samples taken at different stratigraphic levels, and the susceptibility of all samples at each srcp during the natural remanent magnetization (i.irm) thermal treatment. sample 2 lvlmd = 0.325e-01 a/m sample 149 mmax = 0.573é+00 a,h 0 0.00 | 00 a/m j_o examples of irm patterns are shown in fig. 2. sample 148 comes from the cliff and sample 2 from the core, in a layer near the bottom of the lignite. in both cases the acquisition of irm is detected in fields up ro 1 t (tesla) and more than 90o/o of it steeply grows within 0.2 t; the saturation is practically attained ar 0.4 t. the nrm thermal demagnetízation curves show how a relatively strong remanence decays slightly up to 100oc, and more severely beyond this: at 250'c almost 90o/o of it is iost, and at 300oc actually nothing is left (fig 3). the lower diagram of sample 43 is the common curve for samples magnerrzed with a normal polarity; ail samples collected from 25 m above the top of lignite going upward are normally magnefized. the vector directions belong to a single magnetic phase and point straightforward to the origin aiready at 150oc along the characteristic direction of the primary magnetization. samples in and close to the lignite (e.g., sample 25) show a sudden increase berween 300 and 350oc, with a higher intensity ascribed to a new mineralogical phase impiying that magnetite is produced. a possible alteration of clay minerals with temperature could produce moderate increases while the abruptly strong contribution to susceptibility can be ascribed to alteration of sulfides, as these features are more pronounced in the levels closer to the lignite (albianellí, 1994; albianelli er aj,., 1995). in the present materials sulfides are not macroscopically evident and only diffraction anaiysis, made on separates above 63 microns in diameter, has shown traces of pyrrhotite and pyrite. i-ow coercivity and rapid decay of magnetízation, at increasing temperatures, are characteristic of such minerals. this is supported by the behavior in fig. 4, that shows the demagnetízatíon path for a sample taken in the clay levels still close to the lignite and associated to the changes in susceptibility occurring at around 300-350"c. thereafter a strong increase in susceptibility occurs and the directions become unstable. the low temperature trends are interpreted as reprecalibration of an early villafranchian fauna í/|1mor mimo 4 k/ko 3 2 1 fig. 3 two typical demagnetization curues (geographic coordinates) of opposite patterns. in sample 43 (30 m above the lignite) the intensity of normal polarity is progressively de^.,.:-^ --r .l-j00oc ir va_l4/rrré alu arlcr nishes, while in sample 25, f-^l--.^-^ ^-:il:lrom rayers sîil1 rn proxrmlty of the lignite and reversely magnetized, it undergoes a srrdden irrmn rfrer j0ooc as occurring in the anoxic lignite samples. sentative of the primary magnefíz tion, according to the results discussed by albianelli (1,994) and albianelli et ai. (1993;1,995), carrred by magnetite and authigenic sulfides produced during and/or immediately after deposition. the magnetic polarity stratigraphy has been interpreted accordingly (fig. 5). sample 27 t1,3 wj-z sampìe 43 sample 27 ko = 10.65'10e-05 sl unils mo= 2.26'195-95 ort o susceptibility o inlensity r ("c) 500 600 fig. 4 the demagnetization curves for a sample still in the reverse interval shows the removal of a secondary component with normal pol.rrity. in rhe graph on rhe left. up to 200oc treatment. the primary magnetization, with stable direction, slightly decays up to 3o0oc. the successrte rncrease depicts the growth of a new magnetic phase, as funherly remarked by the susceptibility jump after 350oc. 400300200100 0.000 i 0 a/h n/fm!x límox:0.469e-02 a/m n; -z s; +z a. albianelli, a. azzaroli, a bertini, g. ficcarelli, g. napoleone g d. tone magnetic stratigraphy, the whole set of samples shows directions of the paleomagnetic vectors trending consistently from the initial steps of demagnetization. as discussed above, the measured polarities throughout the section yield a reliable reversal sequence. even near the bottom, where the lignite seam has evidence for more reducing conditions, directions remain consistent (albianelli et al., 1993). santa barbara stereooraohic distribution n normal magnetic polarity occurs for more than 1.00 meters in the higher portion of the outcropping beds. in the lower section a reversed polarity takes place near the lignite, followed downwards by a normal one. the composite of fig. 6 contains all the data produced by the overlapping sequences from the two boreholes and the surface section; in its lower portion the composite section extends down to the sandstone basement and the reversed interval is located to cover the upper half of the lignite seam and several meters above it. in the layers close to the top of lignite and just above it, fossils of the mid-pliocene triversa faunal unit were recovered during 1957 -1958 quarrying. the succeeding long normal polarity, persisting through several hundred ka of sedimentation, can be interpreted as belonging to the upper gauss, subchron c2an.1n; this, in addition, can be supported by the fact that it follows a short reversal (fig. 5-6). the latter, therefore, well defined within the two normals, is interpreted as c2an.1r. the presence of the fossil level assigned to the triversa f.u. in a well defined position, within the kaena subchron (c2an.1r), offers firm biostratigraphic constraint and yields an age between 3.10-3.17 ma, according to baksi's scale (1993). palynology sampling and methods. the castelnuovo succession was investigated for palynological analyses since 1990, in a section 2 km distant from that for paleomagnetics. the meleto clays were sampled along a well exposed and continous section in the allori pit of santa barbara mine. in the overlying san donato sands the few silty-clay levels intercalated within sands were sampled in the rena bianca qrrarry. on the whole, 380 samples were collected (bertini,1994). laboratory techniques involve both physical and chemical procedures (hydrochloric acid, hydrofluoric acid, luber method, sodium esametaphosphate, zínc chloride of density 2, potassium hydroxide plus sieving, ultrasound sieving) in order to isolate palynomorphs from other organic components and minerals. a minimum of +00 identified pollen grains were counted for each sample. taxa were organized in 11 groups on the basis of ecologic and climatic requirements of their modern equivalents and represented in a synthetic palynologic diagram (fig. 6). a list of groups is here reported as a caption to the figure: 1. tropical-subtropical elements (sapotaceae, clethraceae-cyrillaceae, etc.) and subtropical,/warm-temperate elements (taxodtum, nyssa, myrica, etc.) demandm normal reversed 29 359" 64' 1 760 -530 4.50 8.60 1 4.9 10.1 r lower hemisphere o upper hemisphere santa barbara 90 lm ctg s vgp lalitude and polarity 0 -90 lou 100 stereographic distribution (bedding corrected) of all selected samples used for build up the vinual geomagnetrc pole path in the meleto clays. the list on the right summarizes the number of samples (n) defining the normal and reversed magnetozone, the mean declination (dm) and inclination (im), the confidence cone (cr95), and the precision parameter (k). the polarity sequence is interpreted on the base of mammal remains located in the lignite seam and just above it, corresponding to the transition from the short reverse interval to the next normal (see also fig.6). circles denote samples from the outcrop, crosses from the co re. o cu o e (ú 'f (g .p^ calibration of an early villafrancbian fauna ll) ding year-round humidify and s/arm conditions which may also be related to local edaphic or/and microclimatic conditions. 2. deciduous elements of warm-temperate and temperate climate: carya, quercus, pterocarya, juglans, engelhardtia, liquidambax, ijlmaceae, etc. 3. cathaya and pinus haploxylon r.ype. 4. pinus and other pinaceae represented !y pooriy preserved grains. 5. mid-altitude conifers: cedrus and tiuga. ap+mp . fi:ffffffffffffi-i ortro .k.(\\\ \ 200 \ 1\\.r ,-\ì:\ q't_-ffif=re1 ei?-:f=-:-e-tr;t_l ?13 lt;zlfsff+f=:+fe-fl .ti .s.: .. f:+l:l;-ì;---q, r;: r'.,7:/7a \+++4 :_' lft i tnii/;777t4f4++4é -tf 6. high altitude elements: picea and abies plus betula and fagus. 7. and 8. taxa lacking climatically significance plus indeterminate and/or indeterminable pollen grains; in group i alnus and salix are evident. 9. mediterranean xerophltes: qwercus ilex type, olea, phillyrea, pistacia, eîc. 10. and ll.contain non-arboreal plants, prevalent herbs: poaceae, asteraceae, ericaceae, etc., including such hydrophilous taxa as nymphaeaceae, potamogetonaceae, sparganiaceae. steppe elements (artemisia and ephedra) are represented in group 11. pollen flora. a rich flora is recorded, composed of eiements nowadays living in different regions in europe, asia and america. the high concentration of weli preserved palynomorphs confirms the occurrence of anoxic conditions on the iake bottom. this also favoured the preservation of leaves, fruits and seeds, which are consisrently present all along the santa barbara section. in the sandy levels of the overlaying san donato sands, carpoflora is dominant whereas remains of leaves are sporadic and poorly preserved. here pollen grains are recovered only in the rare silty-clayey beds. in the overall pollen spectra of fig.6 forest elements sharply dominate over the herbaceous ones, with exception of one spectrum corresponding to the first viable bed of the san donato sands. the five spectra from the main lignite level show a very high percentage of arboreal poilen grains; pollen of taxodiaceae is dominant (taxodiwm/glyptostrobu). alnus pollen grains are occasionally highly abundanr, possibly because of strong fluviatile influx. myrica, llyssa, clethraceae-cyrill aceae, qu erc us, c ary a, e n ge lb ardtia, tilia, magnoliaceae, rhamnaceae, rhoiptelea, sapotaceae, syrnplocos, araiiaceae are also present. the four spectra between the main lignite level and the minor one show an increase of the pinaceae pollen grains percentage, especially of pinus, probably recording a change in the depositional environment. spectra belonging to the levels shortly above the secondary lignite seam do not differ significantiy from the underlying ones. the first of these samples brings a strong increase of catbaya whlle pinus pollen grains are noted; the second one shows a new increase of taxodiaceae and of thermophilous broadleaves pollen grains. spectra from the five uppermost levels of the meleto clays show a decrease of subtropical and warm-temperate elements; the mesophyious element cathaya increases, as well as the microthermal ones, particularly picea. macroflora studies reveal the orominent occurrenu) o (o u) a u) rféffii ì +! r ll,tll i a -- " -:t.tt-.) jjjj j-f i ic 1l tl (d co \ '\. composìte stratigraphic section of the whole casteinuovo dei sabbioni sequence (tickness in meters). the sampled section starts from the base of lignite. "f" indicates the fossiliferous 1evel. from left: magnetic polaritn lithostratigraphy and syntetic pollen diagran that displays the percentate of the pollen groups for which explanations are in tne text. tt) ; o i o fi. a a. albianelli, a. azzaroli, a. bertíni, g. ficcarelli, g. napoleone & d. torre ce of fagus in the same levels (roiron p., pers. comm. iee4). the upper san donato sands is dominated by pollen grains of steppe elements, mainly artemisia and a new increase of forest elements (particularly cathaya and carya) is recorded on its top. paleoenvironmental evolution. the preliminary palynologícal data evidence different vegetational phases which took place during the deposition of the castelnuovo succession. during the development of the peat bog, starting about 3.1 ma, the vegetation is dominated by forest elements such as taxodiaceae, nyssa, myrica, engelhardtia, while pinaceae probably had a restricted distribution. climatic conditions seem characterizedby low seasonality and by high humidity and temperature. a humid subtropical, 'warm-temperate ciimate may be suggested, simiiar to the climate now characterizing south-eastern asia. these conditions are tn agreement with what is known from literature about this period (suc et al., 1995a, 7995b). subsequently the basin underwent a rapid and strong subsidence, the water level rose and a deeper lacustrine environment developed. the pollen flora shows a progressive reduction of tropical-subtropical elements, particularly evident in pollen spectra from the uppermost meleto clays. here a decrease in temperaure is marked by the increase of highland elements (mainly picea), among pinaceae, and by the decrease of subtropical/warm-temperate elements. a forest vegetation, though, seems to have been still dominant. a significant environmental change is recorded during the deposition of the upper san donato sands, characterízed by episodes of ephemeral stream and terminal fans. the first spectrum, characterízed by herbaceous vegetation of steppe type, seems to reflect highly arid conditions. the pollen assemblages from the meleto clays, representative of a forest vegetation, can be correlated with the reuverian (zagwrjn, 1.974; suc and zagwljn, 1983) whereas those of steppe type from the san donato sands can be correlated with the praetiglian (zagwrjn,1960) and the first glacial cycles following the onset of the arctic glaciation dated at 2.6 ma. this event witnessed the development of tundra in northern europe (zagwrjn, 1960) and of steppe in the north-western mediterranean area (suc er al., 19954 1995b). in northern italy a strong increase of highland taxa, mainiy picea, is found in the sections of stirone (bertini ee vannucchi, 1993) and castell'arquato (lona, 1990), of the external apennines. in these sections the absence of a dominantly herbaceous pollen flora could be related to local environmental conditions, although it could have been also produced by unfavourable sedimentary features. in the uppermost part of the san donato sands a ne'w climatic amelioration, reflecting conditions of higher humidity and temperature (increase oí catbaya and carya), could be ascribed either to an interstadial of praetiglian or to the praetiglian-tiglian transition. conclusions the fossil mammals found in the castelnuovo dei sabbioni deposits identify the normal polarity of the paleomagnetic record from the meleto clays with the gauss chron c2an.1n. the continuity of the section, the high reliability of the magnetic signature and the sampling resolution place the local mammal assemblage within the kaena subchron (c2an.1r) dated as 3.1.0-3.17 ma. palynological analysis points to a warm-humid climate drifting progressively to cooler conditions that, near the top, produced a significant increase tn picea, abies and fagus. further upwards, at the top of the san donato sands, two closely spaced layers show two contrasting features, of a strongly arid climate followed by a warmer and humid one. pollen records for the meleto clays appear to have correlatives to the reuverian of the biochronologic scale of zagwtln (1960); those for the upper s. donato sands could indicate either the praetiglian and the transition from praetiglian to tiglian or a stadial-interstadial fluctuation within the praetiglian. new light is shed on the beginning of the villafranchian mammal age. the mammal association of the first sedimentary cycle in the upper valdarno developed in a warm-humid, subtropical climate. since it seems unlikely that the faunal turnover giving rise to the villafranchian fauna could occur in such a climate, the lower boundary of villafranchian preceded the kaena paleomagnetic event. a possible trigger for the turnover could be better identified in the cooling event picked by ò18o anomaly ar ca," 3.3 ma. achnouledgemmts. this work was supported by grants from ministry for scientific and technological research (murst) and national research council (cnr). the national power plant (enel) made available the cores and the facilities in the mine; in particular, dr. g. miotto and his staff kindly provided continuous field assistance. for the measurements at the eth magnetic laboratory and the complete availability of its facilities, useful discussions and advises, dr. f heller is deeply aknowledged. for palynologn many thanks are due to dr. j.p. suc for constructive discussions and to mrs. d. duzer for technical assistance at the laboratory of palynology in montpellier. calibration of an early villafranchian fauna references lt7 agusti j., oms o., garces m., dinares j., peres m. (1995) plio-pleistocene correlations between the mammalian chronology and the magnetic polarity time scale in southern spain. inqua, 14th int. congress, bedin, 314 aug. 1995 (abst.), p. 6, berlin. albianelli a. (1994) proprietà magnetiche e magnetostratigra{iche dei sedimenti anossici fluvio-lacustri e marini coevi. ph.d. thesis, univ. of florence, pp. 190, florenalbianelli a., bertini a., magi m., napoleone g., sagri m. (1995) i1 bacino plio-pleistocenico del valdarno superiore: eventi sedimentari, paleomagnetici e paleoclimatici. il quaternario, v.8, pp. 11-18,. roma. albianelli a., napoleone g., pompeo r. (1993) i1 ruolo dei solfuri nella magîetízzazione residua di sedimenti continentali e sedimenti marini. atti xii cono. gngts, pp. 873-884, roma. baksi a.k. (1993) a geomagnetic polarity time scale for the period 0-17 ma, based ot 40ar/39ar plateau ages for selected field reversals. geoplrys. res. lett., v. 20, pp. l607 -1,61,a, \flashington, d c. bandet y., donville b. ec michaux j. (1978) etude géologi que et géochronologique du site viilafranchien de vialette (puy-de-dóme). bull. soc. geol. france, xx (7) n. 3, pp. 245-251, paris. bertini a. (1,994) palynological investigations on upper neogene and l-ower pleistocene sections in central and northern italy. mem. soc. geol. it., v. 48, pp. 431-443, roma. bertini a. & vannucchi d. (1993) etude palynologique du pliocène de i'italie septentrionale: la section de stirone. xiii symposium ass. palynol. langue fran. (a.p.l.f.),. résumé, p. 33, besangon. boano p. & forno m.g. (1994) carta geologica della successione villafranchiana. dpt. earth sc., univ. of turin. couthure j. & pastre j.f. (1983) contribution à la chronostratigraphie du villafranchien: l'auvergne et le velay lfrance). une série de réíérence du plio-pléistocène européen. actes colloq. le villafranchien méditerranéen, 1982, pp. 1,79-185, liiie. lindsay 8.h., opdyke n.d., johnson n.m. (1980). pliocene dispersal of the horse equus and late cenozoic dispersal events. nature, v.287, pp. 135-138, london. iona f. (1990) late pliocene in northern ltaly. glacial events recognízed by palynological record. in: e. knobloch & z. kvaèek (eds.) proc. symp. paleoflor. paleoclim. changes in the cretaceous and tertrary, pp. 277z/ó. yrasrje. ly meng hout cantagrel j.m., de geer de herve a. & vincent p.m. (1983) révision tephrochronologiques des dép6ts fossilifères des environs de perrier et champeix (puy-de-dóme, france). actes colloq. le villafranchien méditerranéen, 1982, pp. 4a7-422, lrlle. perez b. & soria d. (1992) anfisis de las comunidades de mamíferos del plioceno de layna (soria) y la cùera (teruel). paleontologia i eoolució, v. 23 (1.989-1990), pp. 23 1-238, sabarell (barcelona). sagri m., martini i.p., benvenuti m. e{ magi m. (199a) basin fill architecture of the neogene-quaternary extensional basins in the northern apennines. xv ias regionaì meeting, ischia, pp. 41-74,ischìa. savage d.e. & curtis g.h. (1970) the villafranchian stageage and its radiometric dating. geol. soc. arn. spec. pap., v. 124, pp.2a7-231, boulder, co. suc j.-p., bertini a., comburieu-nebout n., diniz f., leroy s., russo-ermolli e., zheng 2., bessais e. and ferrier j. (1995a) srrucure of the \fest mediterranean vegeration and climate since 5.3 ml acta zool. cracoa., v. 38(1), pp. 3-16, krakow. suc j.-p., diniz f., leroy s., poumot c., bertini a., dupont l., clet m., bessais e., zheng 2., fauquette s., ferrier j. (1995b) zanclean (brunssumian) to lower piacenzian (lower-middle reuverian) climate from 4o to 54o north latitude (vest africa,'west europe and lvest mediterranean areas). meded. geol. sticht., v. 52, pp. 43-56. utrecht. suc j.-p. & zagwiln \7.h. (1983) plio-pleistocene correlations between the northwestern mediterranean region and northwestern europe according to recent biostratigraphyc and palaeoclimatic data. boreas, v. 12, pp. 1,53166, oslo. torre d., albianelli a., bertini a., ficcarelli g., masini e, napoleone g. (1,996) paleomagnetic calibration of plio-pleistocene mammal localities in central italy. acra lool. cracoo., v. 39, n. 1, pp. 559-57a, krakow. zagwln \í.h. (1960) aspects of the pliocene and early pleistocene vegetation in the netherlands. meded. geol. sticht., v. ciii-i, 5, 78 pp. utrecht. zagwrln \7.h. (1974) the piio-pleistocene boundary in western and southern europe. boreas, v. 3, pp. 75-97" oslo. receioed may 17, 1996; accepted october 24, 1996 fernandez 383..400 ��� ����� ��� ���� ��� ������� �� ���������������� ���� � � ��� �� �� ��������� ����� �� ��� � �� ���� �� �� � �� ���� �� ������ �� �� ���� � ���� � � ������ � �� � ��������� �� �� ���� ��������� ��������� �� ��� �� ������ �� !"#$%� �&%#$�'#"( )'*'$ !# * +&� *)"!$ ,'%"!� �"+!$� ,'% -.'!� �-%"#'!"'! ,'%"!� �'/ � !0$+ � 1$%#$.! �$#2&%� ��������� �$3$.'* #$!% 4 � 5$. ,'#2 !"'! ��������� 4. � �"+!$��'%#$**'!$ .$+" ! 6�� �.'!($7 '!0 �'/ � !0$+ '.$' 6� .#-� +'*7 2'3$ /$$! .$3"$5$0� �2.$$ %)$("$% 2'3$ /$$! 0"%#"!+-"%2$0 "! #2$ * 5$.� %# %-/8 !$ 4 #2$ "+8'+ !$ 6�'.3-� �-/8 !$7 9-%# '/ 3$ #2$ / -!0'.& ,'9 ("'! # ,'#2 !"'!� �� ��������� �#-.'!" :�;�<� �� �������� %)� ! 3� :�;�< '!0 �� ������������ %)� ! 3� :�;�<� �! #2$ ,'% -.'! '.$'� ' (2. ! 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"! � ! 3'! $# '*� �gf�k ,$%! % 3 �gf�k ,$%! % 3 � l-#-8 3' �gf�k �'!0 3'* �gf�k ,$%! % 3 (����)���" *�)��" ��� &����+��� ��&�� �����#� ,� !���� .��f� atlantic ocean medit erran ean sea 500 km 10º 0º 40º madrid digne bas auran spain morocco algeria france lisboa p o r t u g a l coimbra lyon cabo mondego torino italy �"+� � � �$ +.')2"('* * ('#" ! 4 #2$ %#-0& '.$'% "! �� �.'!($ '!0 � .#-+'*� stagelower bathonianupper bajocian “marno-calcaires à cancellophycus” “terres noires” fm. 0 4 5 0 2 1 0 2 3 0 2 5 0 2 7 0 2 9 0 3 3 0 3 5 0 3 7 0 3 9 0 4 3 0 1 3 0 1 5 0 1 1 0 0 9 9 468 1 0 2 0 0 5 5 0 0 7 7 0 0 1 0 8 3 0 8 5 0 8 9 0 9 1 0 9 3 3 0 2 9 2 3 2 4 2 5 2 7 2 0 1 9 1 8 1 7 1 6 1 5 1 4 1 3 1 2 11 2 2 3 4 3 5 3 3 3 2 3 1 0 9 5 0 9 7 0 9 9 1 0 1 1 0 3 1 0 5 1 0 7 1 0 9 1 1 1 1 1 3 0 7 3 0 7 5 0 7 7 0 8 1 0 7 1 0 6 9 0 6 7 0 6 5 0 6 3 0 5 9 0 5 3 0 5 7 0 4 7 0 4 9 0 5 1 rb level levels in sturani 1967 1 0 1 3 5 0 m thickness (m) ravin du bès section 0 0 3 3 1 zigzag zone aurigerus zone parvum subzone macrescens recinc. ten. parkinsoni zone bomfordi subzone ravin d’auran section ra level levels in sturani 1967 thickness (m) 0 4 5 0 2 1 0 2 3 0 2 5 0 2 9 0 3 3 0 3 5 0 3 7 0 3 1 0 3 9 0 4 3 0 1 7 0 1 9 0 0 9 4610 0 0 5 5 0 0 7 789 0 0 1 1 0 8 3 0 8 5 0 8 9 0 9 1 0 8 7 0 9 3 3 0 2 9 2 8 2 3 2 4 2 5 2 6 2 0 2 1 1 9 1 8 1 7 1 6 1 5 1 4 1 3 1 2 1 1 3 4 3 5 3 3 3 2 3 1 0 9 5 0 9 7 0 9 9 1 0 1 1 0 3 1 0 5 1 0 7 1 0 9 1 1 1 1 1 3 1 1 5 1 1 7 1 2 1 1 2 3 1 2 5 0 7 3 0 7 5 0 7 7 0 8 1 0 7 9 0 7 1 0 6 9 0 6 7 0 6 5 0 6 3 0 6 1 0 5 9 0 5 3 0 5 7 0 5 5 0 4 7 0 4 9 0 5 1 1 0 1 3 5 0 m 0 0 3 3 bigotites sp. [m+m] protozigzagiceras sp. [m+m] zigzagiceras sp. [m+m] franchia sp. [m+m] bigotites mondegoensis sp. nov. [m+m] limestone marlstone bigotites sturanii sp. nov. [m+m] bigotites diniensis sturani [m+m] �"+� � � �#.'#"+.')2"('* 0"%#."/-#" ! 4 ���������� !����"��"�������� +� ! 3�� (����%�� '!0 /��"�������� "! �'3"! 0? -.'! '!0 �'3"! 0,$a% %$(#" !% 6�.'!($7� �2$ .'!+$ *"!$ 4 ��������� %)� 0"%)*'&% #2$ ((-..$!($ 4 4.'+�$!#'.& %)$("�$!% 4 -!0$#$.�"!'/*$ %)$("$% '!0 #2$ %#.'#"+.')2"( ( !%#'!(& 4 #2"% +$!-% #2. -+2 #2$ %$(#" !� ," %#.'#"+.')2"( * +% 4. � �$.!'�!0$8�� �)$8 �ddh� � �"##' �ggc� �ggf� �ddd7� ���������� ��'�����%������ '!0 �������'�����%������ %2 5 .$*'#"3$*& %"�)*$ %-#-.$% '!0 �'& /$ '(#-'**& ( !%"0$.$0 '% .$).$%$!#'#"3$% 4 #2$ %-/4'�"*& ,"+ #"#"!'$ '!0 4'�"*& �$."%)2"!(#"0'$� ' +. -) 4 1$%# �$#2&'! �$."%)2"!(#"0'$ /.'!(2$0 44 /& ). #$. +$!$%"% 4. � *������%������ :�< � ,#����� ��%������ :�< 4 ( �)*$> %-#-.$%� 52"(2 ((-. "! #2$ ,'9 ("'! �'.'!#"'!' 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'# #2$ $!0 4 #2$ !# +$!"( 0$3$* )�$!#� �"� (. ( !(2% 5"#2 *'#$.'* *'))$#% %2 5 "% ( %#'#$ ."//"!+ 1�� ���# ���%����� ����������� ��� /��"������������ 2����������� ����#� 3�������4 �fc 1 1 1 3 5 7 9 13 15 17 23 3331 413 5 57 7 11 15 19 25 27 29 33 35 37 39 41 43 13 f b f c fd 50 10 15 95 101 107 115 123 133 137 143 153 159 165 173 183 195 199 209 221 0 5 10 15 cadomoceras aff. nepos parona macrescens subzoneparvum subzone zigzag zoneparkinsoni zone lower bathonianupper bajocian stage formation section 90 section 02 thickness (m) thickness (m) level level bigotites sp. [m+m] bigotites cf. diniensis sturani [m+m] bigotites mondegoensis sp. nov. [m+m] protozigzagiceras .g. nov. sp [m] zigzagiceras [m+m] sp. limestone marlstone cabo mondego formation �"+� � � �#.'#"+.')2"('* 0"%#."/-#" ! 4 ���������� !����"��"�������� +� ! 3� '!0 /��"�������� "! �'/ � !0$+ %$(#" !% 6� .#-+'*7� � 0"4"$0 '4#$. �$.!'�!0$8�� �)$8 $# '*� 6�ddj'7� -) # #2$ $!0 4 #2$ !# +$!"( 0$3$* )�$!#� '!0 ' � .$ ( �).$%%$0 %$(#" ! '% 5$**� �2$ �'#-."#& 4 #2$ %)$("� �$!% "% "!0"('#$0 /& #2$ -!( "*"!+ 4 #2$ -�/"*"('* %$'�� '%% ("'#$0 5"#2 '! "!(.$'%"!+ ( �).$%%" ! 4 #2$ 52 .* %$(#" !% "! #2$ �"(. 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(=% "! #2$ �"+!$�,'..$n�$ '.$' 6� -#2� �'%#$.! �.'!($� �$)#� *)$% 0$ �'-#$ �. 3$!($7� ��##� ���� !�#����� :��#�� �j� g���df� � 0$!'� 1$%#$.�'!! �� 6�gcj7 � �2&* +$!"$ 0$. �#$)2'! ($.'#'($'$ -!0 �$."%)2"!(#'($'$ 0$% � ++$.� 1� 3�� .��#� !� #�?����� ��%�� �d�� �����hg� �#-##+'.#� 1$#8$* 1� 6�g�h7 � �#-0"$! 8-. �'*'q !# * +"$ 0$% ! .0 � 5$%#$-. )'q"%(2$! ,'#2 !"$!� !�#����������%���� fh� �ch� �#-##+'.#� high-resolution calcareous nannofossil biostratigraphy across the toarcian oceanic anoxic event in northern italy: clues from the sogno and gajum cores (lombardy basin, southern alps) stefano visentin1 & elisabetta erba1* 1dipartimento di scienze della terra “ardito desio”, università degli studi di milano, via mangiagalli 34, 20133 milano, italy. *corresponding author. e-mail: elisabetta.erba@unimi.it. to cite this article: visentin s. & erba e. (2021) high-resolution calcareous nannofossil biostratigraphy across the toarcian oceanic anoxic event in northern italy: clues from the sogno and gajum cores (lombardy basin, southern alps). riv. it. paleontol. strat., 127(3): 539-556. rivista italiana di paleontologia e stratigrafia (research in paleontology and stratigraphy) vol. 127(3): 539-556. november 2021 abstract: calcareous nannofossil biostratigraphy was conducted across the toarcian oceanic anoxic event (toae) interval cored at colle di sogno and gajum in the lombardy basin (southern alps, northern italy). drilling at both sites resulted in 100% recovery of unweathered material. the sogno and gajum cores consist of pelagic marly limestones, marlstone, marly claystone, and a relatively expanded black shale interval named fish level considered the lithostratigraphic record of the t-oae at regional scale. semiquantitative analyses of calcareous nannofloras allowed to achieve a high-resolution biostratigraphy of the latest pliensbachian-early toarcian time interval. several nannofossil biohorizons were detected, including zonal/ subzonal markers and additional events related to changes in abundance. the nannofossil biostratigraphic correlation of the sogno and gajum cores indicates that, according to their paleogeographic settings, the succession recovered in the sogno core deposited on a pelagic plateau is continuous while a hiatus of ~600 kyrs was detected in the lowermost toarcian in the gajum core located on a slope of a structural high. the njt 5 and njt 6 zones of the standard nannofossil zonation for the mediterranean province were identified in both the sogno and gajum cores. our findings allow an implementation of the reference biozonation with the separation of the njt 6a and njt 6b subzones, and age revision of some secondary events. the zonation established for the lusitanian basin (portugal) is only partially reproducible in the lombardy basin, confirming nannoplankton paleoprovincialism during the early jurassic requiring different zonal schemes in various areas. nevertheless, we underline that the t-oae is unambiguously constrained by the fo of c. superbus crassus and the lo of m. jansae at supra-regional scale. received: january 14, 2021; accepted: june 25, 2021 keywords: calcareous nannofossils; biostratigraphy; toarcian oceanic anoxic event; lombardy basin. introduction the early toarcian oceanic anoxic event (toae) is recognized as one of the most severe and geographically widespread events of oceanic anoxia and organic-carbon burial in the mesozoic (jenkyns 1985, 1988, 2003, 2010). coeval paleoenvironmental perturbations have been linked to volcanism of the karoo-ferrar large igneous province (lip) and its release of volcanogenic co 2 , and/or thermogenic methane (ch 4 ) from sill intrusion into gondwanan coals, and/or biogenic methane from dissociation of sub-seafloor clathrates (hesselbo et al. 2000; kemp et al. 2005; mcelwain et al. 2005; svensen et al. 2007; jenkyns 2010; reolid et al. 2020). this dramatic episode of ecosystem adjustments, global warming and altered ocean chemistry occurred during a crucial time for calcareous nannoplankton diversification as a major speciation episode took place in the late pliensbachian-early toarcian time interval (bown 1987; mattioli & erba 1999; bown visentin s. & erba e.540 et al. 2004; erba 2004, 2006; fraguas & young 2011; menini et al. 2019). new genera and species appeared and quickly evolved allowing a high-resolution biostratigraphy also across the t-oae (mattioli & erba 1999; mattioli et al. 2004; casellato & erba 2015; ferreira et al. 2019). moreover, major changes in abundance of some taxa were proved coeval in the late pliensbachian-early toarcian time interval and probably related to large scale paleoenvironmental stress preceding and across the t-oae (erba 2004; mattioli et al. 2004, 2008; tremolada et al. 2005; fraguas et al. 2012; casellato & erba 2015; clémence et al. 2015; menini et al. 2019; visentin et al. 2021b). calcareous nannofossils were proved to be extremely useful for high-resolution biostratigraphic dating and correlations at low, medium, and high latitudes. two standard biozonations are available for the early jurassic: the one of bown (1987) revised by bown & cooper (1998) established for the boreal realm (united kingdom, germany, the netherlands, central and northern france) and the zonal scheme of mattioli & erba (1999) specific for the tethyan area (italy, southern and eastern spain, south france, hungary, greece). more recently, two zonal schemes were published for the cantabrian range in northern spain (fraguas et al. 2015, 2018) and the lusitanian basin in portugal (ferreira et al. 2019). calcareous nannofossil biostratigraphy across the t-oae has been documented for several outcrop sections (see ferreira et al. 2019 for a review) and a few cores (van de schootbrugge et al. 2019; visentin et al. 2021b). core successions are, in many cases, preferable since unweathered lithologies usually display a much better preservation and sampling can be conducted in much higher resolution resulting in a more resolved taxonomy and biostratigraphy. the sogno and gajum drilling project was, in fact, aimed at recovering high-resolution data from continuous and well-preserved sequences since outcropping sedimentary rocks, and particularly black shales, are commonly badly degraded (erba et al. 2019b). continuous coring is, thus, crucial to recover high-quality fresh material with potentially good preservation. in this work we present a detailed calcareous nannofossil biostratigraphy of two continuously cored successions representing significantly different geological settings within the lombardy basin (southern alps) (erba et al. 2019b). the sogno core was drilled on the albenza plateau (a pelagic structural high) while the gajum core was penetrated in an inner basin along the slope of the mt. corni di canzo structural high (fig. 1). both cores recovered a pelagic succession through the uppermost portion of the domaro limestone (lmst.) and the lower part of sogno formation (fm.) including the fish level, which represents the lithological expression of the t-oae in the lombardy basin (tintori 1977; gaetani & poliani 1978; erba & casellato 2010; erba et al. 2019a, b). the calcareous nannofossil biostratigraphic investigation of the sogno and gajum cores is aimed at building a robust framework to constrain the t-oae paleoenvironmental changes at local, regional, and supra-regional scale. in particular, nannofossil events will be used to date the fish level black shale interval pointing out synchroneity or diachroneity within the lombardy basin, and also for comparison with other basins at different latitudes. analogies and differences will be used to derive reproducibility and lateral extension of calcareous nannofossil events. geological setting and lithostratigraphy during the jurassic, the lombardy basin was a relatively deep area bordered by the canavese zone to the west and the trento plateau to the east (gaetani 1975; 2010; bernoulli & jenkyns 2009). during latest triassic to earliest jurassic times, a rifting phase broke the southern margin of the western tethys into depressions and paleohighs that are, from west to east: monte nudo trough, lugano high, generoso trough, corni di canzo high, albenza plateau, monte cavallo high, sebino trough, botticino high (gaetani 1975, 2010) (fig. 1). in the troughs, lower jurassic partially resedimented sequences may reach a non-decompacted thickness of 3000 m (e.g., in the generoso trough), while condensation and/ or hiatuses distinguish the paleohigh successions, often represented by reddish nodular facies. on the escarpments connecting the highs to the deeper parts, sedimentation was affected by slumps, resedimented intervals and locally submarine breccias, typically interrupting condensed and seldom incomplete successions (gaetani & erba 1990; toarcian oceanic anoxic event in northern italy 541 gaetani 2010) (fig. 1). the lower jurassic successions of the lombardy basin were influenced by local-regional tectonics, but also shaped by global paleoenvironmental perturbations including the toae (erba et al. 2019a). lower toarcian sections contain a distinctive black shale interval that is the sedimentary record of oxygen-depleted seafloor of the tethys ocean. in the lombardy basin the lower toarcian black shale interval was named livello a pesci/fish level (tintori 1977): it generally has a thickness between 0.5 and 5 m, but can reach a few tens of meters in the most expanded sections. black shales are ubiquitous in the deeper settings of the lombardy basin, but are commonly absent on paleo-highs (tops and upper slopes), usually resulting from stratigraphic gaps and/or extreme condensation. the colle di sogno and gajum sites were selected as pelagic records (gaetani & erba 1990; casellato & erba 2015) for continuous coring of the lower toarcian fish level. within the lombardy basin, these sites represent different depositional settings on a pelagic structural high (albenza plateau) and in an inner basin along the slope of a structural high (mt. corni di canzo). at both locations, the fish level is present in the lower part of the sogno formation (gaetani & poliani 1978; gaetani & erba 1990; casellato & erba 2015) providing the opportunity to date, investigate and model the anoxic interval. at colle di sogno, the type-section of the sogno fm. was formalized along the road sp 179 on the northern slope of mt. brughetto (gaetani & poliani 1978). the pelagic calcilutites of the domaro limestone (lmst.) fm. are suddenly followed by the sogno fm. consisting of marlstones, calcareous marlstones, marly limestones, and black shales. the upper pliensbachian-lower bajocian colle di sogno section was characterized stratigraphically through litho-, bio-, chemo-, magneto-, and cyclostratigraphy (gaetani & poliani 1978; jenkyns & clayton 1986; gaetani & erba 1990; hinnov et al. 2000; channell et al. 2010; casellato & erba 2015). the gajum outcrop is located in a small lateral cut of the ravella valley (close to the canzo village), where the carbonate-rich lithologies of the domaro lmst. are followed by dark grey, clay-rich lithologies of the sogno fm. overlain by reddish nodular limestones of the rosso ammonitico lombardo. erba et al. (2019b) described in detail the lithostratigraphy of the sogno and gajum cores. lithologic units were defined taking into account lithological features and sedimentary structures. for each core, at least four dip measurements were taken during lab preparation to calculate the stratigraphic thickness of the drilled sections. the composite sogno core section recovered a complete upper pliensbachian–lower toarcian interval (total stratigraphic thickness = 25.33 m) represented by the uppermost part of the domaro lmst. and the lower part of the no rth am eri ca s. america africa tethysadria 20°n 10°n 10°s 0° paleohigh basin fault sogno core gajum core 0 5 km lecco como monte cavallo albenza corni di canzo co mo l ak e a d d a r iv er sea level albenza plateau monte nudo trough lugano high monte generoso trough corni di canzo high monte cavallo high w. sebino trough botticino high e. sebino trough trento platform 2 km 50 km lombardy basin (jurassic) garda escarpment jurassic drilling areaa b triassic paleozoic fig. 1 location of the sogno and gajum drilling sites relative to a) paleoand b) current geography (after erba et al. 2019b). visentin s. & erba e.542 s 4.82 10.52 9.54 14.55 15.93 21.35 19.10 22.92 24.35 25.47 26.83 11.87 1 2 3 5 6 1.5 7 8 9 10 11 13 14 15 24.99 16.86 20 21 22 23 24 25 26 27 10 3 4 5 6 7 8 9 11 12 13 14 15 16 17 18 19 0 1 2 lithostratigraphic unit sogno core 4 breccia 19.45 breccia 12 fo ld s s s s s s ss fracturedf f f f f f f grey 5y 5/1 black 5y 2.5/1 lig ht b rownish grey 2.5y 6/2 very dark grey 5y 3/1 dark grey 5y 4/1 dark grey 7.5yr 4/0 greyish b rown 2.5y 5/2 dark greyish b rown 2.5y 4/2 grey 2.5y 6/0 very dark greyish b rown 10 yr 3/2 grey 5y 6/1 faultf pyrit e s st yloli t e emerald lamina bi valve biotu rb ation faint bioturb ation laminations legend dus ky red 2.5yr 3/2 dark reddish brown 5 yr 3/3 oli ve 5y 5/3 and 5y 4/3 grey 5yr 5/1 oli ve grey 5y 5/2 and 4/2 lig ht oli ve grey 5y 6/2 reddish grey 10yr 5/1 dark reddish brown 2.5 yr 3/3 dark brown 7.5 yr 3/2 n jt 6 n jt 5 calcareous nannofossil events and zones schizosphaerella recovery (11.08 m) schizosphaerella and m. jansae crises (16.78 m) schizosphaerella decline (25.69 m) watznaueria sp. 1 (9.06 m) m. jansae (11.25 m) c. superbus crassus (17.01 m) d. constans (17.16 m) l. sigillatus (26.69 m) c. poulnabronei (24.27 m) l. crucicentralis (23.65 m) l. velatus (21.46 m) d. ignotus (20.21 m) presence of l. hauffii (26.83 m) samples b s o g n o f or m at io n e ar ly t o a r c ia n d o m a r o li m es to ne la te st p li e n s . a ge s u ni ts hiatus a b a f is h le ve l f is h le ve l fig. 2 lithostratigraphy and biostratigraphy of the sogno core. calcareous nannofossil events in red are the primary events used as zonal and subzonal boundaries. the schizosphaerella decline, crisis, recovery, and m. jansae crisis are reported in blue. toarcian oceanic anoxic event in northern italy 543 sogno fm. (fig. 2). the gajum core recovered the domaro lmst. immediately overlain by the fish level black shales, followed by marlstones of the sogno fm. and the rosso ammonitico lombardo (total stratigraphic thickness = 28.18 m) (fig. 3). materials and methods a total of 154 samples were selected for calcareous nannofossil biostratigraphy of the sogno core. the average sampling rate adopted is 16 cm through the core succession: the lowermost 9 m (core bottom to lower unit 8) and the uppermost 9 m (upper part of unit 4 to core top) were sampled at a slightly lower resolution of 20 cm whereas the fish level and the interval immediately preceding and following (upper part of unit 8, units 7, 6, 5 and lower part of unit 4) were sampled every 10 cm (fig. 2). biostratigraphic analyses were performed on smear slides prepared following the settling boxes method of geisen et al. (1999) that was applied to conduct future quantitative investigations. according to this technique, a basic ph suspension of 0.1 g of dried rock powder and ammoniac water was homogenized and let settled for 24 h on a cover slide in a settling device. the cover slide was recovered, dried, and attached to a slide with the norland optical adhesive. a total of 156 samples were collected from the gajum core. the average sampling rate adopted is 20 cm with the exception of the domaro lmst./sogno fm. lithostratigraphic boundary corresponding to the inception of the black shales of the fish level (lowermost unit 6) sampled every 0.5-1 cm (fig. 3). samples were prepared from limestones and marly limestones of the domaro lmst., limy marlstones, marlstones, marly claystones and black shales of the sogno fm. and nodular limestones of the rosso ammonitico lombardo fm. simple smear slides were prepared following the method of monechi & thierstein (1985): a small amount of rock material was powdered adding few drops of bi-distillate water, without centrifuging, ultrasonic cleaning or settling the sediment in order to retain the original composition. the obtained suspension was mounted onto a slide, covered with a cover slide fixed with the norland optical adhesive. smear slides of both cores were investigated using a light polarizing microscope, at 1250x magnification. calcareous nannofossil abundance and preservation were evaluated by examining at least 6 longitudinal traverses (750 fields of view) in each smear slide. preservation of calcareous nannofossils was characterized adopting the codes described by roth (1983) for etching and overgrowth (see appendices 2 and 3). for each sample, semi-quantitative abundances of individual taxa and total nannofossil assemblages were obtained as number of specimens in a microscope field of view as detailed in appendices 2 and 3. the biozonation scheme adopted is that of mattioli & erba (1999) established for the tethyan realm. calcareous nannofossil taxa recognized in this study are listed in appendix 1. the range charts of calcareous nannofossil taxa from the sogno and gajum cores are reported in appendices 2 and 3. plates 1 and 2 contain the photographs of the most common taxa. taxonomic notes in the present work we followed the recent taxonomic revision for the genus carinolithus conducted by visentin et al. (2021a). specifically, morphometric analyses allowed the separation of c. superbus crassus from c. superbus superbus based on the stem width (sw). the former taxon has a sw > 1 µm whereas the latter is characterized by a sw ≤ 1 µm. as a consequence, visentin et al. (2021a) concluded that c. superbus, whose fo is used as marker for the base of the nj 6 and njt 6 zones in the reference biozonation schemes (bown & cooper 1998; mattioli & erba 1999; ferreira et al. 2019), is indeed c. superbus crassus. visentin et al. (2021a) further demonstrated that the species c. cantaluppii is a diagenetic artefact of carinolithus specimens due to intensive overgrowth (highly calcified c. poulnabronei and c. superbus) and, accordingly, we disregard this taxon. the taxonomic subdivisions of casellato & erba (2015) were applied as follows: a) c. crassus is subdivided into c. crassus (length > 5 µm) and “small” c. crassus (length < 5 µm) ; b) m. jansae is divided into m. jansae and “thin” m. jansae; c) s. punctulata is divided into s. punctulata, “small” s. punctulata and “encrusted” s. punctulata. additionally, similarly to casellato & erba (2015), we consider the taxon watznaueria sp. 1 corresponding to w. fossacincta of mattioli & erba (1999) and ferreira et al. (2019). results calcareous nannofossil preservation and abundance calcareous nannofossil preservation and abundance vary through the studied interval, showing comparable and consistent variations in the two cores. in the sogno core the preservation is generally moderate to moderate/good varying from moderate/poor to moderate in the domaro lmst. and from moderate/poor to good in the sogno fm., with the exception of the fish level where a moderate/ poor to moderate/good preservation was observed. the degree of etching varies from e1 to e2, with stronger dissolution within the fish level, whereas the degree of overgrowth fluctuates between o0 and o2 and is relatively higher in the domaro lmst. similar results were obtained for the gajum core: a generally moderate preservation shows variations from poor to moderate/good in the domaro lmst. and within the fish level, moderate/poor to good in the uppermost part of the sogno fm. and from moderate/poor to moderate within the rosso ammonitico lombardo fm. the degree of etching varies from e1 to e3 and, as for the sogno core, stronger dissolution is observed within the fish level while the degree of overgrowth fluctuates between o0 and o3 with stronger evidence of overgrowth in the domaro lmst. and occasionally in the rosso ammonitico lombardo fm. the calcareous nannofossil abundance is somehow lithology-dependent: nannofossils are rare to rare/frequent and rare/frequent to frequent in the domaro lmst. of the sogno core and gajum core, visentin s. & erba e.544 respectively. within the sogno fm., nannofossils are generally frequent, fluctuating between rare to frequent/common in both cores. a drastic decrease in total abundance is observed in the fish level, characterized by extremely rare nannofossils with a few sporadic samples containing rare/frequent or frequent specimens. the rosso ammonitico lombardo fm. is characterized by rare/frequent to frequent nannofossils. biostratigraphy sogno core a total of nine calcareous nannofossil events were recognized in the sogno core (fig. 2) and the njt 5 (divided into the njt 5a and njt 5b subzones) and njt 6 zones were identified. according to the zonation of mattioli & erba (1999), the lowermost studied sample (26.83 m) is attributed to the njt 5a subzone for the presence of lotharingius hauffii and the absence of lotharingius sigillatus. the first occurrence (fo) of l. sigillatus at 26.69 m (sample s3-c-473) marks the boundary between the njt 5a and the njt 5b subzones. within the njt 5b subzone additional biohorizons were recognized, namely: the fos of carinolithus poulnabronei (24.27 m), lotharingius crucicentralis (23. 65 m), lotharingius velatus (21.46 m), discorhabdus ignotus (20.21 m) and diductius constans (17.16 m). the fo of carinolithus superbus crassus (17.01 m) defines the base of the njt 6 zone. the last occurrence (lo) of mitrolithus jansae (11.25 m) and the fo of watznaueria sp.1 (9.06 m) were detected within the nj 6 zone. following ferreira et al. (2019), we use here the lo of m. jansae to separate the njt 6a and njt 6b subzones (fig. 2). the njt 6/njt 7 zonal boundary was not identified because discorhabdus striatus was not observed in the studied interval. variations in semiquantitative abundance of a few taxa were recorded (see range chart in appendix 2) and previously defined acme-paracme levels were identified. casellato & erba (2015) identified the schizosphaerella decline, crisis and recovery as well as the m. jansae crisis on the basis of absolute abundances obtained from ultrathin section analyses. however, semiquantitative analyses (casellato & erba 2015; visentin et al. 2021b; this work) also allow the identification of the above mentioned biohorizons. in the sogno core, the schizosphaerella decline corresponds to a change from common to frequent abundance at the domaro lmst./sogno fm. boundary (25.69 m); the schizosphaerella and m. jansae crises are marked by a change from frequent and continuous records to rare and discontinuous occurrences and correlate with the base of the fish level (16.78 m). the schizosphaerella recovery has been detected at the level where this taxon returns to be continuous and rare/ frequent above the fish level (11.08 m) (fig. 2). the calcareous nannofossil assemblages of the njt 5b subzone are dominated by schizosphaerella punctulata and m. jansae. among schizosphaerellids, s. punctulata is the most abundant taxon (rare to common) whereas “small” s. punctulata (extremely rare to frequent/common) and “encrusted” s. punctulata (extremely rare to rare/frequent) are subordinated. within genus mitrolithus, m. jansae is slightly more abundant (extremely rare to frequent/common) than “thin” m. jansae (extremely rare to rare/ frequent). within genus lotharingius a slight increase in abundance is observed for l. hauffii, l. frodoi and l. sigillatus in the upper part of the nj 5b subzone. nannofossil assemblages in the lower part of the njt 6a subzone, largely corresponding to the fish level, show a severe reduction in abundance of both schizosphaerella and mitrolithus (schizosphaerella and m. jansae crises). in particular, both s. punctulata and “small” s. punctulata become extremely rare, and “encrusted” s. punctulata is recorded only sporadically. as far as m. jansae is concerned, abundances become rare to extremely rare and discontinuous in the fish level, with a bigger drop in abundance of the normal morphotype. in the fish level lotharingius becomes the dominant genus with l. hauffii, l. frodoi and l. sigillatus as most abundant species. in this interval, an increase in abundance for genera calyculus and carinolithus was also recorded (see range chart in appendix 2). calcareous nannofossil assemblages of the uppermost part of the njt 6a subzone are characterized by a relative recovery of schizosphaerellids (schizosphaerella recovery) above the fish level, although with limited abundance: s. punctulata is extremely rare to frequent and “small” s.punctulata is extremely rare to rare/frequent whereas “encrusted” s. punctulata remains sporadic. through the sogno core, genera biscutum, calyculus, carinolithus and crepidolithus are characterized by a general rare abundance, whereas bussonius, diductius, similiscutum, parhabdolithus and tubirhabdus are extremely rare. toarcian oceanic anoxic event in northern italy 545 plate 1 scale bars represent 2 µm. 1-2 b. dubium, 1) cross-polarized light, 2) quartz lamina, gajum core, sample g c25 995 (26.94 m) 3-4 b. finchii, 3) cross-polarized light, 4) quartz lamina, sogno core, sample s3 c12 330 (17.16 m) 5-6 b. grande, 5) cross-polarized light, 6) quartz lamina, gajum core, sample g c25 994 (26.89 m) 7-8 b. intermedium, 7) cross-polarized light, 8) quartz lamina, sogno core, sample s3 c12 330 (17.16 m) 9-10 calyculus spp. side view (sv), 9) cross-polarized light, 10) quartz lamina, sogno core, sample s3 c29 456 (25.5 m) 11-12 calyculus spp. distal view (dv), 11) cross-polarized light, 12) quartz lamina, sogno core, sample s3 c21 416 (22.95 m) 13-14 c. poulnabronei, 13) cross-polarized light, 14) quartz lamina, sogno core, sample s3 c22 423 (23.42 m) 15-16 c. superbus crassus, 15) cross-polarized light, 16) quartz lamina, sogno core, sample s3 c5 313 (13.42 m) 17-18 c. cavus, 17) cross-polarized light, 18) quartz lamina, gajum core, sample g c16 623 (17.15 m) 19-20 c. crassus, 19) cross-polarized light, 20) quartz lamina, gajum core, sample g c22 841 (23.09) 21-22 c. crucifer, 21) cross-polarized light, 22) quartz lamina, sogno core, sample s1 c32 192 (11.86 m) 23-24 c. granulatus, 23) cross-polarized light, 24) quartz lamina, gajum core, sample g c25 992 (26.85 m) 25-26 d. ignotus, 25) cross-polarized light, 26) quartz lamina, sogno core, sample s3 c14 343 (17.9 m) 27-28 l. barozii, 27) cross-polarized light, 28) quartz lamina, gajum core, sample g c7 275 (9. 15 m) 29-30 l. crucicentralis, 29) cross-polarized light, 30) quartz lamina, sogno core sample s1 c7 35 (4.24 m) visentin s. & erba e.546 gajum core the calcareous nannofossil biostratigraphic investigation of the gajum core resulted in the identification of seven events and of the njt 5 and njt 6 zones (fig. 3). the lowermost investigated sample (31.17 m) is assigned to the uppermost pliensbachian njt 5 zone (njt 5b subzone) based on the presence of l. hauffii and l. sigillatus. the njt 5/njt 6 zonal boundary is correlatable with the base of the fish level, although we precise that the fo of c. superbus crassus was observed 0.8 cm above the base of the lowermost black shale (26.932 m) (fig. 3). the fo of d. ignotus is placed at 26.938 m, in the topmost nj 5b subzone. within the basal part of the njt 6 zone the fos of rare and discontinuous c. poulnabronei, l. crucicentralis, l. velatus and d. constans were detected within a 4.8 cm thick interval. these findings suggest a hiatus at the base of the black shale interval of the fish level eliding part of the latest pliensbachian-early toarcian time interval. in the njt 6 zone, the lo of m. jansae (13.74 m) was detected and used to identify the base of the njt 6b subzone of ferreira et al. (2019) (fig. 3). the uppermost investigated sample (3.08 m) is still included in the early toarcian njt 6 zone due to the absence of d. striatus, which is the zonal marker of the base of the njt 7 zone (mattioli & erba 1999). the schizosphaerella decline was not observed in the gajum core, further indicating the absence of the domaro lmst./sogno fm. boundary interval. the schizosphaerella and m. jansae crises were detected at the base of the fish level (26.94 m). however, as explained above, the lithostratigraphic boundary between the domaro lmst. and the sogno fm. is marked by a hiatus and, consequently, the schizosphaerella and m. jansae crises are presumably apparent and not real events in the gajum core. the schizosphaerella recovery was detected at 12.89 m in the upper part of the fish level where this genus become continuous and rare to frequent. the calcareous nannofossil assemblages of the njt 5b subzone are dominated by s. punctulata and m. jansae. among schizosphaerellids, s. punctulata and “small” s. punctulata are the most abundant morphotypes (rare/frequent to common/abundant and rare/frequent to common, respectively) whereas “encrusted” s. punctulata is subordinated (rare to frequent/common). within the genus mitrolithus, m. jansae is relatively more abundant (rare to frequent) than “thin” m. jansae (extremely rare to rare/ frequent) (see range chart in appendix 3). nannofossil assemblages of the njt 6a subzone corresponding to the fish level show a drastic reduction in abundance of both schizosphaerella and mitrolithus (schizosphaerella and m. jansae crises). in particular, both s. punctulata and “small” s. punctulata become rare whereas “encrusted” s. punctulata is extremely sparse, although in some samples schizosphaerellids displays a frequent/common abundance. similarly to the sogno core, within the fish level m. jansae is equally represented by both normal and thin morphotypes becoming discontinuous and generally rare. in the fish level lotharingius is the dominant genus with l. hauffii as most abundant species. plate 2 scale bars represent 2 µm. 1-2 l. frodoi, 1) cross-polarized light, 2) quartz lamina, gajum core, sample g c22 854 (23.39 m) 3-4 l. hauffii, 3) cross-polarized light, 4) quartz lamina, gajum core, sample g c6 221 (8.14 m) 5-6 l. sigillatus, 5) cross-polarized light, 6) quartz lamina, gajum core, sample g c20 810 (22.19 m) 7-8 l. umbriensis, 7) cross-polarized light, 8) quartz lamina, gajum core, sample g c7 275 (9. 15 m) 9-10 l. velatus, 9) cross-polarized light, 10) quartz lamina, gajum core, sample g c5 160 (6.84 m) 11-12 m. elegans, 11) cross-polarized light, 12) quartz lamina, sogno core, sample s3 c30 466 (26.2 m) 13-14 m. jansae, 13) cross-polarized light, 14) quartz lamina, gajum core, sample g c25 995 (26.94 m) 15-16 “thin” m. jansae, 15) cross-polarized light, 16) quartz lamina, sogno core, sample s3 c24 441 (24.38 m) 17-18 “thin” m. jansae, 17) cross-polarized light, 18) quartz lamina, gajum core, sample g c25 993 (26.87 m) 19-20 m. lenticularis, 19) cross-polarized light, 20) quartz lamina, sogno core, sample s3 c12 330 (17.16 m) 21-22 s. cruciulus, 21) cross-polarized light, 22) quartz lamina, gajum core, sample g c25 995 (26.94 m) 23-24 s. punctulata, 23) cross-polarized light, 24) quartz lamina, sogno core, sample s3 c20 409 (22.5 m) 25-26 s. punctulata, 25) cross-polarized light, 26) quartz lamina, gajum core, sample g c15 599 (16.55 m) 27-28 “small” s. punctulata, 27) cross-polarized light, 28) quartz lamina, sogno core, sample sample s3 c24 439 (24.22 m) 29-30 “encrusted” s. punctulata, 29) cross-polarized light, 30) quartz lamina, sogno core, sample s3 c20 401 (22 m) 31-32 t. patulus, 31) cross-polarized light, 32) quartz lamina, sogno core, sample s3 c13 335 (17.45 m) 33-34 watznaueria sp.1, 33) cross-polarized light, 34) quartz lamina, sogno core, sample s1 c24 164 (10.47 m) toarcian oceanic anoxic event in northern italy 547 plate 2 visentin s. & erba e.548 f f f 4.35 7.11 21.39 25.42 26.94 1 2 3 5 6 7 31.18 3.0 20 21 22 23 24 25 26 27 28 29 30 31 10 3 4 5 6 7 8 9 11 12 13 14 15 16 17 18 19 gajum core lithostratigraphic unit f is h le ve l 10.60 4 s o g n o f or m at io n e ar ly t o a r c ia n d o m a r o l m st . la te st p li e n s b a c h ia n a ge s u ni ts r o s s o a m m o n it ic o lo m b a r d o n jt 6 calcareous nannofossil events and zones schizosphaerella recovery (12.89 m) schizosphaerella and m. jansae crises (26.938 m) m. jansae (13.74 m) c. sup. crassus (26.932 m) d. constans (26.884 m) c. poulnabronei (26.932m) l. crucicentralis (26.92 m) l. velatus (26.92 m) d. ignotus (26.938 m) presence of l. hauffii, l. sigillatus (31.17 m) n jt 5b samples b a fig. 3 lithostratigraphy and biostratigraphy of the gajum core. calcareous nannofossil events in red are the primary events used as zonal and subzonal boundaries. the schizosphaerella and m. jansae crises, and the schizosphaerella recovery are reported in blue. legend as in figure 2. toarcian oceanic anoxic event in northern italy 549 in the uppermost part of the fish level, calcareous nannofossil assemblages are characterized by a general recovery in abundance of schizosphaerellids (schizosphaerella recovery). however, only s. punctulata and “small” s. punctulata were observed (extremely rare to frequent/common) whereas “encrusted” s. punctulata disappears. in the uppermost studied interval represented by the rosso ammonitico lombardo fm., the family calyculaceae shows an increase in abundance of both calyculus and carinolithus, passing from extremely rare to frequent. through the gajum core, genera biscutum and crepidolithus are characterized by rare abundance, whereas bussonius, diductius, similiscutum, parhabdolithus and tubirhabdus are extremely rare. discussion correlation of the sogno core to the colle di sogno section. before coring, the section outcropping at colle di sogno was studied for calcareous nannofossil biostratigraphy and paleoceanography by erba (2004), channell et al. (2010) and casellato & erba (2015). the stratigraphic interval studied in detail by casellato & erba (2015) spans from the upper pliensbachian (domaro lmst.) to the lower toarcian (sogno fm.) and is comparable, but not identical, to the sogno core succession. in fact, the sogno core terminated at a stratigraphic level younger than the colle di sogno outcrop section and, therefore, some nannofossil events were not detected in the present work (fos of calyculus spp., biscutum grande, bussonius prinsii, lotharingius frodoi and bussonius leufuensis). conversely, the upper 5 meters of the sogno core are younger than the top of the colle di sogno section (casellato & erba 2015). as expected, the stratigraphic levels of fos and los found in the sogno core are fully consistent with data documented for the colle di sogno section by casellato & erba (2015). this is the case for the fos of c. poulnabronei, l. crucicentralis, watznaueria sp. 1, the lo of m. jansae, the schizosphaerella decline, crisis, recovery, and the m. jansae crisis. only a few events, such as the fos of l. sigillatus, d. constans and c. superbus crassus, were detected at slightly lower stratigraphic levels in the sogno core relative to the colle di sogno outcrop. relatively higher incongruities regarding the fos of l. velatus and d. ignotus were detected below the fish level in the sogno core but within the fish level in the colle di sogno section. such stratigraphic differences are, presumably, attributable to a better preservation and higher sampling rates of the sogno core, concurring to improved detection of rare taxa. in fact, observed discrepancies mostly depend on the discontinuous range of taxa in their initial ranges (see range chart in appendix 2). moreover, the generally better preservation state of the cored material increases the possibility of finding the most delicate taxa. due to their scantiness, the los of s. cruciulus and m. lenticularis were only tentatively placed in the sogno core in the upper part of the fish level and above the fo of watznaueria sp. 1, respectively, at stratigraphic levels comparable to those documented in the colle di sogno section (casellato & erba 2015). as far as the lo of the standard morphotype of m. jansae is concerned, it was placed at the base of the fish level in the colle di sogno outcrop, based on absolute abundances obtained in thin sections (casellato & erba 2015: fig. 8). this datum was named last common occurrence (lco) of m. jansae and was correlated to similar events at supra-regional scale (casellato & erba 2015). however, sparse specimens of m. jansae were documented within the fish level outcropping at colle di sogno (casellato & erba 2015), consistently with data obtained for the sogno core (this study). the two morphotypes of m. jansae co-occur in the fish level, although rare and discontinuous, and the lo of this species was located above the top of the black shale interval both in the outcrop and in the sogno core. correlation of the sogno and gajum cores calcareous nannofossil preservation, abundance, and biostratigraphy display quite consistent results between the two investigated core successions. the stratigraphic interval spans from the upper pliensbachian (domaro lmst.) to the lower toarcian (sogno fm./rosso ammonitico lombardo fm.), including the lithological expression of the t-oae (fish level) in both cores (fig. 4). the biostratigraphic results indicate that the sogno core visentin s. & erba e.550 recovered a slightly longer interval as documented by the fos of l. sigillatus in the lowermost part and watznaueria sp.1 in the upper part of the sogno core. according to their geological settings, the successions recovered in the sogno and gajum cores are characterized by a continuous and incomplete record, respectively (fig. 4). in fact, the fos of c. poulnabronei, l. crucicentralis, l. velatus and d. ignotus were detected at different stratigraphic levels in the sogno core, but in a few cm thick interval in the lowermost part of the fish level of the gajum core, revealing the presence of a hiatus. this is further evidenced by the absence of the schizosphaerella decline and the occurrence of the schizosphaerella and m. jansae crises in correspondence of the base of the black shale interval. figure 4 illustrates the biostratigraphic correlations of the two cored sequences, providing the estimate of the interval missing in the gajum core. calcareous nannofossil events, however, suggest that the onset of the fish level is most probably preserved. conversely, the lowermost toarcian (and possibly the topmost pliensbachian) is not recorded in the gajum core due to the first co-occurrence of d. ignotus, c. poulnabronei, l. crucicentralis, l. velatus and in the basal part of the fish level. indeed, the domaro lmst. is immediately overlain by black shales of the fish level, while in pelagic complete sections, as for instance the sogno core (erba et al. 2019b) and the colle di sogno section (gaetani & poliani 1978; casellato & erba 2015), the fish level is preceded by the lowermost part of the sogno fm. we estimate that ~ 600kys are missing adopting the zonal scheme of mattioli & erba (1999) and the time scale of gradstein et al. (2012) (fig. 5). the detection of a hiatus in the lower toarcian before the t-oae black shales is a rather common feature in western tethyan areas (wignall 1991; morard et al. 2003; röhl & schmid-röhl 2005; léonide et al. 2012; mattioli et al. 2013; pittet et al. 2014; menini et al. 2019; visentin et al. 2021b). the fo of c. superbus crassus shows relatively consistent results between the two cores although this biohorizon was detected at a slightly higher stratigraphic level in the gajum core (within the basal part of the fish level). this minor difference is possibly imputed to the scarcity of this taxon in its initial range or might suggest an earlier onset of anoxic sedimentation at gajum. the schizosphaerella and m. jansae crises were detected at the base of the fish level in both the sogno and gajum cores, although in the latter core this event is presumably apparent (see discussion above). schizosphaerellid abundances are different in the two studied successions: reflecting the relatively higher carbonate content in a few levels of the gajum fish level, abundances of schizosphaerellids vary from absent to common/abundant. the schizosphaerella recovery and the lo of m. jansae (both standard and thin morphotypes) display very consistent results in the two cored successions. comparison with the tethyan nannofossil zonation the zonal scheme of mattioli & erba (1999) proposed as reference for tethyan and lower latitude sections and, more specifically, for the mediterranean province, is used to discuss the nannofossil biohorizons detected in this work (fig. 5). after 20 years of nannofossil biostratigraphic investigations of lower jurassic sections the events and zones proposed by mattioli & erba (1999) have been tested and partly revised (see ferreira et al. 2019 for a synthesis). it must be emphasized that the fundamental structure – consisting in marker events and derived biozones remains, however, confirmed. the nannofossil zonal biohorizons recognized in the sogno and gajum cores are fully consistent with the scheme of mattioli & erba (1999) as discussed below, from bottom to top: * the fo of l. sigillatus defining the base of the njt 5b subzone was detected in the uppermost pliensbachian domaro lmst. in both cores. as discussed by casellato & erba (2015), the original use of this event to define the pliensbachian/toarcian boundary has been revised to a slightly older age within the latest pliensbachian (see also discussion in ferreira et al. 2019); * the fo of c. superbus crassus used to determine the base of the njt 6 zone was found at the onset of the t-oae black shale interval in both cores. we specify here that, according to visentin et al. (2021a), c. superbus crassus corresponds to c. superbus of mattioli & erba (1999); * the lo of m. jansae was detected above and in the uppermost part of the fish level black shales in the sogno and gajum cores, respectively. although originally this biohorizon was proposed by mattioli & erba (1999) as a secondary event to be toarcian oceanic anoxic event in northern italy 551 fig. 4 calcareous nannofossil biostratigraphic correlation of the sogno and gajum cores showing that the lowermost toarcian sogno fm. is missing at gajum. in the lower part of the figure, the sogno and gajum cores are contestualised on the albenza plateau and on the flank of the corni di canzo high (after erba et al. 2019b). 1 2 3 5 6 7 8 9 10 11 13 14 15 lithostratigraphic unit sogno core 4 12 c. sup. crassus l. sigillatus l. velatus d. ignotus 20 21 22 23 24 25 26 27 10 3 4 5 6 7 8 9 11 12 13 14 15 16 17 18 19 0 1 2 s o g n o f or m at io n e ar ly t o a r c ia n d o m a r o li m es to ne la te st p li e n s . a ge u ni t n jt 6 n jt 5 b a ** * *** 1 2 3 5 6 7 gajum core lithostratigraphic unit f is h le ve l 4 a ge u ni t 20 21 22 23 24 25 26 27 28 29 30 31 10 3 4 5 6 7 8 9 11 12 13 14 15 16 17 18 19 s o g n o f or m at io n e ar ly t o a r c ia n d o m a r o li m es to ne la te st p li e n s b a c h ia n r o s s o a m m o n it ic o lo m b a r d o n jt 6 c. poulnabronei l. crucicentralis l. velatus d. ignotus l. sigillatus n jt 5b *** ** d. constans watznaueria sp. 1 c. sup. crassus b a b a d. constans m.jansae c. poulnabronei l. crucicentralis f is h le ve l 0 5 km 100 m 0 1 km 250 m gajum core sogno core m.jansae visentin s. & erba e.552 confirmed, its reproducibility makes this datum fully reliable at low latitudes (casellato & erba 2015) and we use it to separate the njt 6 zone into 2 subzones as suggested by ferreira et al. (2019). the zonal scheme recently proposed by ferreira et al. (2019) for the lusitanian basin is also considered here (fig. 5). events, zones, and subzones are based on detailed investigation of several lusitanian successions, including two gssp sections, located during the jurassic in the western tethys and, thus, acting as a north-south migration pathway for several organisms, including calcareous nannoplankton and ammonites. ferreira et al. (2019) calibrated nannofossil biohorizons against ammonite zones (azs) as well as against carbon and oxygen isotope excursions where possible. moreover, the work by ferreira et al. (2019) includes an updated and revised taxonomy and a comprehensive synthesis of biostratigraphies from the western tethys. as far as the marker events are concerned, the fo of c. superbus crassus (= c. superbus in ferreira et al. 2019) and lo of m. jansae are comparable with our findings and the zonation of mattioli & erba (1999) (fig. 5). conversely, ferreira et al. (2019) placed the fo of l. sigillatus within the njt 4 zone (base of their njt 4e subzone) of late pliensbachian age (margaritatus az), thus, at a definitively older stratigraphic level relative to mattioli & erba (1999) and our data from the sogno and gajum cores. ferreira et al. (2019) stated that the earliest specimens of l. sigillatus recorded in the upper pliensbachian of the rabaçal section are significantly smaller (average distal shield diameter of 4.34 µm; ferreira et al. 2017) than the size range provided in the original description by stradner (1961) (6-8 µm). nevertheless, since these early specimens show identical diagnostic features and their sizes are not very different from those documented in the emendation by goy (1981), ferreira et al. (2019) considered them within l. sigillatus. we notice that ferreira et al. (2019) report the first common occurrence (fco) of l. sigillatus at the stratigraphic level of the fo of l. sigillatus of mattioli & erba (1999) and as found in the sogno and gajum cores (fig. 5). the fo of l. crucicentralis is also older in the zonal scheme of ferreira et al. (2019), who placed this event within the lowermost emaciatum az at the base of the njt 5b subzone (late pliensbachian). these authors specified that several specimens of l. crucicentralis, although with diagnostic features matching those described by medd (1971) and emended by grün & zweili (1980), are significantly smaller in size. in this study, we adopted taxonomic features of l. sigillatus and l. crucicentralis consistent 184 183 181 182 d. ignotus l. velatus l. crucicentralis l. sigillatus c. poulnabronei la te p li e n s . d om ar o lm st . gajum core (this study) d. striatus w. fossacincta w. colacicchii d. ignotus l. velatus l. crucicentralis c. superbus l. sigillatus c. poulnabronei calyculus spp. l. barozii l. hauffii m. jansae e ar ly t o a r c ia n la te p li e n s . sp in at um te nu ic os ta tu m se rp en tin us a ge a m . z on e standard tethys biozonation (mattioli & erba, 1999) n an no fo ss il z on e d. constans d. ignotus l. velatus l. crucicentralis l. sigillatus c. poulnabronei e ar ly t o a r c ia n la te p li e n s . a ge d om ar o lm st . s og no f or m at io n li th os tr at ig ra ph y watznaueria sp.1 sogno core (this study) l. hauffii schizosphaerella and m. jansae crisis schizosphaerella decline d. constans f is h d. striatus c. superbus m. jansae portugal biozonation (ferreira et al., 2019) l. crucicentralis e ar ly t o a r c ia n la te p li e n s . em ac ia tu m po ly m or ph um le vi so ni a ge a m . z on e m ar ga rit at us l. hauffii z. erectus w. fossacincta d. ignotus d. ignotus fco l. velatus l. sigillatus fco d. constans n jt 5 n jt 6 6a 6b 5c 5b c. sup. crassus e ar ly t o a r c ia n s og no f m . r a l schizosphaerella recovery a ge li th os tr at ig ra ph y n jt 6 5b 5a n jt 5 n jt 4 4b n jt 6 5b 5a n jt 5 n jt 4 4b 5a c. poulnabronei m ar ga rit at us c. sup. crassus n an no fo ss il z on e n an no fo ss il z on e 6b 6a m. jansaem. jansae l e ve l * ** *** * ** *** ** *** fig. 5 comparison of the results obtained for the sogno and gajum cores with the zonations of mattioli & erba (1999) and ferreira et al. (2019). zonal and subzonal events are reported in red. toarcian oceanic anoxic event in northern italy 553 with those of ferreira et al. (2019), without using differentiation based on coccolith size. therefore, we conclude that the fos of l. sigillatus and l. crucicentralis are diachronous, being older in the lusitanian basin relative to the mediterranean province in the tethys ocean. among the secondary events, the fo of c. poulnabronei detetcted in the sogno core is consistent exclusively with that of mattioli & erba (1999). ferreira et al. (2019), indeed, placed this event within the uppermost polymorphum az, just after the fo of c. superbus crassus. it is possible, however, that this discrepancy is mainly the result of the discontinuous occurrence of this taxon in its initial range rather than diachroneity. the fo of c. poulnabronei, indeed, was reported by ferreira et al. (2019) only in the peniche section and looking at the range charts provided in the supplementary material c. poulnabronei occurs sporadically, thus not excluding a potentially older first occurrence. conversely, the fo of d. ignotus detected in our study shows consistency with the biozonation of ferreira et al. (2019), while mattioli & erba (1999) assigned a younger age to this event, between the fos of c. superbus crassus and the lo of m. jansae. the reason of this discrepancy is also probably attributable to scantiness of this taxon. in fact, according to mattioli et al. (2013), d. ignotus has a lazarus behaviour, first occurring in the earliest toarcian, being absent from the sediments corresponding to the t-oae and subsequently occurring consistently from the end of the t-oae upwards. this distribution was observed not only at peniche, but also in sections of the mediterranean province, namely, valdorbia in central italy (mattioli et al. 2013), amellago in marocco (bodin et al. 2010) and la almunia in south-eastern spain (menini et al. 2019). although our findings confirm that the fo of d. ignotus precedes the onset of the t-oae black shales within the njt 5b subzone of mattioli & erba (1999), it is possibly coeval with the datum of ferreira et al. (2019) within their njt 5c subzone. the stratigraphic level of the fo of d. ignotus of the mattioli & erba (1999) zonation appears to be equivalent to the fco of this taxon in the zonal scheme of ferreira et al. (2019) (fig. 5). as far as the fo of l. velatus is concerned, our finding is inconsistent with the zonal schemes of ferreira et al. (2019) and mattioli & erba (1999) who proposed an older (shortly before the fo of z. erectus) and younger (between the fos of c. superbus crassus and d. ignotus) age, respectively. the fo of d. constans was not considered in mattioli & erba (1999), and our results fit well with the datum proposed by ferreira et al. (2019). in our study, the fo of watznaueria sp. 1 corresponds to the fo of w. fossacincta in both the zonations by mattioli & erba (1999) and ferreira et al. (2019). the taxonomic characterization of watznaueria sp. 1 described by cobianchi (1992) and casellato & erba (2015) are applied here. differently from the biozonation of mattioli & erba (1999), the fo of w. colacicchii, reported between the lo of m. jansae and the fo of watznaueria sp. 1 (= w. fossacincta), was not found in the sogno and gajum cores as this taxon is absent in the investigated interval. also casellato & erba (2015) did not observe w. colacicchii in the colle di sogno section. ferreira et al. (2019) place the fo of w. colacicchii in the lower part of the njt 7 zone at the base of the bifrons az, thus at a stratigraphic level younger than the zonation of mattioli & erba (1999) and the interval here investigated. nannofossil events reported by ferreira et al. (2019), but not by mattioli & erba (1999), and not identified in the sogno and gajum cores are, from the oldest to youngest, the fos of axopodorhabdus atavus, zeugrhabdotus erectus, ethmorhabdus spp. and lo of mazaganella protensa. this major difference is presumably linked to nannoplankton provincialism during the early jurassic which further supports the necessity of two biozonation schemes applicable in the lusitanian basin and the mediterranean province, respectively. summary and conclusions high-resolution calcareous nannofossil biostratigraphy carried out in two cores drilled in the lombardy basin allowed the identification of nine (sogno core) and seven (gajum core) events in the uppermost pliensbachian domaro lmst. and the lower toarcian sogno fm. and rosso ammonitico lombardo fm. as expected, relative to the colle di sogno outcrop calcareous nannofossils are better preserved in the sogno core and the high-resolution sampling resulted in improved detection of events. the correlation of the sogno and gajum nannofossil biostratigraphy shows that, according to their geological settings, the succession recovered in the visentin s. & erba e.554 sogno core (located on the albenza plateau) is continuous while a hiatus of ~600 kyrs was detected at the base of the fish level in the gajum core. however, nannofossil data suggest that the black shale interval is complete whereas the lowermost toarcian basal portion of the sogno fm. (and possibly the topmost pliensbachian domaro lmst.) is missing in the gajum core. following the standard nannofossil zonation for the mediterranean province in the tethys ocean (mattioli & erba 1999), the njt 5 and njt 6 zones were identified in both the sogno and gajum cores. furthermore, as proposed by ferreira et al. (2019), we adopt the lo of m. jansae to separate the njt 6a and njt 6b subzones. in the sogno and gajum cores, the fish level, regionally considered the lithological expression of the t-oae, results to be largely constrained between the fo of c. superbus crassus and the lo of m. jansae (njt 6a subzone), although black shales continue for a short interval above the lo of m. jansae in the gajum core. therefore, nannofossil biostratigraphy suggest that anoxic conditions did not cease simultaneously in nearby sites within the lombardy basin. other nannofossil events useful for the biostratigraphic characterization of the t-oae are the fo of d. constans and the schizosphaerella m. jansae crises at the base of the fish level and the schizosphaerella recovery following the lo of m. jansae. the comparison between the results of our investigation and the zonal schemes established for the tethys ocean (mattioli & erba 1999; ferreira et al. 2019) revealed consistencies and differences. confirming the mediterranean affinity of the studied sites, calcareous nannofossil biohorizons identified in the sogno and gajum cores are mostly coherent with the biozonation of mattioli & erba (1999), that is, however, here improved separating the njt 6 zone into two subzones. the zonation of ferreira et al. (2019) established for the western tethys lusitanian basin is only partially reproducible, in practice only for the njt 6 zonal markers, thus confirming the supra-regional value of the fo of c. superbus crassus and the lo of m. jansae. another event traceable from the mediterranean area to the lusitanian basin is the fo of watznaueria sp. 1 in the njt 6b subzone. some discrepancies might be explained by the abundance/rarity of taxa in different areas. in fact, ferreira et al. (2019) report the fco of l. sigillatus at the stratigraphic level of the fo of l. sigillatus of mattioli & erba (1999) and as found in the sogno and gajum cores. similarly, we detected the fo of d. ignotus within the njt5b subzone of mattioli & erba (1999) at a stratigraphic level comparable to that of ferreira et al. (2019), but definitively before the datum of the mattioli & erba (1999) whose fo of d. ignotus appears to be equivalent to the fco of this taxon in the zonal scheme of ferreira et al. (2019). contrarily to mattioli & erba (1999), we did not observe w. colacicchii in the ntj 6b subzone as in the zonation of ferreira et al. (2019), who report the fo of w. colacicchii in the lower part of the njt 7 zone at the base of the bifrons az, thus at a stratigraphic level younger than the investigated interval. our finding of the fo of l. velatus is inconsistent with the zonal scheme of ferreira et al. (2019) while mattioli & erba (1999) proposed a younger age between the fos of c. superbus crassus and d. ignotus. this disparity along with the absence of several marker species used in the zonation of ferreira et al. (2019) support the existence of a prominent nannoplankton provincialism between portugal and the mediterranean province during the early jurassic and, therefore, justify the use of partially different zonal schemes. we underline, however, that at supra-regional scale the t-oae is unambiguously constrained by the fo of c. superbus crassus and the lo of m. jansae. acknowledgments: the associate editor isabella raffi, the reviewers angela fraguas and emanuela mattioli are warmly acknowledged for their constructive detailed criticism that much improved the quality of the manuscript. the t-oae coring project in the lombardy basin derived from fieldwork and stratigraphic characterization of jurassic successions with maurizio gaetani. the sogno and gajum coring campaign was funded by miur-prin2011(2010x3pp8j) awarded to ee., while the nannofossil biostratigraphic characterization was conducted within the prin 2017rx9xxxy awarded to ee. references bernoulli d. & jenkyns h. c. 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(1991) model for transgressive black shales. geology, 19(2): 167-170. rivista italiana di paleontologia e stratigrafia volume 117 no. 1 2 pls. pp. 105-114 april 2011 lercaritubus problematicus flügel, senowbari-daryan & di stefano and vangia telleri (flügel): two problematic organisms from the permian jamal formation of shotori mountains, northeast iran baba senowbari-daryan 1 & koorosh rashidi 2 received: february 20, 2009; accepted: november 22, 2010 1 geozentrum nordbayern, department of paleontology, university of erlangen-nürnberg, loewenichstr. 28, 91054 erlangen, germany. e-mail: basendar@pal.uni-erlangen.de 2 university of payame-e noor, ardakan/yazd, iran. e-mail: koo.rashidi@gmail.com . key words: lercaritubus, vangia, problematica, permian, jamal formation, shotori mountains, iran. abstract. lercaritubus problematicus flügel, senowbari-daryan & di stefano and vangia telleri (flügel) and enigmatic calcareous fossils, known from the sicily, guadalupe mountains, usa and oman is described from the permian jamal formation of shotori mountains, northeast iran. the new genus name vangia is introduced for uvanella? telleri flügel. the systematic position of vangia telleri (flügel) nov. comb. as possibly cyanobaterium and its relationships with bacinella irregularis radoicic, an abundant enigmatic fossil in jurassic-cretaceous shallow water deposits, is discussed. riassunto. vengono descritti lercaritubus problematicus flügel, senowbari-daryan & di stefano e vangia telleri (flügel), fossili calcarei di natura enigmatica conosciuti in sicilia, guadalupe mountains, usa e in oman e ora rinvenuti nella formazione jamal del permiano delle montagne shotori, nel nordest dell’iran. viene introdotto il nuovo nome generico vangia per inquadrare la specie uvanella? telleri flügel. sono anche discussi la posizione sistematica di vangia telleri (flügel) nov. comb. quale possibile cianobatterio ed i suoi rapporti con bacinella irregularis radoicic, un fossile enigmatico abbondante nelle rocce giurassico-cretacee di acque basse. introduction lercaritubus problematicus flügel, senowbaridaryan & di stefano, a tube-like organism was originally described as “microproblematicum a” by flügel (in flügel et al. 1984) from the middle permian of slovenia, and later as lercaritubus problematicus by flügel et al. (1990), from the lower permian reef boulders embedded within the siliciclastic deposits of the lercara formation in sicily. it was reported also from the middle permian reefs in guadalupe mountains of texas and new mexico by senowbari-daryan & rigby (1996) and from the permian reef limestones of oman by weidlich (1992). lercaritubus occurs within the reef or reefal carbonates of jamal formation (lower and middle permian) in the shotori mountians, of northeast iran. it seems to be an index fossil limited to permian time. the occurrence of lercaritubus in sicily, oman, iran, and in usa indicates to possibly its cosmopolitan importance. vangia telleri nov. comb., originally described as possibly chambered sponge uvanella? telleri by flügel (in flügel et al. 1984) from the permian of slovenia, is another problematic and aggregate-building organism (cyanobacteria?). it is composed of numerous irregularly arranged cavities, separated by thin and compact walls. the wall appears dark micritic in transmitted light. vangia telleri occurs as isolated aggregates in sediment or rarely incrusts other organisms. geographic position of studied localities permian deposits of central iran, called the jamal formation (stöcklin et al. 1965), crop out in several localities in the shotori mountains, northeast iran. lercaritubus problematicus was found in two sections of permian jamal formation, located about 65 km and 45 km north of the town of tabas (fig. 1). these localities 106 senowbari-daryan b. & rashidi k. are described briefly below. 1. deh-e mohammad locality (fig. 1, locality 1): this locality lies about 65 km north of the town of tabas, about 5 km northeast of the small village of deh-e mohammad, about 1 km from the road tabasboshruhe, in an area called agheldun (fig. 1, 57o 01` 46`` e and 33o 59` 46.6`` n). here the permian deposits are 290 m thick and overlie the carboniferous sardar formation, which is about 26 m thick at this locality. the lower and middle 247 m of the permian deposits were sampled. the upper 43 m of the formation is dolomitic and was not sampled. 2. bagh-e vang locality (fig. 1, locality 2): this section of the kuh-e bagh-e vang is located 45 km north of the town of tabas, near the town of shirgesht (geological map 1:100.000 of shirgest completed by ruttner et al. 1968). a section of the permian jamal formation crops out on the western and southern flank of the kuh-e bagh-e vang (33˚ 58’ 60” n, 56˚ 47’ 66” e, fig. 1). also in this section the permian sediments overlie the carboniferous sardar formation, which includes siltstones, shales and sandstones. the permian deposits are overlain by the lower triassic sorkh-e shale formation. the permian section of the jamal formation in the bagh-e vang locality reaches a thickness of 293 m (ruttner et al. 1968) to 300 m (leven & vaziri mohaddam 2004), but one of the authors (kr) measured it on the south flank of this mount as about 320 m thick. generally the jamal formation in this locality is composed of sandy limestone with some olistoliths, dark shales, and mediumto thin-bedded marly limestones. partoazar (1995) introduced the name bagh-e vang member for the 60 m of the lower part (being asselian–sakmarian in age) of the jamal formation of this section. the middle part of the section is characterized by medium-bedded limestone intercalated with chert layers. the summit of the section is covered by massive carbonates (fig. 2), which is overlain by the lower triassic sorkh-e shale formation. based on fusulinids, leven & vaziri mohaddam (2004) recognized 10 units within the jamal formation in kuh-e bagh-e vang. the sponges of this locality were studied by senowbari-daryan et al. (2005, 2006) and the bryozoans by ernst et al. (2006, 2009). thin sections, containing the described specimens of lercaritubus problematicus and vangia telleri (flügel) nov. comb. are deposited in “geozentrum nordbayern, department of paleontology, university erlangen-nürnberg” (“material: senowbari-daryan: permian iran, bagh-e vang”). systematic paleontology family uncertain genus lercaritubus flügel, senowbari-daryan & di stefano, 1990 type species: lercaritubus problematicus flügel, senowbari-daryan & di stefano, 1990 original diagnosis: “sessile, tube-like, multi-branched organism with a calcareous skeleton, composed of thick wall segments put into one another. the aperture of the tubes widens distally and is fig. 1 geographic position of the studied sections in northern area of the town of tabas, shotori mountains, northeast iran. 1) locality near the town of deh-e mohammad, 2) bagh-e vang locality. two problematic organisms from the permian jamal formation of shotori mountains, northeast iran 107 characterized by a distinct collar. the outer surface of skeleton shows small polygonal depressions. the interior of the tubes is subdivided by perforated “tabulae” and shows an additional calcareous tissue. some specimens show small pores within the outer wall of the tubes” (flügel, senowbari-daryan & di stefano 1990: 361). lercaritubus problematicus flügel, senowbari-daryan & di stefano, 1990 pl. 1, figs 1-6 1984 microproblematicum a flügel (in flügel et al.), p. 197, pl. 30, figs. 13-14. * 1990 lercaritubus problematicus n. sp. flügel, senowbaridaryan & di stefano, p. 361-362, pl. 1, figs. 1-16, text-figs. 2-3. 1991 lercaritubus problematicus flügel, senowbari-daryan & di stefano flügel et al., p. 176. 1992 lercaritubus problematicus flügel et al.weidlich, p. 4445, pl. 16, fig. 3. 1996 lercaritubus problematicus flügel, senowbari-daryan & di stefano senowbari-daryan & rigby, p. 23, figs. 3.1-3.7, 4.1-4.8. 2001 lercaritubus problematicusweidlich, p. 342. 2005 lercaritubus problematicus flügel, senowbari-daryan & di stefano senowbari-daryan et al., p. 401, figs. 13.8/l, 15.8-9. material: numerous specimens in thin sections from the bagh-e vang section and from the section near the town of deh-e mohammed. for the number of illustrated thin sections see explanation of plates. description. sessile, tube-like, single or branched organism composed of several segments inserted into one another. each individual segment shows a distinct collar around the aperture of that segment. this feature was observed only in some iranian specimens. tube walls of iranian specimens, like specimens from sicily and guadalupe mountains, are strongly re-crystallized. individual crystals appear light to yellow-brown in transmitted light and are relatively large. flügel (in flügel et al. 1984) gives the size of the crystals as up to 750 µm. strong re-crystallisation of the tube walls suggests the primary aragonite skeletal mineralogy. in cross sections, the tube interior is circular, but the outer surface is distinctly wavy (pl. 1, figs 3-6) to spiny (pl. 1, figs 1-2). internal surfaces of the tubes are smooth, but outer surfaces are honeycomb, appearing wavy or with spine-like elements in sections. these elements are the edges of polygonal depressions of the tube surface. a construction of the tube is given by flügel et. al. (1990) tube interiors of the iranian specimens are usually without any internal infilling structure, but some specimens, like most specimens of the type material from the lower permian of sicily and specimens from the middle permian of upper capitan limestone of the guadalupe mountains in new mexico, usa, contain “tabulae”-like or concentric structures. in some specimens in the type material from sicily (flügel et al. 1990: pl. 1, fig. 7) the “tabulae” are perforated, but perforation was not observed in the iranian material. tubes reach lengths of up to 30 mm, with an outer diameter of up to 10 mm, an inner diameter up to 6 fig. 2 view from west to east (west flank of the mount) of the studied section of the carboniferous sardar and permian jamal formation in kuh-e bagh-e vang. the massive carbonate at the top is still permian, overlain by the triassic sorkh-e shale formation, which is exposed in the east flank and not visible from this view. 108 senowbari-daryan b. & rashidi k. mm, and thicknesses of the walls up to 1 mm. weidlich (1992) gives the wall thickness of up to 1.7 mm in material from oman. associated organisms. lercaritubus problematicus occurs in reef or reefal biotopes, like other reef builder (e. g. sponges). it is associated with hypercalcified sponges (senowbari-daryan et al. 2005, 2006), rarely corals, abundant bryozoans (ernst et al. 2009), rare green algae such as anchicodium sp., epimastoporella sp., and imperiella sp. associated problematic organisms include: tubiphytes obscurus maslov and tubiphytes carinthiacus flügel. fusulinids of the locality were described by leven & vaziri mohaddam (2004). almost all specimens of lercaritubus were incrusted by archaeolithoporella, or different types of other microbial crusts. discussion. the systematic position of lercaritubus is uncertain. flügel et al. (1990) compared lercaritubus with bryozoans, annelids, cribricatheans, and with the jurassic problematic alga “bankia” (campbelliella), but because of different morphological features the affiliation of lercaritubus to these groups of organisms seems to be questionable. the systematic position of lercaritubus is still uncertain. occurrence and stratigraphical range. lercaritubus problematicus is known from the lower and middle permian reef boulders of sicily (flügel et al. 1990, 1991), middle permian of guadalupe mountains (senowbari-daryan & rigby 1996), upper permian of oman (weidlich 1992, 2001), and now from the jamal formation (lower and middle permian) in the shotori mountians, of northeast iran. it is restricted to the tropical reef or reefal deposits and seems to be an index fossil of permian age. vangia nov. gen. type species: uvanella? telleri flügel (in flügel et al. 1984) derivatio nominis: named from the type locality kuh-e bagh-e vang (bagh-e vang mount) in the shotori mountains, northeast iran. diagnosis: nodules of aggregates, which are composed of irregularly chambers. chamber walls are thin and imperforate, appearing dark micritic in transmitted light. some chambers contain vesiculae-like structures. comparison: see discussion after description of the species. vangia telleri (flügel in flügel et al. 1984), nov. comb. pl. 1, figs 7-9; pl. 2, figs 1-8; text-fig. 3 ? 1981 bacinella sp.vachard & montenat, p. 33, pl. 2, figs 1112. 1984 uvanella? telleri n. sp. flügel (in flügel et al.), p. 205, pl. 37, figs 4-7. 1991 tubiphytes forming a net-like structure flügel et al., pl. 40, fig. 5. 1992 uvanella telleri flügel weidlich, pl. 16, fig. 2. 2002 spider-web-like microproblematicum wahlman, fig. 15. 2005 “aggregates or lumps” senowbari-daryan et al., fig. 8.7. material: numerous specimens in several thin sections. for the number of thin sections containing the illustrated specimens see plate explanations. description. aggregates of this organism reach dimensions of several mm and are composed of numerous circular, oval, tube-shaped or irregularly cavities or “chambers”, separated from each other by thin walls. not only the shape of cavities, but also their size is very variable. tube-shaped cavities were observed in almost all specimens. aggregates of vangia telleri occur either fig. 3 vangia telleri (flügel). drawn from pl. 2, fig. 1 showing details of the specimen imbedded in micritic matrix. the compact “chamber” walls appear dark micritic in submitted light and are without any perforation. two problematic organisms from the permian jamal formation of shotori mountains, northeast iran 109 as isolated “lumps” within the micritic matrix (pl. 1, figs. 7; pl. 2, figs. 1-2, 7-8) or they rarely incrusts other reef organisms (pl. 1, fig. 8; pl. 2, figs 4, 6). the abundant occurrence of vangia telleri in micritic matrix indicates such to be its growth position, possibly on the sediment surface. it also occurs as a nodular organism that was rarely grown around other organisms and stabilized them to form larger and compacted aggregates. large cavities (e. g. pl. 1, fig. 7c, 9c; pl. 2, figs. 3, 5, 6), filled with sparry calcite cement occur with aggregates of vangia telleri and were formed by the activity of this organism, but do not belong to it. discussion. aggregates of vangia telleri which are composed of chambers with thin, compact and micritic walls, were described as uvanella? telleri from the middle permian of slovenia by flügel (in flügel et al. 1984). uvanella with the type species u. irregularis ott is a sphinctozoan sponge with mg-calcite skeletal mineralogy and was originally described from the ladinian-carnian of the northern calcareous alps (austria) by ott (1967). detail descriptions of all uvanella species, with their stratigraphic ranges and geographic distributions are given by senowbaridaryan (1990). the following characteristics of vangia telleri (flügel) do not justify an affiliation of this fossil with the sphinctozoan sponge uvanella: a) wall of cavities or chambers. the chamber walls in uvanella are distinctly thick and pierced by opening or pores. the wall in vangia telleri is much thinner without any openings or pores. the “very scarce connecting pores”, noted by flügel (in flügel et al. 1984: 203) were not observed within the wall in iranian materials. some small and white-appearing points within the wall of vangia telleri are interpreted, in agreement with flügel (in flügel et al. 1984), as sedimentary particles or grains. b) small, circular cavities within the walls of vangia telleri (see flügel et al. 1984: pl. 37, fig. 7: arrows), which are also present in iranian material, are not known in specimens of uvanella species. c) large cavities indicated with c in pl. 1, figs 7, 9, pl. 2, figs 3, 5, 6 and produced by the activity of vangia telleri were not observed in specimens of the triassic genus uvanella. d) the skeleton of uvanella is composed of mg-calcite (senowbari-daryan 1990). sphinctozoan sponges with mg-calcite mineralogy appeared in the middle triassic (anisian) and became extinct at the end of triassic (senowbari-daryan & rigby in press). sphinctozoan sponges with mg-calcite mineralogy are not known in the permian record. e) uvanella seems to be limited to the middle and upper triassic and is not known from the lower triassic and permian time. general shape and the morphology of vangia telleri are similar to that fossil named bacinella irregularis by radoicic (1959) and which is known from the upper triassic? jurassic to the tertiary of numerous localities on the world. the validity of bacinella as an independent genus or its synonymy or consortium with lithocodium elliott (1956) has been discussed by numerous authors, and finally by banner et al. (1990) who synonymized bacinella with lithocodium. bacinella was accepted as a synonymous of lithocodium by later workers (e. g. neuweiler & reitner 1992). koch et al. (2002) accept not only the synonymy of both genera, but they presume that other genera (e. g. bacinellacodium dragastan, 1985 or radoicicinellopsis banner, finch & simmons 1990) are also synonymous with lithocodium. most authors (e. g. schmid & leinfelder 1996; cherchi & schröder 2006; védrine et al. 2007, and schlagintweit 2010) do not accept that bacinella and lithocodium are synonymous. radoicicinellopsis was established by banner et al. (1990) designating bacinella? sterni radoicic (1972) as type species. other known species of bacinella, including b. ordinata pantic (1972), b. bicellularis sadati (1981), and b. crispa eliasova (1981) were not treated by banner et al. (1990). bacinella crispa is similar to lithocodium/bacinella irregularis, but b. bicellularis and b. ordinata are different and may be were erroneously assigned to bacinella. all three species are different and a comparison with vangia telleri (flügel) is urgent. lithocodiun was interpreted as a codiacean alga by elliott (1956). following this idea, banner et al. (1990) classified lithocodium as a green alga (chlorophyceae family codiaceae, subfamily lithocodiaidea). lithocodium was interpreted as association of cyanobacateria/microbes/porostromata by maurin et al. (1985) and camoin & maurin (1988), cyanobacteria/ algae/foraminifera (leinfelder et al. 1993), as foraminifer by schmid & leinfelder (1995, 1996) or as possibly “colonies of calcified cyanobacteria” by cherchi & schroeder (2006). recently schlagentweit et al. (2010) discussed the systematic position of lithocodium attributing it to “filamentous-septate heterotrichale ulvophyceaen alga”. assigning to “endolithic ulvophycean alga” these authors separate bacinella from lithocodium, which was synonymed by banner et al. (1990). with separation of lithocodium and bacinella the systematic classification proposed by schlagintweit et al. (2010) is followed here to describe the permian species from bagh-e vang in iran (see above). vangia telleri occurs always separate and never together with lithocodium like the type species of bacinella – b. irregularis radoicic in mesozoic deposits. because of the compact wall of the “chambers” a sponge interpretation for bacinella is urgent. an 110 senowbari-daryan b. & rashidi k. interpretation of bacinella as a foraminifer – interpretation of some worker for lithocodium/bacinella – seems to be also unlikely. most probably bacinella should be classified as cyanobacteria. the systematic position of lithocodium/bacinella, however, as an animal (e. g. possibly sponge: koch et al. 2002 or hydrozoas: turnsek & buser 1966), foraminifer (schmid & leinfelder 1995, 1996) or plant (algae: elliott 1956; radoicic 1959; schlagintweit et al. 2010; lithocodium as juvenile stage of bacinella irregularis: fenninger 1972, bacinella irregularis as juvenile stage of lithocodium: segonzac & marin 1972), cyanobacteria; cherchi & schroeder 2006) remains still uncertain. this mentioned difference justifies the establishment of a separate genus for this permian organism, named vangia nov. gen. the twice mass extinctions (at the boundaries of permian/triassic and triassic/ jurassic) and their strong influence during the time interval of permian and jurassic-cretaceous supports the separation of bacinella-like organism of permian time as vangia. because of the compact wall of the “chambers” a sponge interpretation for vangia is uncertain. occurrence. vangia telleri (flügel) is known from the permian of karawank (kochansy-devidé (1970), slovenia (flügel in flügel et al. 1984), oman (weidlich 1992, 2001), sicily (flügel et al. 1990, 1991: pl. 40, fig. 5) and now also from the permian jamal formation of northeast iran. it was first reported as “aggregate or lumps” from iran by senowbari-daryan et al. (2005). wahlman (2002: fig. 15) described vangia telleri from the lower permian (wolfcampian) of west texas as “spider-web-like microproblematicum”. in fact, the irregular aggregate of his “microproblematicum” or of vangia is similar to some fistuliporid bryozoans colonies, but details of bryozoans, like apertures lack in vangia. vachard & montenat (1981: pl. 2, figs. 11-12) described from the upper permian of afghanistan an organism as bacinella sp. which is similar or almost identical to v. telleri (flügel). acknowledgements. the investigations were carried out in frame of the research project “se 416/17” supported by the deutsche forschungsgemeinschaft (dfg) to b. senowbari-daryan. field work was done by one the authors (kr). we are grateful to j. keith rigby (brigham young university, provo, utah) for his linguistic help and useful comments. thanks are addressed to felix schlagintweit (munich) for review of the first draft of the manuscript. valuable comments from pedro cózar as journal reviewer improved the manuscript. remarks of d. vachard, as the second reviewer could not be followed. plate 1 lercaritubus problematicus flügel, senowbari-daryan & di stefano (1-6) and vangia telleri (flügel) (7-9) from the permian jamal formation of bagh-e van, shotori mountains, northeast iran. scale: 1 mm. fig. 1 longitudinal section through a specimen showing the thick wall with smooth internal, but wavy external wall. the wall is strongly re-crystallized in all specimens and is composed of large crystals. dm27/1. fig. 2 oblique section clearly showing the wavy external wall. dm27/1. fig. 3 longitudinal section of a broken(?) specimen with wavy outer surface. dm27/1 fig. 4 longitudinal section of a specimen, which shows an open end (left in photograph). 7/1. fig. 5 longitudinal to oblique section. the specimen is colonized by a small specimen (right in photograph). 7/1. fig. 6 longitudinal section of a specimen with budding at the left side. 7/1. fig. 7 section through an aggregate showing numerous irregularly cavities. c indicates a cavity formed by the activity of vangia telleri, but does not belong to this organism. 12/3 fig. 8 section through a spherical specimen, which is growing on a tubiphytes carinthiacus (flügel). the boundary between tubiphytes carinthiacus (flügel) and vangia telleri (flügel) is marked with a white dotted line. t. carinthacus grew around some chambers of vangia telleri. bs38 fig. 9 section through an aggregate with numerous cavities. arrows indicate two specimens of tubiphytes obscurus maslov. for explanation of the letter c see fig. 7. bs58. plate 2 vangia telleri (flügel) from the permian jamal formation of bahg-e vang and deh-e mohammed section in shotori mountains, northeast iran. scale 1 mm. fig. 1 section through an aggregate with irregularly chambers. almost all chambers are filled with calcite cement. bm58; fig. 2 section through several(?) specimens with irregular and tube-like chambers. bm58; fig. 3 similar to fig. 2. for explanation of the letter c see plate 1, fig. 7. bm58; fig. 4 aggregate of vangia telleri (flügel) incrusts other organism marked with a. for explanation of c see plate 1, fig. 7. bm58; fig. 5 an organism (a) is incrusted by vangia telleri (flügel) with several tube-like chambers. for explanation of the letter c see plate 1, fig. 7. bm58; fig. 6 similar to fig. 4. a indicates an organism surrounded by vangia telleri (flügel). for explanation of the letter c see plate 1, fig. 7. bm58; fig. 7 similar to fig. 2. bm58; fig. 8 similar to fig. 7. bm58. two problematic organisms from the permian jamal formation of shotori mountains, northeast iran 111 plate 1 112 senowbari-daryan b. & rashidi k. plate 2 two problematic organisms from the permian jamal formation of shotori mountains, northeast iran 113 banner f. t., finch e. m., & simmons m. d. 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# "������ $!�?9 rivista ltaliana di paleontologia e stratigrafia volume 104 numero 3 tavole 1 pagine 381-390 15 dicembre 1998 upper triassic amber from the dolomites (northern italy). a paleoclimatic indicator? piero gianolla*, eugenio f*agazzi** & guido roghi* receioed march 4, 1998; accepted may 20, 1998 key-aords: amber, upper triassic, dolomites, infrared spectroscopy, palynolo gy, paleoclimate. riassunta. in strati terrigeno-carbonatici della formazione di dùrrenstein delle dolomiti è stata rinvenuta ambra sotto forma di goccioline di coiore giallo-rossastro. ii ritrovamento di ammonoidi e di palinomorfi permette di assegnare l'età dell'ambra al più alto julico, in prossimità del limite con il tuvalico (cìrca 225 ma). sono state determìnate le caratteristiche fisico-chimiche dell'ambra; la spettroscopia alf infrarosso (ir) evidenzia ie tipiche bande delle resine fossili. larea caratterizzante dello spettro mostra peculiarità riferibili ad un'unica specie, diverse da quelle delle resìne {ossili conosciute. la grande abbondanza dì microspore teniate bisaccate, nei campionì contenenti ambra (41olo), suggerisce una relazione con la specie produttrice di resina, probabilmente una conifera. oltre a que11i delle dolomiti, sono noti ritrovamenti di ambra anche nel triassico superiore della svìzzera, delle alpi calcaree settentrionali e dell'arizona. la sostanziale isocronia delf intervallo stratigrafico in cui è stata trovata i'ambra e la costanza delle caratteristiche paleoambientali dei siti (fluviale, manno màrgrnale) suggerisce una possibile relazione tra la produzione e conservazlone dell'ambra e un evento climatico umido verificatosi in tali reeioni durante il carnrco, al pass,rggro tr.r julico e tuv:rlrco. abstract. amber in triassic deposits in the dolomites is demonstrated for the first time. the amber-bearing deposits beiong to the middle part of the diirrenstein formation, referred to uppermost julian (lower carnian, about 225 my). chemico-physical features of amber, which occurs as small yellow to reddish droplets, have been determined. infrared (ir) spectroscopy shows typical bands of fossil resins; the "fingerprint" region of the spectrum presents a unrque prttern th.rt cannot be referred to any other known fossil resin. palynological investigation of amber-bearing layers shows a large prevalence of bisaccates and circumpolles. particularly, the tàeniate bisaccates are frequent (+lv") and suggest a correlation with the amberproducing species. amber production and preservation is possibly related to a humid climatic event. lntroduction. in the past, ijpper triassic amber has only been reported in northern calcareous alps (pichler, 1868; yavra, 1984; poinar et a1., 1,993), rn switzerland (soom, 1984) and in arizona (lits/in & ash, 1991). as far as we know, amber has never been described in triassic deposits in ita1y, except for a generic mention of "fossil resin" by zardini (1973). in the present study, we report the occurrence of amber in the dùrrenstein formation in the dolomites (fig. 1). the age of the amber-bearing rocks is latest julian (early carnian). since amber can embed and preserve organisms, such as arthropods and soft-body animals, as well as parts of plants and pollen, it provides a reiiabie tool for paleontological studies. the possibility of preserving nulocation map of upper triassic amber investigated in the present study. '. deprrtment of geology, paleontology and geophysics, university of padova, via giotto i,35137 pado".r (itrly) e-meil : piero@geol.unipd.it 'r'f depanment of pharmacologn university of padova, italy 382 p. gianolla, e. ragazzi & g. roghi santa croce / heiligkreuz section z j l f limestones and biocalcarenites dolomites siltstones and marls sandslones conglomerates unconformity stromatolite cross bedding oolite ammonoid palynomorph z i l --) fig.2 stratigraphic section of the dùrrenstein formation at santa crocelheiligkreuz (badia/abtei valley). cleic acids and other organic molecules is still a matrer of debate (brown & brown, 1,994; cano, 1996; austin et al., 1997). occurrence of triassic amber. triassic amber was first collected ín 1823 in the fossil plant-bearing beds in the schilfsandstein ar neuewelt, near basei. the sample was described by soom (1984) and is kept at the museum of natural history in basel. the schilfsandstein is a fluvial deposit, widespread throughout continental europe, that deeply cuts the underlying sequences (aigner & bachmann, 1992). a re-interpretation of palynological data by leschik in krausel & leschik (1.956) and scheuring (1970), suggests a latest julian age for the amber-bearing layers. the first pubiished reporr on triassic fossil resin is by pichier (1868), who described smal1 resin droplets in the "oberen schichten der cardita crenata" of kot;--n tv i i llr:fi r-r--j l/l -7 -7-l tr--ltl |; -;l l-" ò l ,t\ e, o chental (tyrol, austria). the author gave this resin rhe name "kochenit" and produced a complete chemicophysical analysis. the fossil plant-bearing beds belong to the 1st shale (|erz, 1966; r. brandner, pers. comm.) of the raibler schichten in the northern calcareous alps (nca), which typically includes importanr plant remains associated with coal layers. as far as 'w"e know, these amber-containing levels correspond to those in mt. leitnermoos, near schliersee (bavaria, germany), in which amber collected and studied by poinar et a1. (1993) was found. the amber-bearing beds were deposited in a marginal t'luvial environmenr (bechstadt & schweizer, 1991). according to kavary (1966, 1972) and jelen & kusej (1982), these beds are julian in age. as the 1st shale of the raibler schichten bears carnites flo. ridus (wuifen) and a microflora including patrnaspontes justusklaus, in our opinion its age is latest juiian. triassic amber was also reporred near lunz (niederósterreich, austria) by sigmund in 1937 (quoted by yàvra,7984) and was named "copalin". the amber-containing strata belong to the lunzerschichten, interpreted as pro-delta and delta deposits (tollmann, 1976), drectly correlated to the schilfsandstein (behrens, 1973; aigner er bachmann, 1992). once again this unit is particularly rich in fossil piants (cf. krdusel, 1949; dobruskina, 1988) and is iate julian in age (klaus, 1960; dunay & fisher, 1978). litwin ec ash (1991) also pointed out upper triassic amber in the lower member of the petrified forest formation (chinle group) in the petrified forest national park (arizona). the amber is not associated with petrified wood, but rather with carbonized leaf debris in paper coal. according to litwin et al. (l99i), the age of the amber-bearing strata is late carnian. amber in the dolomites. during the revision of the ljpper triassic stratigraphy in the dolornites, some layers bearing small amber droplets were discovered. all of the material was collected in the dùrrenstein formation (pisa et al., 1980; de zanche et al., 1993) in various localities: santa crocelheiligkreuz in badia valley, rumerlo and rifugio di bona near cortina d'ampezzr.. the di.irrenstein formation is a complex carbonate-terrigenous stratigraphic unit consisting of two distinct interfingered lithozones. the carbonate one is the most typical and widespread, made up of intra-supratidal well-bedded dolomites with stromatolitic, birds-eye and tepee structures. the terrigenous lithozone is characterized by sandstones, conglomerates and hybrid arenites; coquinas are widespread in this part of the unit. the vertical evolution of this lithozone shows a decrease in siliciclastic content and consequent increase in carbonate supply. t a6 0m ljpper triassic amber front the dolomites 383 the depositional environment is mostly a wide tidal flat, sometimes interrupted by lagoons (pisa et al', 1980; de zanche et ai., 1993). from these facies, reptile footprints have been reported (de zanche et ai., 1993), suggesting the existence of a nearby emerged land. the terrigenous sediments are restricted to ephemeral distri butor channels, controlled by tidal currents' and represent the dispersal of fluvial supply in a marginal marine setting. the dúrrenstein formation corresponds to the ultimate phases of the carnian regression: in fact, at that time, triassic basins in the dolomites were more or less completely filled. according to the sequence stratizone (krystyn, 1,973).in the amber-bearing beds, cropping out about 10-15 m beloq the miospores spiritisporites spirabili.s scheuring and patinasporites justus kia:us, have been found. s. spirabilis is assumed to be a typical tuvalian element (scheuring, 1970; blendinger, 1987), as it has never been found in julian layers. f{owever, as far as we know, a direct fitting with ammonoids does not exist. therefore, we don't know if the appearance of s. spirabilis coincides with the base of the tuvalian or if it is a little older. help is given by a sequence stratigraphic correlation between the dolomites and the nca. as the car 3 depositionai sequence ar l:unz (fig. 3) is parmorphology and chemico-physical properties of amber of lhe dolomites. amber found in the described triassic formation occurs as spherical-ovoidal granules (fig +)' generally with diameter of about 2-5 mm; occasionally specimens of a larger size (up to 3 cm) have been found. the colour is from light yellow to reddish, with resinous luster, and conchoidal fracture. the amber droplets are minutely and pervasively fractured. microscopic observation of the droplets shows a rough surface. flardness is 2-3 and specific gravíty slightly greater that 1 (about 1.08). exposed to flame, amber burns producing a resinous odour. particulate samples are almost insoluble in alcohol or ether, and slightly soluble in acetone. after age dolomites eastern nca (lunz) western nca (amder from kochental) neuewelt (switzerland) ne arizona srd-order depositional sequences z z é, o 2 raibl fn,4. ;ì oppontfzer*' schichten rai blersc hi chte n ftoreb schtchten') shales 2b-2c untere kieselsandstein painted desert t4b. sonsela i\,18. hst car 4tst lst hst car 3 * oùrnerusretlt i tti lunzer)* schichten w *o,ro*ourra" * schichten raiblerschichten ('cabdita *. schrchfen') * s shales 1a-1b-1c-2a rote wand hauptstein. mergel t\ *\t schilfsandstei n ittitnrili i rl i o bluei'esamb tdesa redondo fi.,,|, shinarut,4p fm. z f tst lst :) ? il * s. cnsstntlo fm. * gostlinger kalk hst car 2 wetfersteinkalk gipskeuper * ammonoids palynomorphs €' amber -sb -'-'-'-'tlpc ---mfs hiatus fig. 3 sequence stratigraphic correlation of the upper triassic amber-bearìng unlts. graphic interpretation by de zanche et al. (1993), the tly julian, due to the presence of ammonoids of the auwhole dùrrenstein formation corresponds m the car 3 striacum zone (krystyn, 1'991), it is highly probable 3.d-order depositional sequence. that the lower part of the dùrrenstein formation is also amber has only been found in a narrow interval upper julian. of siliciclastics lying in the middle portion of the dùrrenstein formation (fig. 2). although different in details, the amber-bearing layers consist of grey-yellowish hybrid sandstones, rich in plant remains and coal. marine invertebrates, mainly large bivalves and gastropods, are abundant. teeth of fossil fish and fragments of terrestrial reptile bones have been detected' on the whole, the environment can be interpreted as marginal marine. the age of these layers is latest julian, possibly close to the julian/tuvalian boundary (about 225 my according to gradstein et al., 1994).in the upper part of the dùrrenstein fm., in correspondence to the maxi mum flooding surface of the car 3 depositional sequence, the ammonoid shastites cf. pilari (diener) has been found (fig. 2), belonging to the lower tuvaiian dilleri 384 p. gianolla, e. ragazzi & g. roght evaporation of the acetone, amber fragments are covered by a whitish crust. infrared (lr) spectroscopy. ir spectroscopy has been suggested as a merhod ro characterize fossil resins (beck et al., 1964; langenheim & beck, 1965). in the present study, small specimens of amber droplets obtained from different localities were removed from their matrix, avoiding contamination with matrix particles. spectra were obtained in the solid state: after grinding in agate mortar, samples were included in potassium bromide pellets. ir determinations were made just after the preparation of the pellets, to avoid an increase of hydroxyl stretching and bending absorptions due ro kbr hygroscopicity (beck, 1986). a perkin elmer 1600 series ftir spectrophobmeter, within a range of 2.5-1,5.5 pm (4000-645 cm-1), was used. typical ir spectra of two samples of amber are shown in fig. 5. additional spectra of samples from different localities in the doiomites ranged between the two in the figure. the observed differences were of quantitative type and possibly due ro different degrees of preservation. the prominent carbonyl band near 5.8 pm (about 1700 cm-1), which is recognized as uniformly typical of all fossil resins and due to the stretching of carbon-oxygen double bonds (langenheim & beck, 1968; beck, 1986), is present at 5.9 pm (1690 cm-1) (band b). the stretching of carbon-hydrogen bonds produces absorptions near 3.4 pm (2950 cm1) (langenheim & beck, 1968) (band a); bending motions of these same bonds produce absorption near 6.8 prm (1470 cm't) and 7.2 1tm (1380 cm-l) (langenheim & beck, 1968) (bands c and d). fiowever, the spectrum does not show a sharp band near 11.3 pm (885 cm1), characteristic of out-offio 4 amber droplets (a) in hybrid rrenire (rurnerlo area). thrn sectron (6, 3x). plane bending of the two hydroben atoms of a terminal methylene group, also considered as a feature of resin acids (i.e. agarhic and copalic acids) isolated from recent resins (langenheim & beck, 1968; beck, 1986). on the other hand, langenheim & beck (1968) suggest caution when interpreting the absence of this band in fossrl resins, since rermrnai methylene groups are easily oxidized. the region between 815 pm shows a unique morphology. with r main band ar 8 prm (band e in fig. 5) and other minor bands, which may include those of carbon-oxygen single bond(s) at 8-10 pm (1250-1000 cm'1) (langenheim, 1969). the absence of the shoulder, known as "baitic shoulder" (beck et a1,., l964; beck, 1986), in the "fingerprint" region between 8 and 8.5 pm (1250 and li75 cm1), excludes a similarity with baltic amber specrra. palynology. black siltstones without amber and biocalcarenites including amber from different localities within the dùrrenstein formation have been processed. the palynological methodology consisted of a standard preparation procedure (hcl, i#, hnor and 15 pm-mesh sieving). the samples supplied a rich palynoflora consisting of spores (evigate, verrucate), pollen (monosaccate, bisaccate and circumpolies) and abundant cuticular fragments. the assemblage includes typical uppermost julian-tirvalian elements, such as spiritisporites eirabilk scheuring, vallasporites ignacii icschlk, patinasporites justus klars, psewdoenzondlasporites surnmus scheuring, samaropo l/enites speciosars goubin, infemopollenites paruus scheuring, "paracirculina " aerrucosa praehauser-enzenberg dwplicisporites co nttnuus praehauser-enzenbery paracirculina maljawkinae klaus, dwplicisporites peryucosus (leschik) scheuring, dwplicisporites granulatus (leschik) scheuring (pl. 1). as shown in fig. 6, amber-bearing sandy biocalcarenites (sample rumt) and biack siltstones without amber (samples hel1 and scs16) underwent a quanrirarive palynological analysis. a large amount of alete bisaccate pollen has been detected in all samples; other forms show different percentual distribution. the taeniate bisaccate pollen lunatisporites acutus leschik and lueckisporites sp. present an unexpectedly high peak of frequency (a total of 41o/o, berng single values 38% and 3o/o, rumerlo santa croce/heil igkreuz d e c b a upper triassic amber from tbe dolomites 4000 2500 2000 1 500 1 000 cm-t 6 6 7 i i 10 11 12 13 14 um 385 fis. 5 infr.rrerl snect r: of rmber in rhe dolomites. upper spectrum from rumerlo area; lower spectrum from santa croce/heiligkreuz section. a-e: marn brnds (see text ior detail$. a comparison with the triassic amber found in arizona (litwin & ash, 1991) shows, in turn, marked differences. the authors suggest that the botanical source of the arizona fossil resin could be araucarian gymnosperms, because of araucariaceae-like ir spectrum. their spectrum however has very few diagnostic bands, which may be the result of a different taphonomic history rather than of a different botanical source. amber in the doiomites also displays a wide range of band amplitude, possibly as a result of different degrees o[ preservation. regarding the botanical source of amber in the dolomites, quantitative palynologicai data support the possibility of a con ifer oriqin the unusual .rbundance o[ lunatisporiles acutus and lueckisporites sp. corroborates this hypothesis. in australian triassic sedimenfs, lunatisporites acutws has been found .rssociated in sttu with gymnosperm cones (townrow, 1,967; balme, f995) and tentatively referred to the genus rissikia, which represents the oldest certain podocarpacean remains (stewart, 1,983). lweckisporites sp. is associated with the conifer majonica alpina klement-\flesterhof, a permian majonicaceae found in the butterloch area (balme, 1995); another affinity is between this pollen and the conifers pramelrautbia bober' felneri (krasser) found in the triassic ar luîa. flowever, attribution of a triassic fossil resin to a specific taxon is hazardous, because during that time the major groups of coniferales had just begun to differentiate from voltziales (milier, 1977 ; ste:svart, 1983). palynostratigraphic and sedimentological analysis of amber-bearing layers (dunay ec fisher, 1928; klaus, 1960; jelen & kusej, 1,982;praehauser-enzenberg, 1970; scheuring, 1970) suggests a narrow correlation between amber occurrence in several triassic localities (fig 7) along 1oo-30o paleolatitudinal belr (gilbert smith et al., 4 respectively) in correspondence to amber level (sample riimz), thus suggesting a link with the amber-producing species. discussion. comparison of the ir spectrum of our samples to spectra of other fossil resins (langenheim & beck, 1965, 1968; langenheim, t969; beck, 1986) suggests that this resin is a unique kind of amber, in spite of the fact that the spectrum has similarities ftand at 8 pm, 1250 cm-1) with those of present-day pinaceae resins (i-angenheim & beck, 1965). a similarity can also be found with spectra of cretaceous amber in \íashington, d.c. and alaska, interpreted as deriving from a complex forest of coniferales, including podocarpaceae, araucariaceae, pinaceae and cupressaceae-taxodiaceae (langenheim & beck, 1968). 386 p. gianolla, e. ragazzi €r g. roghi 1973; yeevers, 1.994). the first surprising result is the substantial synchroneity of the amber-bearing interval. secondiy, the consistency of the correlation is enhanced by a similar depositional typology (fluvial, marginal-mafig.6 qu.rntitarive p.rlynologìc.rl analysis of 3 samples from rhe amber-belrìng inrerurl: hel 1 and scs 16 from santr croce/heìligkreuz aref,i rum 7 from rumerlo (cortina d'ampezzo). samples hel 1 and scs 16 do not bear rmber. r : rare, from o to 5%; c : common, from 5 ro l5o'o and a : abundanr, 1.5yo. rine) throughout different triassic basins, attesting to a sudden and strong increase in immature siliciclastics, in turn documenting an increase in runoff and therefore in rainfall. although rejected by some researchers, on the basis of a detailed quantitative palynological analysis (visscher et al., 1994), in our ooinion r he idea of a time-limited humid event, which during late triassic broke the prevalent and generalized artd climate (simms & ruffel, 1989r manspeizer, 1994; simms et al., 1995) could be once more taken into consideration. in the dúrrenstein formation, the presence o[ spores belonging to different species suggests the existence of a vegetation of hygrophitic type, thus corroborating the idea of a humid event. above data pose new problems which require further investigations. why does amber only occur in such deposits? why are they so relatively abundant? vhat are the relationships between amber and the assumed humid climatic event? can amber production and/or fossilisation be tentatively related to it? the abundance of amber droplets in the sediments may be the consequence of litoraneal replate 1 i) spiritisporites spirabilis, scheuring 1970, (42 pm); slide scs16 iii,145/3;2) patinasporitesjustzs, klaus 1960, (54 pm); slide scs16 i, k32;3) vallasporrtes ignacii, leschik 1956, (36 pm); slide scs16 i, 528/3; +) enzonalasporites úgms, leschtk, 1956, (38 pm); rum/ ix, n40; 5) samaropollmites speciosus goubin, i965, (ength 56 pm); scs16 ii,c32;6) lueckisporites sp., 1954, qength.72;lm); rumz x, p35;7) lunatisporites acutuslesch;k, 1956, (length 47 pm); rumz ix,p28/2; 8) " paracirculina" oerrucosa praehauser-enzenberg, 1,97q, (45 pm); scs16 i, n31;9) paracirculina tmebrosa c^l^"-:-^ 1a7^ i1a "*\'eas16 i, k30/3; 1.0) duplicisporites continuus praehauser-enzenberg, 1970, (a0 pm); scs16 iv, k35; 11) pseudoenzomla-tsrra/, j\ sporites surnmus scheuring, d7a, (46 plm); scs16 ii,p48/4:12) paracirculina maljaakinae klaus, 1960, (36 pm); scs16 i, n33. coordinates of the figured specimens were taken with the england finder using leitzrf/etzlar n. 5345 with attached camera. all the slides housed in the dipanimento di geologia, paleontologia e geofisica of the university of padova. 40 s30 20 t0 0 alete bisaccates taeniate bisaccates monosaccates md vescicates ovalipolliscircmpolles triletespores monosaccates and ._a*. r l;iòhì[ il igà'usel & lesdi[]036* secatus leschik in krausel & leschik. 1956 krausel & leschik. i in krausel & lesch pl. i upper triassic arnber from tbe dolom.ites 187 r-t 388 p. gianolla, e. ragazzi g g. roghi sin deposition due to a decrease of water salinity, foliowing important rainfall evenrs. the lowered density of marine water may have allowed sinking and deposition of the resin. furthermore, a rapid change in climate may have induced stresses in the plants and therefore an increase in resin secretion. although in the other fluvial and/or marginalmarine triassic deposits in the southern alps fossil rel a t i t u d e frg.7 occurrences of upper triassic amber. the distribution of the sites is scattered along the tropical belt. carnian pangea after gilben smith et al. (1973). a) dolomites; b) nonhern calcareoirs alps and switzerland: c) arizona. sins have not yet been mentioned, the occurrence of amber in the carnian cannot be a fortuitous event. further investigations will establish the consisrency of these considerarions. acknouleclgements. authors thanks proff. vittorio de zanche and paolo mretto for helpful discussion and criticism; thanks are due also to j. a. van konijenburg-van cittert, utrecht, and to an anonymous referee for critical reading and suggestions. authors are grateful to paolo fedele (cortina d'ampezzo), who collected and gave us arnber samples. .{/e thank also dr. adriana chilin for rnfrared determinations. thìs research was sponsored by the c.n.r. cenrro di studio per la geodinamica alpina (padova) and by the m.u.r.s.t. (ex 40%, 1996, resp. prof. i. dieni). references aigner t. & bachmann g.h. 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(1989) syncroneity of climatic change and extinctions in the late triassic. geolog, v. 17, pp. 265-268, boulder. simms m.j., ruffel a.h. & johnson l.a. (1995) biotic and ciimatic changes in the carnian (triassic) of europe and adjacent areas. in n.c. fraser and h.d. sues (eds.) in the shadow of the dinosaurs. early mesozoic tetrapods. cambridge university press, pp. 352-365, canbridge. soom m. (1984) bernstein vom nordrand der schweizer alpen: benstein kettery stutt. betr. naturkd., s. c, n. 18, nn l\-/() \fìrîtùîrt t(' " --t "'""b* ' stewart \f.n. (1983) paleobotany and the evolution of plants. cambridge university press, 405 p., cambridge. tollmann a. (1976) analise des klassischen nordalpinen mesozoikums. franz deuticke, 580 p, vien. townrow j.a. (1967) on rissikia and mataia podocarpaceous conifers {rom the lower mesozoic of southern lands. pap. proc. r. soc. tasmania, v. 101, pp. 103-136, hoban. vóvra n. 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(1973) geologia e fossili attorno a cortina d'ampezzo. ed. ghedina, pp. 26, cortina d'ampezzo. rivista italiana di paleontologia e stratigrafia volume 117 no. 1 2 pls. pp. 161-172 april 2011 morphologic variations of the trace fossil rutichnus in cm-thick turbidites from the verghereto formation (northern apennines, italy) paolo monaco received: october 20, 2010; accepted: december 20, 2010 dipartimento di scienze della terra, piazza dell’università, 06100, perugia, italy. e-mail: pmonaco@unipg.it key words: ichnology, rutichnus, taphonomy, trace fossils, turbidites, miocene, apennines. abstract. the analysis of 56 specimens of the branched, walled post-depositional trace fossil rutichnus, sampled from 31 thin-bedded, overbank turbidites from the verghereto formation, romagna apennines, indicates high morphologic variability of this ichnotaxon in hypichnial preservation. the variability concerns the arrangement of outer and inner walls, number and shape of annulations, distribution of pustules, the (false) branching arrangement and the general external shape. highest variability has been observed mainly in the rutichnus rutis ichnospecies, clearly less in the r. irregularis ichnospecies that is rare. the dino-lite microscope camera analysis of oriented thin sections indicates that endichnial preservation exhibits many structures due to burrowing in soft substrate, destroying laminae of sand. consequently, rutichnus probably was produced by a worm or arthropod, inducing strong deformation in turbidite sand during the feeding activity on phytodetritus and organic matter transported by the turbidity flow. riassunto. vengono analizzati 56 campioni della traccia fossile post deposizionale rutichnus provenienti da 31 livelli centimetrici (da 1 a 5 cm di spessore) di calcareniti torbiditiche di overbank intercalate nelle marne della formazione di verghereto, affioranti presso la località di ville di montecoronaro (verghereto, appennino romagnolo). l’esame mostra una grande variabilità morfologica in questo ichnotaxon, specialmente nella preservazione hypichnia. la variabilità nella specie rutichnus rutis è stata analizzata nella forma esterna, nelle pareti esterna ed interna, nel numero e nella disposizione delle annulazioni, nell’aspetto delle (false) ramificazioni e infine nella distribuzione delle pustule. lo studio della variabilità morfologica è stato meno agevole nell’altra specie r. irregularis, a causa della scarsità degli esemplari raccolti. l’analisi mediante camera microscopica dino-lite in sezione sottile ha permesso di evidenziare anche una forte variabilità nella preservazione endichnia; si osservano molte deformazioni avvenute durante la fase post-deposizionale nelle lamine di sabbia alla base delle calcareniti e nella disposizione dei granuli associati, prodotte dalla ricerca e dall’ingestione di fitodetrito e materiale organico da parte di organismi vagili, forse vermiformi (ma senza escludere artropodi), trasportati dai flussi di torbida. introduction for a branched, walled and meniscate trace fossil d’alessandro et al. (1987) introduced the ichnogenus rutichnus, which typically displays wrinkles that are distributed in the exterior of the wall. these authors analyzed many samples from the permian of east greenland and from the miocene flysch (gorgoglione) of southern italy remarking peculiar morphologic characteristics of this walled ichnogenus: external roughness, branching and the occurrence of menisci. branches and menisci are typical and well preserved mainly in the permian specimens (d’alessandro et al. 1987, figs. 6-7). some similarities concerning the wall ornamentation have been found mainly in epichnial nereites, although in hypichnial neonereites the wall is thicker and formed by pustules without short annulations, while in rutichnus annulations and roughness are more regularly arranged and the wall is isopachous. samples of one of the two ichnospecies, rutichnus irregularis, formerly classified as radionereites irregularis by d’alessandro (1982), has been collected as hyporeliefs from soles of turbidites in the gorgoglione flysch deposits, basilicata region, southern apennines, while the holotype of the other ichnospecies, rutichnus rutis, comes from the permian of east greenland (domkirken mount, northern scoresby; d’alessandro et al. 1987). 162 monaco p. this paper addresses to describe an unusual concentration of rutichnus trace fossils that is otherwise rarely known (uchman & demircan 1999; demircan 2008). the highly abundant rutichnus has been found in several 3-5 cm-thick calcarenitic turbidites in marly deposits of the verghereto formation (miocene) of the romagna apennines (ville di montecoronaro, verghereto area). these calcarenitic turbidites probably represent distal fringe (overbank?) sediments deposited in the marginal marnoso-arenacea basin at the transition to the verghereto high (milighetti et al. 2009; monaco et al. 2009). these deposits belong to an area that expresses an excellent preservation and abundance of trace fossils with 50 different ichnotaxa in the ichnofossil-lagerstätte of poggio alto (monaco & checconi 2010). this paper is directed to describe features of hypichnial rutichnus that had not been described before, focusing some taphonomic aspects that can help in the definition of palaeobiological and palaeoenvironmental setting of very thin-bedded turbidites. trace fossils have been sampled bed by bed and have been observed in variably cut surfaces which were photographed. analysis of endichnial preservation was performed mainly using the microscope camera dino-lite at 10x and 20x magnification, in order to observe textural arrangement of grains close to the burrow. investigated locality and geological setting the verghereto formation, miocene in age, consists of thick, rhythmically-arranged gray marl deposits (up to 600 m thick towards ne at m. fumaiolo) and sporadic calcarenitic turbidites. the verghereto formation was deposited in the marnoso-arenacea basin during a multi-phase tectonic activity that in the lowermiddle miocene (serravallian?) produced ridges and elongated basins in a nw-se oriented foredeep system of the northern apennines. in the verghereto area, that represents probably one of the most important of these intrabasinal highs, fringe and overbank deposits are represented by thin-bedded and fine-grained turbidites that reached the marginal sectors of the marnoso-arenacea basin (see the extensive geological description in monaco et al. 2009; monaco & checconi 2010). in the poggio c site (n 43°76’93,63”; e 12°03’55,83”), that is located about 1 km at south of poggio alto ichnofossil-lagerstätte (monaco & checconi 2010), about 8 m of section have been logged (fig. 1). thirtyone calcarenitic beds, from 1 to 5 cm thick, have been analyzed bed by bed for the ichnofaunal content (fig. 2). seven of them exhibit rutichnus rutis which is very common in thicker turbiditic beds (see five asterisks in fig. 2). in the locality of poggio b (n 43°77’14,24”; e 12°03’06,05”), just 250 meters to the fig. 1 localization of studied sections. (a-d) marls and cm-thick calcarenite beds in the poggio c section. numbers in beds are explained in figure 2. (e) epichnial community (mainly scolicia prisca, sp). rutichnus from the verghereto formation (n. apennines, italy) 163 north of poggio c, eleven samples of rutichnus rutis and six samples of rutichnus irregularis have been collected (fig. 1). the poggio b section requires further analyses for its very rich assemblage of trace fossils, and therefore its log and ichnologic data will be provided separately. in the locality of poggio c calcarenitic beds of 1 to 3 cm thick dominate, but the thickest of them (levels 5, 13, 27), are those with the highest abundance and diversity of ichnofauna (fig. 2). calcarenites consist of rhythmical sharp-based beds, finingupward disposed and brown to yellowish in colour. quartz, mica flakes, feldspar, lithic fragments, calcium carbonate and clay suggest immature arenites (milighetti et al. 2009). the basal grain size remains in the range of medium sand while the top is silt or mud; ripple structures are frequent in thicker beds 5, 13 and 27, disposed internally, more rare at the top. both base and top are strongly bioturbated (monaco & checconi 2010). the cm-thick calcarenites correspond to the distal expression of the facies f9b, that is a variety of the facies f9a in the model of mutti (1992), probably an immature deposit of very small-volume flows from low-density turbidity currents in marginal and very distal environments (see discussion in monaco 2008; monaco & checconi 2010). the material studied was collected at south of the village ville di montecoronaro at the foot of a hill that is located about 2 km to the north of the canili-piantrebbio exit in the e45 orte-ravenna road (romagna apennines). trace fossil assemblage the trace fossil assemblage of the poggio c section consists of rich hypichnia and epichnia in addiction to endichnia that are considered separately (fig. 2). hypichnial assemblage is dominated by rutichfig. 2 the poggio c stratigraphic column with ichnologic assemblage. numbers in circle are thin-bedded calcarenites. asterisks indicate thicker beds where the most abundant concentration of trace fossil rutichnus occurs. 164 monaco p. nus rutis and r. irregularis, and biramous meandering graphoglyptid desmograpton that in this area is the most abundant among graphoglyptids (monaco 2010). in the calcarenitic levels 5, 13, 15, 18, 19 and 27 desmograpton can reach a very high ichnodensity, up to four specimens/20 cm2. the dominant ichnospecies is d. dertonensis while d. ichthyforme and d. alternum are less abundant. other graphoglyptids include megagrapton, while net-shaped paleodictyon is scarce (p. strozzii and p. minimum). bergaueria-like structures and many indeterminable small plug-shaped structures dominate in thicker levels 5, 9, 13 and 27, while sporadically they occur in thinner ones. these plug-shaped forms are frequently fluted mainly in thicker levels (5, 13 and 27). among string-shaped forms helminthopsis (mainly h. tenuis), and protovirgularia are the dominant ichnogenera, while scolicia strozzii, which is abundant in the ichnofossillagerstätte of poggio alto, in the poggio c and b sections is very rare or absent (1 dubitative specimen at level 5). the epichnial community of thicker levels 5, 13 and 19 is similar to that described in the ichnofossillagerstätte of ville di montecoronaro (poggio alto) (monaco & checconi 2010), while some differences exist in thinner levels (see fig. 2). in fact, while scolicia prisca and nereites isp. or n. missouriensis are the dominant forms in thicker and rippled beds (monaco & checconi 2010), the epichnial community is dominated by delicate forms as chondrites (c. intricatus, c. targionii), pilichnus and phycosiphon that are abundant in the sharp top of thinner calcarenites that not exceeding 1-2 cm in thickness (fig. 2). fig. 3 thin sections of rutichnus in dino-lite microscope camera. (a-f) endichnial features are irregular deformations in the sand laminae which are curved or irregularly arranged bands forming micro-convolute structures or micro-patchy structures. further explanations in the text. rutichnus from the verghereto formation (n. apennines, italy) 165 morphologic variations and taphonomic aspects of rutichnus in endichnial and hypichnial preservations in order to characterize the morphologic variation and taphonomic aspects of rutichnus rutis, 56 samples have been analyzed (fig. 2). the ichnospecies r. irregularis from the poggio b locality has not been considered in the morphologic analysis (tab. 1). since each studied rock sample usually exhibits different single specimens of r. rutis (from 1 to 8 see column 2 in tab. 1), a total number of 174 single structures are present at sole of calcarenitic beds (tab. 1). they are studied in hypichnial preservation, since the endichnial structures show a strong morphologic variability and must be analyzed separately, adopting an approach that consists of the newly available dino-lite microscope camera. this camera obtains a continuous spectrum of photographs from 5 to 100x magnifications, and can be very useful in cases of structures that were produced by displacement of sand grains induced by burrowing. image contrast and other settings may be enhanced by photoshop. to study endichnial deformations numerous thin sections were produced and analyzed, oriented transversally and longitudinally (e.g. samples mv339, 345, 349, 389, fig. 3). all studied samples are in the repository of the biosedimentary laboratory of earth science department of the perugia university. the deformation structures affect the lower part of a calcarenite (usually 2-3 cm in thickness from the base) but do not correspond to those preserved at the base of a bed. endichnial features consist of irregular deformations in the sand laminae that appear as curved or irregularly arranged bands forming micro-convolute structures (sample mv339, fig. 3a arrow). above the concavity, which demarcates the exterior wall of a hypichnion, a characteristic cutting edges of sand laminae is present which bend upward and return gradually to flat position at the top (fig. 3a). the core structure, which resembles a bean placed vertically in the central position, is deformed itself by the biogenic compaction during the burrowing (fig. 3a). laminae may show a micro-patchy arrangement (samples mv345, fig. 3b). vertical dislocations can be seen in the sample mv349 (fig. 3c). analysis by dino-lite camera confirms that the endichnial preservation of rutichnus rutis never shows a well-defined, repetitive structure but can easily vary from case to case. in contrast hypichnial preservation exhibits quite different characteristics such as variations in diameter, external and internal shape, wall preservation and wrinkle disposition (tab. 1). the variability of rutichnus rutis is very high, mainly in the marker level 5 (fig. 2), where the ichnodensity is highest, reaching up to 10 specimens for a square decimetre, and decreases in other beds (see asterisks in the log of fig. 2). the diameter of this ichnospecies is usually about 10 mm (occasionally 12 or 13 mm, tab. 1); a narrowing of the tube has been commonly observed (7-9 mm), while expansions of tunnels (up to 15-17 mm) represent probably abortive branches (mv343, 384). in rare cases the point of expansion can be flattened (mv340, 343). external diameter appears much more constant in rutichnus rutis than in r. irregularis that occurs in bundles and shows a complicated arrangement in hypichnial preservation. fig. 4 graph showing morphologic characteristics of the endichnial rutichnus rutis in 56 samples. 166 monaco p. it shows a great variation in morphology with ringshaped structures (sample mv370, 374, pl. 2c), which are developed in different part of a single structure. the ring arrangement in r. irregularis has been also shown in the study of d’alessandro et al. (1987, figs 2-5). the shape variation of tunnels of rutichnus rutis may changes varying from straight (pl. 1c-d), displaying a winding course (pl. 2b), or curved (pl. 1b, h; pl. 2a). branching in particular is typical of 60-70% of specimens, as indicated by d’alessandro et al. (1987), although probably may be commonly a false branching (see discussion). tunnels usually depart at angles of 30° to 40° (pl. 1b), reaching occasionally higher values (80° to 90° in a few specimens, pl. 1d). the branching is commonly dichotomous. while a no preferred direction has been noted in curved and straight specimens, in up to 30% branched specimens the disposition of tunnels is roughly radial (pl. 1h). in 25% of specimens with dichotomy, the disposition of tunnels is side by side, running tangentially with sudden changes in directions and also in diameter (e.g. sample mv324, pl. 1l). in this case narrowing and enlargement of tube segments are frequent. overcrossing is another interesting aspect that has been noted in 23 samples (tab. 1). this feature consists of superimposition of many knobby parts over smoother ones (e.g. samples mv365, 332, pl. 1a). usually, the lower trace fossil in the hypichnial assemblage is bent downward, overlapping to a previous one; the geometry of the tube and the shape of tunnels seem indicate original overcrossing of burrows without compactional features (e.g. sample mv365). the ending of the tube is another very typical feature of rutichnus rutis; it consists of a short and thin bulge, usually 4-5 mm wide, that expands sharply in two rings 10 mm wide, made of elongated pellets (samples mv324, 333, 362, pl. 1e-f). the end has been found in almost 40% of samples and this feature has figured also in the holotype of the east greenland (d’alessandro et al. 1987). the ending structure in rutichnus irregularis is similar, although much more gradual and without sharp enlargements of rings of pustules (pl. 2e, arrow). the outer wall surface of rutichnus rutis is another very interesting aspect, with changes in their preservation (fig. 4). three types have been observed: (1) annulated with oriented and elongated pustules (type 1), (2) poorly annulated with casually oriented pustules (type 2) and (3) smooth to lumpy with few and scattered pustules (type 3) (tab. 1; fig. 4). (1) in the type 1, annulations are well developed (pl. 1a, b, h) and form somewhat imbricated folds 2-4 mm thick (sample mv326, pl. 1h), with usually equidistant rings of pustules (2 3 rings, rarely more rings/ cm2). in some samples, annulations are rhythmically samples rutichnus individuals tf arrangement lenght cm (mean) diameter (mean) n° annulations/ cm2 distribution of pustules n° pustoles/ cm2 longitud. striation type of relief concave relief inner wall branching mv324 7 radiated 2 to 5 1.10 3.00 fractal 32.00 y convex y smooth 60° mv325 3 radiated 2 to 4 1.20 2.00 fractal 38.00 y convex 80° mv326 6 radiated 4 to 7 1.40 2.00 oriented 36.00 y convex 90° mv327 1 straight 1 to 2 1.00 2.00 fractal 34.00 convex mv328 1 straight 5 1.00 5.00 oriented 28.00 y convex y smooth mv331 5 curved 2 to 6 1.30 2.00 fractal 39.00 convex y smooth 40° mv332 4 branched 3 to 7 1.20 2.00 oriented 26.00 y convex y smooth 40°-90° mv333 3 straight 2 to 6 1.30 3.00 oriented 30.00 y convex y irregular 40° mv334 2 irregular 3 to 5 1.00 3.00 oriented 32.00 y convex y smooth mv335 4 irregular 2 to 3 1.10 2.00 fractal 33.00 convex mv336 2 irregular 2 to 3 1.50 2.00 fractal 27.00 convex y irregular mv337 2 curved 8 1.10 2.00 oriented 32.00 y convex y smooth 30° mv338a 2 side by side 5 to 7 1.00 2.00 oriented 33.00 both y smooth turn mv338b 2 straight 1 to 4 1.20 2.00 fractal 34.00 convex mv339 6 straight 2 to 7 1.10 3.00 oriented 35.00 y convex y irregular mv340 5 branched 1 to 5 1.20 3.00 oriented 35.00 y convex y 90° mv341 2 spoon-shaped 4 1.10 2.00 28.00 concave y ridges mv342 1 curved with ending 9 1.20 3.00 oriented 29.00 y convex turn mv343a 1 curved, enlarged 6 1.20 3.00 oriented 26.00 y convex mv343b 2 ending, irregular 1 to 2 1.10 2.00 oriented, elongated 28.00 convex flat mv344 2 straight 2 to 8 1.20 2.00 oriented, elongated 27.00 both y smooth mv345 5 curved, aligned 3 to 4 1.10 2.00 oriented, elongated 32.00 y convex turn mv346 5 irregular 2 to 5 1.00 2.00 oriented, elongated 31.00 y convex y irregular 40° mv347 1 enlarged 8 1.30 2.00 oriented 32.00 y both y irregular mv348 3 curved, side by side 3 to 5 1.00 2.00 oriented, elongated 30.00 y both y irregular mv349 4 curved, side by side 3 to 8 1.40 2.00 fractal 28.00 both y irregular mv350 2 curved 1 to 2 1.10 2.00 oriented 38.00 y both y smooth 40° tab. 1a, b main morphologic variations analyzed in 56 specimens (174 single structures) from poggio c and b sections, verghereto marls formation (romagna apennines). rutichnus from the verghereto formation (n. apennines, italy) 167 arranged showing thickest rings alternated with thinner ones (see sample mv324, pl. 1b). in some specimens rings are obliquely arranged (e.g. sample mv326, pl. 1h). the distribution and shape of pustules in type 1 are variable: in the majority of analyzed forms (70%), pustules are elongated and oriented orthogonally to annulations, while in few cases (30%) they are short even if oriented (pl. 1a, f, arrows). (2) in the type 2, annulations are very poorly developed and pustules are never oriented, usually knobshaped and larger than type 1 (up to 1 mm thick, samples mv333, 334, 345, 365, pl. 1e, g, l); in many cases they appear with different sizes producing a fractal disposition (up to 80%). (3) in the type 3, the outer wall surface is usually fairly smooth and suddenly becomes lumpy with few and scattered knobs (pl. 2a, b). the outer wall surface of type 3 resembles the exterior surface of spirophycus bicornis, although the spiral whorl forming a convolute ring has never been found in the studied specimens from the verghereto area (monaco & checconi 2008). the three types may occur in the same sample. occasionally a transition from type 1 to type 3 has been observed in the same sample (e.g. mv332, pl. 1a), as gradual transitions from type 1 to type 2 and vice versa can exist (sample mv324, pl. 1b). in all three types the number of pustules for square centimetre has been measured, even if the measure is easier in type 1 and type 2 than in the other type (tab. 1). the analysis indicates values of 21 to 38 pustules/cm2, with 5 to 7 pustules of type 1 that occur in the same line; frequently, the arrangement of pustules in type 1 produces elongated and oriented lines of pustules that are concentrated in 3 rings/cm2 (tab. 1). in other specimens they are arranged in two rings/cm2. in all these cases longitudinal alignments have been observed showing a high degree of regularity. an alignment has never been observed in the type 3. the distribution of relief (full or semi-relief) has also been analyzed: in the 66% of studied samples rutichnus rutis occurs in convex full relief, while in 33% of samples it appears in concave semi-relief. both types of preservation can be present in the same sample and usually they are randomly distributed (50%). in the mv379 2 straight 2 to 7 1.20 2.00 smooth prevalent 25.00 convex mv380 2 curved 8 to 10 1.40 2.00 smooth prevalent 22.00 both y smooth 50° mv381 3 straight 4 to 6 1.50 3.00 fractal 28.00 both y smooth mv382 3 curved, branched 4 to 8 1.40 2.00 oriented, elongated 35.00 y convex y smooth 70° mv383 3 curved, ending 3 to 5 1.10 2.00 irregular 22.00 both y irregular mv384 2 branched, ending 4 to 7 1.20 2.00 irregular 20.00 both y smooth 30° mv385 1 curved 7 1.10 2.00 irregular 21.00 convex total: 174 samples rutichnus individuals tf arrangement lenght cm (mean) diameter (mean) n° annulations/ cm2 distribution of pustules n° pustoles/ cm2 longitud. striation type of relief concave relief inner wall branching mv351 3 spoon-shaped 2 to 4 0.90 2.00 28.00 concave y smooth mv352 1 ending, curved 6 1.20 3.00 oriented 32.00 y convex mv353 3 side by side, tangential 4 to 6 1.30 2.00 fractal 27.00 convex y irregular mv354 1 spoon-shaped 4 1.10 2.00 oriented, elongated 26.00 convex y smooth mv355 4 spoon-shaped 1 to 2 1.10 2.00 28.00 concave y smooth mv356 3 spoon-shaped 2 to 8 1.00 2.00 oriented, elongated 27.00 y both y smooth mv357 3 side by side, tangential 2 to 4 0.90 3.00 oriented 33.00 convex mv358 1 straight 5 0.80 2.00 28.00 concave y irregular mv359 4 curved, branched 4 to 6 1.10 3.00 fractal 35.00 convex 50° mv360 1 straight 8 1.00 3.00 oriented, elongated 29.00 y both y irregular side mv361 1 enlarged 5 1.50 2.00 oriented, flattened 25.00 convex mv362 3 spoon-shaped, ending 3 to 5 1.60 2.00 oriented 33.00 y convex y smooth mv363 7 ending, irregular 2 to 6 1.30 2.00 oriented, elongated 35.00 y convex y smooth 40* mv364 1 irregular 4 1.00 2.00 oriented 29.00 concave y smooth mv365 6 spoon-shaped, ending 1 to 4 1.30 3.00 oriented, elongated 31.00 y convex y smooth, ridges 20° mv366 3 spoon-shaped, ending 2 to 5 1.50 2.00 oriented, elongated 26.00 y convex y smooth turn mv367 3 smooth, irregular 5 to 7 1.00 2.00 28.00 concave y smooth 30° mv368 5 side by side, tangential 3 to 10 1.00 2.00 oriented, elongated 35.00 convex y smooth 30° ma73 2 curved 3 to 5 1.20 3.00 fractal 36.00 y convex y smooth 30° mv376 8 irregular 3 to 10 1.00 3.00 oriented, elongated 35.00 both y irregular 30° mv377 3 curved, ending 6 to 9 1.20 2.00 oriented 33.00 y convex 40° mv378 9 curved, ending 3 to 10 1.10 2.00 oriented, elongated 30.00 y both y irregular 168 monaco p. hypichnial concave semi-relief a spoon-like shape (samples mv334, 341, 351, 354, 355, 362) frequently occurs with the interior wall surface usually smooth (about 60% of samples; pl. 1i). longitudinal ridges in the hypichnial concave semi-relief are other very peculiar features. they are 3 to 5 short ridges, 0.2-0.3 mm thick, parallel oriented into direction of the tracemaker locomotion (see sample mv341, 365, pl. 1i, arrow). these parallel ridges are enigmatic structures, probably produced during the contractions of the body of the animal; however, it cannot be excluded that they were formed by a wormlike organism during the movement by anchoring or elongation (d’alessandro et al. 1987). when these ridges are present, an irregular development of interior wall surface can also occur (samples mv349, 368, 376). as the rule an unstructured mud infilling is present that rarely shows longitudinal striations (samples mv332, 333, 339, 345, 354, 378, pl. 1m). the sectioned meniscate interior is very poorly preserved in the studied specimens of the verghereto formation (only three doubtful cases, samples mv356, 367, 381), while it has been commonly found in holotype specimen from east greenland (d’alessandro et al. 1987, fig. 7). the poor preservation of menisci in the apennine specimens resulted probably from diagenetic processes that destroyed the meniscate structures or by the infilling of the gray mud that hides this structure. discussion and conclusions a model for the mode of construction of rutichnus structures has been inferred by d’alessandro et al. (1987). these authors classify rutichnus as a postdepositional feeding structure produced by a worm-like deposit feeder organism, that constructed and reinforced tunnels during their feeding activity in many steps of repeated forward and backward body movements. the meniscate structures represent different phases of anchoring and elongation of the body; menisci in studied specimens of verghereto are very poorly preserved, so it is difficult to understand their origin. on the contrary, the branching is a more clear feature than menisci. in the sketch with a sectional view of rutichnus rutis d’alessandro et al. (1987, fig. 8) show the structure of branching, indicating that it is not typical as the case of many other trace fossils (e.g. thalassinoides, crustaceans), but develops as a false branching with overcrossing of a different tunnel. usually, the new tunnel departs at the branching point forming a different direction and their external wall cuts the previous burrow. this fact is evident in the figure 9 of the same authors where a false branching is clearly visible in the centre of the photograph. the false branching may be really a normal structure in rutichnus rutis when a changing in direction occur, as observed in many of 22 specimens from the verghereto study (see pl. 1b, tab. 1). the mode of construction of external ornamentation is another important element; it was not considered in detail by cited authors, although the anchor pressure of the soft body may produce irregularities of the outer surface and ornamentation, mostly during the backward movement. these cited authors in their historical treatment compared rutichnus to other trace fossils mostly in respect to branching, meniscate and walled aspects, but without showing the meaning of morphologic variations in rutichnus and the mode of the wall exterior fabrication. this latter aspect is fundamental for understanding the ethology of wall construction, as in the case of some hypichnial nereitids (dreginozoum preservation, seilacher 2007), where the burrow exterior shows remarkable similarities with that of rutichnus. concerning the mode of construcplate 1 hypichnial preservation and morphologic variations of the trace fossil rutichnus rutis, poggio c (verghereto marls formation). (a) overcrossing of type 1 (annulated with oriented and elongated pustules) over type 3 (and lumpy with scattered pustules), sample mv332; (b) false branching in walled specimens of type 1; note rhythmically arranged rings of pustules showing thickest rings alternated with thinner ones, sample mv324; (c) poorly annulated specimen with poorly oriented pustules (type 2), sample mv345; (d) some specimens with different preservation of rings of pustules; note imbrication (arrow), sample mv340; (e) different specimens with small pustules that show fractal arrangement (lower arrow) and end of trace (upper arrow), sample mv333; (f) detail of a end with sharp change in diameter (arrow), sample mv362; (g) specimens showing inner concave (black arrows) and outer convex (white arrow) wall surface with widening and irregular pustules; note elongated, parallel ridges in inner wall surface (lower arrow), sample mv334; (h) radiated, annulated specimens with imbricated folds of oriented pustules, sample mv326; (i) inner wall surface of concave specimens with smooth surface and parallel ridges (arrow), sample mv341; (l) tangentially oriented specimens with short ends, sample mv365; (m) specimens with mud fill and longitudinal striae (arrow), sample mv378. plate 2 hypichnial preservation and morphologic variations of rutichnus rutis and r. irregularis trace fossils, poggio c (verghereto marls formation). (a-b) the outer wall of r. rutis type 3 is usually fairly smooth and suddenly becomes lumpy with scattered knobs; note poorly developed annulations in b, samples mv342 and 359; (c-d) specimens of rutichnus irregularis with characteristic central ring (r), sample mv370; (e) irregular arrangement of a dubious specimen of rutichnus irregularis?, sample mv373; (f) massive, irregular occurrence of pustules (patchwork) of rutichnus cf. irregularis, sample mv372. rutichnus from the verghereto formation (n. apennines, italy) 169 plate 1 170 monaco p. plate 2 rutichnus from the verghereto formation (n. apennines, italy) 171 tion of nereitids, seilacher (2007) suggested that the annelid worm can remove the sediment in front but not by radial emersion of a voluminous proboscis, but rather probing and backfilling the rejected sediment laterally around the body. the cited author indicated that it is possible for the worm by bending the head with small proboscis in different directions and at regular intervals producing oriented pellets. arthropods are excluded by cited author because faecal pellets are nearly the size of the body of the worm. in the case of rutichnus from the verghereto area it is not clear if the material is effectively faecal because it is highly variable in size and distribution, frequently fractal (see three types indicated above), and pellets are smaller than the size of the body. the parallel ridges in the concave wall interior, which are impressed also in the mud fill, are similar to scratch marks produced by arthropods (curran 2007; giannetti et al. 2007). the absence of meniscate structures and the high variability in shape and regularity of pellets distribution suggest that arthropods are not excluded, although a worm-like deposit feeder could be most probably the tracemaker. another aspect that should be discussed concerns the causes of unusual abundance of rutichnus at the lower sand/mud interface (hypichnial position) of thicker beds when also strong endichnial activity is common. the most concentration of specimens occur in such beds (3 to 5 cm thick) that exhibit most abundant endichnial structures, indicating an intense burrowing perhaps at different tiers in the sand (intense postdepositional activity); on the contrary, in thinner beds (of 1 to 2 cm) the concentration of rutichnus decreases abruptly (few and scattered specimens) and endichnial activity is shallow and rare; in this case other trace fossils prevail such as graphoglyptids (e.g. desmograpton) and plug-shaped forms. in any case, may be excluded the role of erosion induced by turbidity currents that probably was irrelevant and did not involve rutichnus tracemakers. it is worthy to point out that this ichnogenus was never found in any other parts of the marnoso-arenacea basin, while locally becomes extremely abundant, as at the transition to the verghereto high, when marginal, cm-thick calcarenites were deposited (milighetti et al. 2009; monaco et al. 2009). the unusual abundance can be due to the availability of a large amount of organic matter and bioor phytodetritus that could be accumulated rapidly by thin gravity flow inducing an unusual proliferation of deposit feeders (lutz et al. 2007; uchman 2007; wetzel 2010). sudden fluctuations in the organic matter on the sea floor and changes in detritus abundance induced effects on the benthic habitats as pointed out by wetzel (2010), reflecting differences in the trace fossil distribution. in several beds we can observe upand downward shifting of tunnels with overcrossing of many parts of rutichnus, with sharp changes in direction, in thickness and in morphology at lower surface of beds. this has been observed also in endichnia where different types of sand laminae deformations occur. this trace fossil arrangement seems reflect a frenzied behaviour of opportunistic animals during a very short time, perhaps arthropods or worms, probably transported directly by low-density gravity flows; in this case, the survival of such animals during transport and deposition from nearby areas could not be excluded. in this case calcarenites may not represent a typical distal flow deposit but could be material coming directly from proximal areas. unfortunately, there are no physical evidences (e.g. tool marks) to validate this hypothesis and remains the open question how to estimate the effects of currents on the distribution of organic matter that surely had to be present along the slopes of the verghereto high. ripples at top of calcarenitic beds with widely developed epichnial nereites missouriensis and scolicia prisca (or s. vertebralis) suggest that concentrations in the organic matter persisted even at the top of beds. acknowledgements. the field work was carried out with the help of t. trecci and m. milighetti. many thanks to reviewers a. uchman and a. wetzel for many important suggestions and improvements in the manuscript. thanks also to m. gaetani and c. lombardo for help and advices. this research was supported by research project rb 2008-2010 of the earth science dept. of the university of perugia (p. monaco). 172 monaco p. curran h.a. 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(2010) deep-sea ichnology: observations in modern sediments to interpret fossil counterparts. acta geol. pol., 60(1): 125-138. r e f e r e n c e s boncheva 329..356 ����������� �������� ��� ����� ����� ��� ���� ���������� ���� ������� ���� ���� �� � �� ������� ���� �� ������ ������ � � � ������ ������ ������ � � ��� ����� � �� ���� ������ � ������� ������� ��������� �� � � � � ����� ��������� �� � � �� ���� ��� ���� � ���������� �������� ��!�� �"��"�! #��$������ ���% ��� #��$������ ����"��� ��$�" " �� �� ��������� ����& �� �����& �������� � '���� �� � ����( ���� �� �� �� �" $��� '�$)( � � *! �' � �� ��� � �'��� � �'���� �" �������� ��!� �� ����& �" ������+��)"�' ��� ���������+��)"�' ���� �� �" � ��$ ��� �, �(-��+ �" � ,����� '������� )(�����$ +�, ,�� �' ���!� �" �!������ �� �"�((�. .�� � � )��������( ,�(!����& �" � �� �"���)���!'��, ( , (� .��" '�������� �� �" �������� �!� � '���� ���+��+ �� �+ ���$ ��(��!�������� �� �" ��) �� ���"/����� 0��. � ��� ��� � 1 ������ �� ���� � 2�� �3& �" � �� �'��' ,�� �' ���!� �" ���� ��� ��� )� � �' 1 ��� 4 ���-(,������ ������� �� ��� ����� � ���� '������� 2�� � �� �"��"�! #��$����� ��� !������� �� ���� � ��� ��� � )�����( ���� ��� ��� �� ������ #��$����� "�, � � ��� ����� �& �" � '������� 2�� � �� � )��� � ��� �" ����� ��$ �� �"�� �� �& � '�������( (�$ ���� 1 / � � "���2��� �� ���' ��( �� �" ��!�� � �� � �� . 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acceted december 10, 1998 key-uords: ammonoids, paratirolites, bellerophon fm., upper permian, dolomites. riassunto. viene descritto il ritrovamento, il primo della teti de occidentale, di un esemplare di paratirolites. l'ammonoide, raccolto nel detrito della fm. a bellerophon nei pressi di santa cristina valgardena (bolzano, dolomiti), è conrenuîo in una piccola lastrina di calcare marnoso, nerastro, della facies badiota, con una microbiofacies dorninata da ostracodi. sulla base del litotipo e della biofacies si presume che esso possa derivare dai livelli dell'associazìone ad ostràcodi, sìtuati pochi metri sotto a quelli dell'associazìone a cornelicania e nanhinella. non si può escludere tuttavia una provenienza da livelli più bassi, sempre comunque appartenenti alla facies badiota. questo ritrovamento consente di stabilire, contrariamente a quanto sostenuto da alcuni autori, che l'associazione a comelicania e nanbinella è ptù recente della zont a phisonites comelicania (:gruntalli na) della transcaucasia e che essa è correlabile o è pir\ giovane della zona a paratiro/ires del dorashamiano suo. abstract. the first discovery of paratirolites in the western tethys is here described. the ammonoid was collected in the debris of the bellerophon fm. near santa cristina valgardena (bolzano, dolornites). the specimen is contained in a small slab made of marly dark grey limestone with a biomicrofacies dominated by ostracods. it probably comes from the ostracod assemblage beds (badiota facies), which are situated few metres below the comelicania beds. the occurrence o[ paratirolites allow us to dàte this segment of the upper bellerophon fm. to the late dorashamian, and to deíìne the conteli cdni.l ànd nankinella beds of the dolomites as younger than rhe pbí sonites comelicania (:gruntallina) beds of the transcaucasia (basal dorashamian). these dolomite beds are equivalent ro or younger than the paratirolites zone (ate dorashamian). introduction. in the dolomites, the upper permian is represented by the bellerophon formation which was formed in a shallow marine environment, mostly inhabited by benthic organisms. ammonoids of the bellerophon formation of the dolomites are thus veíy rare. to date, only few specimens of paraceltites sextensis (diener) have been found at the end of the last century in the sesto (sexten) area (diener, 1897). ijnfortunatly, such an endemic species does not al1ow a direct correlation with the ammonoid standard zofiafion of the late permian. the chronological framework of the bellerophon fm. has therefore been made using other taxa, mainly forams and brachiopods, by means of which a correlarion with the ammonoid-bearing sequences has been artempted. however, the ages obtained from these fossils are often approximative and lack a general consensus, among different authors, on their precise chronostratigraphical posirion. the majority of authors consider the bellerophon fm. dzhulfian/changxingian in age, even if the position of the boundary between these two srages has not been detected, as most of the formation was deposited in a restricted shallow mr.rine environmenr in which fossils are rare, and represenred by long-ranging taxa. only in the uppermost metres of the formation did more open marine conditions allow the presence of fusulinids and brachiopods, which have a grearer chronostratigraphical vaiue. flowever, the ages inferred from these taxa are also uncertain. for instance, the age proposed for the comelicania beds ranges from ropmost dzhulfian-basal changxingian (kozur, 1.989), to iower dorashamian (assereto er à1., 1.973), ro upper dorashamran (broglio loriga et a1., 1988; posenato, 1988; etc.). stratigraphic position of the ammonoid. the ammonoid described here was discovered by one of us (h.p.) on the blocks of a large landslide which fell several tens of years ago from pic }l4t. (pizza cuecena in the ladinian language, pitschberg in the german ianguage). this locality (]muèia da insom), about 2 km norths/ard of santa cristina valgardena (fig. 1), is the same as that in which the frsh archaeolepidotus leonardii accordi was found. 'r dipanimento dj scienze geologiche psr@dns. unife.it 'r" via stu[rn 15, oniser, bolzano. e paleontologiche, università di ferrara, corso ercole i d'este 32, 44rca ferrara, italy. e-mail: , 1km . ànnad,?* ortisei tesero 1iu r. posenato & h. prinotb fig. i geographical setting of the locality (asterisk) from which the ammonoid has been collected. the ammonoid occurs in a smail slab of blackish marly limestone, very rich in ostracods, belonging to the badiota facies (fig. 2). this facies repiesents the upper part of the bellerophon fm. in the valgardena area, as weil as in all the western dolomites. in his schematic description of the pic mt. section, accordi (1955) described a unit (unit 6) made up of "calcare argilloso a ostracodi e a piccoli molluschi" (:marly limestone with ostracods and small molluscs"), from which probably the ammonoid derives. flowever, accordi (1955) did not report the thickness of this unit but only that of the whole badiota facies, which is 35 m thick (from unit 3 to unit z). in the valgardena area, recent stratigraphic data on the bellerophon fm. are reported from the seceda section (massari et al., t994), located about 2 km nw of pic mt. in this section the badiota facies is about 35 m thick, as in the pic section, and represents the 5'h and 6'h sequences of third order into which massari et al. (1994) divided the upper permian of the eastern southern alps. the lower part of the 5'h sequence is a transgressive segment made up of dominant subtidal fossiliferous limestones, rich in algae and forams (upper part of the algae assemblage of broglio lnríga et al., 1988). the uppermost, regressive part of the sequence consists o{ silty dolomites and marly limestones, with a biofacies characterized by the mass occurrence of ostracods (ostracod assemblage of broglio loriga et al., 1988). the sequence ends with vuggy dolomite with root traces which suggest subaereal exposures (massari et a1., 1994). this regionally regressive event is about 10-12 m thick (fig 3) the uddermost bellero-"rr"_'_""' nhnn fm ehorrt 1 5 m thick inrrr,t evvs( r,j r the seceda section, records the transgressive event of the sixth sequence which is mainly represented by the basal -werfen fm. ('fesero florizon and mazzrn member). the top of the belleostracod biofacies of the m"trix containing the ammonoid. negative print from acetate peel (x 40). t32 r. posenato & h. prinoth tains three dorashamian genera: paratirolites stoyanoq dzhulfites shevyrev and abichites shevyrev. the latter two genera have been considered by teichert et al. (1973) as synonyms o[ paratirolites. because dzhuffites is slightly older than the other two genera (paratirolites and abichites are associated in the paratirolites beds), a comparison with the genera proposed by shevyrev (1965) is necessary. dzbulfites is characterized by an ovoidal-subrectangular, slightly compressed whorl secrion and a ventral lobe which is simple-pronged and non-denticulated. the dolomites specimen has a whorl section which is sli ghtly depressed, scarcely subpentagonal, with a small denticle on the last adoral ventral lobe as occurs in paratirolites and abichites. fig. 4 paratirolites sp., lateral (a) and ventral (b) views; jmuèia da insom, santa cristina valgardena (bolzano), x 2. (.-_) " abichites has a compressed, rectangular or subquadrate whorl section; the venrer is slightiy convex in the inner whorls, while it becomes flattened in the body chamber. this genus differs from paratirolites by a sharp dorsal shoulder and the absence of ventral nodes on the adult whorl. since our specimen is smaller by half with respect to the smallesr specimen of abichites illustrated by shevyrev (1965), the ontogenetic changes of the whorl shape cannot be used here as an element of comparison even if the lacking of ventral nodes in a. abtchi shevyrev and a. stoyanorai (kiparisova) (see shevyrev, 1965, pl. 34, [ig. 2-4), seems to occur already in the inner phragmocone whorls, a character in common with the dolomites specimen. flowever, abichttes has sharp dorsal and ventral shoulders whereas they are rounded in the dolomites ammonoid, as occurs in some paratiro/lres species (e.g. p. waageni). the dolomites specimen has a slightly depressed whorl section and a low-arched venter with a low crest. a depressed whorl is common among paratirolites species, but a ventral ridge is nor present in the specimens figured by shevyrev (1965). however, according ro tozer (1979), the lack of a ventral ridge in many dzhulfitid species of the type area may be related to an imperfect preservation of the material, as specimens of paratirolites aediensls shevyrev from kuh-e-ali bashi (iran) have one. paratirolites has lrtero-ventral nodes; these are absent in the dolomites specimen, in which only ribs occur. the difference could be related to the differenr ontogeneîic stages of the confronted material illustrated by shevyrev (1.965), as these transcaucasian specimens have sizes which are three or four times larger. f{owever, the lectotype of p. kittli figured by spath (1934, {ig. 135 a, b) has ventral nodes already 19 mm in diameter. the lack of ventral nodes in the dolomites specimen thus raises some doubts as to its attribution to the gen. parat iro i it e s s en s u shetry rev. the specimen described here thus belongs to the fam. dzhulfitidae but, following the shevyrev's classification, it is impossible to determine its precise generlc position as it is too sma1i, with morphological characters in common with different genera^ the lack of ventral nodes is a character of abichttes, whrle the well rounded ventrai shoulder, slightly depressed whorl and 1ow, smooth ventral crest are taxonomicai features of paratirolites. since it shares more characters with paratirolttes than those in common with abichttes, the specimen resembles the former genus more than any other late permian ammonoid. a different classification of the transcaucasian dzhulfitidae was proposed by teichert et a1. (1973), who interpreted a large morphological variability among them regarding whorl section and ornamentation, even if they did not analyse them in detail on account of the incompleteness of the specimens. for this \.f\ shell section (a, x 2) and last (end of phragmocone) suture lines at a whorl height of 7.1 rnm ft, x 7) of paratirolites sp.; jmuéia da insom, santa cristrna valgardena (bolzano). l. 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accepted october 24, 1999 key u,ords: ditrupa, new species, neogene, southern italy, biomer.-. -i"r^"r"""r".o riasswnto. viene descritta la nuova specje ditrupa brnis (polychaera, serpulidae) riscontrata in sedimenri siltosi del pliocene medio di tufara (italia meridionale). d. br*-ìs è stata anche segnalara per il miocene superiore della calabria. il paleoambiente, in base alle faune associate, corrisponderebbe ad un fondale infralitorale o circalitorale superiore. rispetto t d. arietina la specie ha un tubo piìr piccolo, meno affusolato verso i'apice e presenta frecluenti inspessirnenri circolari della parete ("annuli") in prossimità dell'apertura. osseruazioni al microscopio elettronico a scansione su fratture trasversali del tubo hanno evidenziato una microstruttura, comune anclre al genere, caretterizz.ttt dalla presenza di due strati: uno esrerno trasparente con grossr cristalli isoorientati ortogona.lmente al tubo, ed uno interno bianco opaco, più sottile, con cristalli prisnatici di piccole jirncn:ioni, .ì strutturx incrocirtr. i tubi di ditrupa cosrituiscono il subsrrato per numerosi orgrnisn-ri epifaunali; la loro elevata densità. rn un fondale può influenzare composizione e diversità delle comunità di fondo mobile associate. sono ipotizzrre differenti posizioni di vita di ditrupa rn rapporto al substrato, dovuti alla sua capacità di risposta alla dinamica del fondo r-nobile in cui vive. abstract. the new species ditupa brnis (pol1,chaeta, serpulidae) is described from middle pliocene silts of southern italy. it is also reported from upper miocene sediments of southern italy. the associated faunas sugg,est an infralittoral or upper-circalittoral distribution. a r-norphometrical and microstructural analysis of tube was carried out. d. brnis closely resembles ditrupa arietina brt severzl norphometrical differences allow ro discriminate the two specres. the ditrwpa tubes provide a substrate for a diversified epifauna. 'l'heir high density grearly affecrs species composition and diversrtv òf soft-bottorn cornmunitie'. it is concluded tht di*upa can live in various positions with respect to the sediment surface, depending on local sedimenrxrion rare and dynamics. lntroduction. ditrupa berkeley, 1835 is the only serpulid genus characterised by an unattached tusk-shaped rube. it has a long history of taxonomic confusion, early descriptions being poor, ofren based on empry tubes, which are rather featureless. this often led to rnisidenti[y ditrupa as scaphopod rnolluscs and vice versa. ditupa has been associared with many serpulid genera in identification keys (placostegws, vermiliopsis s.1., bonhowrella, sclerostyla, dasynema and marfugia) but it really seems to ser srell apart sysrematically. in some cases, even the genus has been misinterpreted. in the palaeontological literature rhe raxonomic confusion has been greaferi represenratives of rhe genus di*upa often were placed erroneously wirh other serpulids and vice versa. an example is given by the serpulid serpula libera sars, 1835 later placed into the synonymy of d. arietina. ditrupa is known since the tertiary: paleoceneeocene of spain (gaemers, 1978), miocene of taiwan and australia (cheng, 1,974; ten hove & smith, 1990). it is common in mediterranean plio-pleistocene sediments. ditrwpa arietina o. f. miiller, 7776 occurs in the eastern atiantic (f rom iceland to azores, canary islands and senegal) and the mediterranean. ten flove & smith (1990) re-described a distinct species, ditrupa gracillima grube, 1828, from the indo-pacific and on this occasion gave a rhorough review on ditrupa taxonomy, morphology and ecology. f{ere, a new species ol ditrwpa from the neogene of southern italy is described. materials and methods. the examined material of the new species ditrupa brez,is originates f rom miocene and pliocene silty deposits, in southern italy: diparrimento di scienze geologiche, sezione di oceanologia e paleoecologia, università di catania corso italir 55, 95129 cetenir e-rrrril:'.rniiror@mbox.unict.it 456 315 r. sanfilippo fig. 1 location of tufara and bene\t.lre section:. arrons point lavers containingto d. brnìs. benestare some populations of gypstrrì :;;..",,. ditrupa drietina lronr pliocene.sands pleistocene and recent sedimenrs, were also studied: m i d d le p l i o c e n e m0 tufara. this section is ca. 300 m thick and crops out along the eastern edge of the caudina valley, s\l of benevento (fig. 1). it consists of alternating ciayey silts and sands middle pliocene in age (mpl5 zone) (de casrro coppa et al., 1969; violanri, unpubl. data). faunas testify an upper-circalittoral palaeoenvironmenr in the basal part of the section, evolving to infralittoral upwards (barbera et al., 1995). near the contact with the sandy levels, some silty layers with corbwla gibba and ditrwpa breais occur, corresponding to the present-day heterogeneous communities. these layers contain species of different biocoenoses and are characterised by quasi-permanent instabiiity of qualitative and quantitative faunal composition (picard, 1965). the type material of the new species is from these silty beds. the tubes are often broken (more than 1,000 tube fragments). benestare. this secrion is located ca. 1 km nv of benestare (fig. 1). it is a classic locality, originally reported on by seguenza (1879-80) late miocene and probably tortonian in age. the sequence is made of clays, 100 m thick, with silty-sandy layers on its upper part and a fossiliferous level of siliceous sands and messinian gypsum. ditrwpa brevis comes from two silty layers (benestare i and benestare ii) at the top of the clayey bed. fragmented tubes largely prevail. samples also contain abundant corbula gibba and lunulitiform bryozoans, indicating heterogeneous communities (di geronimo et al.. 1.992\. sjlty sands clays diolo. along the stramonte river in vestern emilia, northern ltaly. material comes from .r silty-sendy layer 2 m rhick, wirhin a clayey-silty sequence of middle pliocene age. it contains circalittoral faunas referable to a coastal detritic assemblage (di geronimo et al., 1992). the presence of sedinrentary insrability indicators among molluscs and l:ryozoans, rogether with the dominance of corbwla gibba and d. arìetina, suggesrs an incipient rurbidity (see di geronimo & robba, 1989). lonian sea dredges. gulf of noto (off se sicily): stn. ps/s1 5b (47 m),36"52.98'n, 15'10.48'e; stn. ps/81 5c (34 m), 36'53.30' n, 15'09.82' e. srmples consist of biogenic sands containing faunas referable ro the coastal detritic assemblage. catania gulf dredges, stn. ct/11 (-110 m), 3z'29.15'n, 15'09.05'8. sandy gravels containing wùrmian faunas (di geronimo & li gioi, 1980) including several tubes of d. arietina; acitrezza, srn. ac/50 (50 m) dredged ca. 1 km off the coasrline. volcanic sands containing circalittoral faunas referable to a coastal detritic assemblage swept by bottom currenrs. tyrrhenian sea dredges. off patti, stn. pd4 (50 m), 38'09.40'n, 15"02.19'e. muddy sands with faunas belonging to the coastai detritic assernblage subject to an incipient turbidity. measurements were taken from recent and fossil specimens of ditrupa, mainly following the method proposed by ten hove & smith (1990) (fig. 2). only whole tubes were measured. values and standard deviation of each morphometric character have been compiled (tab. 1). pearson correlation coefficients between data were cluster analysed (figs. 3, 4), using the average linkage method (sokal & sneath, 1963; sneath sc sokal, tlzl;. plate 1 scale bar = fig. 1-3: 0.6 mm; fig. 4: 150 prn; fig. 5, /, 8: 1 mm; fig. 6: 200 pm; fig. 9: 3 mm. fig. 1,) ditrupa breois,holotype (on the right) and parat)'pes. tufara, middle pliocene. fig.2) ditrupa 1zre,--2.s. benestare, miocene. ì.-ig. 3) smallsized ditrupa arietina. ps/81 58, recent. fig. 4) apex of d. brnis. the rirn consists only of the outer hyaline layer. tufara, middle pliocene. ii5. 5) d. arietina. diolo, middle pliocene. fig.6) d. arietina. ps/81 5c, recent. fig. 7) d. arietina. pd 4, recent. fig. 8) large d. arietina. ct/1.1, wùrmian. fig. 9) d. arietina wìth hydroides norvegicus coiling around the anterior ends of rubes. ctl11, wirrmian. pl. 1 a new serpulid from mediterrdnedn neogene 457 458 r. sanfilippo chord (m!) height mean d. d. mouth d. apex arcuation (mm) (%) 0-4.48-8 (sd : 1.8) .02-0.1 3-0.50 0.29-0.60-0.95 0.34-0.53-0.75 0.28-0.37-0.56 1-2.a2-7 (sd = 0.11) r (sd = 0.15) (sd = 0.11) (sd = 0.07) (sd = 1.58) tufara 10 benestarel 2.70-4.68-9'80 (sd = 1 63) 2.60-3.'t 6-3.80 0.05-0.1 1-0.20 0.40-0.55-0.75 0.40-0.49-0.58 0.30-0.36-0.44 1 .70-3.26-5.20 nin t^^ àriaiinà ct/1 1 .199 , 0.55-0.791 .1 0 i (sd = 0.14) i (sd : 0.1!l ' o.oo-0.64-1 .50 (sd = 0.27) , t?i.?strs 0.45-0.65-0.85 0.25-0.79j1 .40 0.30-0.71-1 .70 = o.24) 0.23-o.47-0.68 0.30-o.70-0.80 (sp = 0.1) 0.30-0.73-1 .20 (sd = 0.16) (sd : o.oe) (sd : 2.49) 0.21-0.34-0.38 3.2-4.21-13 €d=005) 1sd=235) 0.20-0.36-0.57 3-6.01-14 (sd=013) (sd=1.2) o.à-o.zo-r 7.so1s-22.s1 , 0.32-o.63-0.97 (sd = 7.1e) (sd = 1.a6) (sd = 0.42) r (sp : o.l1 (sd = 0 71) 0.30-0.62-1 .30 0.20-0.36-0.70 10-10.69-15.13 =n21\ 0.5-1 .59-2 (sd = 0.38) (sd=021) (sd=2.83) 5.50-24.01-40 0.8-3.73-7 ps/81 58 2.90-4.80-10.80 j 0.10-0.32-1 .30 2.60-8.os-'t6 0.10-0.s1-2.30 88 i (s?= 0.291 1 (sq !.1!l diof o 2 ", 2.70-6.24-21 .10 0.27-0.71-9.10 72 (sd = 3.23) (sd = 0. tab. 1 morphometric parameters measured on tubes of d. brnis and d. arietina populations. for each semple rmximum, nrinimum, rne rn values (bold) and standard deviation are reported. the total number of measured tubes for each sample is reported on the left. systematics. genus ditrupa berkeley, 1835 ditrupa brevis sp. nov. (pl. 1, figs. l-9 pl.2, figs. 1-5) derivatio nominis: from latin brevis (= 5[611; type material deposited in the palaeontological museum of the catania university, collections of the departn-rent of geological science. labelled pmc.s1.20-10-99. description. tube nor encrusting. circular in cross-section, scarcely bent and smal1 (mean length 4 mm, mean diameter 0.6 mm). tube opened at both ends, slightly rr^rrowing to apex (almost cylindrical). mean diameter at mouth 0.5 mm, diameter of apex ranging from 0.3 to 0.5 mm. just before mouth, tube narrows forming conical shoulder. rim of both mouth and apex circular and smooth. distal ends often with irregularly spaced annular thickenings ("annu1i" sensu ten hove & smith, 1990) (fig. 5; pl. 1, fig. 1). outer tube surface smooth, except for occasional transversal lines and/or constrictions mainly near the mouth. holotype: (pmc.s1.20-10-1999) length 3.2 mm, diameter of apex 0.4 mm, diameter ar mouth 0.5 mm. miorostructure. transverse fractures of the wal1 show two layers (pl. 2, fig. 3): an inner white one, thick ca. 1/4 of the entire wall and an outer somewhat transparent one rhicker just before mouth, showing still some transparency despite fossilisation. under sem magnification the outer hyaline layer is composed of needle-like crystals arranged in lamellae, perpendicular to the tube surface. the inner opaque layer shows small prismatic crystals, crossiy arranged. the apex is made only of the outer hyaline layer, the inner opaque layer starting at some distance from the apex (pl. 1, fig. 4). at higher magnifications, the outer surface does no longer appear smooth but is faintly dotted, due to the tips of crystals perpendicular to the surface (pl. 2, fig. 5). a discontinuous thin layer with an amorphous cryptocrystalline structure is randomlv dresent on the outer surface. plate 2 scale bar: fig. 1: 200 pm; figs.2,4: 100 pm; fig.3: 40 pm; fig.5:4 plm. fig. i) ditrupa breris; a) anterior end; b) detail of a transversal line corresponding to a beginning of the tube growth. tufara, middìe pliocene. fig. 3) ditrupa bretis, rra.nsverse section of the tube wall showing an inner opaque layer with criss-cross arranged crystals and an outer glossy layer, q.ith needle-like crystaìs, perpendicular to the outer surface. benestare, miocene. fig. 2) d. arietinawith a cylindrical bore-hole by a rnuricid gesrropod. mecanical abrasion trrìces are visible on the outer surface. ps/81 5c, recent. fig. 4) bore-hole due to a naticid gastropod wrth bevelled edge. d. arietina. ps/81 5c, recent. fig. 5) detail of the outer surface of d. brnis, fa.intly dotted. tufara, middle pliocene. 459pl. 2 a new serpulid from mediterrlrnedn neogene +60 r. sanfilippo biometry. the measured tubes of ditrupa are easily classed in two groups (tab. 1): in the first group (d. breais: samples tufara, benestare i and benestare ii) tubes have notably low mean values of chord (3.t0 4.68 mm) and height (0.11 0.22 mm); in the second group (d. arierina.'samples ac/50, pd 4, ps/81 5b, ps/81 5c, diolo 2, ct/1,1) tubes have mean values of chord of +.so 24.01 mm and of heighr of 0.32 3.73 mm. in the first group apices show mean diameters (0.36 0.38 mm) similar to those of the second group, despite in the latter the tubes are generally much larger in size (tab. 1) . in both species the apex evidently remains unchanged from juvenile to adult stage (as showed also by the low values of sd). even between specimens of d. arietinahaving different sizes, the apex diameter seems to be the only parameter morphometrically unchanged, tube accretion being possible only at the mouth. the mean values of the arcuation index 2.82 4.48% also differ in the first group, versus 4.21 l5'h in the second group (tab. 1). arcuation varies considerably within a population, ranging from almost straight (juveniles) to stròngly curved (adults). it may also vary between populations from different sites. for instance, in the d. arietina population from the gulf of catania (pl. 1, figs. 8, 9) the arcuation index is greater than in other recent populations here examined (ps/81 58 and 5c) (pl. 1, figs.3,5, 6, 7). the other examined size-shape parameters of ditrupa also increase notably with growth and can vary considerably between different recent and fossil populations. the differences between the two species are also shown by cluster analysis (fig. 3) where samples of d. breztis (benestare i, benestare ii and tufara) are grouped rogerher. sd values (tab. 1) of the two species are nor markedly different. remarks. d. breois tube is much shorter than those of the other two ditrupa species. compared rc d. arietina and d. gracillima, tube of the new species is aiso scarcely bent and almost cylindrical, slightly narrowing to the apex. both d. arietina and d. brevis show transversal lines and consrrictions on the outer surface, mainly near the opening (pl. 1, fig. 8; pi.2, fig. 1). these are due to a thinning of the external wall layer, as occurring at the mouth rim, and were interpreted by gambi (1986) as markers of previous tube openings. conversely, the socalled "annuli", described for ditrwpa arietina and d. gracillima by ten hove & smith (1990) as irregularities in the tube wall thickness, may occur in d. breois (pi. 1., fìg.2 morphometrics of d. brnis trbe. d=diameter of tube n-routh, d:diameter of lpex, rld=mean wrdth, c:chord (length measured in straight line betr.een orifice ro rpex), h:height (wiclest perpendicular disrance between rnner cur-ve of tube and chord). fig. 1). however, these "annuli" are more frequent and elevated than in d. arietina but less abrupt than in the "monilifera" form of d. gracillima. a ditrwpa tube resembles a scaphopod shell, particularly the curved and smooth shell of gadila. however, microstructural analysis reveals a distinctive twolayered pattern ol ditrupa tube (p1. 2,fig.3): an outer layer somewhat transparent (which may become opaque by abrasion or fossilisation), and an inner bright white layer, of more randomly arranged crystals (ten hove & zibrowius, 1986;ziltrowius er ten fiove, 1982). the distinction between the. two layers is generally evident, although in fossils the tube is entirely opaque, due to recrys tallisation. the outer tube surface is also peculiar. at higher magnification, the seemingly smooth surface can appear faintly dotted (p1.2, fig. 5), similar to the fine honeycomb structure described by ten hove & smith 1t9lo; for d. gracillima, bv. less pronounced. ecology and dlstribution. ditrupa lives with its tube free on muddy sediments on the continental sheif, in areas with high sedips/81 58 ps/81 5c pd4 ac/50 diolo 2 ct/l1 tulara benestare 2 benestare 1 mentation rate and a quasi-permanent turbidity. in such environmental conditions, populations can be very dense (di geronimo et a1., 1989) and tubes ofren constitute a conspicuous component (over 5o%) of the biogenic carbonate sediments (vilson, 1976).it also occurs in circalittoral detritic bottoms with scarce muddy component, although never abundanr. empty rubes, after the death of the animal, are easily preserved in the sedi, ment. numerous dead tubes occur with the living aninrals and may constitute nearly 100"k ol the total number of tubes (light, 1999). forming dense assemblages of live and dead tubes, ditrupa may control the sediment texture. it also provides a substrate for epibionts and rheophile taxa (di geronimo et al., 1989; gambi, 1986; gambi 8c jerace, 1997 ligfu, 1999) and thereby affects species composition and diversity of soft-bottom communities. observations on r. arietina assemblages from 60 to 300 m deep soft-bottoms of lceland reveal that tubes can be densely colonised by brachiopods, solitary scieractinians, barnacles, serpulids, bryozoans, sponges, and foraminiferans. opercula lre ofren encrusted by foraminiferans, hydrozoans and erect flexible colonies of bryozoans. most of dead specimens tubes are inhabited by sipunculids. ditrupa is common in recent mediterranean sediments. in fossil sediments, it can be abundant, often occurring in banks, due to the combined action of gregariousness and selective hydrodynamic transport (ten hove & van den hurk, 1993). it has been used as an indic;rtor of sedimentary instability (di geronimo & robba, 1 d. arietina from pliocene and pleistocene sediments is found together with hydroid.es noruegicus. although common in the recent mediterranean, borh serpulid species are more abundant in pleistocene deposits, where they usually are larger than in the recent. a fine example is given by the \fùrmian ct/1,1 station, where f1. noroegicus and even more d. arietina a new serpulid from mediterrd.rredn neogene distance iv]etric is i-pearson correlation coefficient average linkage i\iethod 0 000 distances 2.000 461 fig. 3 dendrogram of samples of ditrupa based on all five factors (c/h, c/arc., max d of apex, rnx c, arc.), expressing morphomctric characters, specimens have a markedly large size (p1. 1, figs. 8, 9). it seems that both species benefied of the lower water rcrrpcrature. for both localities with ditrupa brevis an upper circalirtoral palaeoenvironrnenr can be hypothesised. like other members of the genus, d. brer,,is can be considered as a species indicating an unstable environment with low diversity and be used to recognise f{eterogeneous communities. comparing sediments containing d. bre.r,ts to recent ones conraining d. ari etina, a similar ecological niche, at least fron-r miocene onwards, can be hypothesised. d. brevis seems ro have been replaced by d. arietina durrng the pliocene. in both studied localities, d. breztis co-occurs with warm water taxa, particularly some bryozoan species extinct in the present-day mediterranean, but still living in tropical atlantic and pacific regions (di geronimo er ai., 1992; rosso, pers. comm.). according to the present data, d. brer.,is ts restricted to the neogene. conversely, d. arietina continues all through the pleistocene till the recent and was especially abund;rnt during cold quaternary phases of the mediterranean. behaviour. many tubes oî d. breois and d, arietina show a circular hole bored by gastropod predators. two types can be distinguished: distinctive countersunk holes vrith ber'elled edges due to naticids, and smaller holes with nearly straight edges, made by mr-rricids (p1. 2, figs. 2, 4). records of borer-attachs upon serpuloidean tubes (due to nudibranch, muricid and thaidid gastropods) are frequent (ten hove. 199,1;. in the examined marerial, drillings mainly occur near the mid point of the rubes, most commonly on its concave side. similarly, the solitary coral caryopbyllia smithi encrusting d. arietina (wilson, 1.976) matnly occurs on the concave side, between the mid point and the anterior end of the tube. these data suggesr that the dttrupa tube lies horizontally on rhe sediment. this is in agreemenr with ten f{ove e smith (1990) observations r.hat d. arietina, when placed on the sediment surface, is 462 r. sanfilippo able to turn over its tube from lying on the right or left sides to lying on the convex side. thus, only the concave side is always available for colonisators andlor predators. notwithstanding this, the same aurhors also observed that ditrupa can be partially buried, with the two tube ends emerging from the sediment. conversely, gambi (1986) assumed that ditrwpa lives vertically buried in the sediment, with only the anterior end of its tube protruding above the surface. wilson (1976) observed that tubes had the apex slight buried and the convex side turned upwards. these latter models do not explain that on live ditrwpa the corals are attached, generally, near the mid point of the tube (settling on the concave side) and that bore-hole by muricids (preying only on epifauna, reyment, 1966; taylor, 1970) occur in the same posirion. the presence of hydroìcles noraegicus, matnly coiled around the anterior end of the tube, suggests that the posterior end of rhe tube was buried within the sediment (obliquely or almost horizontally). observations in aquarium reveal that ditrupa ís a filter-feeder (ten hove & smith, 1975) although observed life position have implied that deposit-feeding might occur (hong, 198a). the worm is able to burrow in fine sediments using the branchial crown, thus varying its posture (lighq 1999). lmm fig. 4 different positions of drtrup.r orr rhc.oir borronr. r, tube n'ith only the anterior end exposed; b) tube horizontally buried with the two end5 expo5ed: c.1 rubc lying horizontally above the sedrítenl. to summarise, it can be assumed that ditrupa can live in different postures: tube with only the anterior end exposed, at more or less steep angle (fig. 4:r) ; tube buried horizontally in the sediment, with the two ends above the surface (fig. ab); tube lying horizontally above the sediment (fig. ac) . the actual position probably reflects the sedimentation rate, suggesting rhe ditrupa's capability of reacting to sediment dynamics. acknooledgements. i am indebted to c. barbera (dipartimenro di paleontologia, napoìi) for loaning toporypic material. i also thank h.a. ten hor.e (instituut voor taxononische zoologre, amsterdam) and h. zibrorius (station marine d'endoume, marseilles) for useful suggcsrions. c.n. bianchi who criticallv reviewed the manuscript. r. la perna for effort in statistical elaboration. o. torrisi (istituto internazronele di vulcanologia, catania) supplied assistancc in sem obsen.-rtions. this paper is financially supporred by 40% and 60% (di geronimo) m.u.r.s.t. grants. re,ferences barbera c., robba e., sanfilippo r., violanti d. (1995) the middle pliocene tufara section (benevento, south italy). in cherchi a. (ed.) proceedings vi palaeobenthos international symposium, 28-30 october 1995, pp. 129' 131, alghero. cheng ym. (1974) paleocurrent direction of the taliao formation (lower miocene) indicated by dìtrupa sp. (serpulidae) shells.,4cta geol. taizu., v 17, pp. 13-22,taiper. de castro coppa m.g., moncharmont zerm., pescatore t., sgrosso l, torre m. (1969) depositi miocenici e pliocenici ad est del partenio e del taburno (campania). .atti acc. gioenia sc. nat., s.7, v. 1, suppl. sc. geol., pp. 479-51.2, catania. di geronimo i. e( li gioi r. (1980) sur un gisement wurmien sous-marin du golf de catane. rapp. comm. int. mer médit.,v.25/26$), pp. 137-139, monaco. di geronimo i. & robba e. (1989) the structure of benthic communities in relation to basin instability. mem. acc. lincei, v.80, pp. 341-352, roma.fig. 5 d. brcuìs: tube fragments of distal ends with "annuli" a neu serpulid frotn mediterranedn neogene 463 di geronimo i., raffi s., rosso a. (1989) dominanza di specie opportuniste nei "popolamenti eterogenei" del pleis tocene inf erio re di mazzarìno, sicilia cenrrale. b o//. acc. gioenia sc. nat., v. 20, n. 331, pp. 1,29-166, catania. di geronimo i ., rosso a., sanfilippo r. (1992) bryozoans as sedimentary instability indicators. ria. it. paleont. strat., v. 98, n. 2, pp. 229-242, milano. gaemers pa.m. (1978) biostratigraphy, palaeoecology and palaeogeography of the mainly marine ager formation (upper paleocene-lower eocene) in the tremp basin, central-south pyrenees, spain. leidse geol. meded., v. 51, pp. 151-231, lleida. gambi m.c. (1986) ecological implications in tube morplrology and epibiota of a population of ditrupa arietina (polychaeta, serpulidae). ropp. comm. int. mer médit., v. 30, n. 2, p. 19, monaco. gambi m.c. 8rjerace s. (1997)epibiosis on the tubes of the polychaete ditrupa arietina (serpulidae) in some populations of the soft bottoms of the southern tyrrhenian sea. biol. mar. medit., v. 4, n. 1, pp. 380-383. hove h.a. ten (1994) the dualistic relation between molluscs and serpulid tube-worms. pp. 65-7a.in coomanseustatia m., moolembeek r., los !( ec prins p (eds.), de horen enzijn echo. stichting libri antilliani,zool, ogisch museum amsrerdam,y. of 279 pp., amsterdam. hove h.a. ten & hurk p van den (1993) a review of recent and fossil serpulid "reefs"; actuopalaeontology and the "upper malm" serpulid limestones in n\7 germany. i ar"veot. en lynjno., v. / 2, pp. zj-bl, amsterdam. hove h.a. ten & smith r.s. (1990) a re-description of ditrupa gracillima grube, 1 b7b (polychaeta, serpulidae) from the indo-pacific, with a discussion of the genus. rec. awstr. mus.,v.42, n. l, pp. 101-118, sydney. hove h.a. ten & zibrowius h. 11986) laminarubus alaini gen. et sp. n. and protis hydrothermica sp. n. (polychaeta, serpulidae) from the bathyal hydrothermal vent communities in the eastern pacific. zool. scripta, v. 15, n. 1, pp.21-31, góteborg. hong j.s. (1984) on two polychaerous serpulids new ro korean waters with nores on the ecological aspects. korean j. zool., v. 1.2, n. 1,, pp. 34-48. light j. (999) the free-living serpulid, ditrupa arietina, on the outer continental shelf. newsletr.er porcup. mar. nat hist. soc., v. 2, pp. 4a-42, london. picard j. (1965) recherches qualitatives sur les biocoenoses marines des substrats meubles dragables de la région marseillaise. rec. trao. stat. mar. end.oume, v. 52, n. 36, pp.1-160, marseille. reyment r.a. (1966) preliminary observations on gastropod predation in the western niger delta. palaeogeogr. palaeoclim. palteoecol., v. 2, n.2, pp. 81-102, amsterdam. seguenza g. (1879-80) le formazioni terziarie di reggio (calabria). mem. rend. acc. lincei, cl. sc. fis. mat. e nat., s.3, v.6, pp. 1-446, roma. sneath ph.a. & sokal r.r. (1973) numerical raxonomy: rhe principles and practice of numerical classification. v ol 573 pp. freeman libr., san francisco. taylor (197a) feeding habits of predatory gasrropods in a tertiary (eocene) molluscan assemblage from the paris basin. palaeontologt, v. 1,3, n. 2, pp. 254-260, london. vilson j.b. (1976) attachment of the coral caryophyllia smithii s. & b. to tubes of the polychaete ditrupa arietina (muiler) and other substrares. j. tnar. biol. ass. u.k., v. 56, pp. 291-303, london. zibrowius h. & hove h.a. ten (1987) neovermilia falcigera (roule, 1898) a deepand coldwater serpulid polychaete common in tl-re mediterranean plio-pleistocene. bull. biol. soc. wash., v. z, pp. 259-27i,,lflashington. torric_kne 201..222 ��� ������ �� � ��� � � ���� ��� ���� �� ���� � � �� ��� ���� �� ���� ��� �� ������� ��� �� � ��� �� ��� �� ����� ����� ������� ����� � � � �� ��� � ��������� ��������� ��� � � ��� ���� ��������� ��� �� ��� ��� �� ��� ��������������� �������� �!!"# #���$!��"!!��" ������ ��!���"�%� �����$����!�� �%�&�#��"�� '�!�������"���"�� �%�"!���� ����!�� ���!�( ���� ���( ��" ��%#� �$ ��!���)����� ��# ��!���"�%� ������ $����!� �"��'"�"# $��) ��" '�!�������"����� �%��"����� �%��������� �� ������ ��# "��)� *����!�+ #��%)"��� �%�"!��� ���")&!��"� ��)���� �&!" �� ����" �%&!���"# $�� ��" �%�� �%$$��"( ���� �" "'�#"��"� ��)� &��"# ��� �"����������� �"���")���! 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r r r r r r r r r dinoflagellate cysts b is a c c a te s s p p . in a p e rt u ro p o ll e n it e s s p p . e c h ip e ri p o ri te s e s te la e c la s s o p o ll is c la s s o id e s r pollen s c h iz o s p o ri s s p 2 v e rr u c a to s p o ri te s u s m e n s is z li v is p o ri s s p p . c o n c a v is s im is p o ri te s s p p . l e io tr il e te s s p p . s c h iz o s p o ri s s p 1 in d e te rm in e d s p o re s r spores a c ri ta rc h s c a ly p to c y s ta e x o le ta legend occurrence ( 1 ) rare ( 5 ) scarce ( 10 ) common ( 20 ) abundant ( 40 + ) ���( 9< � ����" ����� �$ ��!���)����� �"��'"�"# $��) ��" ������ ��� �"�����( �!"'����� �"��"�"��� )"��"� $��) ��" &��" �$ ��" �"�����( � ����#� $�� �" ��-"#( ��" 1"#��"����"�� �"���� �� �"���#"# ����� �����$����! /��" �.;g *b�!!��)� "� �!( ;<>?+( � ����#� $�� �"� ��-"#( "�" #"�"��"#( �����#���!�� /��" 1�.;g ��� &" �#"�� ��$�"# $��) ��)�!" �� d %� ��#�� �"�"�� �� ��" !� "� ���� �$ ��" �"����� � )��" �"�"��� 1�.;;� 1�.;g ������)"�� �� ������"#� #%" �� ��" �&�"��" �$ #"$����" )��-"��( ������ ���� ��� �� �� � ���� � � ����� ���� �"����� �� ��)�!"# $��) � �!������! �%����� �$ ��" �"����� �!����� �!�"�#� #"����&"# &� ����!� "� �!( *9>>h+ ��# #" ���� "� �!( *;<+( �� �"�"��!� ���")&!��"� ��" #�)����"# &� �&%�#��� &�������" ��!!"� ��� �%&��#����" .�!"��"�" #���$!��"!!��" ����� ��# � $" �" ��-"# �"���"�%� �����( ��)" �%�"!��� )��-"�� "�" $�%�# �� ��)�!"� .�� 9:� 9?� 9d� g<� ��)"!� 2� %��)���� 6������� �� ���� �� �������� ����� * ��� ��)�!"�" �"���%!%)+ ��# �� ��0 ���� � ( �� �"���"� &)� ���� ��� * ��� ��%��"� ��,� ��'"&""�#"�"��"#� ��" ��" �$ ��"�" ���")&!��"� �� !�-"!� ���!� �!����"�"� ���� �� �������"�� ��� 1�.;;�;g /�� ��! �����&%���� ��$"��"# $��) �����$����!�( &���� �� � �����'������( �%� �$ ��,�""� ��)�!"� ��%#�"#� ��!� ����" $��) ��" !� "� ���� �$ ��" �"����� ��"!#"# #��������'" ���")&!��"�� �"�"�� ��" )�##!"� %��"� ���� �$ ��" �"����� �� &���"� �� '"�� ���� �$ ���� ��$����!�( �"���"�%� �" ��-��� �� $�"6%"�� ����%���%�� ������%!��!� �&%�#��� �� ��" !� "�� ��)�!" *���( ;<+( ��" ����"�� ��)�!"� ��" .�� 9:� 9?� 9d� �������"���"# �� ������ �� ���!� �" #�;9; lithology tr o in a -t u sa f o rm a ti o n top polizzi fm. sandstone shale marl sample ���( 9; � ������������� !�� �$ ��" ������ ��� �"����� ��� ��)�!" ��������( e le v a ti o n 150m 202m 100m f o rm a ti o n a g e s a m p le s rtr 17 rtr 14 t ro in a -t u s a f m . r u p e li a n rtr 12 rtr 11 rtr 10 rtr 7 rtr 5 section troina rtr a re o li g e ra c f. s e n o n e n s is c ri b ro p e ri d in iu m e d w a rd s ii d e fl a n d re a p h o s p h o ri ti c a d is ta to d in iu m e ll ip ti c u m g la p h y ro c y s ta s p p . in d e te rm in e d d in o fl a g e ll a te s l ic ra c y s ta c o ry m b u s o d o n to c h it in a o p e rc u la ta p o ly s p h a e ri d iu m s p . r o tt n e s ti a b o ru s s ic a s p in if e ri te s ra m o s u s s p in if e ri te s s p p . s y s te m a to p h o ra a n c y re a t h a la s s ip h o ra p e la g ic a a s c o s to m o c y s ti s p o ta n e c ir c u lo d in iu m d is ti n c tu m c y c lo p s ie ll a s p . d in o p te ry g iu m c la d o id e s h o m o tr y b li u m p le c ti lu m h o m o tr y b li u m te n u is p in o s u m s p in if e ri te s p s e u d o fu rc a tu s a c h o m o s p h a e ra a lc ic o rn u c h a rl e s d o w n ie a c la th ra ta c ir c u lo d in iu m b re v is p in o s u m h y s tr ic h o k o lp o m a s a la c ia is a b e li d in iu m s p p . p e n ta d in iu m la ti c in c tu m t h a la s s ip h o ra s u c c in c ta w e tz e li e ll a g o c h ti i c e rb ia ta b u la ta c e ro d in iu m s p p . c o rd o s p h a e ri d iu m g ra c il e c ri b ro p e ri d in iu m te n u it a b u la tu m d e fl a n d re a h e te ro p h ly c ta e n n e a d o c y s ta p e c ti n if o rm is im p a g id in iu m s p p . o p e rc u lo d in iu m m ic ro tr ia in u m p e n ta d in iu m g o n if e ru m p s e u d o c e ra ti u m c f. s e c u ri g e ru m s a m la n d ia c h la m y d o p h o ra s u b ti li s p h a e ra p e rl u c id a g la p h y ro c y s ta c f. s e m it e c ta h e m ip la c o p h o ra s e m il u n if e ra o p e rc u lo d in iu m c e n tr o c a rp u m r e ti c u la to s p h a e ra a c ti n o c o ro n a ta r h o m b o d in iu m d ra c o c h ir o p te ri d iu m g a le a d is ta to d in iu m c ra te ru m h y s tr ic h o k o lp o m a e is e n a c k ii n e m a to s p h a e ro p s is s p . c le is to s p h a e ri d iu m ? s p l in g u lo d in iu m m a c h a e ro p h o ru m s p in if e ri te s m ir a b il is w e tz e li e ll a s p p . r r r r r r r r r r r r r r r r r r r r ? dinoflagellate cysts b is a c c a te s s p p . in a p e rt u ro p o ll e n it e s s p p . c la s s o p o ll is c la s s o id e s r r r r pollen p o ly p o d ia c e o is p o ri te s s p p . z li v is p o ri s s p p . c o n c a v is s im is p o ri te s s p p . in d e te rm in e d s p o re s s c h iz o s p o ri s s p 2 r spores legend occurrence ( 1 ) rare ( 5 ) scarce ( 10 ) common ( 20 ) abundant ( 40 + ) ���( 9g � ����" ����� �$ ��!���)����� �"��'"�"# $��) ��" ������ ��� �"�����( �!"'����� �"��"�"��� )"��"� $��) ��" &��" �$ ��" �"�����( � ����#� $�� �" ��-"#( &� � �%�"!��� ���")&!��" *�"�����"��" �$ .� '� *��� ��# ,� �� � �+ �$$"��"# &� ���"�" �" ��-���( �� $��) ��"��)"�� �$ /� ���� "�" �#"���$�"#� ����� $��) � 6%"������&!" ��" �� .�� 9?� ��!" ���" ��"��)"�� �$ /� �� �)0���������� "�" �&�"�'"# �� ��)�!"� .�� 9? ��# 9d( ���" ��# �����"�"# ���%��"��"� �$ &� ������� � !���"� ���� 9< �) "�" �&�"�'"# �� ��)�!" .�� h� ��!" �� �� � � �� #"�"��"# �� ��)�!"� .�� ? ��# 9?( �� ��)�!" .�� 9d� ��" #����!'"# �����$����! �"� �")&!��� �� ��� �� *����� � �� �&�"�'"#� &%� ���� (�� ����� ���% ��)�� ���� ' �* �)� ������� �����) �%��� +������� ,����;9g elevation 1 5 0 m 2 0 0 m 1 0 0 m formation age zone samples r t r 1 7 r t r 1 6 r t r 1 4 r t r 1 2 r t r 1 1 r t r 1 0 r t r 7 r t r 5 triona-tusa fm. rupelian mnp23 mnp22-mnp23 s e c ti o n t ro in a r t r totalcount:calcareousnannoplankton 3 0 0 totalcount:cretacousreworking 3 0 0 arkhangelskiellaspp. calcidiscusprotoannulus coccolithusmiopelagicus coccolithuspelagicus cretarhabdussurirellus cribrocentrumreticulatum cribrosphaerellaehrenbergii cyclicargolithusabisectuslarge cyclicargolithusfloridanus dictyococcitesbisectus dictyococcitesscrippsae/hesslandii dictyococcitesspp.3-7mu discoasterbarbadiensis discoasterdeflandreigrp. discoasterseptemradiatus eiffellithuseximius ericsoniaspp. helicosphaerabramlettei helicosphaeraeuphratis helicosphaeracf.recta isthmolithusrecurvus miculaspp. reticulofenestradaviesii reticulofenestradictyoda reticulofenestraaff.umbilica smalldictyococcitesspp. smallreticulofenestraspp. sphenolithuspredistentus sphenolithusspp. watznaueriamanivitae watznaueriaspp. zeugrhabdotusspp. cyclicargolithusaff.abisectus eprolithusspp. ericsoniaformosa reticulofenestraumbilica sphenolithusdistentus/predistentus toweiusspp. chiasmolithusspp. cyclicargolithusabisectussmall discoastercf.binodosus discoastertaniigrp. helicosphaeraaff.carteri helicosphaeracf.gartneri pontosphaeraspp. prediscosphaeraspp. reinhardtitesspp. reticulofenestracircus reticulofenestraspp.3-7mu rucinolithusspp. sphenolithuseditus sphenolithusgr.moriformis sphenolithusobtusus stradneriacrenulata toweiuseminens toweiusoccultatus transversopontispulcheroides cretarhabdusspp. discoastermultiradiatus ericsoniaobruta helicosphaeracompacta helicosphaeracf.perch-nielseniae helicosphaerarecta lanternithusminutus neocrepidolithusspp. sphenolithusmoriformis sphenolithusradians transversopontispygmaea aspidolithusparcus ericsoniacava helicosphaeracf.bramlettei toweiuscrassus ericsoniasubpertusa helicosphaeraperch-nielseniae sphenolithuscf.conicus arkhangelskiellacymbiformis cruciplacolithustenuis discoasterspp. reticulofenestrahampdenensis nannoconusspp. r r r r r r r r r r r r r r ? 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(� ,(��� ��� ��!> %� $� #�� � (� � �$��$ "#� � (� �#��#"� �����-�"� $� .�!/#�� 0��� ��&��! �.�&+ �#1+ .�" $���"�' ��! ��� (�/�!�� � � � ��� �8 + � � ��� # � ���%�" � �$$�� ���%�"> �� � ���� � #$ �' ��� ���%�"+ ������ !" #������� $"%" &������ �" ' $������ ("565 radiolarian assemblages at ponte serra (trento plateau, southern alps, italy) plate figure code uaz 95 uaz sa ps 2 ps 7 ps 8 ps 9 ps a ps d ps e ps 11 acaeniotylopsis variatus s.l. (ozvoldova) 1 1, 2 4063 1-8 a-b alievium sp. a sensu conti & marcucci 1991 2 2 4004 8-9 angulobracchia sp. b sensu baumgartner et al. 1995c 1 3 4006 7-9 archaeospongoprunum imlayi pessagno 1 4,5,6 bernoullius dicera (baumgartner) 1 7, 8 3223 3-10 a-f bernoullius rectispinus s.l. kito, de wever, danelian & cordey 1 9, 10 4010 1-9 cinguloturris carpatica dumitrica 2 17 3193 7-11 d-f dicerosaturnalis angustus (baumgartner) 1 20 3082 6-10 b-e emiluvia hopsoni pessagno 1 11 3225 6-15 d-f emiluvia orea s.l. baumgartner 1 12 4069 4-11 c-f emiluvia premyogii baumgartner 113, 15 3210 3-10 a-e emiluvia salensis pessagno 1 14 3215 4-13 eucyrtidiellum nodosum wakita 2 18 3014 3-10 b-e eucyrtidiellum ptyctum (riedel & sanfilippo) 2 19/21 3017 5-11 b-f eucyrtidiellum unumaense s.l. (yao) 2 22 3052 3-8 a-c gongylothorax aff. favosus dumitrica sensu baumgartner et al. 1995c 2 23 3279 7-8 guexella nudata (kocher) 2 24 3061 5-8 a-b haliodictya (?) antiqua s.l. (rüst) 1 16 3243 4-11 haliodictya (?) hojnosi riedel & sanfilippo 1 17 3254 3-10 helvetocapsa lemanensis o'dogherty, gorican & dumitrica 2 25 hexasaturnalis minor (baumgartner) 1 18 b-f hexasaturnalis nakasekoi dumitrica & dumitrica-jud 1 19 homoeoparonaella elegans (pessagno) 1 22 3104 4-10 hsuum speciosum hull 3 23 lanubus cornutus (baumgartner) 27 3166 8-10 d-e levileugeo ordinarius yang & wang 1 21 loopus doliolum martae beccaro 3 17 lpmt c-e mirifusus guadalupensis pessagno 3 -1 3160 5-11 monotrabs goricanae beccaro 1 24 mtgc a-b monotrabs plenoides gr. baumgartner 1 23 3152 5-8 obesacapsula morroensis pessagno 3 2 3266 5-21 orbiculiforma sp. b sensu widz 1991 1 25 palinandromeda podbielensis (ozvoldova) 3 3, 4 3008 5-9 pantanellium riedeli pessagno 1 30 3078 7-12 b-f parahsuum stanleyense (pessagno) 3 24 2023 3-8 paronaella broennimanni pessagno 1 28 3137 4-10 paronaella mulleri pessagno 1 26 3139 6-10 paronaella pygmaea baumgartner 1 27 3133 7-9 e podobursa andreai beccaro 3 5 pdan b podobursa polyacantha (fischli) 3 6, 7 3174 5-8 a-d podobursa triacantha (fischli) gr. 3 8 pdtc b-f podobursa vannae beccaro 3 -10 pdvn c-f praewilliriedellum convexum (yao) 2 26 3055 1-11 b praewilliriedellum robustum (matsuoka) 2 27 3298 5-7 b pseudodictyomitrella tuscanica (chiari, cortese & marcucci) 3 18 pseudoeucyrtis firma hull 3 -19 3176 5-10 a-b pterotrabs arcuballista dumitrica, baumgartner & gorican 2 1 ristola procera (pessagno) 3 -16 3163 5-9 saitoum elegans de wever 3 13 3022 8-21 saitoum levium de wever 3 -11, 12 3024 4-9 d-f saitoum pagei pessagno 3 -10 3020 4-11 sethocapsa sp. cf. s. trachyostraca foreman 2 29,30 3063 7-22 striatojaponocapsa plicarum s.l. (yao) 3 15 3051 3-8 b tethysetta dhimenaensis dhimenaensis (baumgartner) 3 20 4072 3-11 b tethysetta dhimenaensis ssp. a sensu baumgartner et al. 1995c 321,22 4071 3-8 a-b tetradytrima corralitosensis s.l. (pessagno) 2 3 3273 3-10 tetraditryma pseudoplena baumgartner 2 4 3123 4-11 tetratrabs zealis (ozvoldova) 2 5 3121 4-13 b-e theocapsomella himedaruma (aita) 2 28 4038 transhsuum brevicostatum gr. (ozvoldova) 3 26 3181 3-11 a-e transhuum maxwelli gr. (pessagno) 3 25 3180 3-10 triactoma blakei (pessagno) 2 6 3095 4-11 c-f triactoma enzoi beccaro 2 8 tcez b-d triactoma jonesi (pessagno) 2 11 3096 2-13 triactoma mexicana pessagno & yang 2 10 3412 5-9 triactoma parablakei yang & wang 2 9 3413 4-7 b tritrabs casmaliaensis (pessagno) 2 14 3117 4-10 b-f tritrabs ewingi s.l. (pessagno) 2 13 3113 4-22 b-f tritrabs rhododactylus baumgartner 2 15 3118 3-13 e-f williriedellum carpathicum dumitrica 3 29 4055 7-11 b-e williriedellum (?) marcucciae cortese 3 30 4060 4-8 b-c williriedellum yahazouense (aita) 3 28 williriedellum yaoi (kozur) 3 27 zhamoidellum ventricosum dumitrica 3 31 3308 8-11 c-f age of the samples by uaz-sa (beccaro 2006a) age of the samples by uaz95 (baumgartner et al. 1995b) preservation index (kiessling 1996) 7 78 3: good 4: average 5: fair 6: poor 7: very poor a: early-mid bath. early call b: early call.-early oxf. c: middle oxf. d: middle-late oxf. e: late oxf.-early kimm. f: early-late kimm. 6: mid bath. 7: late bath.-early call. 8: middle call.-early oxf. 9: middle-late oxf. 10: late oxf.-early kimm. 11: late kimm.-early tith. b 5-6 4-5 3-4 4-54-5 3-4 5-6 6-7 �#%+ � � � "# �&"#'(��#� $�� "�%� ��! �"#$���#"�#! #8# # ��! � ��""# ��� ��! 0�"�! � ��# �#� ��� (�"! ��'� � #�)1+ �(� �'����� �#"")�!& # !���"��#� ��$� #"� ����� "# �$ �! �#��&#" !�" � #�+ 0�66a�1> (� �'����� �#"")�!& # �� �"�$ ��$� #"� �!���$�$ �! ����#"� 0�44b#1+ �(� #&�� � (� �#�'��� #"� &�7�! %) (� �!� #") ������# ��! c�!�� ��#��&#" !�" � #�+ 0��c6a> �66a%1 #!$ %) (� �!� #") ������# ��! c�!�� -�" �����) #!$ (� ��� (�"! ��'� 0��c���1 �����#"� 0�44b#1+ �(� �"���"7# ��! �!$�8 "#!&�� -"�� � 0&��$1 � 3 07�") '��"1 0?������!& �66b1+ �#�( #8�! �� �#"2�$ %) # ��$� �! �#%+ �� (� �'����� �#"")�!& # !���"��#� ��$� #"� ����� "# �$ �! 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**�� ���5�� ��5$� 2�� �������3� ����� �5� :�� �����,� /��g� � �5�� 2a44c3 ����!�� )� )� ��� ��*��� �� ���� �� � � ����� � �� � ������� *�� ���� ����� �* ���� ���� c&� c da&�dca� ?� �/�,�� rivista italiana di paleontologia e stratigrafia volume 104 numero l pagrne 123-130 aprile 1998 noa breve middle pliocene cetaceans from monte voltraio (tuscany, italy). bio stratigraphical, paleoecological and paleoclimatic observations giovanni bianucci (*), giovanni sarti (**), rita catanzariti (***), ubaldo santini (****) received august 13, 1997; accepted jarutary 19, 1998 key-uord;: odontoceti, cetacea, systematics, stratigraphn biostratigraphy, paleoecology, paleoclimatology, middle pliocene, tuscany, ita.y. riassunto. viene esaminata la collezione storica di odontoceti (cetacea) fossili del monte voltraio vicino a volterra (toscana) e viene condotta un'indagine litostratigrafica e biostratigrafica nella località di ritrovamento. laffioramento di monte voltraio è attribuito al pliocene medio, in particolare alle zone a globorotalia aerniliana e discoaster tarnalis. i resti di odontoceti vengono riferiti alle famiglie kogiidae (kogia pusillq e delphinidae (globicephala? eturiae e dre reperti indeterminati che potrebbero appartenere a hemisyntrachelus e a stenella giulii). la fauna a cetacei del pliocene medio del bacino mediterraneo (associazioni di monte voltraio e di fuo stramonte) include taxa estinti o attualmente assenti in questo bacino. la loro scomparsa potrebbe essere messa in relazione ai deterioramenti climatici pliocenici e,/o quaternari (per esempio, 1a crisi climatica intorno a 2.6-2.4 ma). abstact. the historic collection of fossil odontocetes (cetacea) from monte voltraio, near volterra (tuscann ltaly) has been examined and lithostratigraphical and biostratigraphical investigations on the find locality have been carried out. the monte voltraio outcrop ìs referred to the middle pliocene, in panicular to globorotalia aetniliana and discoaster tamalis zones. the odontocete remains are assigned to the families kogiidae (kogia pusilla) and delphinidae (globí cephala? etruriae and two indeterminate specimens which might belong to hemisyn*acbelus and stenella gìuli). the middle pliocene cetacean fauna from the mediterranean basin (monte voltraio and rio stramonte associations) includes extinct tàxa or extant taxa no longer represented in this basin. the disappearance of these raxa may be linked with the pliocene andlor quaternary climatic deteriorations (e.g. the climatic crisis at *tout 2.6-2.4 m,l). lntroduction. some historic collections and new finds of fossil cetaceans from italian piiocene sediments have recently been examined (bianucci, 1996a-b, 1997a-c). the new systematic interpretation of these records confirmed a basic renewal of the mediterranean fauna during the pliocene with respect to the archaic miocene cetacean association, already pointed out by bianucci & landini (l99i, 1992). these studies also documented the presence during the pliocene of some extinct taxa or extant taxa no longer represented in the mediterranean sea. these taxa disappeared from the mediterranean basin in the course of the pliocene and/or of the quaternary. the evolution of these mediterranean representarives during the last 5 ma is unknown and possible evenrs of extincrion and/or renewal events, at present, are not recognized. this is due to the fact that most of these fossils were collected in the second half of nineteenth century without reliable stratigraphic reference. this paper is a biostratigraphical study aimed to shed new light on the evolution of the pliocene cetaceans of mediterranean basin. materials and methods. the four cetacean fossils examined make part of the lawley collection, which includes many other cetacean remains collected from pliocene sediments of tuscany (pilleri, 1987). these specimens are preserved in the museum of geology and palaeontology of the university of florence (igf). although incomplete and fragmentary these specimens are relatively significant from both a systematic (two of them are the holotypes and the only known records of fossil species) and from a stratigraphical point of view (the fossils are very close in age), these specimens had been collected in the nineteenth century from pliocene sediments outcropping af "la rocca", 1ocality on monte voltraio, a hill 458 m above sea level about 2 km east of volterra (province of pisa) fig. t) the systematic analysis of cetacean fossils was completed by a geotrogical and biostratigraphical study of the monte voltraio outcrop. a section represenrative of the whole depositional sequence of this area was examined and sampled. twenty-nine samples were collected for micropaleontological analysis (foraminifera and calca/'i\ n:-'*:---'^ r: scienze della terra, via s. maria, 53,56126 pisa (italy). ("'f) consulting of emilia romagna region, via amendola, 4,56127 pisa (italy). ('r't'f) consulting of emilia romagna region, via possenti, 10, 56010 asciano, pisa (ita1y). 1*++t) via cristoforo colombo, 25, 55a49 viareggio, lucca (italy). 1,24 g. bianucci, g. sarti, r. fig. i location of monte voltraio cetacean assemblage. a : studied section. reous nannofossils). the small amount of matrix still preserved on the fossil specimens was also sampled for nannofossils. the abundance of the species ol discoaster was noted in the samples in which l0 or more specimens of this genus were counted. at the latest 100 specimens were counted in the samples with very abundant discoasfer (backmann ec schakleton, 1983; rio et al., 1990). the plankrcnic foraminiferal zonal scheme of laccarino & salvatorini (1982) followed in other papers concerning the pliocene sediments of tuscany (bossio et a1., l995) was also followed here. the calcareous nannofossil zonation is by rio et al. (1990). geological and biostratigraphical setting. the extensional basin of volterra is filled by rpper miocene-quaternary sequences (sarti & testa, 1,994; bossio et al., 1995). pliocene deposits outcrop at monre voltraio. they consist of grey claystones (called "pag" on the foglio 12 of the 1/100,000 national geological cartography) at the bottom, and by yellow sandstones at the top. the studied section is exposed to se, of monte voltraio; here the succession has an average thickness of 150 m fig. 2). the first 100 m at the base of the section are grey massive silty claystone. moving upward the sand content increases rapidly, and after few alternating sand and clay layers yeilow medium-fine sands outcrop at the top of the sequence. three decimetric levels of weil cemented grey bioclastic (bivalves and gastropods mainly) calcarenites are present in the uppermost part of the section. the foraminiferal assemblage is suggestive of a transition from an outer-inner neritic zone (basal-medium part of the section) to a littoral zone (upper part of the section). in particular the associations are diversified and abundant in the samples collected from the lowest part of the section (p\b ratio 1\10-1\5). in the samples from the middle and upper part there is a progressive replacement of planktonic foraminifers and of benthic taxa from the deepest environcatanzariti, u. santini ments by littoral benthic assemblages. therefore the section represents a coarsening and shallowing upward sequence. the genera sclpbosphaera and pontospbaera, both belonging to the family pontosphaeracae, are parricularly well represented in the nannofossil assemblage. these taxa indicate neritic and emipelagic environments (perch-nielsen, 1985), which is consistent with the foraminiferal datum. from a biostratigraphical point of view, the presence in the studied section of globoroulia crassaformis crassaformis associated wíth globorotalia bononiensis and globorotalia aemiliana in the claystone part of the sequence suggests a reference to the lower part of globoro, talia crassaformis crassaformis subzone (upper part of globoroulia aemiliana zone) and therefore this part of sequence is dated middle pliocene. the calcareous nannofossil association is referable to the discoaster tamalis zone. in fact, in the samples where the discoaster is relatively frequenr, the identified species of this genus (d. swrculus, d. tamalis, d. asymmetricus, d. brouweri, d. intercalaris, d. pentaradiatus, d. triradiatu) show abundance comparable to those of di scoaster tamalis zone of the western mediterranean basin reported by rio et al. (1990). zonai markers are lacking in the upper part of section where littoral conditions are reached. detailed regional biostratigraphical studies carried out in tuscany (bossio et ai., 1995) show that the regressive upper parr of the pliocene deposits can be entirely included in globorotalia aemiliana zone. the upper pliocene deposits (globorotalia inflata zone) are indeed lacking in tuscany because of a general uplift of the whole area. the calcareous nannofossil assemblage contained in a sample collected from the sediment preserved on the fossil delphinid igf 1562v is referable to rhe same zone (discoaster tamali) identified in the studied section of monte voltraio. the samples collected from the sediment on the others fossil cetaceans either are sterile or bear biostratigraphicaly insignificant caicareous nannofossil associations. systematic palaeontology. class mammalia linnaeus, 1258 order cetacea brisson, 1762 suborder odontoceti flower, l867 family k o g i i d a e (g1ll, 1871) mìller, 1923 genus kogia gray, 1846 kogia pusilla (pilleri, 1987) (fig. 3ar-az) 1893 placoziphias, v beneden capellini, pp.287-288. 1,987 hyperoodon pusillus pilleri, pp. 35-36, fig. 11, pl. 13. 7997c kogia pusilla (pìllert) bianucci, pp. 172-173, {tg. 6. 1,25middle pliocene cetaceans from monte voltraio €. èrl .--,; t.g no é9x o* rca sè òo = ú; '= g 6 dc 2\? é ^r ., o qi ", d >'1, 'e o o b ll"ò ,>tr o'ú tr cè^lî,a ph -gjh ! e .9 op >\ o :'^ îu* ú.;1,x tr.tr iú' --9x ào;ì f = 'n = i àbf" oòt iú n @ ui o plpln3alund c sllc swolossbj? s wajjea whalqo j paìromal sja)sooss p ldlol (tii) ss:{n)i3ihj iù & = tsa o z o f z l a fo & a zz z 0661 lo l8 oru +:f òe sinnn 9è 3è i, i nnil q9 tnnt{ l oh 9+dd qo a 09 i nnt'.l ;€ s;:v rnîr )g6t àrìnb rt opdìo i olll o zt n 3 z t 1.6 | lu!.p dt't @ zz r zz -t--zlz zz =z 'ooo o; !@ d dd slè'r bl punal ? | d6 : i qr: i ì è.5 3 i ds : àhr s"j ruao ounlojoqoln d ?of@?túd o!, lb lo b |o?j ét déi €è s1 p{e 3è. 3'go ol!c'pueua 9161 d j l 6 f c o naèhj ssnvc i lusattc ^1 tè\.lod ti rrtrlrtlrtìltttttlll úobo .:dr;+ ahdvuc i.ivuls o nouh 3 nvtsvtlgl nvrzr.l]cvraluv:rcruvz fnlcottd t26 g. bianucci, g. sarti, r. catanzariti, u. santini the skull (igf1540v) is slightly deformed and lacks of part of the right side and of ventral portion of braincase. this specimen 'was originally referred to a ziphiid by capellini (1893) and pilleri (1987), but actually belongs to the kogiidae (bianucci, 1,997c).it represents the only preserved fossil skull of pigmy sperm whale of the genus kogia. it is the holotype and the only known record of the extinct species kogia pwsilla (pilleri, 1987). kogia pusilla differs from living k. simus and k. breviceps in having a more elongated rostrum, a smaller antorbital process, and an apparently more marked dorsal asymmetry. the posterior portion of the cranium is also slightly more uplifted. a detailed description of this species will appear in a future paper (bianucci, in preparation). family d e i p h i n i d a e gray, 1825 genus globiceohala lesson. 1828 globicephala ? etruriae (pilleri, 1987) (fig. 3br-b3) 7987 globicepbala etruriae pilleri, pp. 28-30, p|. 6. 79961: globicepbala? etruriae (pilleri) bianucci, pp. 98-99, fig. 33, pl. 9, fig. 3-s . the incomplete mandible lacks the posterior parts of the rami but still bearing 30 teeth in place (igf 1675v). pilleri (1982) choose this specimen as holotype of the extinct delphinid species globicepbala euuriae. this mandible (the only known record of g. etruriae) differs from the mandible of living pilot whales (globicephala melas and g. macrorlrynchu) in its smaller size and in larger number of teeth. the monte voltraio fossil may belong, however, to a different genus from that of the two living species (bianucci, 1996b). delpbinidae indet. 1987 tì.trsíops sp. pilleri, p. 44. fourteen complete teeth and seventeen fragments of teeth (igf 1562v) were referred to thrsiops sp. by pilleri (1982), but are morphologically and dimensionally similar to the teeth of hemisyntrachelus, a delphinid which lived in the mediterranean sea during the pliocene (bianucci, 1996b, 1997a, 1997b). isolated teeth are however inadequate for generic determination. delphinidae indet. 1987 odontocett gen. et sp. indet. pilleri, pp. 47-48, ftg. 13. seven cervical and the first two thoracic vertebrae, belonging to of the same animal (igf 15489, all lack the neural and transverse processes. pilleri (1987, pp.4748) referred this find to "odontoceti gen. et sp. indet" (slc). the atlas and the axis are fused and the other cervical vertebrae have small and very flatten corpora; the thickness of the corpora increase in the first two thoracic vertebrae. these vertebrae are illustrated by pilieri (1.987, fig.13) although he misinterpreted the fused atlas and axis as the first cervical vertebra (and consequently in pilleri's iliustration there is only one thoracic vertebra). the atlas is similar to but larger than that of stenella sp. from the pliocene of tuscany (bianucci, 1996b). these vertebrae might belong to the relatively large-size dolphin stenella giulii, an extinct species represented by only four finds from pliocene sediments of orciano (tuscany). the calcareous nannofossil content of the sediment preserved on three of these fossil specimens of stenella giulii (one, the holotype, has been lost) was analysed. the sediment on one of these specimens (igf 1541v) contained a biostratigraphically insignificant nannofossil assemblage, while the assemblages of the samples from the other t.wo specimens (mc cf1, igf i545v) are attributable to the same nannofossil zone (discoaster tama/i) found in the monte voltraio outcrop. these biostratigraphical data support the hypothesis of the presence of stenella giulii ín the monte voltraio sediments. discussion and conclusion" the bathymetric analysis of foraminiferal and nannofossil associations indicate the transition from the inner-outer neritic to the littoral zone. there is an apparent discrepancy between the environmental conditions documented by the plankton and the pelagic habitat indicate by kogia and the delphinid near globicephala. the recent species of these cetaceans live on the continental siope and feed in the bathyal environment. i-ong drifting of the carcasses from pelagic water to near coast environment may have occurred after death. prolonged carcass transportation has actually been observed in present-day cetaceans (shafer, 1972); such an explanation has already been advocated to explain the odd presence of a fossil pelagic whale in a shallow'water environment (bianucci, i996a). the odontocete cetacean assemblage from monte voltraio can be re{erred to a restricted time span in biostratigraphical terms, ranging from approximately 3 to 2.6 lr/.a. this is the same interval in which the clayed marls of rio stramonte in piacenza province were deposited (monegatti & ranieri, 1987; monegatti, pers. com.). these clays yielded fossil remains of cetaceans in the past (bianucci, 1.997b; cigala fulgosi, 1990; del prato, t897,1900). the two associations together shed new light on the middle pliocene cetacean fauna of the mediterranean basin (tab. 1). middle pliocene cetaceans frorn monte voltraio 127 'locm odontocete remains from middle pliocene sediments of monte voltraio (tuscany). ay kogia pusilla (pllleri, 1987), incomplete skull in dorsal view; a2, the same in lateral view; br, globicephala? eturiae (prlleri, 1987), incomplete mandible in dorsal view; bz, the same in ventral view; b:, the same in lateral view. fr : frontrl, la : lacrimal; mx : maxilla; n : externàl nares, pmx : oremaxilla. br 1,28 g. bianucci, g. sarti, r monte voltraio rio stramonte suborder odontoceti family delphinida globice phala? etruriae he mi syntrac helus coftesi i hemisyntrachelus sp. stenella giulii family kogiidae kogia pusilla suborder mysticeti family balaenidae balaena paronai tab. 1 middle pliocene cetaceans from monte voltraio (tuscanv) and rio stramonte (emilia romagna). the three families recognized are still living, although two (kogiidae and balaenidae) are no longer represented in the mediterranean sea. the listed taxa include both living (kogia, balaena and probably stenella) and extinct genera (hentisyntracbelws and a delphinid near globicephala). kogia and balaena, as are their families, are missing today from the mediterranean sea. all of these raxa are extinct at the specific levei. these data indicate that the mediterranean cetacean association of middle pliocene differed from the extant one in composition and structure. pliocene and/or quaternary phases of climatic deterioration (and consequently changes in water circulation, in the water productivity and in the food availability) might have caused the extinction of some middle pliocene cetaceans. the effects of climatic changes and of other correlated factors are already associated with variations in cetacean distribution (barnes, 1974; davies, 1963; fordyce, 1980, 1984; fordyce & barnes, 1994; gaskin, 1926; \x/hitmore, 1,994).in particular, food avaiiability seems to have an important control on the structure and composition of cetacean communities (fordyce, 1996; lìpps & mitcheil, 1976). as regards, the co-occurrence of a delphinid near globicephala and of kogia (both fed on squids in the bathyal environment) in the middle pliocene monte voltraio association, indicate that the teutophag trophic resource was probably larger than that existing today. the analysis of isolated periotics from lower-middle pliocene sediments of tuscany confirms the presence of kogiids and teuthophag delphinids (probably globicephala and grampus, see bianucci, 1996lr) in the mediterranean earlier than the late pliocene. kogia today shows an almost cosmopolitan distribution (caldwe1l & caldwell, 1989) and its absence from the mediterranean might be due to the scarce food supply and to the competition with teuthophag delphinids (grarnpus, globicephala and pseudorca). the climatic deteriorations in the mediterranean basin have been dated and calibrated on the basis of excatanzariti, u. santini tinction events of molluscs (raffi & monegatti, 1992; raffi et al., 1985) and benthic foraminifera (sprovieri, 1986). a possibie change in the mediterranean cetacean communities may have occurred at the time of the intense climatic crisis which is detected at about 2.6-2.4 ma (approximately correlatable with the upper deposition boundary of monte voltraio and rio stramonte outcrops). according to landini ee menesini (1988), this climatic deterioration caused the extinction of bregmaceros (teleost, fish) from the mediterranean basin. this extinction episode of molluscs, foraminifera and fishes has been correlated with the first major glacial event of the northern hemisphere (thuneil & williams, 1983). a better calibration of cetacean extinction events with and climatic changes will be possible when systematic and biostratigraphical studies of other mediterranean cetacean assemblages (particularly from the late pliocene and quaternary) are carried out. acknouledgements. \le are grateful to p. mazza (museo di geologia e paleontologia, university of florence) for grating us access to the monte voltraio cetacean collectron and for revising the english. many thanks to \7. landini (dipartimento di scienze della terra, university of pisa) for the interesting discussions. 'we also wish to thank d. j. bohaska (u.s. national museum) for critically reading rh" -".".".i.t references backman j. & shackleton n.j. (1983) quantitative biocronology o{ pliocene and early pleistocene calcareous nannoplankton from the atlantic, indian and pacific oceans. mar. micropaleont., v. 8, pp. 14i-170, amsterdam. barnes l.g. (1974) outline of eastern north pacific fossil assemblages. syst. zool., v. 25 (a), pp. 321-343, new habianucci g. (1996a) a new record of baleen whale from the pliocene of tirscany (italy). atti soc. tosc. sci. nat., mem. (ser. a), v. 102, pp. 101-104, pisa. bianucci g. (1996b) the odontoceti (mammalia, cetacea) from italian pliocene. systematics and phylogenesis of delphinidae. palaeontograpbia ltal., v. 83, pp. 73-167, pisa. bianucci g. (1,997a) hemisyntrachelus cortesii (cetacea, delphinidae) from pliocene sediments of campore quarry (salsomaggiore terme, italy). boll. soc. paleont. iral., v. 36 (1., 2), modena. (in press). bianucci g. 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(eds.) contributions in marine mammals paleontology honoring frank c. $lhitmore jr. proc. san diego soc. nat. hist., v.29, pp. 233-227, san diego, california. rivista italiana di paleontologia e stratigrafia volume 117 no. 1 pp. 3-13 april 2011 a new species of cyrtospirifer (brachiopoda) from the middle devonian of the western sahara (northwestern africa) mena schemm-gregory received: december 10, 2010; accepted: february 23, 2011 centro de geociências e departamento de ciências da terra da universidade de coimbra, largo marquês de pomal, 3000-272 coimbra, portugal; phone: 00351-239-860-571, fax: 00351-239-860-501. e-mail: mena.schemm-gregory@dct.uc.pt key words: cyrtospirifer, brachiopoda, phylogeny, palaeobiogeography, givetian, devonian, western sahara. abstract. a new species of cyrtospirifer is described from the middle to upper givetian of the western sahara (northwest africa). cyrtospirifer tindoufensis n. sp. differs in its smaller number and coarser medial and flank plications and equibiconvex shell profile from the other givetian species of cyrtospirifer that all occur in europe and to which the new species probably gives rise. the new implications of the proposed phylogeny of the earliest cyrtospiriferids and their origin from the western sahara are discussed. the palaeogeographic distribution of the cyrtospiriferids during the givetian and frasnian is shown and its migration ways are described considering the global transgression and regression cycles. riassunto. in questo lavoro viene descritta una nuova specie appartenente al genere cyrtospirifer proveniente dal givetiano mediosuperiore del sahara occidentale (africa nord-occidentale). cyrtospirifer tindoufensis n. sp. differisce dalle specie europee del genere cyrtospirifer, alle quali ha dato origine, per il profilo equi-biconvesso e per il numero minore di plicae mediane e sui fianchi, le quali risultano inoltre più robuste. vengono inoltre discusse le implicazioni della filogenesi proposta per i cyrtospiriferidi più antichi e la loro possibile origine nel sahara occidentale. vengono infine descritti la distribuzione paleogeografica dei cyrtospiriferidi durante il givetiano e il frasniano e i possibili modelli di migrazione tenendo conto dei cicli trasgressivi e regressivi globali. introduction taxa attributed to the genus cyrtospirifer nalivkin in fredericks, 1924 have been studied since the 19th century (e.g, murchison 1840; quenstedt 1871; gosselet 1894) and are used as important index fossils for the upper devonian due to their distinctive morphology and global pan-tropical distribution. as a result, almost every upper devonian spiriferid with fine medial plications historically has been assigned to cyrtospirifer. however, the oldest species of cyrtospirifer first evolved by the mid-to-late givetian (upper middle devonian) (e.g., paeckelmann 1942; brice et al. 1976, 1979; brice 1982, 1988, 2003). in recent studies focused on revisions to key cyrtospiriferids ma & day (1999, 2000, 2003, 2007) and ma et al. (2003) revised cyrtospirifer. in their 2003 revision of cyrtospirifer they highlighted the fact that historically cyrtospirifer has been used as a taxonomic garbage can for upper devonian cyrtospirifer-like taxa resulting in assignment of what have been demonstrated to be distinct genera and species (see examples cited by gratsianova et al. 1989; ma & day 2003). ma & day (2003: 270) proposed an evolutionary tree for the early cyrtospiriferids of the givetian and frasnian. however, the oldest species shown in their tree is probably younger than the new species from north africa described in this work, according to brachiopod data (schemm-gregory & jansen 2005). the primitive morphologic characters of c. tindoufensis, such as coarse and few plications, help to fill in the record on the earliest-oldest known genuine species of cyrtospirifer in the tenticospirifer-cyrtospirifer evolutionary lineage proposed by ma & day (2000). geological setting during field sampling conducted as part of the german-moroccan co-operation “genesis of devonian reefs” new specimens of cyrtospiriferid brachiopods were collected from givetian-age carbonate 4 schemm-gregory m. and clastic rocks at the southeastern flank of the tindouf syncline close to the military town of smara in the western sahara (northwest africa) (fig. 1). the middle devonian beds in the study area are dominated by sandstones, siltstones, and shales; thin beds of limestones and associated bioherms and biostromes at several levels occur, too. the homoclinal structure of this area, the lack of vegetation, and the removal of the surrounding shale created unique exposures revealing the structure of reefs and associated off-reef sedimentary successions. between 1930 and 1950 spanish and french geologists started to examine the western sahara for the purposes of petroleum exploration after the second world war (such as jacquet 1936; hernández-pacheco et al. 1949). dumestre & illing (1967) described the development of middle devonian reefs of this region and compared them with reefs of equivalent age in the rainbow area of northwestern alberta (canada). the study area itself lacks a modern study of geology and paleontology, thus the lithology is still undescribred and strata lack formation names, how opods were collected from givetian-age carbonand clastic rocks on the southeastern flank of the tindouf syncline close to the military town of smara in the western sahara (northwest africa) (fig. 1). the middle devonian beds in the study area are dominated by sandstones, siltstones, and shales; thin beds of limestones and associated bioherms and biostromes occur at several levels. the homoclinal structure of this area, the lack of vegetation, and the removal of the surrounding shale created unique exposures revealing the structure of reefs and associated off-reef sedimentary successions. between 1930 and 1950 spanish and french geologists started to examine the western sahara for the purposes of petroleum exploration after the second world war (for example jacquet 1936; hernández-pacheco et al. 1949). dumestre & illing (1967) described the development of middle devonian reefs of this region and compared them with reefs of equivalent age in the rainbow area of northwestern alberta (canada). the study area itself lacks a modern geological and paleontological study, thus the lithology is still undescribed and strata lack formation names. however, correlation with western european and north african strata based on faunal content is possible and biostratigraphical assignment are done with different fossil groups. in the recent years, a few studies of the bryozoan and stromatoporid faunas have been started (scholz et al. 2005; königshof & kershaw 2006). the brachiopod faunas recovered from the study area indicates correlation of the reefs to the middle-upper givetian (upper middle devonian) and lower frasnian (lower upper devonian) (schemm-gregory & jansen 2005). conodont samples did not give sufficient data for stratigraphic assignment (p. königshof, pers. comm. 2010). recent palaeobiogeographic studies on the genus paracrothyris (wu in wang et al. 1974) discuss the global faunal exchange between nevada (western usa), south china, and the western sahara during this time interval (schemm-gregory & jansen 2008). fig. 1 map of the western sahara region of northwest africa showing location of the study area. fig. 2 geographical grit showing locations of fossil localities plotted. a new species of cyrtospirifer (brachiopoda) from the middle devonian of the western sahara 5 the studied outcrops form a belt in a northwestsoutheast direction consisting of bioherms of different sizes. in previous works, the largest of the studied reefs was named “sabkhat lafayrina” situated in the se, a smaller one “gor-al hessen” is located in the nw of the study area (königshof et al. 2003). the other reefs are numbered 1 to 10 (fig. 2). cyrtospirifer tindoufensis was found in proximal off-reef strata near the reefs and, in rare cases, within reefs themselves. material and methods specimens of cyrtospirifer described herein are usually preserved as disarticulated internal and external moulds (figs. 3a, b, 4 a, b). articulated shells and rare isolated shells were also recovered. because of weathering the shell material is poorly preserved and not well suited for taxonomic studies. latex casts of internal and external moulds were made to study the internal features of the new taxon (figs. 3c, 4c) as well as the micro-ornamentation (fig. 5). due to the extraordinary preservation of internal molds, serial sections of articulated specimens were omitted. measurements were taken with a digital caliper and rounded to 0.1 mm. drawings were made with the help of a camera lucida. specimens were coated with magnesium oxide prior to being photographed. institutional abbreviations smf: senckenberg forschungsinstitut und naturmuseum, frankfurt am main, germany. systematic palaeontology order spiriferida waagen, 1883 suborder spiriferidina waagen, 1883 fig. 3 morphological terms of internal ventral mould of cyrtospirifer tindoufensis; a: smf 66595b, ventral view. b: smf 66081a, posterolateral view. c: smf 66081c, holotype, plan view on latex cast. scale bar =10 mm. fig. 4 morphological terms of internal dorsal mould of cyrto spirifer tindoufensis, smf 66081b; a: dorsal view. b: oblique posterior view. c: plan view of latex cast. scale bar= 10 mm. 6 schemm-gregory m. superfamily cyrtospiriferoidea termier & termier, 1949 family cyrtospiriferidae termier & termier, 1949 subfamily cyrtospiriferinae termier & termier, 1949 cyrtospirifer nalivkin in fredericks, 1924 type species: spirifer verneuili murchison, 1840: 252, pl. 2, figs. 3a-e. diagnosis: medium to large-sized cyrtospiriferids with well developed delthyrial plate, plate thickened to various degrees, ventral interarea low-concave to high-straight; delthyrium wide, sometimes with pseudodeltidium in the upper third of the delthyrium; with micro-ornamentation (where preserved) of minute pustules in grooves and on plications: 8 to 30 sinal plications differentiated into a medial and two lateral regions; dental plates well developed. [after ma & day (2003)]. discussion. this study shows that the diagnosis of cyrtospirifer has to be emended to include shell features observed in the new material enhanced by the number of sinal plications, it decreases in c. tindoufensis n. sp. and increases in c. syringothyriformis. the ventral interarea may also be catacline and flat. however, a revision of cyrtospirifer is far beyond the scope of this work. stratigraphic distribution. middle givetian to lower famennian (upper middle devonian to lower upper devonian). geographic distribution. cosmopolitan. cyrtospirifer tindoufensis n. sp. figs 3-7, tab. 1 holotype: internal mould of ventral valve, stored in the senckenberg forschungsinstitut und naturmuseum under the inventory number smf 66081c (figs. 6.1a-c, 9). length 23.5 mm and width 41.5 mm. derivation of name: after the tindouf syncline on which se flank the type locality is situated. type locality: reef no. 4 (n 26° 41,636, w 11° 39,904‘), western sahara (northwest africa). type horizon: beds of upper givetian (upper middle devonian). stratigraphical distribution: middle givetian to ?lower frasnian (upper middle devonian to ?lower lower devonian). geographic distribution: tindouf syncline, western sahara (northwest africa). material: reef no. 4: loc. 12 (n 26° 41.636‘, w 11° 39.904): 4 ventral internal moulds (smf 66081a, c holotype, d, e); 1 dorsal internal mould (smf 66081b); few single dorsal and ventral internal moulds (un-numbered). loc. 13 (n 26° 41.646‘, w 11° 44.120‘): 2 ventral external moulds (smf 666595a, b); 5 ventral internal moulds (smf 66603, 66605a-c); 1 dorsal exfoliated shell (smf 66604); 1 dorsal external mould (smf 66607); few single dorsal and ventral internal moulds. reef no. 5: loc. 15 (n 26° 40.397‘, w 11° 44.028‘): numerous single dorsal and ventral external moulds. loc. 16 (n 26° 40.229‘, w 11° 44.027‘): 2 ventral external moulds (smf 66083, 66598); 4 ventral internal moulds (smf 66594, 66596b, 66600, 66601); 1 dorsal external mould (smf 66597); 1 dorsal internal mould (smf 66596a); 1 articulated internal mould (smf 66606); numerous single dorsal and ventral external moulds (un-numbered). loc. 17 (n 26° 40.221‘, w 11° 44.090‘): numerous single dorsal and ventral external moulds (unnumbered). loc. 18 (n 26° 40.216‘, w 11° 44.104‘): 1 exfoliated dorsal shell (smf 66602); numerous single dorsal and ventral external valves (un-numbered). remarks: the locality numbers refer to the expedition locality numbers. further details may be obtained by the curator of the brachiopod collection of the senckenberg forschungsinstitut und naturmuseum. diagnosis: cyrtospirifer with medium-sized shell and an equibiconvex to ventribiconvex profile. megathyrid, in rare cases brachythyrid outline. dental plates extrasinal, 8 to 12 plications in the sulcus. c. tindoufensis n. sp. c. verneuili (murchisoni, 1840) c. verneuiliformis (paeckelmann, 1942) c. syringothyriformis (paeckelmann, 1942) size medium to large medium to large medium to large large outline megathyrid to brachythyrid megathyrid megathyrid strongly megathyrid curvature in longitudinal section equibiconvex to ventribiconvex ventribiconvex ventribiconvex dorsibiconvex ventral interarea moderately high apsacline to catacline curved moderately high apsacline to catacline curved high catacline sometimes curved high catacline straight dental plates extrasinal extrasinal extrasinal intrasinal sulcus and fold in cross section flattened to rounded flattened rounded rounded posterior, flattened anterior number of plications in sulcus 8-12 14-20 15-20 20-30 cardinal extremities convex convex ? convex and laterally flat bifurcation of plications on flanks + fold strongly elevated strongly elevated hardly elevated moderately elevated crural plates + + ? stratigraphic distribution upper givetian upper givetian upper givetian to frasnian upper givetian to frasnian tab. 1 comparison of species of cyrtospirifer nalivkin in fredericks, 1924 occurring in the givetian of western europe and north africa. a new species of cyrtospirifer (brachiopoda) from the middle devonian of the western sahara 7 description of the holotype (smf 66081c) form and size the ventral internal mould is transverse, semielliptic in outline, and megathyrid with small mucronations. the specimen is convex in longitudinal section. exterior of ventral valve the ventral umbo extends to posterior over the hinge line. the ventral interarea is high, apsacline, and curved. the delthyrium is open. deltidial lamellae very fine. interior of ventral valve ventral muscle field extends to posterior over the hinge line. lateral apical cavities reaching almost to the posterior end of the muscle field. hardly any shell material developed in apical region. a small and narrow delthyrial plate is posterior to the ventral muscle field from which it is separated by a noticeable constriction on the internal mould. ventral process small and broad leaving a distinct groove on the internal mould at the posterior end of the muscle field. the myophragm is very thin, almost not recognisable leaving a very fine furrow through the whole ventral muscle field on the internal mould. the muscle field is elongated, pyriform in outline, and not impressed into floor of valve. diductor scars are impressed as subradial striae in the centre and as longitudinal striae in the anterior part of the muscle field on the internal mould. adductor scars elongated, thin, situated admedially to the myophragm, and enclosed completely by the diductor field; posterior and anterior pair of adductors gently impressed into shell. free portions of dental plates short, leaving small slits on either side of the posterior half of the ventral muscle field on the internal mould. gonoglyphs are not preserved. sulcus deep, with steep flanks, and rounded in cross section. impressions of 10 simple and fine plications are countable in the sulcus. impressions of 18 plications on each flank are preserved, their expression diminishes towards the lateral margins. plications are separated by narrower furrows; furrows and plications are rounded in cross section. impressions of growth lamellae faintly preserved in the anterior half of the internal mould. description of paratypes form and size shells medium-sized and wider than long. ventribiconvex profile and semicircular to semielliptic in outline. megathyrid, in rare cases brachythyrid, with small, thin, and convex mucronations that are rarely preserved with their lateral terminations on the internal moulds. maximum width 51.6 mm, maximum length 25.0 mm. exterior of ventral valve ventral interarea apsacline to catacline, in the upper part curved with transverse growth lamellae. delthyrium open, restricted by a pair of fine deltidial lamellae. sulcus conspicuous, moderately broad, with steep flanks, and weakly rounded to flat at the bottom. exterior of dorsal valve interarea lower than ventral one and anacline to orthocline. notothyrium open. fold strongly elevated, conspicuous, with steep flanks, and weakly rounded to flat on top. macro-ornamentation flanks, sulcus, and fold are covered by fine plications that are rounded in cross section and separated by narrower furrows. plications in the sulcus and on fold bifurcate close to the anterior margin. eight medial plications present in the sulcus, 7 on the fold before bifurcation on the external mould, increasing through bifurcation towards the anterior margin to 12 plications in the sulcus and 11 on the fold; up to 26 flank plications on the largest paratype; on the external dorsal mould are 2 plications less. fig. 5 latex cast of external ventral mould showing the microornamentation of cyrto spirifer tindoufensis, smf 66083; a: dorsal view. b: oblique posterior view. c: plan view of latex cast. scale bar 10 mm. 8 schemm-gregory m. micro-ornamentation micro-ornamentation fimbriate, with a single row of pustules at the edge of each growth lamellae (fig. 5). interior of ventral valve fillings of the umbo and the lateral apical cavities extend posterior over the hinge line. lateral cavities open. a distinct delthyrial plate is developed that is separated by a profound indentation from the ventral muscle field (figs. 3a, b; 6.3, 4a-d). ventral muscle field longer than wide, pyriform to subrhombic in outline and not, in some specimens weakly, embedded into shell material (figs. 3b; 6.1a, 3, 4a, c). a small ventral process leaves a short groove on the internal fig. 6 1-9 cyrtospirifer tindoufensis sp. nov. all figures x 1.0, if no other indication. stratum: upper givetian (upper middle devonian). leg. u. jansen 2002. 1 smf 66081c, holotype. locality: 12. ventral view of internal mould (a). latex cast of ventral internal mould, upper (b) and oblique posterior (c) views of ventral interior. 2 smf 66081b. locality: 12. dorsal internal mould, dorsal (a) and oblique posterior (b) views. latex cast of dorsal internal mould, upper view of dorsal interior (c) and cardinalia (d). scale bar 10 mm. 3 smf 66594. locality: 16. ventral internal mould, oblique posterolateral view. 4 smf 66081a. collecting locality: loc. 12. ventral internal mould, ventral (a) and oblique posterolateral (c) views. latex cast of ventral internal mould, upper (b) and oblique posterolateral (d) views of ventral interior. 5 smf 66595a. locality: 15. upper view of ventral external mould (a) and latex cast (b). 6 smf 66596a. locality: 16. upper (a) and posterior (b) views on dorsal internal mould. 7 smf 66597. locality: 16. upper views on dorsal external mould (a) and latex cast (b). 8 smf 66598. locality: 16. upper views of ventral external mould (a) and latex cast (b). 9 smf 66081. slab with holotype. scale bar 10 mm. a new species of cyrtospirifer (brachiopoda) from the middle devonian of the western sahara 9 mould. the ventral process gives rise to a low and thin myophragm, extending anterior of the muscle field expressed as a fine furrow on the internal mould. flanking the myophragm are the thin and elongated adductor scars that are weakly embedded into the shell. the diductor scars enclose the adductors and are preserved as radial striae in the centre of the muscle field and as longitudinal striae at the anterior margin of the muscle field (fig. 6.1a, b). anterior margin of the muscle field inconspicuous, except in rare cases when a fine muscle bounding ridge leaves a very thin furrow on the internal mould. dental plates extrasinal or positioned at sinal border, short to moderately long and thin, in adult specimens sometimes wedge-like. they enclose or bound the first half to two thirds of the lateral border of the ventral muscle field. external macro-ornamentation weakly impressed on the internal mould anterior of the ventral muscle field. bifurcation of plications in sulcus or in fold normally not visible on floor of valve. internal expression of plications diminishes towards the lateral margins. weak impressions of growth lamellae at the anterior margin. interior of dorsal valve dorsal umbo extends slightly posterior of the hinge line. ctenophoridium pointed at posterior widens anteriorly, situated perpendicular to the commissural plane beneath the dorsal muscle field, up to 24 lamellae are visible, the number of lamellae varies with the state of preservation. notothyrial shelf usually lacking, sometimes indicated. ctenophoridium bordered laterally by two very thin furrows expressed as a short ridge on either side on the internal mould (figs. 4; 6.2a). dental sockets short, cone-shaped, rounded in cross section, and pointing in apical direction. brachiophores relatively thick and curved over the dental sockets (fig. 6.2b-d). crural bases (sensu ma & day 2003: 273, referring to rudimentary crural plate and crural base) are preserved as small indentions above the ctenophoridium. dorsal muscle field elongate, lentiform in outline, and indented at its posterior margin by a small dorsal median process, giving rise to a thin myophragm bisecting the adductor field. adductor scars extend laterally and are slightly impressed on the shell floor of the flanks. the posterior half is bordered by a low ridge expressed as a small furrow on the internal mould that diminishes anteriorly (fig. 6.2a, b). lateral of the muscle field a few widely spaced gonoglyphs are preserved as lowelongate bulges on the internal mould oriented almost parallel to hinge line. fold moderately broad, strongly elevated, and rounded in cross section with steep flanks and flattened on top. impressions of plications are preserved anterior of the dorsal muscle field. medial plications of fold finer than flank plications. bifurcating plications on fold not visible on the internal mould. impressions of growth lamellae at the anterior margin. discussion. cyrtospirifer tindoufensis differs from other givetian cyrtospirifer species by its shell outline, curvature of its interarea, and number of plications in the sulcus. table 1 shows the morphologic features of c. tindoufensis and other closely related givetian cyrtospiriferid species. cyrtospirifer tindoufensis has the lowest number sinal plications (ranging from 8 to 12), whereas c. verneuili (murchison, 1840), (figured by brice 1988: pl. 44, figs. 1-10; and ma & day 2003: figs. 6.1-14) has 14 to 20. cyrtospirifer verneuiliformis (paeckelmann, 1942, pl. 2, fig. 3) is similar to the latter species with 15 to 20 sinal plications and c. syringothyriformis (packelmann, 1942) with 20 to 30 as illustrated in brice (1988: pl. 44, figs. 11-13). shell profiles of c. tindoufensis are equibiconvex to ventribiconvex, whereas most shells of c. verneuili and c. verneuiliformis are ventribiconvex in longitudinal section. cyrtospirifer syringothyriformis is the only species with a dorsibiconvex shell profile in the middle-late givetian. the new species and c. syringothyriformis both have intrasinal dental plates, in contrast to the other two species that have extrasinal dental plates. cyrtospirifer tindoufensis and c. verneuiliformis are both characterised by convex cardinal extremities and a moderately high, catacline to apsacline, and concave ventral interareas. the ventral interareas of both c. verneuiliformis and c. syringothyriformis are high and catacline, and rarely concave. only c. syringothyriformis shows concave cardinal extremities. the fold and sulcus of c. syringothyriformis and c. verneuiliformis are rounded in cross section, whereas in c. verneuili and c. tindoufensis they are flattened and rarely rounded. cyrtospirifer verneuiliformis is the only taxon with bifurcating flank plications which are also the finest of the four species compared here. in the other three species, shell plications are only observed to bifurcate in the sulcus and on the fold. cyrtospirifer tindoufensis has the coarsest plications. cyrtospirifer syringothyriformis is the only species lacking crural plates, in the others, crural plates are distinctly developed. cyrtospirifer verneuili and c. syringothyriformis both range into the frasnian, whereas c. tindoufensis and c. verneuiliformis became extinct in the givetian. stratigraphic interpretation the new faunas collected from the western sahara range in age from middle givetian in the se to early frasnian in the nw (schemm-gregory & jansen 2005). correlations of the brachiopod fauna found with the blacourt formation in ferques (boulonnais, northern france), the portilla formation (cantabrian mountains, northern spain), and the hamilton group (eastern usa) show that the c. tindoufensis-yielding 10 schemm-gregory m. beds have a middle to late givetian age but may reach to the lowermost frasnian (unpublished data). low diversity coquina assemblages with cyrtospirifer tindoufensis occur in either sandstones or limestones consisting almost entirely of shells of c. tindoufensis (fig. 7). other species found associated with cyrtospirifer in the coquina shell beds include: meristina sp., arcuaminetes cf. scitulites sensu racheboeuf (1981), xystostrophia cf. umbraculum (von schlotheim, 1820), ?adolfia sp., schizophoria sp., and desquamatia sp. the occurrence of arcuaminetes cf. scitulites and xystostrophia cf. umbraculum suggest a middle givetian age. other known deposits with arcuaminetes cf. scitulites sensu racheboeuf (1981) include the “calcareous limestones” in the cantabrian mountains (spain) and the kerbelec and the lanvoy formations in the armorican massif (france) which are correlated with the varcus conodont zone (middle givetian) (racheboeuf 1981). this form also resembles to arcuaminetes scitulites (cooper, 1945) from the north american hamilton group, however, this form is not suitable for refined correlation because of its long stratigraphic range. in north america it is reported from the late eifelian of the iowa and michigan basins by day & koch (1994) and koch & day (1996) and the entire givetian in the appalachian basin of eastern north america (bizzarro 1995). the genera xystostrophia havlíček, 1965 and meristina hall, 1867 are restricted to the givetian, but are not reported from the frasnian. cyrtospirifer tindoufensis occurs together with nalivkinaria issoumourensis drot, 1971 which is assigned to the upper givetian (p. sartenaer, pers. com. 2004). in the northwest part of the study area c. tindoufensis was found close to beds of early frasnian age defined by the occurrence of douvillina dutertrei (murchison, 1840), however, the exact stratigraphic position of the c. tindoufensis in comparison to d. dutertrei is not clear so that the ongoing of c. tindoufensis into the early frasnian remains speculative. early phylogeny of cyrtospirifer the new species contributes to the knowledge of the stratigraphic range and palaeogeographic occurrence of one of the oldest known (middle devonian) species of cyrtospirifer to the early part of the phylogenetic tree of cyrtospirifer proposed by ma & day (2003: fig. 2), however, the authors of that paper recommend a careful restudy of the eurasian species attributed to cyrtospirifer in the older literature. cyrtospirifer tindoufensis new species displays primitive characters including its medium-sized equibiconvex shell, the small number of lateral and medial plicafig. 7 smf 66599. rock slab of cyrtospirifer tindoufensis coquina. scale bar=50 mm. a new species of cyrtospirifer (brachiopoda) from the middle devonian of the western sahara 11 tions, the low apsacline ventral interarea, and flattened ventral sulcus and dorsal fold. later in the phylogeny, shells of cyrtospirifer become larger and the maximum width shifts to anterior to a megathyrid state. the shell profile tends to be either ventribiconvex or dorsibiconvex, and fold and sulcus are rounded in lateral cross section. another trend is the increase in height of the ventral interarea and in the variation in its degree of curvature from apsacline and curved to catacline and straight. the majority of frasnian cyrtospirifer species known in eurasia and western north america are alate, whereas the european and north african forms are still compact but record a trend towards enlargement of mucronate cardinal extremities. discussion of the origins of cyrtospirifer from an older middle devonian species of tenticospirifer tien, 1938 (as revised by ma & day, 2000) requires the study of larger material of both genera but this is far beyond the scope of this work. givetian-frasnian palaeogeographic distribution of cyrtospirifer the occurrence of c. tindoufensis suggests that the initial endemic centre from which cyrtospirifer dispersed from may have been the middle givetian shelf area of northern gondwana (northwest africa) instead of western europe (france, germany) as proposed by ma & day (2003). because of their close palaeogeographic proximity, dispersal of cyrtospirifer from northern gondwana to western europe could have been easily accomplished (jansen 2001; schemmgregory 2008a, 2008b), supported by widespread occurrences of frasnian species of cyrtospirifer in france and other areas of western eurasia. descendants of c. tindoufensis such as c. verneuili, and c. syringothyriformis became widespread in the early frasnian in eurasia and by the middle frasnian in western north america (fig. 8). the known distribution of the oldest north american species of cyrtospirifer indicates that the initial migration of members of two different species groups (see ma and day 2003) into western (northwest territories, canada) and eastern north american (appalachian foreland basin, new york) shelf areas took place during the initial major marine transgression of the middle frasnian transgressive-regressive (t-r) cycle iic of johnson et al. (1985) during the punctata zone. a second major north american migration into continental margin and epeiric carbonate platform habitats occurred during the late frasnian marine sea level rise of devonian t-r cycle iid (montagne noire zone 11 of klapper, 1989). the late frasnian migration from western canada into most north american carbonate platforms is evidenced by occurrence of widespread species of cyrtospirifer in the great basin (idaho, nevada, utah), the southern laurussian ouchita continental margin (new mexico), and epeiric carbonate platforms (arizona, iowa). the faunal exchange between south china and europe/north africa began during the late emsian (late early devonian) as it is seen in delthyridoid spiriferids (schemm-gregory 2009). evidence of later migrations includes the common occurrence of the terebratulid genus paracrothyris in south china and the western sahara indicates a faunal connection between these two regions during givetian time (schemm-gregory & jansen 2008). acknowledgements. special thanks go to gerhard plodowski for initiating this project, organising the expedition to the western sahara, and for fruitful discussions of the cyrtospiriferids. i am grateful to ulrich jansen for giving me access to the brachiopod collection. many thanks are due to juliane eberhardt and erika schellerwagner for technical help, especially for preparing the latex casts. peter königshof, eberhard schindler, and achim wehrmann (all forschungsinstitut und naturmuseum senckenberg, frankfurt am main, germany) gave information on the studied reefs and their facies. i also thank mike reich for access to the brachiopod collection and frank langenstrassen (both geowissenschaftliches zentrum der universität göttingen, museum, germany) for discussing middle devonian brachiopods from ferques, (boulonnais, france). jed day (illinois state university, normal, usa) kindly improved the english of the manuscript and gave fruitful critics to an earlier version of the manuscript. fig. 8 palaeomaps showing the distribution of cyrtospirifer plotted. a: palaeogeography during the givetian (middle devonian). b: palaeogeography during the frasnian (late devonian). 12 schemm-gregory m. lucia angiolini (università degli studi di milano, italy) provided the italian translation of the abstract. the present study is part of the german-moroccan co-operation programme “genesis of devonian reefs” conducted by the following institutions: senckenberg forschungsinstitut und naturmusem, frankfurt am main (germany), ministère de l’energie et des mines, rabat (morocco), institut scientifique, département de géologie, rabat-agdal (morocco), and centre régional de la géologie de laayoune (morocco). the expedition to the western sahara (2002) was funded by the paul ungerer foundation, senckenberg forschungsinstitut und naturmuseum. r e f e r e n c e s bizzarro m. 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(1942) beiträge zur kenntnis devonischer spiriferen. abh. reichs. bodenforsch., 197: 1-188. quenstedt f.a. (1871) die brachiopoden. petrefactenkunde deutschlands, 2: 1-748. racheboeuf p.r. (1981) chonetacés (brachiopodes) siluriens et dévoniens du sud-ouest de l’europe (systématique phylogénie biostratigraphie paléobiogéographie). mém. soc. géol. minér. bretagne, 27: 1-294. schemm-gregory m. (2008a) a new species of filispirifer (brachiopoda: delthyridoidea) from the dra valley, morocco (lower devonian). zootaxa, 1739: 53-68. schemm-gregory m. (2008b) a new terebratulid brachiopod species from the siegenian (middle lower devonian) of the dra valley, morocco. palaeontology, 51: 793-806. schemm-gregory m. (2009) frequentispirifer, a new spiriferid genus and its phylogenetic position within the delthyridoidea (brachiopoda, lower devonian). neues jahrb. geol. paläontol.-abh., 251: 53-70. schemm-gregory m. & jansen u. (2005) middle and upper devonian brachiopods from the western sahara (morocco). fifth international brachiopod congress: abstracts, 27, copenhagen. schemm-gregory m. & jansen u. (2008) first report of the stringocephalid genus paracrothyris (brachiopoda, middle devonian) from north africa. bull. geosci., 83: 169-173. schlotheim e.f. von (1820) die petrefactenkunde auf ihrem jetzigen standpunkte durch die beschreibung seiner sammlung versteinerter und fossiler überreste des thier und pflanzenreiches der vorwelt erläutert. v. of 387 pp. beckersche buchhandlung, gotha. scholz j., ernst a., batson p. & königshof p. (2005) bryozoenriffe. denisia, neue serie, 16: 247-262. termier h. & termier g. (1949) essai sur l’évolution des spiriféridés. notes mém. serv. géol. (rabat), 74 : 85112. tien c.c. (1938) devonian brachiopoda of hunan. palaeontol. sinica, new series b, 4: 1-192. waagen w.h. (1883) salt range fossils, part 4 (2), brachiopoda. palaeontologia indica. memoir series 13, 1(2): 391-546. wang yü, yu changming & wu qi (1974) advances in the devonian biostratigraphy of south china. mem. nanking inst. geol. palaeontol., acad. sinica, 6: 1-71. [in chinese.] ziegler w. & sandberg c.a. (1990) the late devonian standard conodont-zonation. courier forschungsinstitut senckenberg, 121: 1-115. rivista itaiìena di paleontologia e stratigrafia brackish marsh benthic microfauna and paleoenvironmental changes during the last 6.000 years on the coastal plain of marathon (se greece) maria v triantaphyllou" kosmas pavlopoulos" theodora tsourou' & michael d. dermitzakis' recei'-ed apríl 23, 2aa2; acceptecl januatl 22, 2003 | 1pr l;;,' key uords: benthic foraminifers, ostracodes, microfaunal biotopes, marsh paleoenvironments, holocene, marathon plain. abs*act. the present 5ssi1., based mainly on rhe analysis of toraminifers and ostracodes, provides evidence of paleoenvironmental changes on the coastal plain of marathon (e. greece) during rhe last 6.000 vrs. three sedirnentary units lagoonal formatìons were recognized and identified as a, b and c. they range in time bers.een before 55oobp 3500bq 3500bp-25008p and 25oobprecenr, respectiveiv the study of the brackìsh marsh microfauna of the marathon plain holocene sediments reveals the presence, durine the last 55oo )'rs, of three distinct biofacies in the sedin-rentary units alre:dy e:tablished. alternating mesohaline oligohaline (mo), oligohaline fresh water (ofv) and mesohaljne oligohaline to olieohaline fresh water (mo-of\í) biofacies in the framework of the sedimentary units indicate a general trend iandnard along the plain suggesting a slowìng of sea-level rise probabll. correlated with a relevant tectonic uplift. one prominent feature of this study is the clarification of the ecological preference of the species tricholrya/us aguayoi (bermudez, 1935), which is dominant in oligohaline conditions under an influence of fresh $'ater input (salinity less than 15 %,). riassunto. questo studio è basato soprattutto sull'analisi dj fcraminiferì ed ostracodi e consente di valutare le variazioni ambienrali durante gli ultimi 6.000 anni-nella pianura costiera di maratona (grecìa). sono state distinte tre unità sedimentarie di ambiente lagunare, definite a, b, e c, che si dìstribuiscono ln tempo tra 55008p 350088 3500bp-2500bp and 250obp-arruale, risperrir amente. i-o studio della microfauna della palude salmastra nei sedimenti olocenìci di maratona rivela la presenza di tre biofacies alf interno delle tre unità sedimentarie precedetentemente stabilite. iialternanza di biofacies mesohaline oligohaline (mo), oligohaline acqua dolce (ofv) e da mesohaline oligohaline a oligohaline acqua dolce (mo-ofv), nel contesto delle tre unità sedìmentarie, indica la tendenza verso i'incremento dell'area emersa della piana, ottenuta mediante un rallentamento dell'innalzamento del livello del nr:re colhgaru con urr signi[icarivo soller.amento strutturale. ljn aspetto irnportante di questo studio è i'indiv:iduazione delle preferenze ecologìche del foraminifero tricbohyalus aguayoi (bermudez, 1935), che domina nelle condizioni oligohaline sotto l'influenza di apporti di acqua dolce (salinìtà inferìore al 15 %").trio topographic map showing the location oi trenches (1,4,10,1 1) and boreholes (6,2). 1 universitv of athens, dept. of geologv, section of historical geologv-paleontology, panepistimiopolis 15784, athens, greece e-mail: mtriant(qì geol.uoa. gr 2 harokopio universitl., faculty of geographl., z0 el.venizelou str., 1761, athens, greece. 510 m.v triantaphyllou, k. par-,lopoulos, t. tsourou e. m. d. dermitzakis stratigraphic correlation of the boreholes and trenches studied, location of the samples analyzed and position of sedimentary units and biofacies (mo:rresohaline-oligohaline, ofw:oligohaline-fresh water, mo-o\wf:mesohaline-oligohaline to oligohalinefresh water). legend: 1. artificial fill, 2. topsoil, 3. sand, 4. sand with gravels and pebbles, 5. silt, 6. clay, 7. silty sand, 8. sandy silt, 9. clayey silt, 1 0. sandy clay, 1 i . siltv clay, 12. sandy clay with sub-rounded gravels, 13. palustrine mud, 14. lagoonal mud,1 5. peloidal algal mud, 1 6. mudcracks, 12. rootlets, 18. charophytes, 19. pellets,20. gastropods, bivalves,21. bioturbation,22.pert,23. algal pear,24. hard horizon. peat ages are calìbrated (in yr. b.p). trench i 2.o ?.p c) e -c .c .a c) i |,0 trench 4 trench i0 borehole ó borehole 7 t8-7 t8-ó t8-5 t8-l mo tg-2 tl 0t1 il 0-4 tii-i borren borren unit c 24008p-preseni unii b 3500-2400 bp unit a before 5500-3500 bp 4..ró_p.l.l tó.pia' t6-p2a tvo tvo tó-fossl t7-5 tegend ffifl w e!.e!@ fri:íl t:111"í.:....'] r-----l ]_ t:iii:ii:il fnnnl f:: i:::i:: ì r flq'' e ? l-----1rn ? f ::-:t i l ffia-l-.:-:-l^ v, f.__ltr, , i--:-t-|. o, f-.:-r-rl t4, a fro eeee ffi i ffi lîl:lt1 17. 18, 19, 20. 21 . 22, 23, fic -\bsolùte sample altitutc (m) 0.25 -1.33 -2.00 -2.33 trn.jch ] tl-l 1.30 borei]ole 6 t6-pìb -1.29 t6-pia -r..1 2.12 t6-p2a -1.53 2.62 t6-foss1 2.20 3.20 borihole 7 t7-9 t7-8 t7-l trencii .1 t8-7 t8-ó t8-5 t8-3 t8-4 t8-1 rg-z trench ii) tl0-l t10-3 t10-4 tl0-5 trench i 1 sih) p.al contaìnrng teuestrial gastfopods, charoph)tcs and tòramìn'1èral tìagments. pcllclcd mud containing nmlluscs. :.,rj) .1.) ..'r'r' ' ': .l'.'r.plir:.. dcpth (m, sedimcnî description , , n : ' rj1 cljr , rr :rrnin: j(11 rrrr fiegnlenls and gàstropods. palc j'.cllow (-5y v8i c2) palusbine nnd contaming charophytes olivehfo$n(2.5y q!4g!!qsp4!v,1rc2) color units (\4unsell\crlct broracres j,t;i',""t:j:ì, dark grai l2.5 y, i ) bafren ""*llìlì1"' of\\ gfa) (5y vsicì) l\lo ofw gastropods eild bi\ilvcs l2l pallrstdne mud includìng yeìlon'ish reducllon spols. 2-32 pelleted algal nud i r'. 1r' rjj 'ntair'r u .hr.op.1 t'.1 uu lrrrnpoiì' rn.l brrr rtr lcì cr.r .rlr.rì r'..j . jlorì1.! -1j f lnsroplrvt(s ùlrri gj\llùf'ì'll\ olrve (5y vsrcl) mo b 1: . tr'' ui, tut r ,la_,,t s a pale oìj\.e (5y v6icl2) mc) a ìightgrar(l0yr\r7,tl) ofw a l.l< 0._5 \-.ll\ nrd ml-.r n, ,\.,ror,t)r(. .rn.t t;, ;-.,.15\ r: r ., g3r ron^o( q,95 i1.95 silty sand. gray (sy v5.c i ) 0 5) l 1.5 sllty sand. light gra) {5y v7ic2) _0.62 2.55 palustrine nud irulirdng rcduction lrlhrbrotrf (-.5yr llots. \i6ic.1) _t ,i i ti pr rsrint nrr.l including rcducrion lìghlbfown(7.5yr.fo,.. \ " a.i) -t ro ì ln l.rgl',r.r rrud containìng charoph)les. gfa-vish bro*n (2.5 y 8/'.rolod..nd b \î 'c. i: a: -2.02 3.92 l-agoonal mrid conreining gastropods. light gray (5y \r7lc l) ,,rrt ofw of'w \,jo n40 n4() lvlo c c c b b c' 0 7q 0.70 clayel .silr. 0.50 0.95 clltrllr fllr: 0.20 l20 sand), silt. -l00 1.30 lagoonal md. ìight grav (5\' v7[2) brrren grar (5y \5,c1) !q].-l llght gray 15y v7rc2) ofw c gfîvish bror,n (2.5 t tnto : \j5rc2) introduction the elongated, ne-s\í oriented, marathon plain is located in east attica, e. greece (fig.t). the broader area surrounding the marathon plain consists of the "ne attica" geotectonic unir, represenring a "relatively autochthonous" metamorphic sequence (lozios 1993). the quaternary is represenred by various rypes of alluvial deposits formed mainly b1' the inois river, ivhich divides the plain into two parrs. pleistocene and flolocene talus cones and screes cover steep valley sides as well as fault zones. the marsh of marathon is extended at the eàstern side of the plain and separated from the sea by a barrier beach with low sand dunes, morphologically resembling the typical coastal plains of greece according to kraft (kraft et al. 1975,1977). baeteman (1985) conducted a systemaric drill hole studv of the area and presented detailed informarion on the stratigraphic sequence of the plain. pavlopoulos et al. (in press) determined the sequence of depositional environments and the climate and sea let'el chanees recorded in the area since about 6000 years bii using rnicron.rorphological and micropaleontological methods in addition to radiocarbon dating of seyeral horizons, including peat lalrers. they recognised three sedimentary units, nan-ìed a, b and c respectively (fig. 2), on the basis of the features of each sedimentary phase. sedimentary unit a (before 55oo to 3500 bp) is generally composed of fossiliferous bioturpaleoent,ironmental changes on the coastal plain 541 tab. 1 descriptìon of tbe samples studicd and their relation with sedimentary units and biofacies. bated lagoonal mud with occasional peloidal charophytic mud, algal peats and macrophytic peats. unir b (35002500 bp) consists of mixed carbonare and siliciclastic palustrine mud and the uppermost sedimenrary unit c (2500 bp-present) represents mosrly fluvial deposits. the marathon coastal plain is famous for the ancient battle of 490 bc, fought between the athenians and the persians. this area ràras proclaimed a national park by the greek government after athens won rhe comperirion for the organization of the 2oo4 olympic games. fiowever, a ros/ing center has been under construction since early 2001. given the great environmenral and archaeological importance of the area, great inrerest developed for understanding the palaeoenvironmental conditions of this plain in the past. this paper presents a detailed micropaleontological study of the subsoil of marathon coasral plain. taking into account the benthic foraminiferal and ostracode assemblages (together with sedimentological data; pavlopoulos et al. in press) the analysis arremprs ro determine the paleoenvironmental changes during the last 6.000 yrs on the coastal plain of marathon, based on the study of the brackish marsh benthic microfauna. materials and methods in order to obtain information about the holocene stratigraphy under the recent alluvial cover rwo boret6 foss-l t6 pl b t6 p2 a î7-1 '7 ostracodes cyprldels ?òfosa (jonet 97% cyprinotus salínus (baaov) cyprinotus sp. 0.50% i ^w^î^^.h) ò11;^1;.. 21, i.av. atv" 82% 6% 89.7% 74.5% 85% b./" 8.3% 17.4ó/ó 12% 2.9% 2.7% 12% aaa/. 3.6% 1.6% 3% 69.1% 1.8%298% 9% 5.7% 2.5% cpn!ptqps9g!.9;;; . i:ll"t) l other candoninae 12% 2.5ok 111.k 2o.b% 6% 10 .7'/. q 3%-, o 5x '16.9% 3.40/a 1% tzóà 0.5% 24.8./" 14.5% 13.8% 6.5% limnocythere ínopínata (berno) limnocythere sp. 0.6% 1a/. 16% 0.5% 1.44/. a7% 3 b% 6.8% 44l 115% | glbba t2% 55.20/. 724/.(reuoorc) llyocypri:trp] daruinula stevensani (bnnov & roaenrsor) herpgtggyp!$pbenthic foraminifers ammonia beccaii (ltnne) haynesína germanica haynesina depressula (walker & jacob) trichohyalus aguayoi (bern.4udez) elphidium granosum 0.5% 0.5% 6% 0.9% 9a.4% 96.4% 3.670 1 00% 28.3% 2% 93.5%95% 3.6% 1.5% 4yó 35.2% 6.5% 1.à 36.5% :\+ / m.v. triantaphyllou, k. paalopoulos, t. tsourou u m. d. dermitzakis tab. 2 relative percentage abundances of species in at least 3oo counts (ostracodes) / 2oo counts (benthic foraminifers), in the samples studied were treated with h,o, to remove the organic matter, and then washed through 63, l25,250, 500 pr.m sieves and dried in an oven at 50c c. qualitative analysis was performed on all samples, and seventeen samples presented a foraminiferal and ostracode abundance which was sufficient for the quantitative analysis. a subset of each sample (fraction of 1 2 5 p,m i b enthic f oraminif e rs, 2 5 opcm/os rracodes ) was obtained using an otto microsplitter untiì aliquots of at least 2oo benthic foidminifers and 300 osrracodes, respectively, remained. a scanning electron microscope analysis (sem jeol jsm 5600) was used to assist in the identifications. the taxonomy of benthic foraminifers in this paper is based upon loeblich & tappan (1988, 1994) and bronnimann et al. (1992). taxonomic references of the ostracodes include mainly athersuch et al. (1989) and additionally mazzrni et al (1999), sun er al. (1999). the relative abundance of the species was treated in a q-mode hierarchical analysis, performed to study similarities between samples, which is based on euclidian metric distance and was carried out using statgraphics plus 4.0 statistical software. fig. 3 distrìburronef p:rrern of benrhìc forrminifcr. \r, and o.lrrcodes (b) in borehole /. holes were driiled -with a portable drilling setand 4 trenches were excavated (fig. 1). the description of the sediments in the sequences and their stratigraphic correlation are presented in table 1 and fig.2 . twenty-one samples of about 50 g each were collected from all the boreholes and trenches (fig. 2). they benthic foraminifers and ostracode distribution and microfaunal biotopes the species diversity of benthic foraminifers is very low. only six species were actually identified in seventeen samples. concerning the ostracode fauna, twelve taxa were identified and counted (table 2). ammonia beccarii generally follows haynesina spp. frequency pattern (figs. 3a, 4a,5a, 6a) except for sample t8-3 where it shows an opposite trend, towards haynesina germanica (fig. aa). additionally, a. beccarli shows a negative trend towards trichobyalus aguayol in sample t6-p2a (unit a, borehole 6) . concerning the relative frequency patterns of ostracode species, a *"'s .r.c *."u,".a"j ]ilirli ll-l r'"è' _.3 t ui l^l l_.1 .9^ .d """d -."e.nt* ò\ bg sdmples î *i l::i f] h tl i'l ! h ".pljnils sqmples (j "d os s' -uu .''l.o'";s' lll lf lrt rrl ll' i i ' i 'r, ,ro . .r, i ri---: ò--,0 do o 5 fig. 4 dirtributional patlern ol bcnrhic ior:minr[er. 1:; :nd o,rracodes (b) in trench 4. negative trend emerges in all the units between cyprideis torosa and cyprinotws salinus, darwinula ste-,tensonì, limnocythere sp., cand,ona neglecta and llyocypns spp. (figs. 3b, 4b, 5b, 6b). the frequency data indicate that several changes in the composition of the benthic microfauna took place in time and space on the coastal plain of marathon. these changes were confirmed when qmode cluster analysis was applied to the data set to define areas of similar environmental conditions. in the resulting dendrograms (fig. z) the clusters are regarded as biotopes and are interpreted as representing different ecological conditions. these biotopes together with the environmenral interpretation are: cluster i. tricbobyalws agwayoi, candona neglecta, other candoninae, limnocythere inopinata, danainwla s t ev en s o ni, i i iy o cy p r i s bra dy i, i ly o cyp ri s gìbóa as s embla ge. shallow oligohaline fresh water biofacies paleoenr:irctnmental changes on the coastal plain 543 (ofw). the first cluster considered (fig.z) is represented by samples t11-1, t10-4, t8-5, t8-6, t8-z (belonging to unit c); t6p1b (belonging to unit b); and t7-5,t6p2a (belonging to unit a). they are characterizedby the total absence of benthic foraminifers (except for the remarkable monospecific presence ol t. aguayoi, as a brakish water species (bronnimann er ai. nez\, ìn sample t6p2a). c. neglecta, d. stez,ensonì, l, inopinata, i. bradyi, i. gibba are indicators of shallow freshwater and oligohaline envìronments (athersuch 1,979; grafenstein et al. 2o0o; mazzini et a\.1999; athersuch 1979;sokac 1978). this assemblage consisting of dominant freshwater ostracode species, associated with brackish water species (c. torosa, c. salinws) indicates the persistence of a restricted temporary communication vrith the sea (c1avé et al. 2001). it suggests a salinity of less than l5"/oo (neale 1988) and fits well vrith high-middle marsh environmenrs (scott et al. 1979), indicating an approximate elevation of 20 cm above the mean paleo-sea level. cluster ii. ammonia beccarii, haynesina germanica, haynesina depressula, cyprideis torosa, loxoconcha elliptica, cyprinotus salinus assemblage. shallow mesohaline-oligohaline biofacies (mo). the second cluster considered (fig.z) is represented by samples t8-1, t8-2, t7-1, t6foss1, t10-5 (belonging to unit a) and t8-4, t8-3 andtt-9 (belonging to unit b), which are characterized by the high percentages (rising up to 100%) of benthic foraminifers a. beccarii or h. germanica and h. depressula. a. beccarii -an euryhaline species living in a wide range of different environments (murray 1991; alve 1995) is characteristic of the part of the lagoon which is more affecred by marine warers. whereas haynesina is found in the internal portions of the lagoon affected by anthropogenic events (serandrei barbero et al.1997). both species characterize the lower intertidal environments (cundy et a1. 2000).the ostracode fauna is marked by the dominance oî c. torosa, rising up to 97o/o, l. elliptica and c. salinws, which are typical brackish species (gliozzr &'mazzint 1.998;mazzrni et ai. 1999). this assemblage suggests a salinity of more than 15 %' (neale 1988) and fits well with low marsh environments (scott 1977;scort et ai.1979; petrucci et al. 1983), indicating approximately the mean paleo-sea level. cluster iii. cyprideis torosa, cyprinotus salinus, candona neglecta, limnocythere sp., dalwinula steztenson i assemblage. shallow mesohaline-oligohaline to oligohalinefresh-water biofacies (mo-of\q. as shown in fig. 7, sample tz-8 (belonging to the upper part of unit a), which is characterized by the absence of benthic foraminifers and the balanced presence of brackish shallow-water and shallow freshwater ostracode species, suggests an interrnediate mesohaline-oligohaline to oligohaline-fresh-water lagoonal environment. 544 m.v. triantaphyllou, k. pavlopoulos, t. tsourou g. m. d. dermitzakis -$ "ts od òoii \t\t .-.d * .r.-uu ""s ...s .,"-*.o.. ;'s i i ehll i'itil fis. 5 distributional pattern of benthic forarninifcrs (a) and ostrecodes (b) in borehole 6. paleoenvironmental interpretation and site evolution the biofacies documenred appear in all trenches and boreholes ttable l. fig. 2 t. and therefore they characterize all the three different units. the mo biofacies (a. beccarii, haynesina spp., c. rorosa, l. elliptica, c. salinus assemblage) of unit a (5500-3500 bp) is present in trenches 10, 4 (located in the southern parr of the marathon plain) and in the lower parts of boreholes z and 6, indicating a brackish water marsh. the documentatron of of\l and mo-ofv biofacies in boreholes 7 and 6, at levels where macrophytic and algal peats and characean oogons intercale rvith palustrine mud, suggesrs a brackish water marsh rvith fresh water inflow, probably associated ro several shallowine cycles detected in the inner parts of the plain. unit b (3500-2500 bp) has similar characteristics, since the of\{/ biofacies is present only in borehole 6. pavlopoulos et al. (in press) suggested frequent exposure of the basin during this time interval, which is definitely confirmed at the northern side of the plain by the terrestrial environment corresponding to substratum 25oobp (documented in sample t1-1 (trench 1) which conrains only verv few -probably not aurocrhonousostracode valves). the biofacies distribution is totally different in unit c (25oo-present). of\f is the only biofacies detected at the southern part of the plain (trenches 11, 10, a) indicating the dominance of fresh warer ro oligohaline marsh environments. as pavlopoulos et al. (in press) sugqesred, the change from palustrine to alluvial environment happened repeatedly, implying a very shallow and frequently exposed plain without any connection to the sea. fig. 6 distributional pattern of bcnthic foraminifers (a) and ostracodes (b) ìn trench 10. hence, towards the landward part of the marathon coastal plain a general trend was documented along the transect a-b (fig.8). as shown in fig.8 during 55oo2500bp time interval, the southern part of the plain is considered to represent a low marsh environment, whereas the rest of the area was a high-middle marsh. on the contrary the paleoenvironmental conditions changed drastically during the last 2500 yrs, as the high-middle marsh conditions were restricted to the sourhern part of the plain, while the northern part towards the landward area was probably exposed frequently. this is probably the result of a small apparent tectonic uplift the area underwent during the 55oo1300bp time interval, albeit ar slower rates (0.4-o.5mm/ q-rlode hierarchjcal analvsis ieading to the distincrìon of three nrain clusters corresponding to different biofacies. :-o[oonoine loil-8 oligoholine íiesh woter -biofocies imo-ofw) t7-r l8-3 t8-4 tó-fos5] tj-g mesoholine oligoholine biofociés i8-2 (tvlo) t8 l tl 0-5 -l-tl a i tw oligoholine' fresh wolef bìofociès(of\tl tó.p] b t6-p24, l8-7 r8-ó 17-5 r8-5 il 0-4 nl-l f rg,. 7 unit c 2500bp-presenl unit b 3500-2500 bp unit a beforè 5500-3500 bp yr; pavlopoulos et al. in press) than the isostatic rate (0.6a.7mm/yr, lambeck 1996), which caused apparenr coasral stability from the classical times onward (kraît 1972). conclusions the sedimentary sequences of the middle-late holocene of the marathon plain are represented generally by lagoonal formations related to a slowing of the sea-level rise. the micropaleontological analysis at the holocene coastal plain of marathon revealed the absence of agglutinated foraminifers and hence the absence of a sait marsh paleoenvironmenr (scorr et al. 1979; cundy et al. 2000). one prominenr featllre of the present study is the clarification of the ecological preference of the species trichohyalus aguayoi (bermudez, 1935), (p1. i, figs 1, 2), paleoenoironmental changes on the coastal plain 515 fi-. i schem.rric inrernrcrrrion oib'" paleoenvironmental conditions of the study area, according to the biofacìes ìdentified in the sedimentary units a, b, c. (t: trench, b: borehole) which is generally regarded as a brackish species (bronnim.lnn cî rl. lgg)r. scor. ^* ^l r roto\ j^^,..-led il underuv!uillllll the name dicorinopsis aguayoi in recenr coasral marshes of \il{ greece. our results clearly indicate that it dominates in oligohaline conditions (salinity of less than 15 "/"'). three biofacies were recognized in sedimentary units (pavlopoulos et'al. in press) of the marathon plain during the last 55ooyrs. the alternation of mo, ofv and mo-of\f biofacies in the framework of the sedimentary units indicates a seneral trend towards the landward area of the plain, suggesting a slowing of sea-level rìse probably correlated with a relevant tectonic uplift. akno'oletlgments. \fle thank dr. m. dimiza for her help in treating the data st:rtistically and taking the sem micrographs. the authors also would like to thank d. besso, a. bossio and m. geetani nhose incisive and constructive comnrents greatly improved the nrrnuscrìpt. references alve e. 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(1999) cenozoic ostracoda and palaeoenvironments of the northeastern tarim basin, western china. palaeogeogr., palaeoclim., palaeoecol., 148 37 -50, amsterdam. plate i fig. 1 t)"ichohyalus aguayoi (bermtdez). umbilical vierv, sample t6p2a. fig.2 hichohyalus aguayoi (bermudez). dorsal vier., sample t6p2a. fig.3-ammoniabeccarii (linne). umbilicalview,samplet6ijossl. fig.4-haynesinadepressula(\falkertrjacob).sideview,sanrple t6foss1. frg 5 chara sp. sample t1o-3. fig. 6 daruinula stevensoni (brady & robertson), san.rple t /-5, left valve, exrernal vien. fìg 7 loxoconcha elliptìca brady, sample t6 foss1, right valve, external view. frg. 8 candona neglecta sars, sample t6 p1b, right valve, external vieu'. ftg.9 cyprinotus salinus (brady), sample t z-5, right valve, external view. fig. 1o ilyocypris brarlyi sars, sample t6 plg, 1.ft valve, external view. fig. 11 ilyocypris gibba (ramdohr), sample t6 p1b, left valve, external view. frg. 12 cypr:ideis torosa (jones), sample t6 foss1, female, left valye, externàl vierr. pttleoert'-ìrotnnental cbanges ctn the coastal plain 517 bona 323..330 ����� ������� � � ����� ������ �� � ��� ��� ��� ������ �������� � ������ � ���������� ���������� ������ ���� �� ��� �� � ���� ��� � � ��������� ��� �� ��������� ����� ����� ���� ��������� �� ��� �� �� � ��������� ����� �� �� ��� ������ ��� �� ���� � � ����� �������� ���� � �� ��� � ��� ��� �� �� �� � �� � ����� � � ���� ! ��������! 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(/,,-* � *�� � ������ � % �� 7 ���� ��� � ������� �� ���� '���� � � � ���� % ()� � � ���� *� �� ���� � ��� ��� � ���� �$ ! ���� 2�� +��� #������ -,.(�*� 1+-�10/! << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /all /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /warning /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /syntheticboldness 1.00 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true 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/pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /fra /enu (use these settings to create pdf documents with higher image resolution for improved printing quality. the pdf documents can be opened with acrobat and reader 5.0 and later.) /jpn /deu /ptb /dan /nld /esp /suo /ita /nor /sve /kor /chs /cht >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [595.000 842.000] >> setpagedevice rivista italiana di paleontologia e stratigrafia volume 106 numero 3 received march 1 3. 2000. key-oords: ostracoda, obliacythereis (paleoblitacythereis), taxonomy, miocene, southern italrv riassunto.il genere obliacythereis e i due sottogeneri inclusi, oblitacythereis e paleoblitacytherels, sono stati descrittì da benson (1977).tuttavìa, la diagnosi originale dì paleoblitacytherezl e la designazione della sua specie tipo si basano su esemplari erroneamente attribuiti da benson (1977) a carinocythereìs rugierii russo, 1966. gli esemplari illustrati da benson (1977) sono, in realtà, chiaramente rjferibili ad un'altra specie segnaiata in letteratura, oblitacythereis sp.3 russo er bossio, 1926, come già affermato da bonaduce & russo (1985). in conseguenza di questa erronea attribuzione il sottogenere paleoblitacythereis risulta soggetto a potenziale instabilità. la sua specie tipo dovrebbe, pertanto, essere considerata come una nuova specie a sé stante e carinocythereis ruggíerii russo (specie che realmente appartiene al sottogenere pa/eoblìacytbereis) dovrebbe essere rivista. lo studio di esemplari ben conservati, provenienti da alcune formazioni mioceniche dell'italia meridionale, ha permesso di reaiizzare un'analìsi dei rappresentanti italiani del sottogenere paleobliucythereis. come risultato di questo studio viene qui proposra una revisione tassonomica delle seguenti speúe: oblitacythereis (paleoblitacytberei) rugierii (russo, 1966) e oblitacytbereís (paleobliucytbereis) bossioi n. sp. (: oblìtaclthereis sp. j russo & bossio, 1976), che viene qui proposta come nuova specìe tipo del sottogenere paleoblitacytherets. ljna forma considerata nuova per la, letterattra, oblitacythereis (paleoblitacythereis) apula n. sp., viene inoltre descritta ed illustrata. per ciascuna specìe sono fornite una sinonimia aggiornata, un attento confronto con le altre specie note, le segnalazioni precedenti e la distribuzione bio-cronostratigralìca nell'ambito delle sezìoni esaminate. vengono, infine, esposte alcune considerazioni sul significato paleoecoìogrco di cìarcun: 'pecie. abstract. the genus oblitacythereis, type species oblitacythereis (oblitacytbereis) mediterranea benson, 1.977, and the subgenera oàlitacytl.,ereis and paleoblitacythereis are well defined by benson (1927), who designated carinocythereis rugierii russo, 1966 as type species of paleoblitacytherels. the specimens figured and described as carinocythereis ruggierii by benson (1922), however, clearly differ from russo's species, and coincide well with oblitacythereis sp. 3 russo & bossio, 1976, as stated by bonaduce & russo (i985). as a consequence of this rnisidentification the subgenus paleob/itacytbereis is subject to uncertainty and potential instability the type species designated by benson should be considered as a ne\il.nominal species and carinocythereis rugierii russo, which actually belongs to paleoblitaqttbereis, should be revised. the study of well-preserved specìmens from some miocene formations in southern italy prompts the author to propose herein a systematic revision of the itaiian representatives of the subgenus paleoblitacytherels. three species are discussed. these are: o blitacytherei (paleoblitacythereis) ruggieril (russo, 19 66), ob lipagine 391-398 dicembre accepted. jwly 20, 2000 tacythereis (paleobìitacytbereis) bossioi n. sp. (oblitacythereis sp. 3 russo & bossio, 1976), here proposed as the new nominal type species of paleoblitacytbereis and, ítnally, oblitacythereis (paleobliacythereis) apwla t sp., described as new. the systematic notes of each species are given with the bio-chronostratigraphical distribution resulting from the present study together with some palaeoecological remarks. introduction. the genus oblitacythereis, type species oblitacythereis (oblitacythereis) mediterranea benson, 1977, was erected by benson (1977), who described its diagnostic features and drew attention to its stratigraphical and palaecological value. according to benson (1976, 1977,1978) oblitacythereis is a typical inhabitant of the llpper bathyal zone with water temperature ranging from 1o"c and 1.6-20"c and esrimared depth range of 300 to 1000 meters. benson (1977) subdivided oblitacythereis into two subgenera (oblitacytberels benson, 1977 and paleoblitacythereii benson, 1,977), based on the structure of the anterior reticular system and the development of the three major longitudinal ridges. the original diagnosis of paleoblitacythereis and the designation of its type species are based on spanish and sicilian specimens, which were incorrectly attributed by benson (1977) to carinoqtthereis rwggierii russo, 1966 (see detailed discussion in the systematic section). thìs misidentification produces uncertainty and potential instability in the common usage of paleoblitacytbereis (iczn art.z0) and the type species designated by benson should be considered as a new nominal species (iczn art. 70, c). since paleoblitaqtthereis is of considerable palaeoecological and stratigraphical significance for the mediterranean miocene, a critical taxonomic revision of its ltalian representatives is herein presented. three species are discussed, of which two are proposed as new. for each species an updated synonymy, detailed comparison with allied species, the bio-chronostratigraphical revision of the ostracode subgenu s pale o b litacyt herei s benson . 1,97 7 barbara dallantonia'i '' dipartimento di scienze della terra, universitl' of pisa, via s. maria 53,i56126, pisa, italy. e-mail: barbarad@dst.unipi.it 392 distribution in the examined sections, and the previous records are g;\-en. their palaeoecological significance is also briefly discussed. material. the present study is based on some miocene sections from the tremiti islands (northern margin of the apulian platform) and the hyblean plateau (southeastern sicily). the material used in this paper was dated by both planktonic foraminifera and calcareous nannoplancton. the planktonic foraminiferal zonal scheme followed herein is that of iaccarino & salvatorini (19s2) and iaccarino (1985), recently emended for the middle miocene by foresi et al. (1998). ten sections from the miocene sedimentary sequence of the tremiti islands, comprising the cretaccio formation (selli, 1971) and the calcari di s. nicola formation (selli, tlzl;, have been examined for the present study. they are those studied and described by hccarino et r1. iin progress), to whom the present atlthor refers for a detailed description of lithology and chrono-biostratigraphy. the investigated sequence ranges from lower langhian (praeorbulina glomerosa s.i. zone) to messinian (non-distinctit,e zone). the sicilian material comes from four miocene sections comprising the irminio member of the ragusa formation (rigo & barbieri. 1959) and the lower part of the tellaro formation (rigo & barbieri, 1959). the examined sections are entirely of langhian age and extend from the praeorbulina glomerosa sicana subzone to the base of the orbulina universa subzone (dall'antonia et al., in progress). in addition, some material previously collected by a. bossio as well as samples from the langhirn strrtotype have also been examined. systematic descriptions. all the illustrated specimens are housed in the ostracoda collection of prof. a. bossio (c.o.b. 15-24), department of earth sciences, university of pisa, italy. the taxonomic classification and the terminology of the extern.rl carapace features followed herein (fig. 1) are those proposed by benson (1977). the author refers to benson (cited op.) for diagnosis and detailed descriptìons of the genus oblìtacythereis and the subgenera o b litacy th ereis and pale ob litacytb ereis. as a consequence of the above mentioned misidentification by benson, all the forms reported but not figured as c)blitacytbereis ruggieril (russo) by benson (1,976,1928), benson et al. (1991) and berggren et al. (1976), are considered of doubtful attribution and are not dealt with in the present paper. fam i ly trachyleberid idae si i vesterbradley i 948 subfamily trachyleberidinae silvester-bradley, 1948 genus oblitacyth ereis benson, 797 7 subgenus paleoblitacythereis benson, 1.977 oblitacythereis (paleoblitacythereis) apula n. sp. (fig. 1a; pl. 1, fìg.3, a,7) 1976 oblttacythereis sp.1 russo & bossio, p.220, pl. 1,ftg.7,7a material. more then 7o valves and 10 carapaces. etymology. f-rom latin apulus/apula : inhabitant of apulia. holotype. a left valve (c.o.b. 15) figured in fig. 1a and in pl. 1, tls. j../. type-level. serravallian (neogloboquadrina continuosa stbzone) of the cretaccìo formatron. type-locality. north-eastern area of s. nicola island (tremiti islands); section 8 (sampìe n. 177 ) ìnlaccarino et aì. (in progress). paratypes.,l valves and 1 carapace (c.o.b. 16-20), of these a right valve (c.o.b. 16) is figured in pl. 1., fig. 4. dìagnosis. a species characterized by strong ornament with a marked coalescence ol the fossae in the posterior region; the muri in the anterior area are poorly developed and a simple transverse rib extends from the position of the absent eye tubercle to the median longitudinal rib, anterior to the position of the frontal scar. description. in lateral view the left valve is sub-rectangular, thick-shelled and strongly ornamented. the regularly rounded anterior margin is denticulete, especially in the anteroventral region. the dorsal margin is straight but overhung by the dorsal rib. the ventral margin has a distinct antero-ventral keel. the ventral and dorsal iines slightly converge backward. the posterior extremity is sub-acuminate, with xpex xt mid height. the posteroventral margin is denticulrre. the three rypical longitudinal ribs are well-defined. in the posterìor region, rhe fossae tend to align horizontally and to join vertically and two subdued, iongitudinal secondary ribs are present; a lower one, which divides the/ossae of no series from the fossae oí the p-v series and an upper one, which separates the fossae of k series from the/ossae of the l-m series. in the anterior area, there are some vesrigial parts of the primirive anterior marginrl ridge. nevertheless, a poorly organised system of nodes and short muri, which might represent an initial stage on the way toward the formation of the interoconcentricwm (benson, i977,p.30) is present (fig. 1). this consists of rhe following srructures: i ) a simple rrrnsverse ridge. which joins pore-conulws scorpio to the blind ocular tubercle passing through pore-conulws capricornws;2) a short oblique and arched rib, separating fossa c1-c2 from fossae b 1 and b2 and connecting p ore-conwlws afa to the former transverse rib; 3) a short rib, jorningporeconulus alpba and pore-conulws beta and reaching the median rib. internai features typical of the genus. b. dall'antonia t h e o strac o d. pale ob litacy th er eis 393 fig. 1 reticular silhouettes and rìdge patterns oí (a) oblita cytb er e i s (.pale o b litd cyth er eis) apula n. sp.; (b) oblita cyth ere is (pal e o b lìtacyth ere is) bossioi n. sp.; (c) oblitaryrhcrc i s 1pa i cobl ira cy t here ì s 1 ruggierii (russo). the fossal patterns and the positions o{ roms of rhc nr.rin por.conuli are indicated accordìng to benson (1972, 1977). strong ornament and in the tendency of the fossae to coalesce. ì-l --. ...: .irffer in thelrlc lwu )pqtrtrò urrr lateral outline, the former being higher nirh a prominenl antero-ventral keel. they also differ in some details of morphology and di'tribution o[/os.rrre postero-dorsally, but mainly in the different arran€lement of the ribs e,nd muri in the anterior area. in oblitacythereis (paleoblitacythereis) apula n. sp. /oss/1e d7, k2 and e11 are very depressed and tend to coalesce and a simple transverse rib joins the blind oclrlar tubercle to pore conulus capricornus. in oblitacytbereis (paleoblitacytbereis) bossioi n. sp., however, the structure separating/ossd d1 from/ossae k2 and e11 is an important feature of the anterior area and the same transverse rib bifurcates before reaching the median rib. moreover, the two species clearly differ rn the morphology ol fossae c1 and c2 and in the development of the muri which define them. the species [igured bv russo & bossio (1976) as oblitacythereis sp. 1 clearly resembles oblitacythereìs (p.) apula n. remarks. the proposed new species is very/ similar to oblitacythereis (paleoblitacythereis) bossioi n. sp. in its sp. in presence of a completely depressed area just posterodorsal to pore-conulus capricornws. oblitacytbereis sp. 1 slightly differs from the present new species, however, only in having fossae c1 and c2 not completely coalescent, but distinctly separated. even though the illustrations given by russo & bossio (1976) are not very clear, due to the preservational quality of the figured specimens, a direct examination of the maltese collection leaves no doubt that the material is conspecific. size (in mm). holotype (lv) t e. oth height 0.500.8 1 hg"..d i'."t)". (r\r) 0.8 0 0.50 n-1. "f obrerr ed \pecimens 0.76-4.84 0.48-0.52 394 the present author is conscious that the main differences between oblìtacythereis (p) apwla n. sp. and oblitacytbere,s (p.) bossioi n. sp. are confined to the region around pore-conwlws capriconws, adjoinìng fossae d3 and e1o. nevertheless, it must be emphasized that according to benson (1977, p. 13), this is one of the regions where the major changes in the structural evolution of oblitacytherezs seems to be focused. furthermore, in all the specimens the present author encountered, no gradual passage from one species to the other could be detected. previous records. aquitanian ("2 th interval" of giannelli & salvatorini, 1972) of the maltese archipelago (russo & bossio,1976). occurrence in the examined sections. the proposed new species is quite common in the tremiti islands, where it occurs rather constantly from the lower langhian (praeorbwlina glomerosa sicana subzone) to the uppermost serravallian (top of globorotalia menardii subzone). it has also been recovered in the lower langhían (praeorbwlina glomerosa sicana subzone) of the hyblean plateau. palaeoecological remarks. the species has been reported for the first time from deep thermospheric assemblages of the maltese archipelago (russo & bossio, 1.976).in the sections from the tremiti islands the species is well represented in deep associations referable to the langhian-serravallian interval. it is present, even if with low values of abundance, also in assemblages characterized by the occurrence of the psychrospheric genus agrenocythere benson, 1,972. according to these data the species seems to have been able to extend down towards the deeper and cooler environments of the psychrosphere. oblitacythereis (paleoblitacythereis) bossioi n. sp. (fig. 1b; pl. 1, fig.5,6,8) 1976 oblíucytbereis sp. 3 russo & bossio, p. 221, pl. 2, tìg. 1,2 1977 oblitacythereis (paleoblìtacythereís) ruggìeríi benson, p. 34, fig. 4c-e, 6a-c; pl. 1, fig. s, 6, 8; pl. 2, îrg.5-7; pl. 3, fig. 2, 3, 8 1980 oblíacythereis sp. (: oblìtacytbereís sp.3 in russo ec bossio, 1976) ciampo, p. 10, pl. 1, fig. 10 1981 oblitaq,thereìs sp. (: obhtarythereis sp. in ciampo, 1980) ciampo, p. 56,62. material. 3 valves and 6 carapaces; of these a left valve (c.o.b. 21) and a right valve (c.o.b.22) are figured. this material, added to that synonomized above is regarded as sufficient to erect the new taxon. size (in mm). !:.slh height range of observed specimens ,,.. _ 0.76-a.78 0.43-0.45 etymology. in honor of prof. alessandro bossio. holotype. the right valve (usnm 190828) figured by benson (1e77, pl. 1, fig. 6). type-level and type-locality. early langhian oi the northwestern area of sutera (agrigento, sicily). paratypes. the disarticulated valves figured by benson (1922, pl. 1, fig. 5, 8; pl. 2, fig. 5-7r pl. 3, fig. 2,3, 8) from the miocene of spain and sicilv. diagnosis. as for benson (1.977, p. 3a). description. as for benson (197, p.34). remarks. the species has been illustrated but not described as oblitacythereis sp. 3 russo 8r bossio (1976), from the upper langhian-serravallian of the maltese archipelago. benson (1977) described and figured it as type species of the subgenus paleoblitacythereis but erroneously referred it to carinocytbereis rugierii russo, 1966. according to benson, his attribution was based on a previous identification by ruggieri (cited in benson, 1977, p.34, as personal comm.), who misapplied russo's (1966) nominal species. later, ciampo (1980) figured it from the tortonian of the hyblean plateau and mentioned benson's incorrect attribution. finally, bonaduce 6c russo (1985) confirmed ciampo's affirmation and stated that "the specimens described by benson as p ruggieril are identical to those illustrated by russo & bossio (1976) and by ciampo (1980) as oblitacythereis (paleoblitacythereìs) sp. 3, that probably are a new species". as ciampo (19s0) noted, carinocytbereis ruggierii russo and oblitacytherels sp. 3 russo & bossio are quite different in their reticular siihouettes and fossal patterns. the latter is, in fact, more massive and characterized by a simpler reticular pattern with a marked fusion of the fossae rn the posterior region. moreover, the anterior structure of oblìtacythereis sp. 3 is more disordered and primitive, almost completely lacking the interoconcentricum (fig. t). for detailed comparison wrth oblitacytbereis (paleoblitacytbereis) apwla n. sp. (this paper) see under that species. b. dall'antonia plate 1 f'ig. 1,2, 9 oblitacytbereis (paleoblitarythereis) rwgierìl (russo); tremiti islands, serravallian. 1: left valve, c.o.b. 23, x 80. 2: right valve, c.o.b. 24, x 80. 9: detail of the anterior area, same specimen of fig. 1 c.o.b. 23, x 110. fig. l,4,7 -oblitacythereis(paleoblitacytbereis)apulan.sp.;tremitìislands,serravallian.3:leftvalve,holotypec.o.b. 15,x80.4:rightvalve, paratype c.o.b. 16, x 80. /: detail of the anrerìor area, same specimen of fig. 3 c.o.b. 15, x 150. fig.5,6, 8 oblitacytherek (paleoblìtacytherek) bossíoi n. sp.; hyblean plateau, langhian. 5: left valve, c.o.b. 21, x 80. 6: right valve, c.o.b. 22,x8a.8: detail of the anterior area, same specimen of fig. 5 c.o.b. 21, x 150. the ostracod paleoblitacytbereis 395 396 b. dall'antonia as there is no doubt as to the identity of the form designated by benson (1977) as type species of the subgenus paleoblitacythereis, the present author proposes oblitacythereis (paleoblitacythereìs) bossìoi n. sp. (: oblitacythereis sp. 3 russo ec bossio) as the new nominal type species of paleoblitacytbereis. previous records. upper langhian-serravallian ("middle-upper part of the 6th interval and zth-8th intervals" of giannelli & salvatorini, 1975) of the maltese archipelago (russo & bossio,1976). lower langhian-tortonian of sicily and tortonian-messinian of andalusía (benson, 1977). serravallian and tortonian (globorotalia acostaensis acostaensis zone sensu d'onofrio et al., 1,975) oî rhe ragusa area, hyblean plateau (ciampo, 1980, 1981). occurrence in the examined sections. this species occurs in scattered samples only in the lower langhian (praeorbwlina glomerosa s. 1. zone) of the hyblean plateau. palaeoecological notes. in the hyblean plateau oblitacythereis (p.) bossioi n. sp. occurs in typical deep thermospheric associations and also, although always represented by few specimens, in assemblages characterized lry the presence of the psychrospheric genus agrenocytbere. in the literature it is mainly reported from thermospheric associations. the available data, although not highly significant and its strong affinity n,ith oblitacytbereis (p.) apula n. sp. support the supposition that this species might have lived in a wider bathymetric and thermal range then that typical of the lower thermosphere. oblitacythereis (paleoblitacythereis) rug gierii (russo,1966) . (fig. lc; pl. 1, frg. 1,2,9) 1961. bradleya sp. oertli, p. 28, pl. 5, írg. 47 1.966 carinocytheren ruggierii russo, p.242, p|.11, fig. 4; pl. a5, rrg. r, r3-o ? 1969 bradleya (?) rp. russo, p. 23, p|.2, fig. /a-b non 1976 oblìtacytherek rugierii berggren, benson, hag, riedel, sanfilippo, schrader & tjaslsma, p. 224, pl. 6, tig. 5 1976 oblitacythereis ruggierii russo tr bossio, p. 22a, p|. 1,, ttg. 4 non 1977 oblitacytbereis (paleoblítacythereis) rugierii benson, p. 34, fig. 4c-e, 6a-c; pl. 1, fig. 5, 6,8; p|.2, fig. 5-7; pl. 3, fig. 2, 3, 8 1985 paleoblitaq,thereis rugierii bonaduce er russo, p. 130, pl. 3, fig. 4a-c material. more then 50 valves and 8 carapaces, of these a left valve (c.o.b. 23) and a right valve (c.o.b. 2,1) are figured. size (in mm). length height remarks. carinocythereis rugieríi russo, 1966 bas been reported from the middle miocene of various ltalian localities. however, it has never been either clearly illustrated by previous authors (owing to the poor state of preservation of the material at their disposal) or compared with the other representatives of the genus. the examination of well preserved material allowed the present author to reveal a surprising affinity between russo's species and oblitacythereis (oblitacytbereis) mediterranea benson (1,977, p. 33, fig. 3, 4a, 5b; pl. 1, fig. 1-4 pl. 3, fig. 1). they are very close in terms of their general slender aspect and the organ;zation of the reticulwm (fig. 1). despite this similarity, carinocythereis rwgierii russo must truly be assigned to paleoblitacytbereis, as suggested by bonaduce & russo (1985), due to the following features: 1) the basic pattern of the interoconcentricum is present, but it is discontinuous and does not form a separate structure from the rest of the reticular muri; the three small ribs extending distally from the interoconcentricum to the anterior outer margin are poorly developed and the whole reticulum of the anterior area is rather depressed; 2) the three longitudinal major ribs are not ponticulate as in oblitacythereis (o.) mediterranea and there is very little difference in height between them and the underlying mwral network. russo's species dìffers lrom oblitacythereis (o.) mediterranea benson also in its different lateral outline, the former being less elongate with a more arched longitudinal dorsal rib, a less sinuous ventral margin and a tendency to expansion of the anterior cardinal area. moreover, in oblitacytbereis (paleoblitacythereis) ruggierìi fossae c1-c2 are not completely coalescent. being separated by a low, but well-defined structure. as previously discussed, the form described and illustrated by benson (1977) as oblitacythereis (paleoblitacytbereis) rwggierii (russo) is not conspecific with russo's species and is herein described as oblitacythereis (paleoblitacythereis) bossioi n. sp.. the right valve illustrated by berggren et al. qlle) as oblitacythereis ruggieril (russo), from the el cuervo section belongs neither to oblitacytbereis (paleoblitacythereis) bossìoi n. sp. nor oblitacytbereis (paleoblitacythereis) ruggierii (russo) . in 1977 benson himself figured a left valve from the el cuervo section and assigned it to another species which he left in open nomenclature. although benson (1977) did not mention any relationship between this form and the one previously figured from the same section, they clearly belong to the same species. the el cuervo form differs from oblitacythereis (paleoblitacythereis) bossioi n. sp. in its different lateral outline, the former being stockier with a higher height/length ratio, and also in the clearly different distribution of the anterior fossae and muri. from oblitacytb ereis (pale oblitacytb ere is) rwggierii (russo) it differsrange of observed specimens 0.86-0.94 0.50-0.58 in its more massive appearance and the more obvious fusion of the posteromedian fossae. in the presenr author's opinion the form figured and described by russo (1969) as bradleya (i) rp. may possible be referable to oblitacythereis (p.) rwgierii, neverrheless rhe available illustrations do not allow a firm attribution. previous records. aquitanian-burdigalian (from nn1 to nn4 nannofossil zone sensu martini, 1,971) of sardinia (bonaduce & russo, 1985). burdigalian ("3th interval" of giannelli & salvatorini, 1972) of the maltese archipelago (russo & bossio, 1926). langhian of the langhian srratotype (oertii, 1961) and the northern apennines (russo, 1966). tortonian of the northern apennines (russo, 1969, questionable presence). occurrence in the examined sections. the species occurs in the tremiti islands from the lower langhian (praeorbulina glomerosa sicana subzone) to the tortonian (g lobigerin o iches extremus g lobigerinoide s obliquus refer benson h. r. (1972) the bradleya problem, with description of two new psychrospheric ostracode genera, agrenocythere and poseidottamicus (ostracoda: crustacea). smithsonian contr. paleobiol., v. 12, 138 pp., tvashington. benson h. r. (1976) miocene deep-sea ostracodes of the iberian portal and the balearic basin. mar. micropal.,v. 1, pp. 249 -262, amsterdam. benson h.r. (1977) evolution oî oblitacythereis from paleocosta (ostracoda: trachyleberididae) during the cenozoic in the mediterranean and atlantic. smithsonian contr. paleobiol.. v. 33, 47 pp., \x/ashington. benson h. r. (1978) the paleoecology of the ostracodes of dsdp leg 42 a. initial reports. dsdp, v. 42, n. 1, pp. 777-787, (u. s. governement printing office) vashtngton. benson r. h., rakic el bied k. & bonaduce g. (1991) an ìmportant currenr reversal (influx) in the rifian corridor (morocco) at the tortonian-messinian boundary: the end of the tethys. ocean. paleocean., v. 6, n. 1, pp. 1 6 4 1, 9 2, ril/as hin gt on. berggren \( a., benson r. h., hag b. u., riedel \( r., sanfilippo a., schrader h.j. & tjaslsma r. c. (1976) the eì cuervo section (andalusia, spain): micropaleontologic anatomy of an early late miocene lower bathyal deposit. mdrine micropal., v. 1, pp. 195-218, amsterdam. bonaduce g. & russo a. (1935) the miocene ostracodes of sardinia. boll. soc. paleont. ital., v. 23, pp. 421-437, modena. 397 subzone), with a wide gap in its distribution from the upper part of praeorbulina glomerosa sicana subzone to the lower part of globorotalia praemenardii-globorotalìa p eriph eroronda subzone. palaeoecological notes. on the basis of the studied sections the species is common in deep thermospheric assemblages, while it is poorly represented or absent in associations characterized by the occurrence of the psychrospheric genus agrenocythere.in the literature oblitacythereis (o.) rwggierii has never been reported with certainty (see introduction ro systemaric descriptions) associated with psychrospheric ostracods. this evidence supports the supposition that this species was a typical inhabitant of thermospheric waters and was probably not able to survive in colder and deeoer environmenrs. acknot'uledgements. the author is grate{ul ro prof. a. russo and prof. g. bonaduce for their useful suggestìons and critical reading. i am deeply indebted to prof. a. bossio for provision of the material and for his invaluable support. special thanks are expressed to prof. r. c. vhat.ley and prof. g. ciampo for their consrructive reviews of rhe manuscript. ences ciampo g. (1980) ostracodì miocenici (tortonianomessiniano) della regione di ragusa (sicilia). boll. soc. paleont. itdl., \. 79, pp. 5-20, modena. ciampo g. (1981) ostracodi fossili (oligocene superioreserravalìiano) del monte cammarara (sicilia centrooccidentale) & del ragusano (sicilia sud-orientale). boll. soc. paleont. ital., v. 2a, pp. 53-72, modena. dall'antonia b., di ste{ano a., foresi l. m. (in progress) integrated micropalaeontological study (ostracods and calcareous plankton) of the langhian western hyblean successions. d'onofrio s., giannelli l., iaccarino s., morlotti e., romeo m., salvatorini g., sampò m. & sprovieri r. (1975) planktonic foraminifera of the upper miocene from some italian sections and rhe problem of the lower boundary of the messinian. boll. soc. paleont. ital.,v. 14, pp. 177-196, modena. foresi l.m., iaccarino s., mazzei r. & salvatorini g. (1998) new data on middle to late miocene calcareous plankton biostratigraphy in the mediterranean area. ria. ital. paleont. strat., v. 104, pp. 95-1.14, milano. giannelli l. & salvatorini g. (1972) i foraminiferi planctonici dei sedimentr terziari delì'arcipelago maltese. i. biostratigrafia del "globigerina limestone". atti soc. tosc. sc. nat., mem., ser. a,v.79, pp. 49-74,pisa. giannelli l. & salvatorini g. (1975) i foraminiferi planctonici dei sedimentr terziari dell'arcipelago maltese. ii. biostratigrafia di "blue clay", "greensands" e "upper coralline limestone".,4ttì soc. tosc. sc. nat.. mem.,ser. a, v. 82, pp. 1-24, pisa. the ostracod pal eobli ucythereis 398 t h e o straco d pale o b litaqttb ereis iaccarino s. 11985) mediterranean miocene and pliocene pianktic foraminifera. in bolii, h. m., saunders, j. b. & perch-nielsen, k. (eds.) plankton stratigraphy, pp. 283 -3 1 4, cambridge university press. iaccarino s., foresi l. m., mazzei r. ec salvatorini g. (in progress) calcareus plankton biostratigraphy of the miocene sediments of the tremiti islands. iaccarino s. ec salvatorini g. (1982) a framework of planktonic foraminiferal biostratigraphy for early miocene to late pliocene mediterraneaî ^íea. paleont. stratigr. eool., v. 2, pp. 1,15-125, roma international commission on zoological nomenclature (1985) international code of zoological nomenclatllre. international trust for zoological nomenclature, pp. 1-338, london. martini e. (1971) standard and tertiary and quaternary calcareous nannoplankton zonation. proc.2th plankt. confer., v. 2, pp. 739-785, roma. oertli h. j. (1961) ostracodes du langhien-type. riv. ital. paleont. strat.,v. 67, pp. 1,7-44, milano. rigo m. & barbieri f. (1959) stratigrafia pratica applicata in sicilia. boll. serv. geol. 1r., v. 80, pp.351-44i, roma. russo a. (1966) ostracodi langhiani del pescale (appennino settentrionale modenese). boll. soc. paleont.1r., v. 3, pp. 227-251,, modena. russo a. (1969\ ostracodi tortoniani di montebaranzone (appennino settentrionale modenese). boll. soc. paleont. it., v. 7, pp. 6-56, modena. russo a. & bossio a. (1976) prima utilízzazione degh ostracodi per la biostratígrafia e la paleoecologia del miocene dell'arcipelago maltese. boll. soc. paleont. it., v. 15. pp. 215-227, modena. se1li r. (1971\ isole tremiti e pianosa. note illustrative della carta geologica d'italia alla scala 1:100.000. foglio 156 "s. marco in lamis". setu. geol. ial.roma. sylvester-bradley p c. (1948) the ostracod genus cythereis. j owrn. paleont., v. 22, pp. 792-797, tulsa. leven 153..162 ���������� ��� ���� ��� � �� �� ������ �������� ���� � ��� ��������� ��� ��� ��������� ����� � � �� ����� �� � �� �� ��� � ����� � �� ����� ��� �� �� � ���� �� ����� � ��������� ���� �� � � ���� �������� ����� ��� ��� ��� ������ �������� ����������� ��������� !"� #���� ����� ��� �!� $����� %�&� &' �!� �&��!������� (&����� ���� &' #��� (&������ &' &''�!&�� ��� ��� �� )���� �� &���� �!�� �((�* ������� �� �!� $����� $���� ��� )!�(! )��� ��+��,�����" &+��(��� &��& �!� ������� ������ &' #���� �!� &����� �� &���� &' �!� $����� %�&� ��� �!� ����" ������&��� &' �!� -����� .��� �!�� !�/� +��� ����� �� ����" �& ���� ������� &� 0����+��� +" ��''����� )&�1���� ��(����" ���(&/���� &��(�& � &' �!� -����� .��� '�&� �!� �&��!��� $����� ���� ��� ��(!�" '&�����'��&�� ��(������ �!� &((�����(� &' '�* �������� �� �!��� �� ����� &��(�& �� �!� '��������� �� �������� ��� �)& � �(��� &' ���������� * � ��+����� &' � � � � ��� � � � � � � ��� ���������� ! 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>> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /fra /enu (use these settings to create pdf documents with higher image resolution for improved printing quality. the pdf documents can be opened with acrobat and reader 5.0 and later.) /jpn /deu /ptb /dan /nld /esp /suo /ita /nor /sve /kor /chs /cht >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [595.000 842.000] >> setpagedevice stratigrafiarivist.r ìraliana di prleontoloera e settembre 2001 nota breve-short note the fossil vertebrate database of the natural history museum of florence and high-resolution magnetic stratigraphy in the upper valdarno basin, as a clue to date old collections elisabetta cioppi i ce gtovrnni napoleone 2 receioed. febrwar la, 2000; accepted october 1 1, 2a00 key-words: fossil vertebrate collections, data base catalogue, magnetostratigraphic geochronologl-, mid-late plìocene, upper valdarno. norrh.rn apennine.. riassunto. la sequenza del villafranchiano in italia venne fissata per una parte abbondante sulie associazioni faunistiche prolenientì dalle collezioni del museo di soria naturale di firenze. lc antiche collezioni del valdarno superiore (vs) contengono per 1o più raccolte di fossili di provenienza stratigrafica indeterminata, o con qualche possibilità di ricostruire la posizione dalle condizioni geologiche dei dintorni. la recente ricostruzione map;netostratigrafica della sequenza sedimentaria del vs ha permesso di riferire i reperti più antichì a ca. 3.0 ma nel pliocene medjo e quelli più recenti (finora) dei siti di matassino e poggio rosso ed altri della serie della successione di montevarchi, a ca.2.0-1.8 ma ìn corrispondenza del crono olduvai che m:rrca il pliocene finale. gli ultimi reperti del villafranchiano del vs costituiscono i'unità faunistica tasso e sono assegnati al pleistocene inferiore, ma non sono calibrati con la magnetostratigrafia: tale calibrazione diviene ora possibile anche per le collezioni antiche e mal datate, ed è facilitata dalle informazioni controllabili con l'automatiz.zazione del cataloso del museo che fornisce una serie di controlli incrociatì di rapida effettuazione. in tal modo, la fauna di faella e la sezione di cava faella sono state le prime ad essere controllate, quando è risultata chiara l'ìmporatanza del ruolo che il catalogo può rappresentare, specialmente per le collezioni di età vicìna all'olduvai, poiché in questo intorno sono raggiungibili gradi di definizione estremamente elevati. la datazione numerica è in grado di permettere risoluzioni comparabili tra le vecchie e le nuove collezionì giacchè entrambi i tipi di reperti si inquadrano in uno schema unirrrio rappresentato dalla cronologia delle faune, dall'evoluzione sedimentaria deì bacino e dallo sviluppo delle variazioni clìmatiche regolato dalla scansìone dei ternpi della magnetostratìgrafia. abstract. the well established biochronologic sequence of the villafranchian stage in italy is mainly based on {aunal associations from the upper valdarno (uv), mostly collected since the late 18 hundreds, and housed in the natural history museum of florence. the old collections were assembled from mostly unidentified stratigraphic levels, and their position possibly reconstructed from the surrounding geologic features. the recent magnetostratigraphic assessment o{ the sequence marked the earliest finds at about 3.0 ma in the mid pliocene. the end of the pliocene was recorded by the olduvai magnetochron, in the matassino and poggio rosso sites, and by other sparse assemblages. the tasso faunal unit, assembled in the uy is assigned to the pleistocene, yet to be clarified by magnetostratigraphic data. this calibratìon of old, poorly timed faunas was greatly facilitated by the computer-automated catalogue of the museum. the possibility of numerous feed-back controls enhanced any contradictory information in fossil collections and made them most fruitful for paleomagnetic calibration: the faella fauna and the faella main outcrop are the ones that will be first reexamined. it is ìn fact now evidenced the potential role of the catalogue for accomplìshing the calibration of old findings timed around the olduvai chron, with an accuracy depending on the available record of their inferred stratigraphic level. numerical dates will make the old collections comparable îo the new ones, as both fitting into a comprehensive framework of faunal chronologies, sedimentary evolution of the uv basin, and development of the pliopleistocene climate changes. introduction. the first fossil collection of the natural history museum of florence was catalogued in 1845 with the 356 finds from the upper valdarno (uv) vertebrates (azzaroli et a|.,1992). while the collection continued to grow, its main purposes remained since focused on determination of taxa and exhibition of the best samples, and minor attention was payed to their geographic and stratigraphic distribution. in the mean time paleontologists, sedimentologists, and geologists found the uv an apt site for development of their theories in reconstructing bulk structures (meria, 1951) and sedimentary evoiution of subsiding basins (sestini, 1936). aì1 reconstructions of that evolution (fig. t) directly involved the biochronologic succession of the fossil remains assembled in the museum collections, last formalized in the villafranchian stage by azzaroh (1,977) and defined in the mammal neogene (mn) age classification for the western europe. 1) museo di storia naturale, sezione di geologia e paleontologia, università di 2) dipartimento dì scìenze della terra, università di firenze, via la pira 4, i firenze, via la pira 4, i 50121 firenze, e-mail: cioppi@unifi.it 50121 firenze, e-mail: napo19@unifi.it 298 e. cioppi € c. napoleone new initiatives involved three separate aspecrs: 1 development of the computerized data-base catalogue for the more than 2o,o0o individual fossil vertebrates in the museum (cioppi et ai., 1996) which permits automated extraction of data frorn the entire collection, and provides a ready means of integrating new finds such as those retrieved from the matassino clay pit in 1964-65 and casa frata in 1928 (borselli et al., 19go), and the latest one ar poggio rosso in 1995: sites precisely located and completely excavated (mazzini et ai., 1.999). 2 assessment of the magneric polarity srrarlgiraphy for the uv conrinental sequences, first tested on the sediments containing the matassino deposits (torre et al., 1993), and then extended down to the lower sequence ar the onser of the sedimentation; now the entire uv sequence rs going to be analysed, in order to accornplish the other major goal of extending it to the whole 1.5 my rimc span rhere recorded. 3 continued discoverics of fossil deposits, mosrly made by amateurs, witl.r 'u'hom a more than 2o year collaboratior-r provided quick information and help in identifying and re covering important finds thar in rccenr years mainly came from the clay pits used by brick factories and for lignite exploitation. the upper valdarno faunas and their significance to the villafranchian stage. the faunas used for setring up rhe biochronologic sequence of the villafranchian stage are the classical ones from the museum collections. the sequence, recenrly updated by azzaroli et al. (1986) and gltozzr et fig. 1 geologic sketch map of the northern apennines, and tectonic setting of the extensionai basins along the belt, since the late miocene (rr-rostly marine basins, in the western areas), and pliocene and pleistocene (continental basins, to the east). the n\fse trend is their typical feature (modified, after martinì er sagri, 1993). the continental basins were generally activated in more recenr :rges, their deposition having started since the gauss chron. ai. (1997), begins with the faunal unit (f.u.) of tiversa (near asti, north-wesrern ltaly), which in the uv lacustrine sediments is represented by the local mammal fauna (l.m.f.) of castelnuovo dei sabbioni in the homonymous sequence (fig. 2). the following f.u. of montopoli (in the lower valdarno, downstream from florence) is not represenred in the uv but has a key position in the museum collection. also the nexr sr. vallier f.u. and costa s. giacomo f.u. are not represented. the new mammal unit showing a major faunal turnover is the olivola f.u. (established from the olivola locality in the norrhwestern tuscany), which n-rarked the onset of the late villafranchian: in the u! its affinities with the matassino l.m.f. were recognized by azzaroli (1977). the last f.u. is that of tasso, esrablished in the uv by the old findings and roughlv reconstructed by inferences from the geologic notes on the catalogue; it was collected frorn various localities in the upperrnost levels of the montevarchi sedimenrary sequence, but mostly in unidentified positions. the fossil deposit discovered in 1978 ar casa frata (borselli et a1., 1980), was thought to conrain close affinities with the tasso f.u. (de giuli er masinr, 1986). the biochronologic assessmenr made by azzaroli e t al. (1986) confirmed the uv villafranchian succession with its main evenrs, and the menrioned site of casa frata was d:rted prior ro the tasso f.u., while in the next updating by gliozzi er al. (1997) the olivola f.u. differentiated the affinities of the matassino local fauna by requalifying this latter as transitional betv.een tasso and olivola faunas. although olivola was retained in the base of the pleistocene, thc matassino assemblagc was magnetostratigraphically calibrated as latest pliocene (torre et al., 1,993). noteworthy for the uv biochronology and the museum collections is the significance of the poggio rosso large assemblage (recovered in 1995), pending a detailed taxonomic study of this fauna: its stratigraphic position between the olivola and tasso faunas identifies it as belonging to the general transition interval between the pliocene and pleistocene -an interval of particular interest for the uv new magnetochronologic dating of the upper valdarno sequence. briefly, the geochronologic updating of the uv mammalian site ages and the environmental evolution of the main sedimentary units follow the three well-established tectono-sedimentary phases, castelnuovo, montevarchi, and monticello cycles (fig. z). the earliest sedimentary cycle contains also the onset of the villafranchian fauna, whose remains were mainly assembled from the lignite-bearing seam in the meleto clay at an age close to 3.0 ma (albianelli et al., 1997).the meleto clay accumulated at changeable rates of continuous deposition almost until the end of the gauss chron (fig. 3), missing its final record until the gauss-matuyama (g/m) boundary ^r. 2.58 ma by only 60 ka, as calculated by the magnetocyclic analysis there applied (napoleone ee albianelli, 1998). during that time the climate underwent a progressive deterioration, from warmer and humid conditions to more temperate ones, starting at about 2.85 ma, at which spectral analysis of the magnetic signature showed a transition from a fio, 299 stratigraphic sketch of the uv i illing. r' eraged r[ter thc sections that crop out on borh rir er .ide' (m"dif ie.. after martini tr sagrì, 1993) prevailing precessional signal to the onset of obliquity driven climates. no fossil remains in the uv documented this climate crisis. in the lower valdarno the apennine uplift was less effective and the first fauna, the triversa f.u., showed there a slow evolution toward the faunal assemblage of the following montopoli f.u. (dominici, 1994; benvenuti et al., i995). this latter, in contrast, was so sharply defined as to represent the onset of the main turnover in the villafranchian (azzarol| 1977); the fossil site was caiibrated at the mid/late piiocene boundary as it was located just some meter passed the g/m magnetochronologic boundary (lindsay et al., 1980). in the second sedimentary cycle the matassino 1.m.f. represented a rather similar variety of species with the olivola f.u. (azzaroii, 1,977), taken sufficient to mark the beginning of the late viliafranchian and that of the pleistocene. the matassino fauna was calibrated in the paleomagnetically surweyed section (torre et al., 1,993), and assigned to the late olduvai (ca. 1.8 ma). therefore, in the uv fossil record, some 0.2-0.8 my time span is missing after the g/m boundary, to be added to the almost 0.4 ma in the late gauss unfossiliferous sequence in the upper meleto clay. more discussion on that will be reported, while new sites in the uv are examined with the aim to fill the gaps in the magnetostratigraphic reconstruction that presently interrupt the most important continental time series in the northern apennines. at the olduvai time the record of fossil vertebrates became rich. besides the matassino fauna (and the impressive poggio rosso deposit, recovered on 1.995 higher up in the same clay pit), the nearby faella area fossil ,t,ertebrate database and magnetic stratigraplry in valdarno 300 upper e. cioppi & g. napoleone valdarno sequence gpts magnetostratigraphy composite section r.3 4 fig.3 -magnetochronologic composite of the pliocene sediments from the uv the s. barbara section of lacustrine clays, starting frorn the basal gravels (ca. 3.3 ma), contains fossil remains of the castelnuovo dei sabbioni l.m.f. (1) in rhe ìignire-bearing layer (ca.3.0 ma); no finds are reported until its end (modified after albianelli et al., lqq7r. the contpo.ite section of ler-ane and matassino (modified, after torre et al., 1993) yielded the matassino l.m.f. (3) and the site was calibrated to the initial olduvai. in the same section the poggio rosso locality (4) was recovered in 1995 almost on top of the normal polarìty. the next section at the faella clay pìt contains the first [o'.il [ind 127. dated i nrl later than the last castelnuovo find. it is marked on the 1og, but does not represent the newly fixed assemblage of the faella l.m.f. because this covered the whole \equence. the section labeled as tasso has the officlàl njnle or rne t.rrro railroad tunnel, not to be related to the san.ìe name earlier used for the tasso f.u. assembled in the museum. no fossils r.ere there found but it exhibited the split terminal olduvai. the casa frata l.m.f. is not yet calibrated. the or errlì corttpo.ire . 'ummarìzing the present magnetochronologic assessment of the upper valdarno sequence, is reported along u irh rhe cpts "[ berggren et aì. (1995). lithology i;r -=-= --o i ot,,.tof ;] t"'.i ? li s^"af:!tu l-atcrillal2 ura\ (r è :11_ = =_ol iol iol 9i et q d) () o(j a ti z d i e i -i|lil i irt unio l -. knenct i.0.t yielded the older collection to which recent finds were added, such as the deer anrler of april 1999. in the matassino pit, moreover, new finds were recovered stratigraphically above the poggio rosso site, until julyaugust 1.999,they were collected from layer beds which can now be correlated with the ones defined in the faella clay pit, recently surveyed for sedimentology and magnetostratigraphy along new cuts in the section ralbianelli et al. 2000). the succeding tasso f.u. still belongs to the last deposits of the montevarchi sedimentary cycle ìà/ith samples collected from the areas around the tasso creek. but the variety of samples assembled in the museum old 3 .22 lfannoth collectìons were also with poor paleontological investigations. such fossil remains, as iisted in the catalogue, came from a number of localities whose sediments are generally made by the upper sandy members of the montevarchi sequence. whereas some markers suggest that the time span involved could have been rather short, the tasso f.u. is not as yet calibrated. the third sedimentary cycle, monticello, accumulated by alluvial fans produced by the arno river meandering when the sedimentary activity moved towards the basin fill; continuity in deposition is difficult to establish. most sediments accumulated in the basin upstream, and fossil remains were assembled from ma o u o@ e1 fl 2i èl 'lel 6i oi 60 capitone (s. etru.) 111 86 74ca senèze (s. etru.) med 134.91 109.91 84.42 65.63 min 128 104.95 78.11 60.16 max 150.88 117.8 91.5 71.3 cagnes sur mer (s. etru.) 123.64 107.49 76.26 62.12 vergranne (s. hund.) 130 115.5 80 59 116ca 80 71 untermassfeld (s. hund.) 120.8 108 81.5 75 108.8 73 mosbach (s. hund.) 127 115 83 72.8 114 85 65 tab. 5 comparative dimensions of scapulae from pirro nord (* = data from mazza et al. 1993), pietrafitta (data from mazza et al. 1993) and of s. etruscus (s. etru.) (data from guérin & heintz 1972; mazza 1988; garcía-fernández et al. 2001) and s. hundsheimensis (s. hund.) (data from fortelius et al. 1993; guérin 1983; kahlke 2001; lacombat 2005). dtd = breadth of distal epiphysis; tdcs = breadth of collum scapulae; lg = length of the glenoid cavity; bg = breadth of the glenoid cavity. fig. 4 bivariate diagram of scaphoids from pirro nord and pietrafitta (data from mazza et al. 1993), s. etruscus (s. etru.) (data from mazza 1988) and s. hundsheimensis (s. hund.) (data from kahlke 2001; lacombat 2005). lpb = length of the proximal articular surface; bpa = breadth of the proximal articular surface. all the rhinoceros remains should be ascribed to stephanorhinus sp. cava redicicoli (rome, central italy) the site of cava redicicoli, discovered by blanc in the 1950s (blanc et al. 1955), may be chronologically correlated to the colle curti fu. in this site, caloi et al. (1979) listed a few rhinoceros remains, including a skull fragment, a fragmentary mandible and some teeth. these remains were ascribed to dicerorhinus sp. (rhinoceros sp. in blanc et al. 1955) and were later reported as s. hundsheimensis (caloi & palombo 1988; di stefano et al. 1998; palombo et al. 2002). the remains from cava redicicoli have morphological characters similar to s. hundsheimensis, such as the very small difference in height between the bottoms of the valleys in the lower molars (mpur 1956 r45a, b), the relatively large and robust horizontal ramus of the mandible (mpur 1956 r45), the profile of the occipital crest of the skull (mpur 1956 s.n.), convex in posterior view and slightly concave in dorsal view (pandolfi, unpublished data). the faunal list of the site includes typical villafranchian taxa and taxa more frequently recorded in galerian faunas or closely related to the galerian ones. due to the composition of the redicicoli fauna, caloi et al. (1979) and di stefano et al. (1998) suggest the presence of two different faunal assemblages, which are believed to have come from two different levels. the first level is correlated with the pirro fu and the second one with the isernia fu. according to di stefano et al. (1998) the rhinoceros remains come from this second level. caloi & palombo (1995, 1997), pa lombo et al. (2002) and milli & palombo (2005) suggest that the taxa are coeval and that the redicicoli faunal assemblage should be ascribed to the colle curti fu only. unfortunately, the fossiliferous deposit does not exist anymore and the problem is difficult to resolve. 180 pandolfi l. & petronio c. imola basin (imola, northern italy) rhinoceros remains discovered in the imola basin (referred to the late villafranchian-early galerian) were ascribed to rhinoceros leptorhinus by falconer (1868) and to dicerorhinus cf. etruscus by azzaroli & berzi (1972). later, they were reported as s. cf. hundsheimensis (palombo et al. 2002). the upper toothrow, preserved in the museum of imola and figured in falconer (1868) and in azzaroli & berzi (1972), has a number of common characteristics with s. etruscus and s. hundsheimensis. however, some characters allow to the remains to be ascribed to falconer’s species, such as the general profile of the vestibular wall in the molars and premolars, the absence of the crista and the presence of a single crochet in p3/, the presence of a crista, a single crochet and a mesial cingulum in p4/, the absence of the antecrochet and the presence of a protocone constriction in m1/, a well developed single crochet and the absence of the antecrochet in m2/ (see also lacombat 2006). in addition, the upper toothrow shows many similarities in morphology and dimension with s. etruscus from capitone (terni, central italy) (fig. 9). mugello basin (mugello, central italy) specimens with s. hundsheimensis affinities were reported in the mugello basin; in particular, a deformed and damaged skull from grezzano (igf 12728) preserved in the museum of geology and paleontology of the university of florence (mazza 1988). the characters of the teeth and the skull from grezzano are more similar to toula’s species, even if the dorsal profile angle (n) is more quite similar to that of s. etruscus (n = about 150) than s. hundsheimensis (n >170) (see also lacombat 2005). also, in the skull of grezzano the anterior border of the orbital cavity overlies the anterior half of m2/ and the posterior border of the narial notch lies at the level of p4/, but these characters are similar to that of the skull of s. etruscus from capitone (in which the anterior border of the orbital cavity lies at fig. 5 ratio diagrams for mc iii from pirro nord (fi = preserved in the museum of palaeontology of florence, data from mazza et al. 1993), pietrafitta (data from mazza et al. 1993), s. etruscus (s. etru.) (data from guérin & heintz 1972; mazza 1988) and s. hundsheimensis (s. hund.) (data from fortelius et al. 1993; kahlke 2001; lacombat 2005) (standard diceros bicornis). abbreviations in tab. 4. stephanorhinus etruscus from pirro nord and other late early pleistocene rhinoceros remains of italy 181 the level of m2/ and the posterior border of the narial notch at the anterior half of p4/). unfortunately, the remains were found in fluvio-lacustrine deposits with a very long chronological range and they have no direct stratigraphic control. from the same localities, typical early galerian mammal remains have been found as well (masini et al. 1994; abbazzi et al. 1995). discussion and conclusions the proportions, morphology and dimensions of the rhinoceros remains from latest villafranchian sites in italy are comparable to those of s. etruscus. according to guèrin (1980), during the late villafranchian, this species is inclined towards a general variation in body size and proportions. guérin (1980) considers the populations of this time span as “second evolutionary stage”. the same proportions and size of the latter are evident in the capitone sample and in the remains from senéze, well as in the specimens from pirro nord. in italy, s. etruscus is reported from several villafranchian sites (ambrosetti 1972; ambrosetti & cremaschi 1975; mazza 1988; petronio et al. 2002); according to various authors, its last occurrence is at the beginning of the late villafranchian (tasso-olivola fus; gliozzi et al. 1997). in the present study, however, the occurrence of the etruscan rhinoceros is confirmed until the villafranchian-galerian transition. during the latest villafrachian and earliest galerian, the presence of s. etruscus has been reported from several european sites, for example cueva victoria, incarcal, venta micena, huéscar-1, cúllar de baza 1, atapuerca (gran dolina td4-8 and sima del elefante te14), la sartanette, blassac-la-girondie and peyrolles (heintz et al. 1974; cerdeño 1993; van der made 1998; rosas et al. 2001; garcìa-fernandez et al. 2003a; van der made et al. 2003; palombo & valli 2003; lacombat 2005) (fig. 10). stephanorhinus etruscus was also initially reported from the spanish sites of fuente nueva-3 and barranco león-5 (1.3-1.1 ma), where lithic artefacts have been found (turq et al. 1996; martínez navarro et al. 1997; gibert et al. 1999; oms et al. 2000). the rhinoceros remains of these sites were later ascribed to s. cf. hundsheimensis (martínez navarro et al. 2003). in europe, the first occurrence of s. hundshefig. 6 ratio diagrams for the mc iv from pirro nord (fi = preserved in the museum of palaeontology of florence, data from mazza et al. 1993), pietrafitta (data from mazza et al. 1993) versus s. etruscus (s. etru.). (a) (data from guérin & heintz 1972; mazza 1988) and s. hundsheimensis (s. hund.) (b) (data from kahlke 2001; lacombat 2005) (standard diceros bicornis). abbreviations in tab. 4. 182 pandolfi l. & petronio c. imensis is reported during the latest early pleistocene and early middle pleistocene. it is present in sites such as vallonnet, sainzelles, tour de grimaldi, ceyssaguet, saint-prest, durfort, untermaßfeld, dorn-dürkheim 3 and trlica (guérin 1980; radulescu & samson 1985; codrea & czier 1991; codrea & dimitrijević 1997; franzen et al. 2000; kahlke 2001; palombo & valli 2003; guérin et al. 2003; lacombat 2005) (fig. 10). recently, s. hundsheimensis was reported from the spanish site of vallparadís (1-0.800 ma) (alba et al. 2008). thus, during the latest early pleistocene – earliest middle pleistocene, s. etruscus and s. hundsheimensis seem to be coeval in europe. however, in agreement with van der made (2000), the typical early middle pleistocene s. hundsheimensis appears to be more primitive than the early pleistocene stephanorhinus; this last is represented by an evolutionary stage of s. etruscus (second evolutionary stage). s. hundsheimensis may have been an immigrant that replaced s. etruscus at the end of the early pleistocene. this is in line with the model of fortelius et al. (1993), which does not derive s. hundsheimensis from s. etruscus, rather than with guérin’s (1980) model. stephanorhinus hundsheimensis probably reached europe from asia between 1.3-1.1 ma and gradually replaced s. etruscus; at the same time, populations of the latter species were fragmented into isolated demes. in italy, the occurrence of s. hundsheimensis in the late villafranchian sites (pirro nord, madonna della strada) is not considered reliable at present time. in the lignitiferous deposit of pietrafitta (perugia, central italy), chronologically earlier than pirro nord site (farneta fu), several rhinoceros remains were found as early as 1900. at first, ugolini (1921) ascribed some teeth to rhinoceros etruscus and later, other rhinoceros remains were ascribed to dicerorhinus etruscus (ambrosetti et al. 1987) and s. cf. hundsheimensis (mazza et al. 1993). some morphological and morphometrical considerations of the rhinoceros remains from pie fig. 7 ratio diagrams for mt iv from pirro nord (data from mazza et al. 1993), pietrafitta (data from mazza et al. 1993). s. etruscus (s. etru.) (data from guérin & heintz 1972; mazza 1988) and s. hundsheimensis (s. hund.) (data from fortelius et al. 1993; kahlke 2001; lacombat 2005) (standard diceros bicornis). abbreviations in tab. 4. stephanorhinus etruscus from pirro nord and other late early pleistocene rhinoceros remains of italy 183 trafitta, listed by mazza et al. (1993), are reported as evolutionary trends of s. etruscus by guérin (1980: 599-603). furthermore, the small late early pleistocene rhinoceroses, very likely identical to that was assigned to s. cf. hundsheimensis by mazza et al. (1993), are reported from spain as s. etruscus (cerdeño 1993; van der made 1998). however, the remains from pietrafitta are proportionally more similar to those of s. etruscus than s. hundsheimensis (for example see figs 6-8). besides this, some morphological and metrical differences in the pietrafitta remains may fall within the intraspecific range of variation of the etruscan rhinoceros; they may also be the result of adaptation to a particular habitat (see also garcía-fernandez et al. 2001). however, a more detailed analysis is needed. other rhinoceros remains discovered in italian sites chronologically related to the villafranchiangalerian transition (mugello basin) have close affinities with s. etruscus. also, remains attributable to s. hundsheimensis from the earliest galerian sites (colle curti, castagnone) are uncertain and scarce, and no more data is available on the origin of the remains of cava redicicoli. thus, s. hundsheimensis seems to be present in italy only after 1 ma (slivia fu). this species is certainly present during the middle galerian (middle pleistocene). it is found in cesi (about 0.7 ma) (mazza 1996; ficcarelli et al. 1997), in the lower levels of ponte galeria (about 0.8 ma) (petronio 1988; petronio & sardella 1999; pandolfi, unpublished data) and in isernia la pineta (about 0.6-0.55 ma) (sala & fortelius 1993; coltorti et al. 2005). the dispersal event of s. hundsheimensis into europe and italy may be related to the climatic deterioration of the latest early pleistocene. the same may be true of the variations in proportions and size of s. etruscus. in fact, during this time, the passage to a phase characterized by long glaciations with 100.000-year cycles occurs. the difference between the temperatures fig. 8 ratio diagram for mt ii from pirro nord (data from mazza et al. 1993), pietrafitta (data from mazza et al. 1993). s. etruscus (s. etru.) (data from mazza 1988) and s. hundsheimensis (s. hund.) (data from kahlke 2001; lacombat 2005) (standard diceros bicornis). abbreviations in tab. 4. 184 pandolfi l. & petronio c. during glacial and interglacial phases becomes more pronounced (leroy 2007). in italy, more arid conditions, during the pirro fu, are testified to by the occurrences of bison degiulii and a lightly build, medium-sized horse, equus altidens. also, typical african taxa, such as hystrix refossa, megantereon whitei and theropithecus sp. disperse on the peninsula (petronio et al. in press). concurrently, the disappearances of leptobos and eucladoceros were recorded. however, at pirro nord, woody areas and humid patches must have been still present at some point, as indicated by the presence of apodemus flavicollis, as well as the presence of amphibians and insectivores. at other sites related to the pirro fu, in which s. etruscus was discovered, lacustrine basins or ponds were present (for example in the scoppito site; maccagno 1962). later, a very arid-steppic phase is recorded during the colle curti fu. in the ranica site in northern italy, a cold, steppic phase with cervalces latifrons is well documented (ravazzi et al. 2005). nevertheless, trees remain present at the foot of the alps even during this phase. in contrast, in central italy, the pollen diagrams of colle curti and cesi (bertini 2000) show high percentages of herbaceous forms, a clear evidence of more open vegetation. among the mammals, presence of a cold microfauna represented by prolagurus pannonicus and predicrostonyx sp. is reported in the blue-clay layer of the ponte galeria area (roma, central italy) (kotsakis et al. 1992). these finds testify a very cold phase, correlated with the marine isotopic stage 24-22. during this time, s. etruscus disappeared from italy and the occurrence of s. hundsheimensis is reported. other asian taxa, such as praemegaceros verticornis and bison schoetensacki, co-occur with s. hundsheimensis. the diffusion of s. hundsheimensis can be also related to a greater capacity of the species to adapt to the climatic conditions and diet (with highly flexible feeding, kahlke & kaiser in press) than s. etruscus. moreover, the distribution of the latter seems to be more influenced by humidity (guérin 1980). in italy, the absence of remains definitely attributable to s. hundsheimensis before 1 ma may be due to the conformation of the italian peninsula. this may have played a decisive role in the delay of the dispersal event when compared to the rest of continental europe. in fact, delayed dispersal events are common in fig. 9 upper toothrows of s. etruscus from the imola basin (a) (after azzaroli & berzi, 1972; not in scale) and capitone (b) (scale bar: 5 cm). fig. 10 location map of selected late early pleistocene sites of europe and italy with remains of s. etruscus and s. hundsheimensis. sites with s. etruscus: 1 = pirro nord (1.3-1.6 ma); 2 = scoppito (1.3-1.1? ma); 3 = imola basin (latest early pleistocene); 6 = lézignan-le-cèbe (about 1.5 ma); 8 = atapuerca gran dolina td4-8 (about 0.800 ma) and sima del elefante te14 (1.4-1.1 ma); 9 = cueva victoria (about 1 ma); sites with s. hundsheimensis: 4 = vallonnet (about 1 ma); 5 = soleilhac (about 0.750 ma); 7 = fuente nueva-3 and barranco león-5 (1.3-1.1 ma); 10 = dorn-dürkheim 3 (about 0.800 ma); 11 = untermaßfeld (about 1 ma); 12 = trlica (latest early pleistocene). stephanorhinus etruscus from pirro nord and other late early pleistocene rhinoceros remains of italy 185 italy during the pleistocene, in particular for species that arrive from asia (for example coelodonta antiquitatis is not present before mis 4 and praemegaceros solilhacus is present during the isernia fu, about 0.6000.550 ma) (petronio et al. in press). some african or southwest asian taxa seem instead to be present first in italy and later in central europe. this is the case of dama clactoniana and perhaps homo. acknowledgements. we thank prof. l. werdelin and an anonymous reviewer for useful comments on the manuscript. also we thank dr. v. mecozzi, c. salari and dr. l. salari for english review. abbazzi l., benvenuti m., rook l. & masini f. 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�� ���� �� +��� � ������ 0$", �#+*' 9$"-'#)�. �#� �"*#3 �. �. #1 �"#3 �. �. 7�1(8 � �5* �') $�/�'%�( 0$", �+� �$' ((') "0 �+'# . ���� 7�&�)� )�� ��� ���� @� �����. m�# �+'� #�(�n. , s. majiashanensis. rivista italiana di paleontologia e stratigrafie volume 1 1o pp. 3-4 april 2004 the 6t" international symposium on the jurassic system and its proceedings since the first edition of the international symposium on the jurassic system held in erlangen in 1984 and mainly devoted to ammonite biostratigraphy, more and more the isjs became a wider aimed meetins where the focus is a time period with all its aspects. this makes scientists from very different fields of earth sciences to meet in an international and multidisciplinary environment, resulting in lively and fertile discussions. the 6'1'isjs was oreanized by giulio pavia (torino university) in mid-september 2002. more than 200 people convened to mondello, near palermo in sicily, both to join the large field trip program and to contribute in the scientific sessions that were hosted at the centro congressi la torre from 16 to 19 september. the participants had in fact the possibility to spend four sunny days on the blue sea of the mondello bay. in many occasions they also enjoined outstanding cultural attractions and the sicilian cuisine that was sublimated in some country lunches and in the social dinner organized in the fascinating atmosphere of the old palermo. from the scientific point of view, the numerous field trips were orsanized in two intervals. the pre-symposium fieldtrip was aimed to introduce the jurassic geology of western sicily giving an overview of the different paleogeographical domains of such a western tethys sector. four post-symposium fieldtrips were also organized, ts/o focussed on the trapanese and saccense domains, other two visitine some classical jurassic sections in central apennines and southern alps. a great effort resulted in the publication of a volume edited by massimo santantonio (roma university) that was distributed to all the participants. the field trip volume is a huge collection of mostly unpublished data, that derive from researches of a national group coordinated by g. pavia and funded by the itaìian university ministry. part of these data senerated some of the notes collected in the present volume; some others need widenins for future publication. as to the scientific sessions, more than 200 abstracts were submitted and compelled the organisation of three parallel sessions and two periods of poster display. the scientific program was organized in six topical sessions, that highlighted the multidisciplinary character of the meeting: (1)jurassic tectonics and sedimentation. (2) taphonomy, facies and paleoenvironmental analysis. (3) jurassic organisms in space and time. (4) integrated stratigraphy. (5) paleoceanography and paleobiogeography. (6) geoconservation: protecting jurassic fossils, sites and science. four plenary sessions were dedicated to lectures of outstanding scientists concerning aspects ofjurassic stratigraphy. jobst wendt (tùbingen) gave a fascinating and unforgettable picture of how sicilian geology and society appeared in the sixties. raimondo catalano (palermo) illustrated the complex structural arrangement of the sicilian chain. john callomon (london) made the point of two centuries of progresses in jurassic bio-chronostratigraphy. gerd westermann (hamilton) discussed on ammonoid biogeography integrating ecology, phylogeny and oceanography aspects. the president of the 6'hisjs would conclude this presentation first stating that the symposium can be confidentially introduced as a highly positive scientific performance; this is proved, if further necessary, by the collection of the papers published in this volume of the prestigious italian periodical. second, it must remind that the meeting would not have been possible without the activity of the organizing committee among which the coordinators rodolfo coccioni, stefano cresta, piero di stefano, luca martire, umberto nicosia, guido parisi. giulio pavia president of the 6"'i sj s this special issue of the rivista italiana di paleontologia e stratigrafia consists of 45 original papers presented at the 6th international symposium on jurassic systen.r. these papers can be erouped in two catesories: in fact the first part of the volun.re deals with topics regardine the geo-paleontological knowledees of the jurassic period, rvhilst the second part of the volume is focussed on geoconservatio of jurassic geo-paleontological sites. the subjects dealt by italian and foreigner authors involve muldisciplinary approach to strati!íraphy with notable contributions on new biostratigraphic dirta, biozonations, stratieraphic correlations, boundaries definitions, palaleobiogeography and palaeoenvironnlents of several geological areas in europe, america, asia and africa. this volume represents an importrnt tool for global correlations based on biostratigraphic, sedimentological and geodynamic data emphasiz-ing new paleoreographic features. these several contributions, accompanied by wide and new iconographic documentation, represent an indispensable synthesis for a modern integrated biostratigraphic approacn. one of the main developed topic of this volume regards the definition of the main boundaries of jurassic period dealt with biostratigraphic, physical and chemic,rl methods. furthermore, several papers give new data on paleoecological and paleoenvironmental reconstructions of the different jurassic facies. the last part of the volume is dedicated to the inventory of representative documents, modelling and characterisation of the essential controlling factors, inventory of representative sites for the studied seoevents, selection of geosites, proposal of olistic formats for a typologic inventory and for evaluation of the them,rtic geodiversity. this is linked to the geosites iugs program, which develops a geoconservation strategy aimed to the protection of "geolouical nronuments". first of all we whish to thank the reviewers of the manuscripts: aberhan martin, berlin; atrops francoise, lyon; aurell marc,zarasoza; bloos gert, stuttgart; bosence dan, london; bortolotti valerio, firenze; bown paul, london; brancucci gerardo, genova; callomon john, london; caracuel jesus, alicante; cariou elie, poitiers; carter elisabeth, portland; catalano raimondo, palern.ro; cecca fabrizio, paris; christ hans a., basel; cirilli simonetta, perugia; clari pierangelo, torino; coccioni rodolfo, urbino; conti maria alessandra, roma; cope john, cardiff; cresta stefano, ron.ra; d'argenio bruno, napoli; d'arpa carolina, palermo; di stefano pietro, palermo; dietl gerd, stuttgart; dinis joree, coimbra; dun.ritrica jud ruth, bern; elmi seree, lyon; enay raymond, lyon; erba elisabetta, milano; feist-burkhardt susanne, london; fernandez lopez sixto, madri d;fozy istvan, budapest; fursich franz, \wurzburg; galacz andras, budapest; garzanti eduardo, milano; glowniak eva, warsawa; gorican spela, ljublf ana; guex jean, lausanne; hart malcolm, plymouth; henriques maria helena, coimbra; hewaidy abdel galil, cairo; hillebrandt von axel, berlin; hughes geraint, dhahran; jansa luba, halifax; jenkyns hugh, oxford; khrishna jay, varanasi; lozar francesca, torino; martire luca, torino; masetti daniele, trieste; mattioli emanuela, lyon; matyia bronislaw, warsawa; meister christian, genève; melendez guillermo, zaragoza; monaco paolo, perugia; morton nicol, london; nicosia umberto, roma; oloriz federico, granada; page kevin, plymouth; palfy joszef, budapest; pavia giulio, torino; remane jurgen, neuchatel; rettori roberto, perugia; riccardi alberto, la plata; ruiz ortiz pedro, jaen; srntantonio massimo, roma; schweigert gunter, stuttgart; simone lucia, napoli; smith paul, vancouver; stampfli gerard.m., lausanne; surlyk finn, copehagen; thierryjacques, dijon; voros attila, budapest; \warrington geoffrey, nottingham; wendt jobst, tubingen; westermann gerd, hamilton; wierzbowski andrej, warsawa; zeiss arnold, uttenreuth. the guest-editor appreciates the patience of the authors in waiting for the publication of this volume, but believes that it can be justified by the quality and sisnificance of the final product. thanks are due to the editorial board of the rivista italiana di paleontologia e stratigrafia for its support and accepting for publication the proceedings of the symposium. in particular thanks are due to maurizio gaetani and cristina lombardo (university of milano) for their kind availability. many thanks are due to the dipartimento di scienze della terra of the university of perugia (italy). ve whish to thank roberto rettori (university of perueia) for his co-operation, gloria andreini, massimiliano bilotta and fabio masini (university of perueia) for contribution to the manasing the papers. special thanks are due to kevin page (university of plymouth) for his great and kind contribution in the editing the part of the volume dedicated to geoconservation subjects. guido parisi, guest editor rivista italiana di paleontologia e stratigrafia voìume 104 numero 1 tavote l-l pagrne 131-138 aprile 1998 noa breve tube morphology and structure of the bathyal mediterranean s erpulid h y alo p om atu s v ar i o r ug os us ben-eliahu & fiege, 1996 (annelida, polychaeta) rossana sanfilippo* receioed september 23, 1997; accepted february 13,1998 key-uords: polychaeta, serpulidae, tube morphologn tube structure, recent, pleistocene, mediterranean, deep-sea benthos. riassunto. il polichete serpulide hyalopomatus oariorugosus, recentemente descritto da ben-eliahu & fiege (1996), mostra una peculiare rugosità sulla superficie esterna del tubo, a differenza d,elle a\tre specie cogeneriche. tale c^ratreíe presenta una notevole variabilità, anche lungo uno stesso tubo. osservazioni al sem rivelano che 1a rugosità è data dalla presenza da "scaglie", irregolari per forma e dimensione, disposte lungo le strie di accrescimento. viene discusso il valore sistematico di questo ed altri caratteri morfologici. da osservazioni su fratture, la parete del tubo presenta cristalli di carbonato di calcio equidimensionali, ad habitus prismatico. questi hanno una struttura omogenea microcristallina granulare e sono disposti in strati che si sovrappongono durante l'accrescimento intorno al lume del tubo secondo superfici coniche. una pellicola discontinua di cristalli a struttura amorfa criptocristallina può ricoprire la superficie estèrna del tubo. tubi fossili di tale specie non sembrano presentare modrfìcazioni diagenetiche dal punto di vista strutturale. lr specie, di minuscole dimensioni, colonizza piccoli substrati duri sparsi su fondali fangosi de1 piano batiale. può anche essere gregaria; in questo caso i tubi incrostano microcavità come l'interno dei calici di coralli. sono, infine, riportate nuove segnalazioni di h. variorugosus in tanatocenosi profonde del mediterraneo occidentale ^lt'. "h" i. depositi batiali pleistocenici dell'italia meridionale. abstract. the species studied differs from the orher hyalopo. matus species by having a peculiar rugose sculptured outer tube surface. this character is markedly variable; it can be more or less developed and may also vary along the same tube. observed by sem, this rugosity proves to consist of irreguìar flaps, roughly following the grofth lìnes. sem obsenations on transverse fractures show a homogeneous conposition of the tube wall, consisting of calcium carbonate crystals. these crystals show a prismatic habitus and are arranged in a homogeneous granular microcrystalline structure. new records of h. variorugosus are from deep water stations in the western mediterranean and from pleistocene bathyal deposits of southern ltaly. the discussion covers aspects of tube characters of use for taxonomy. introduction. expeditions lirania 92, eocumm 94 and eocumm 95 in the southern tyrrhenian sea provided material of the recently described serpulid hyalopomatws aariorugosus ben-eliahu ec fiege, 1996. the species has also been found in pleistocene pelitic sediments from southern kaly (di geronimo et a1., in press). previously tubes of this species had been menrioned as "serpulidae sp." in barrier er al. (1939) and as protis sp.\ in di geronimo et al. (1995). these records were from mediterranean bathyal thanatocoenoses and pleistocene deposits in southern ltaly. ben-eliahu & fiege (1996) described h. vartorugosus [rom the levant basin. in the present paper, the tube morphology and structure of h, rtariorugosus are investigated in more detail. the tube is characterised by a rugose microsculpture, which allows an easy identificarion even of empty tubes (dead or fossil specimens). serpulid tube morphology and strucrure have rarely been investigated with modern techniques. some more recent works insist on the importance of tube characters (bubel et a1., 1983; ten fiove & zibrowius, 1986; zlbrowius 6c ten hove, 1987; ten flove & smith, 1990; nishi, 1993;ptiiai 6c ten flove, 1994; aliani et ai., 1995, sanfiiippo, 1996). the study of h. oariorugosus provides the opportunity to investigate the reliability of tube morphology and structure as diagnostic features for taxonomy. examined material. recent (southern tyrrhenian sea) (fig. 1): urania cruise, july 1992, "del thoro" canyon, stn. dg03, 38o43.78'n, 8020.59'8, i95 m: many empty tubes on fossil oyster shells of wùrmian age; a few tubes and fragments free in the mud. eocumm 94 exp., july 1994. stn. 9, 38'34.39'n, 14"29.56'f-,359 m: a few tube fragments free in the mud. stn. 18, 38'31.59'n, 1,4"53.57'f.,3oo m: a few 'r istituto policattedra di oceanologia e paleoecologia, corso italia 55,95729 catania. ). tyriìhenian se s n + i i 0 100 knr . eolian islands stn. 18, 300 m r eolian islands stn. 27, 350 m i eolian islands stn. 26, 945 m a eolian islands sur. 9, 959 m o eolian lslands sll. 7. i197 m r eolian islands stn. 328, \219 nt [ "dcltbro"ciuryon sm. dg03, 195 m a lazzàro, brly to middlc pleistocene marls o archi, elrly to middle plcistoccne marls sicily 14 1,32 n" r. sanfilippo l6 tube fragmenrs. stn. 27,38"2t.45'n, 15o46.55'e, 350 m: five tube fragments free in the mud. eocumm 95 exp., june 1995. stn. 7, 38o34.09'n, 1.4"25.01'f-, ll97 m: one complete empty tube on a pteropod shell fragment; some broken tubes on pteropod shells and on pumice. stn. 26, 38o38.30'n, 14o57.53'e, 945 m: one broken tube on a pteropod shel1 fragment; one distal end. stn. 32b, 38'33.53'n, 15'05.33'e 121,9 m: two complete empty tubes on a small piece of volcanic rock; one incomplete tube on a pteropod she1l. pleistocene (southern calabria) (fig. 1): lazzàro, early to middle pleistocene bathyal marls rich in scleractinians and isidid gorgonians, various samples: one tube on a scleractinian, some tubes inside lophelia pertusa calices; one distal end of tube; many tube fragments. archi, early to middle pleistocene bathyal marls, various samples: many distal ends and tube fragments. methods. the deep water material was collected by means of dredge (stn. dg03), grab (stn. 27) or box corer (stn. fig. 1 location of samples. 7,9, 18,26,328). the fossil material was collected by means of bulk-samples (di geronimo & robba, 1976). except for part of dgo3, in which serpulids were found encrusting oyster shells, the samples were treated as routinely done for macrofauna studies, i.e. washed and sieved. h, aariorugoszs tube fragments s/ere found in the fine fractions (500 pm and 250 pm). some tubes were brokeú, in order to show the inner structures. before coating with gold palladium, tubes were treated with concentrated fizoz to eliminate organic matter from the surface. results. tube morphology and structure. lvhereas the original diagnosis of h. variorugosus was mainly based on the anatomy of the soft body (ben-eliahu & fiege, 1,996), the present description focuses on tube morphoiogy and structure. the tube is white, opaque and small-sized, up to 15 mm in length (p1. 1, fig. 1,2), often more or less coiled, folding on itself in the attached part and distally becoming straight and rising from the substrate (p1. 1, fig. 6a, b, c). plate 1 hyalopomatus oariorugosus ben-eliahu ec fiege, 1996. scaie bar : 500 pm (figs. 1-2), 100 pm (figs. 3-6). fig. 1 tubes cryptic in a small cavity of the substratum. lazzà,ro. fig. 2 tubes gregarious in inner pan of a scleractinian calìce. lazzàro. fig. 3 tube orifice with a circular smooth peristome. eolian islands, stn.27. fig. 4 distal ends of tubes. lazzaro. fig.5 attached pan of the tube with a smooth peristome directed towards the tube orifice. "del thoro" canyon, stn. dg03 fig. 6 distal erecr parts of tubes, rising from the substratum, more (6a) or less (6b, 6c) rugose. lazzà;o (6a), archi (6b, 6c). 133pl. 1 tube motpbology and structure of hyalopomatus aariorugosus t34 r. san/ìlippo the cross section is circular (p1. 1, fig. 1,2,3), the outer diameter relatively constant aiong the tube, ca. 170 ytm (pl. 1, fig. 3), and the wall thickness reaching ca. 20 pm (pl. 2, fig. 3, 5). sometimes there are peristome collar rings, slightly flaring and directed distally (p1. 1, fig. 5). the tube orifice may be surrounded by a smooth peristome, slightly flaring (pl. 1, fig. 4, 5. pl. 2, fíg. 3a). like in other serpulids, the early part of the tube, corresponding to the juvenile stage (sanfilippo, 1993), is smóoth. it is separated from the adult portion by a peristome (pl. 2, fìg. 1,). being very thin and fragile, the early portion is often lost in the adult specimens and in fossil material. the outer surface of the tube is rugose. sem examination shows the rugosity to consist of transverse flaps, more or less frayed out and posteriorly directed (pl. 1, figs. 4, 5, 6; pl. 2, fig. 2), roughly following the growth line pattern. sometimes these flaps are few and weakly developed (pl. 2, fig. 3c), the tubes appearing almost "smooth" (pl. 1, fig. 4; pl. 2, fig. 3b). in other cases, the flaps are well-developed and numerous (pl. 2, fig. 2), giving the tube a "scaly" appearance. both "scaly" and "smooth" tubes may occur in the same sample. the sculpture may also vary along the same tube, as already stressed by ben-eliahu & fiege (1996). sem observations on transverse fractures of recent material evidence a homogeneous composition for the tube wall, consisting of calciurn carbonate crystals (pl. 2, fig. 3, 6). in longitudinal section the crystals are arranged in layers that dip towards the opening at an angle of ca. 30" (pl. 2, fig. 5). the layers overlay each other during growth, according to a conical plane. this type of microstructure and the layered pattern are common to most of the mediterranean serpulids (sanfilippo, t993, t996). at higher magnification all crystals show a simiiar size q-3 pm in length), and they have a prismatic habitus (pl. 2, fig. 6). they are arranged in a criss-cross orientation, in a homogeneous granular microcrystalline structure (see hall, 1980) (p1. 2, fig. 6). the opaque appearance of the tube is due to the randomly arranged small crystals, as reported also for other serpulids (ten hove & zibrowius, 1986;zrbrowius & ten hove, 1987). crystals and their arrangement are also visible on the outer surface of the tube. rarely, the surface may be covered by a discontinuous film of very small crystals, organised in amorphous cryptocrystalline masses (pl. 2, rtg. / ). the microstructure appears not to be affected by diagenetic modifications, being unchanged in fossil tubes. distribution and ecology. in the tyrrhenian stations (muddy bottoms, depth 200-2000 m) fl oariorugosus occurs rogether with other small-sized serpulids, í.e. metaoermilia multiuistata (philippi), sernioermilia agglutinata (marenzeller), fllogranula stellata (southward) and filogranula gracilis langerhans. in the early to middle pleistocene marls of lazzàro (barrier et al., 1996), h, aariorugosus co-occurs with f, stellata, s. agglutinata and 44. mwlticristata. boulders in these marls are encrusted by serpulids l(vitreotubus digeronimoi zlbrowius, newermilia falcigera souie), placostegus tridentatu.s (fabricius)1, isidid gorgonians and scleractinians. many species of these palaeocommunities are no longer part of the mediterranean fauna g.e. v digeronimoi and l/. falcigera). both the marl and the boulder faunas indicate a depth of about 600 m. the archi section (di geronimo et a1., in press), consisting of a pelitic early to middle pleistocene sequence, supplied several tubes ol h. aariorugosus. benthic assemblages point to a depth of 500-1000 m and are marked by an atlantic affinity. among serpulids n. falcigera and f. stellata are common. these ne'q/ records ol h. aariorugoszzs extend its previously known geographical distribution. ben-eliahu tr fiege (1996) reported living specimens of this species from the levant basin (s of crete, s-sv of cyprus, off israel, off egypt) and from the ionian sea (guif of taranto), depths 792-2009 m, in crevices of hard substrates, such as coal, slag, hardened sediment crusts, and on pteropod shells. plate 2 hyalopornatus aariorugosus ben-eliahu 6c fiege, 1996. scale bar : 100 pm (fig. i, 2, 4), 5 ;tm (fig. 3, 5, 6, 7). arrows indicating direction of growth. fig. 1 micromorphology of outer surface consisting of transverse flaps. lazzàro. fig.2 initial pan of the tube, smooth, no ornamentations. eolian islands, stn.7. fig. 3 transverse section of tube wall, outer surface continued into flaps. archi. fig. 4 crowded high transverse flaps, frayed out and turned. lazzàro. fig. 5 longitudinal section of tube. vall consisting of layers dipping towards tube mouth. eolian islands, stn. 27. fig. 6 tube microsrmcture. criss-cross arranged crystals of prismatic habitus, with a homogeneous microcrystalline structure. eolian islands, stn. 27. fig,7 outer tube surface, covered by an irregular film of very small crystais forming amorphous cryptocrystalline masses. eolian islands, stn. 27. pl. 2 tube morphology and structure of hyalopomatus oarìorugosus 135 i )t) r. sanfi.lippo therefore, h. oariorwgosus can be regarded as a mediterranean species, although a record from the atlantic (halfway between madeira and gibraltar) q" browius, pers. com.) should be noted, suggesting a possibly wider atlantic distribution. the new recent material was found on a variety of both smooth and irregular substrates, e.g. pteropod shells, oyster shells, pumice and slag. in fact, its small size particularly enables the species to colonize interstices and microhabitats. the new fossil material was generally cryptic, within small crevices of hard substrates or inside the calices of scleractinians. detached and fragmented tubes (especially disal ends), secondarily free, were found in muddy sediments. branching tubes are interpreted as a proof of scissiparity, as it had also been reported in other genera (filogranula langerhans, josephella caullery & mesnil, rhodopsis bush, spiraserpula regenhardt, filograna ber' keley; ten hove, 1979; ben-eliahu & ten hove, 1989; plllai,1993; pillai & ten hove, 1994). discussion. the finding and description of h. aariorugosus were carried out relatively late, owing to its ecological, biogeographical and stratigraphical range. its scanty occurrence could be also due to the inadequate routine methods: the very small-sized tubes are not found until the finest granulometric fractions of sediment (250 and 500 prm) are examined. the scarcity of deep-sea benthos in the mediterranean sea (bellan-santini et al., 1992) ís another reason why h. oariorugosus has seldom been recorded. records of other species of hyalopomatus from deep mediterranean 'waters are rare (zibrowius, 1969; 1977) and of uncertain specific attribution (ben-eliahu ec fiege, 1996). the other nominal species of the worldwide distributed genus hyalopornatws are morphologically close to each other, and also iive in bathyal and abyssal depths" h. variorwgosrzs is similar to some other hyalopomatus species (ben-eliahu & fiege, 1996). however, it markedly differs from all other species in its rugose sculpture. in most descriptions of recent serpuiid species no attention is paid to the tube structure. nevertheless, it may provide useful characters for identification. however, the knowledge of variability of these tube structures, for instance caused by external factors, is still insufficient for an optimum use. this might be corroborated by bornhold & milliman (1973) who mention a temperature-dependent mineralogy. in fact, the shape and size of the tube wall crystals seem not useful for generic nor for specific determination. these characters are not related to the size or to phylogenetic position of the worms; they seem to have evolved independently (nishi, 1993). a few occasional studies, especially when using sem techniques, have shown a considerable diversity in the tube wall structure, such as homogeneously opaque onejayered tubes (e.g. vermiliopsis saint-joseph, metavermilia bush, filogranula langerhans, filograna berkeley, protula risso) and twojayered tubes, the outer layer of which is transparent (ditrupa berkeley, laminatubus, regenhardt, galeolaria lamarck, neoaermilia day, pseudochitinopoma zibrowius). some other genera (placostegzs philippi, vitreotwbus zibrowius) are characterised by an entirely transparent tube. small serpulids such as filogranula, filograna, protis and josepbella possess tubes with thin and entirely opaque walls, as shown in this paper for h. aariorugosus. in other genera tube characters may not be constant within the genus. thus, shape and other morphological features are available for determination at specific level but, till noq it has not been possible to find tube attributes characterising the various genera. the same problem exists in hyalopornatus. although the tube of h. variorugosus has a morphological character distinctive at specific level, it is hardly possible to refer the tube to the genus hyalopomatus without characters of the worms themselves. a morphological tube feature, diagnostic at a generic level, apparently has not yet been found for the genus hyalopoma' tws. thus, tubes lacking the specific rugosity, could be misidentified {or protis or filograna species, wich have similar generic tube characters. in the eyes of a paleontologist, this may be caused by the strong systematic weight given to characters of the worm by neontologists. the paleontological approach is clearly based on different morphological features which may lead to different systematic views. the calcareous tubes are the fossilizable remains palaeontologists have access to whereas the classical taxonomy of serpulids by biologists is widely based on morphological features of the soft-bodied worm. in view of a better coordination between the classical "soft-body" taxonomy and the palaeontological approach, it is necessary to carefully study the macroand microstructures of the tubes of all recent serpulid species. in summary despite the recent interest in tube taxonomy, with promising results, further studies in this field are needed. aclenouledgements. i am grateful to h. zibrowius (stat. mar. endoume, marseille) and h.a. ten f{ove (instituut voor taxonomische zoòlogie, amsterdam) for their critical review of the manuscript; to s. di geronimo (istituto policattedra di oceanologia e paleoecologia, catania), p and e.s(/ knight-jones (department of zoology, swansea) for useful suggestions; to c. corselli (dipanimento di scienze della terra, milano) for loaning some material; o. torrisi (c.n.r., catania) and m. canzanella (dipanimento di scienze della terra, napoli) kindly supplied assistance in sem obserwetions. this paper is financially supponed by 40o/' (rosso) and 600/o (di geronimo) m.u.r.s.t. grants. tube rnorphology and structure of hyalopomatus ztariorugosus references r37 aliani s., bianchi c.n., de asmundis c., meloni r. 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"��� #��$-��� �# ����� �� ���� ����6�� %�� � �����7�� ���� $(��%( �� ������,-���3 � ���$ 7��� ��� � �� ��� �,�+ �# ��� %( �� ����3 � %( �� ��$(�(���� ������ �7�����3 ��� ���-��� %��� ���$(�� ��� %( �� ����� �� ���� 7���� 0��! ��� ��� -��,����$� �� #���(��� ��� ��6� 7� ��#�� *�4@ 8 �� )���������� �,! #��� +! , -���� � �!�@8 museum number skeleton element locality attribution by pasa (1947) new attribution v. 10924 m1 inf. sx viatelle felis leo cfr. spelaea homotherium sp. v. 10929 frag. humerus dx viatelle felis sp. panthera cf. fossilis v. 10949 4pm inf. viatelle felis sp. canis cf. c. mosbachensis v. 10920 caudal vertebra m.te zoppega felis leo spelaea? felidae ind. (dim. homotherium vel pan v. 4128 frag. ileum dx sentiero felis leo spelaea panthera cf. p. fossilis v. 4130 paw sx sentiero panthera pardus panthera cf. p. fossilis v. 10917 frag. scapholunar sx castello felis pardus cfr. antiqua felidae ind. v. 4463 lumbar vertebra castello felis leo spelaea felidae ind. v. 4464 lumbar vertebra castello felis leo spelaea felidae ind. v. 4465 frag. prox. tibia dx castello felis leo spelaea felidae ind. (dim. homotherium vel pan v. 4466 calcaneum sx castello felis leo spelaea felidae ind. (dim. homotherium vel pan v. 4468 lower jaw dx juv. castello felis leo spelaea panthera cf. p. fossilis ? v. 4469 2 phalanges castello felis leo spelaea felidae ind. v. 4461 frag. dist. femur sx m.te tenda felis pardus indet. v. 4462 scapholunar sx m.te tenda felis pardus panthera cf. p. fossilis "�%! � � ��� �# ��� �,� �-��� ��(���� �� ���� ,�,�� ��� ������ �� ��� �(��� /��� � �� ������ ���(���� �� *�����! ����� � ������ � "�� �,� �-�� *8�c4 �� �� �� �-,���� ��#� ,�7 �-,���� %� ��� (%���� ��� �� � ��� ��c� ��� ��8� ����� #���� ,�����$�� ��� �7� ����-���� 1��$! ��� "�%! 2! "�� 1��$! ��2� ��� ,��+�-�� �,�,����� �� ���$���� �����$(��� 7��� ��� ���� (������ #� ��� #�� ��c 7��� ,��� ������3 ��� ��7�� #� �� �� �� (��� 7������ ��� (,,�� ��� �� �� ����,�� ���$��(������� ������,��! "�� ���,����� �� ,��+�-���� ���$���� �����$(��� ��� �������� ������� ���� ���,������ ��,������! "�� ������ �,�,����� �� ��� 7��� ,�������� %(� �� � 7���� �� #�������0�! 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image resolution for improved printing quality. the pdf documents can be opened with acrobat and reader 5.0 and later.) /jpn /deu /ptb /dan /nld /esp /suo /ita /nor /sve /kor /chs /cht >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [595.000 842.000] >> setpagedevice in memoria carla rossi ronchetti (1,916-2000) il 29 luglio 20oo carla rossi ronchetti ci ha lasciato. lentamente. negli ultimi anni si era distaccata dalla rivista e dalla sua redazione, anche se tutte le volte che la si incontrava non mancava di inreressarsi a quella intrapresa cui per tanti anni aveva dedicato tempo ed energie. aveva ormai altri orizzonti. nata a ghiffa (nor.ara) nel 1916, laureata in scienze naturali a milano nel 1938 con una tesi sui brachiopodi del carbonifero de1 fezzan (libia), era presto divenuta prezioso collaboratore di a. desio. le prolungate assenze del direttore connesse con le spedizioni in karakorum e hindu kush, la portarono a costituire la presenza certa e costante in istituto, ad esserne il punto di riferimento. in cattedra dal 1962, assunse 1a direzione dell'istituto di paleontologia sino alla sua confluenza nel dipartimento di scienze della terra nel 1982. maestra atrenra agli a1tri, ma esigente, fondò la scuola di paleontologia di questa università. quanti geologi che si sono fatti onore nella professione e nella ricerca la ricordano, chi magari per i'impegno dell'esame, mitigato dalla nostalgia dei vent'anni, ma tutti con gratitudine, perché la preparazione paleontologica ottenuta rimase a lungo nel loro bagaglio cuitura1e. gli allievi a lei più vicini con affetto, perché il suo insegnamento superò i semplici aspetri scientifici, ma fu più profondo, scuola di vita. era schiva e riservata, e so che non avrebbe amato panegirici. la lista delle pubblicàzioni che qui segue, meglio di tante parole illustra la sua personalità ed i suoi interessi scientifici, dalla libia alle a1pi, dal karakorum all'hindu kush. i molluschi mesozoici e cenozoici costituirono il principale campo di artività. lr sua memoria con 1a revisione della collezione brocchi credo sia uno dei volumi più diffusi della paleontologia italiana. voglio invece spendere qualche parola sulla sua attività per la rivista italiana di paleontoiogia e stratigrafia, di cui carla rossi ronchetti ne fu il direttore scientifico per oltre 30 anni. penso che tutti i paleontologi ed una parte degli stratigrafi italiani di una certa età abbiano pubblicato almeno un articolo sulla rivista. flanno così avuto modo di sperimentare la precisione, carla rossi ronchetti left us on jwly 29, 2000. oper the pdst feut years she became less and less inook',ed in editing the rivista, though sbe continued to show a strong interest in the swbject to uthich she bad devoted her time and energt for so many years. sbe w,as gazing no,(e at different borizons. she r.uas born in ghtffo (nooara) in 1916 and received ber degree in natwral sciences from the university of milan in 1938, presenting a thesis on the brachiopods of the carboniferous of fezzan (lybia). she soon became an intalwable collaborator of prof, a. desio, director of the geolog institute. during prof. desio's prolonged expeditions to tlce karaleorum and tbe hindw kusb, she became a steady and notable presence at the institwte, after becoming full professor in 1962, she ,uas appointed director of the paleontolog, institute until the department of earth sciences was formed in the 1982. an attentive and demanding teacher, she was the founder and leader of the paleontologt scbool of the unit,ersity of mìlan. many presently successful geologists will remember her, some perhaps reitb a viz,id flashback to her exactingpaleontologt exam, since melloued by the nostalgia of their tzaenties, but all feeling grateful for the bnor.e;ledge that became an integral part of tbeir scìentific background.. her c/osest students remember ber zaìth affection, her teaching extendìng beyond the scientific, to deeper aspects of lfe experience. sbe zaas a reseroed person zoho would not bave liked a long euloglt. the following list of pwblications better reflects her persona/ity and scientijic interests, from lybia to tbe alps, from tbe karakorum to the hindw kush. mesozoic and caenozoic mollusbs formed tbe bwlb of her studies. tbe memoria containing ber rersision of the brocchi collection is probably one of the most popular volumes in italian paleontolog. i would like to address in more detail her influence on tbe rivista ltaliana di paleontologia e stratigrafia as scientific director for more tltan 3a years, i believe tbat all italian paleontologists and a swbstantial nwmber of stratigra?bers of a certain generation published at least one paper in the rìoìsta. they all experienced her precision, ber heen remarìes and the swbstantial improaement her critique memorte i'acurezza delie osservazioni e il grande contributo che ella dava a1 miglioramento dei manoscritti. la sua preparazione era messa a generosa disposizione. nel triennio in cui fu presidente della società paleontologicaltaliana, profuse ia sua capacità editoriale anche per il bollettino della società. per quesra artivirà ne venne poi nominata socio onorarío. la rivista rimase rra 1e sue occupazioni predilette, e noi che oggi ne conrinuiamo l'opera, amiamo ricordarla così, 1a "signora". brought to tbeir manuscripts. she zoas always generous luith her knoruled.ge and during her three year tenure as president of the società paleontologica ltaliana, sbe extencled her editorial experience to the bulletin of the society as well. for her commitment she r,tas rezaarded with the title of honoraw member. editorial work of the rivista was always one of her favorite interests, and rue of the editorial staff who are carrying on her legacy like to remember her just as she ruas known, that is "la signora". maurizio gaetani bibliografia rossi c. (1939) fossili carbonici del fezzan. ann. museo libico st. nat., v. 1, pp. 158-251. milano. rossi c. (1941) fossili miocenici del sottosuolo della gefara tripolina (libtt).ann. museo libico st. nat.,v.2 qgaa), pp.211-249, milano. rossi c. (1942) su alcuni fossili della dolomia principale dei dintorni di scutari (albania). ric. ital, paleont., v. ,18, n. 1, pp. 17 -21, miìano. rossi c. (1942) faunetta eocenica del serir tibesti (sahara libico). ann. museo libico st. nat., \.3 (1941), pp.971ob, milano. rossi c. (1942) molluschi paleogenici della sirtica.. ann. museo libico st. nat., v 3 (1941), pp. 109-193, milano. rossi c. ( 1943) rer.isione e critica della exogtra oterwegi r.on buch (maestrichtiano). riz.,. ital. paleont., v 49, n. 3-1, pp. 1-22, milano. rossi c. (1914) ricerche sul genere fistulana bruguiere. rlz. ital. paleont., v. 50, n. 1, pp. 3-10, milano. rossi ronchetti c. (19a5) diagnosi di forme nuove. rlv. itdl. paleont., r'. 51, n. 2-4, pp.38-41, milano. rossi ronchetti c. (1946) su1la presenza di argille fossilifere del pliocene nel sottosuolo di cadorago (como). àlz. ital. paleont., v. 52, pp. 1-8. milano. rossi ronchetti c. (1946) revisione del sottogenere veneri. cardia nel neocretacico della l1bia. rili.. ital. paleont.,v. 52, n. 1, pp. 1-35, miiano. rossi ronchetti c. (1947) revisione della fauna neocretacica della libia: gen. ptychocerar, sortogen. parapachydiscus. riv. ital. paleont., v 53, n. 2, pp.33-39, milano. rossi ronchetti c. (9a7) revisione della fauna neocretacica della libia, gen. bostrychoceras. riv. ital. paleont.,v. 53, n. 3, pp. 101-110, milano. rossi ronchetti c. (1947) appendice alla revisione dei gen. nautilus, indoceras e baculites del neocretacico della libia. rlo. ital. pdleont., v. 53, n. 4, pp. 153-155. milano. rossi ronchetti c. (1951-1956) i tipi della "conchiologia fossile subapennina" di g. brocchi. riz,. it. paleont. .ltrar., suppls. at voll. 57-62; pp. 1-343, milano. questi supplernenti sono poi raccolti nella memoria 5 della rivista ltaliana di paleontologia e stratigrafia,. ion lo stesso titolo e gli stessi numeri di pagina. rossi ronchetti c. & brena c. (1953) studi paleontologici sul lias del monte albenza (bergamo): brachiopodi dell'hettangiano. ri.o. ital. paleont. strat.,y.59, n. 3, pp. 111-134, milano. rossi ronchetti c. (1953) revisione del1a fauna neocretacica della libia: fa.m. ostreidae.ann. mus. libico st. ndt.,y. 4, pp. 111-19,{. roma. rossi ronchetti c. (1954) nuova fauna malacologica continentale associata a fauna marina nel miocene della libia. report of the algère international geological congress. part 2; p. 159. rossi ronchetti c. (i954) revisione critica del nemocardium cyprium (brocchi 181a). àlu. itd,l. pllleonr. strat.,t. 60, n. 1, pp.21-28, milano rossi ronchetti c. (1955) revisione della faunp neocretacica della libia. coralli: fam. smilotrochidae, carlophylliidae, parasmi/iidae, eupsammidae. rili.. itdl. paleont. strat.,v.61, n.3, pp. 101-123, milano. rossi ronchetti c. (1955) nuova segnalazione di elephas ttntiquus falc. nellavalle latina. rir:. ital. paleont. strat., v. 61, n. 3, pp. 127-135, milano. rossi ronchetti c. (1955) i foraminiferi del deposito elveziano di dogliani (cuneo). rir:. ital. paleont. strat. v. 61, n. 4,pp. 171-177, milano. rossi ronchetti c. (1955) sopra un singolare caratrere morfologico dell'l/rsus spelaeus ros. rlz,. ital. pdleont. strat.,v. 61, n. 1, pp. 181-181, milano. rossì ronchetti c., farioii mirelli a. (1958) rudiste e nerinee nella creta del pakistan nordoccidentalc. f.eports of the new mexico international geological congress. pp. 355-360. rossi ronchetti c. (1958) i mammiferi quaternari delle grotte della lombardia. ric. ital. paleont. strat.,v. 64, n. 4, pp. 303-345, milano. rossi ronchetti c. (1959) revisione della fauna neocreraclca della libia; fa:m. trochidae, columbellinidae, olioidae. ri'u. ital. paleont. strat.,v.65, n. 1, pp. 55-68, milano. rossi ronchetti c. & farioli mirelli a. (1959) rudists and nerineids of north-west pakistan creraceous. rio. ital. paleont. strat., v. 65, n.2, pp.91-96, milano. rossi ronchetti c. (1959) 11 trias in lombardia (studi geologici e paleontologici); i, lamellibranchi ladinici del gruppo delle grigne. rir,. ital. paleont. strat.,v.65, n. 4, pp.271-315, milano. memorî.e rossi ronchetti c. tl9b0) il trìas in lombardia lstudi geologici e paleontologici); ii, cefalopodi ladinici de1 gruppo del1e grigne. ria. ital. paleont. strat., v. 66, n. 1, pp. 1-49, milano. desio a., rossi ronchetti c. & invernizzi g. (1960) 11 giurassico dei dintorni di jefren in tripolitania. rir,. ital. paleont. strat., v. 66, n. 1, pp. 65-113, milano. desio a.er rossi ronchetti c. (1960) sul giurassico medio di garet el-bellaa (tripolitania) e sulla posizione stratigrafica della formazione di tacbal. riv. ital. paleont. strat., v. 66, n. 2, pp. 173-19a, milano. desio 4., rossi ronchetti c. & viganò p-l. (1960) su1la "tntionfi. j"l rils in tripolitania e nel sud-tunisino."'_--_b'_"* rio, ital. paleont. strat., v. 66, n. 3, pp. 273-318,mi1ano. rossi ronchetti c. & fantini sestini n. (1961) la fauna giurassica di karkar (afghanistan) . ri"r. ittll. paleont. strat.,v. 67, n.2, pp. 103-140, milano. rossi ronchetti c. & albanesi c. (1961) fossili cenomaniani del gebel tripolitano. rio. ital. paleont. strat.,v. 67, n. 3.pp. 251-3a7, milano. rossi ronchetti c. (1961) fossili cretacei di pull-i-khumri (afghanistan). rio. ital. paleont. strat., v. 67, n. 1, pp. 341-363, milano. rossi ronchetri c. (1962) variazioni e accrescimento relativo in exogtra or.,erwegi von buch del maestrichtiano libico. rio. ital. paleont. strat., v. 68, n. 2, pp. 193-243, milano. rossi ronchetti c. (1965) rudiste e nerinee del cretaceo di yasin (pakistan nord-occidentale). in: italian expeditions to the karakorum (k2) and hindu kush, scientific reports (ardito desio, leader) .part 4,v. 1, pp. 229272. l. brtll ed., leiden. rossi ronchetti c. (1965) sulla presenza di ofiuroidi nella formazione di gorno (trias superiore della val camonica (lombardia orientale) . rend. ist. lombardo sc. lett.,b,v.99, n. 1, pp. 51-63, milano. rossi ronchetti c. (1965) new contriburiorì to the knowledge of the jurassic fauna of karkar (afghanistan). in: fossils of north-east afghanistan. in: italian expeditions to the karakorum (k2) ) and hindu kush, scientific reports (ardito desio, leader). part 4, v.2, pp. 4374. e.l. brill, leiden. rossi ronchetti c. & allasinaz a.(1965) il trias in lombardia (studi geologici e paleontologici); xi, curionia, nuovo genere di lamellibranco eterodonte triassico. rlz. f . | ^ |ital. lhleont. strat., v.71. n.2, pp. 351-j93, mrlano. rossi ronchetti c. & allasinaz a. f1966) i1 trias in lombardia (studio geologici e paleontologici) ; xx, 'pseudomyoconcha' ntovo genere triassico di lamellibranchi eterodonti. rio, ital. paleont. strat.,v. v2, n.4. pp. 1083-1114, miiano. desio a., rossi ronchetti c. & pozzi r. (1966) osservazioni alla nota di p f. burollet 'remarques sur ìa strrtigraphie du jebel nefusa'. ria. ial. paleont. strat., v.72, n. 4, pp. 131,9-1322,milano. rossi ronchetti c. (1967) mollusks from the upper cretaceous at burji-la (baltistan, central asia). rlz. ial. paleont strat., v. 73, n.3, pp. 811-827, milano. rossi ronchetti c., fantini sestini n. & gaetani m. (1968) su una fauna liassica del dintorni di zandobbio (bergamo); nota preliminare. boll. soc. geol. |t,,v.87, n.4, pp. 743-746, roma. rossi ronchefii c. (197a) megafossils of some localities of north-eastern afghanistan. in: italian expeditions to the karakorum (k2) and hindu kush, scientific reports (ardito desio, leader), part 3, v. 3, pp. 509-520. e. j. brill, leiden. desio a., premoli silva l, rossi ronchefit c. (1977) on the cretaceous outcrop in the chumarkhan and laspur valleys, gilgit-chitral, n\f pakistan. rit:.. ital. paleont. strat., v. 83, n. 3, pp. 561-57 4, milano. senow 357..368 ������� ������� � ��� ������� �� � ��� ������ ��������� �� ��������� �� ���������� �� ��� ��� ���� � ���� ���� ������������� ���� ��������� ����� � ��������� �� ����� ��� � ��������� �� ��� �� ����� ��������� ��� ��� ���� ��� ������ � ����������� �������� ��������� �� � �������� ��������� !��� ������� ����������� ��� ���� �!�� �"� �##�$%$& '�(� ���� � )(�#�!�* � ������* �+�,������� ���$ "�%"�!$�( ,�"!�-� �% -!�, ��$ ��� ��� !$$%�,$ ���$ ���$ �##�$%$& '�(����� � ��!��$!� )(� #�!�� .$ "!�/$. � ������� �""����#� ��� /� �"��%0� (���!� ��$ "����/�"�$!��� ���(!$ ���� �!#��� ,* �� � ��� � )�.!�0�� � �!�#���%%� ���(#�� /� �"��%0* ��! ��� ,�"!�-� �% � �""(!��# �� �!�� �" ��%%�1 1��$! 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"���������� ;*�����! 9����� <� ���;b� �"# "#( << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /all /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /warning /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /syntheticboldness 1.00 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo true /preservecopypage true /preserveepsinfo true /preservehalftoneinfo false /preserveopicomments false 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(http://www.color.org) /pdfxtrapped /unknown /description << /fra /enu (use these settings to create pdf documents with higher image resolution for improved printing quality. the pdf documents can be opened with acrobat and reader 5.0 and later.) /jpn /deu /ptb /dan /nld /esp /suo /ita /nor /sve /kor /chs /cht >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [595.000 842.000] >> setpagedevice rivista italiana di paleontologia e stratigrafia volume 117 no. 1 1 pl. pp. 189-196 april 2011 regeneration and abnormality in benthic foraminifer rosalina leei: implications in reconstructing past salinity changes k.r. sujata, rajiv nigam, rajeev saraswat* & v. n. linshy received: october 25, 2010; accepted: january 18, 2011 micropaleontology lab, geological oceanography division, national institute of oceanography, dona paula, goa403 004, india * corresponding author. e-mail: rsaraswat@nio.org; phone: 91-(0) 832-2450489; fax: 91-(0) 832-2450602 key words: culture experiment, rosalina leei, salinity, dissolution, regeneration. abstract. a laboratory culture experiment has been conducted to assess the response of marginal marine benthic foraminifer rosalina leei to salinity and associated ph changes. live specimens of rosalina leei were subjected to a range (10-35 psu) of salinity. it was observed that hyposaline condition leads to dissolution of the calcareous tests. however, if the hyposaline condition persists only for a short period, then even after considerable dissolution, specimens were able to regenerate the dissolved part of the test. additionally, in all the specimens subjected to lower than normal salinity, the regenerated chambers were abnormal. the abnormalities included smaller or larger chambers and addition of new chambers in planes different than the normal plane of the tests. the regenerated specimens, however, attained a final size almost equal to that of control specimens that were not subjected to hyposaline conditions. the differential response of r. leei was attributed to decreased seawater ph under hyposaline condition. the findings can help understand the increased abundance of abnormal specimens under ecologically stressed environments. riassunto. è stato compiuto un esperimento con cultura in laboratorio per verificare la risposta delle variazioni in salinità e ph di un foraminifero bentonico di ambiente marginale, rosalina leei. esemplari vivi di rosalina leei sono stati sottoposti a variazioni di salinità, da 10 a 35 psu. si è osservato che le condizioni isoaline portano alla dissoluzione del guscio calcareo. tuttavia, se le condizioni isoaline persistono solo per un breve periodo, in seguito anche se vi è stata dissoluzione significativa, gli esemplari sono stati in grado di rigenerare la porzione di guscio dissolta. inoltre, in tutti gli esemplari portati a salinità inferiore al normale le camere rigenerate avevano caratteri anormali, quali dimensioni variabili o aggiunta di nuove camere su piani diversi dal normale piano di avvolgimento. gli esemplari rigenerati tuttavia raggiungevano una dimensione finale confrontabile con gli esemplari di controllo, non sottoposti alle variazioni di salinità. la risposta differenziale di r. leei è stata attribuita alla diminuzione del ph in condizioni ipoaline. questi risultati possono aiutare nel comprendere il significato dell’incremento di esemplari in condizioni di ambiente sottoposto a stress. introduction shallow marine waters are potential sites for high-resolution paleoclimatic studies, due to relatively higher sedimentation rate. huge aeolian and riverine flux and proximity to the land lead to high sedimentation rates in the shallow marine water, especially in regions where rivers meet the ocean. benthic foraminifera, preferentially marine microorganisms with a hard calcareous or agglutinated exoskeleton known as the test, are one of the most abundant groups of microorganisms in the shallow marine waters. they are very sensitive to the slightest changes taking place in the ambient environment and have preservation potential. therefore, the characteristics of benthic foraminifera have often been used to reconstruct past climatic changes from the shallow water regions (nigam et al. 1992, 1995; nigam & khare 1999; robinson & mcbride 2008; rossi & vaiani 2008; kemp et al. 2009). in order to decipher past climatic changes from the benthic foraminiferal characteristics, it is necessary to understand the factors affecting benthic foraminiferal distribution in the shallow marine waters. various factors including water depth, sediment type, organic matter flux, availability of oxygen, seawater temperature, salinity, distance from the river mouth, extent of bioturbation, etc. have been proposed to affect benthic foraminiferal distribution in the shallow water regions (mendes et al. 2004; bouchet et al. 2009; murray 2006; scott et al. 2001). out of these several biological and physico-chemical factors affecting the benthic foraminiferal distribution in the marginal marine areas, fresh water runoff related salinity changes are especially effective in shallow water areas (bouchet et al. 2009; 190 sujata k.r., nigam r., saraswat r. & linshy v.n. samir et al. 2003; hromic et al. 2006; horton & murray 2007; eichler et al. 2008; frezza & carboni 2009). changes in abundance, species assemblage, size and even the number of dissolved and distorted benthic foraminiferal tests have been assigned to various ecological parameters including salinity changes (boltovskoy et al. 1991). however, specific effect of salinity changes on benthic foraminifera is not well understood, as it is difficult to delineate the effect of a particular parameter, from the field studies. if such specific effects of salinity changes are known, it can help decipher changes in the monsoon intensity during the geologic past. we have monitored changes in salinity at a shallow marine location off goa for a period of two years (fig. 1). the location is directly affected by the fresh water influx from the nearby land during the monsoon season. additionally, we also recorded the changes in seawater ph at the same location. it was noted that the hyposaline waters are relatively less alkaline (fig. 2). we opined that the change in seawater ph as a result of decreasing salinity might be one of the causes for dissolved benthic foraminiferal tests reported from the shallow water regions. therefore, we decided to understand effect of seawater salinity related ph changes on benthic foraminifera. it is difficult to understand the effect of only salinity changes on benthic foraminifera, from field studies as many factors operate and simultaneously co-vary in the field. the changes in benthic foraminiferal characteristics might be the result of any one or a combination of a few of the physico-chemical parameters changing along with salinity. on the other hand, the laboratory culture studies can help to understand the foraminiferal response to varying seawater salinity and associated ph changes. the results of such studies can then be applied to the sediments collected from the field. however, so far, very limited attempts have been made to understand the effect of salinity changes on benthic foraminifera, in laboratory culture (bradshaw 1955, 1961; stouff et al. 1999a, 1999b; nigam et al. 2006, 2008). such studies on benthic foraminiferal species from indian waters can help to reconstruct past changes in indian monsoon. the present laboratory culture experiment was carried out to understand the effect of hyposaline water on benthic foraminiferal species rosalina leei and its capability, (if any), to overcome the adverse effects of hyposaline conditions. here our objective is to find out the effects of seawater salinity and associated changes (ph) on the hard part of the foraminifera. materials and methodology samples containing live specimens were collected from the waters off goa, where the salinity shows large seasonal fluctuation, varying from 11 psu to 36 psu (fig. 1). the location has two major estuaries namely zuari and mandovi draining huge amount of fresh water during southwest monsoon, which leads to large-scale changes in the seawater salinity within short time period. the floating as well as attached (to rocks submerged in seawater) algal material was collected and transferred to plastic tub having filtered seawater. the algal material was shaken vigorously to detach foraminifera. after vigorous shaking, complete material was transferred on to the sieves of size 1000 mm to get rid of extraneous material and subsequently over to 63 mm sieve to remove finer material including clay and silt. the >63 mm material was collected in beakers containing seawater and brought to the laboratory. live specimens of r. leei were picked under reflected light microscope. a total of six sets, each consisting of 6 specimens (total 36 fig. 1 location of sampling station. regeneration and abnormality in benthic foraminifer rosalina leei 191 specimens) of living r. leei were used for the experiment. out of the total six sets, two sets were maintained at 35 psu salinity, the same salinity as that at the time of collection of samples from the field. these were considered as control sets (cs-a, cs-b). the salinity of the control sets was maintained constant (35 psu) throughout the experiment. remaining four sets of specimens, considered as experiment set (es-1 to 4), were subjected to salinity varying from 10 to 35 psu. in each of the four experiment sets, salinity was gradually decreased from 35 psu to 10 psu, in steps of 3 and 2 psu, every second or third day. once the effects of lower than normal saline water became evident, the salinity was increased gradually again, till it reached to the initial level (35 psu, same as at the time of collection of material from the field). all the culture trays were maintained at 25° c temperature under incubators and under 12 hour light -12 hour dark condition throughout the experiment. in order to avoid evaporation, culture trays were wrapped in thin polythene film, immediately after changing the culture media. culture media was changed every alternate day. food was added in the form of diatom navicula sp. the change in salinity and ph of the media after adding food was negligible. the ph of culture media was routinely measured, before and after changing the media, with the help of labindia µp controlled ph analyser. the seawater of different salinity was prepared either by diluting the seawater with distilled water (to get seawater with salinity lower than that of the field) or by evaporating the seawater at 40° c temperature (to get seawater with salinity higher than that of the field). the salinity was measured with atago hand refractometer. growth, abnormality of test, if any, and pseudopodial activity were observed every alternate day. growth was estimated by measuring the average maximum diameter fig. 2 relationship between seawater salinity and ph as observed in the field. fig. 3 changes in average maximum growth of control and treatment set specimens in response to salinity changes. the control sets specimens showed continuous growth as they were maintained at constant salinity (35‰). in experimental sets, growth took place initially and subsequently when the salinity was lowered below 17‰, the tests started dissolving, as evident from the downward trend in growth. later on, when the salinity was increased again, the average growth also increased. the vertical lines are the standard deviation of growth as calculated from the growth between two consecutive readings. 192 sujata k.r., nigam r., saraswat r. & linshy v.n. of the specimen under inverted microscope (nikon eclipse te 2000u) connected to computer by using act 2u (auto camera tame to you / utility) software. though, occasionally, the tests were enclosed in a cyst made up of food provided to the specimens, the outline of the test was still visible. therefore, even in case of specimen with test enclosed in cyst, it was possible to measure the diameter and thus the growth of the specimen. results and discussion from the beginning of experiment till the salinity was lowered to 23 psu, all the specimens were very active showing visible signs of being alive. growth was observed in all the specimens of both controls as well as treatment sets till 23 psu salinity (fig. 3). once the salinity was lowered below 23 psu, it resulted in decreased pseudopodial activity and dissolution of foraminiferal tests (fig. 3). however, the effect was not uniform on all the specimens. test of 8 specimens (2 in es-1, 3 in es-2, 1 in es-3 and 2 in es-4) started dissolving once the salinity was decreased to 20 psu. a further decrease in salinity to 17 psu resulted in dissolution in 9 more specimens (1 each in es-1 and 2, 5 in es-3 and 2 in es-4). test of additional 6 specimens (3 in es-1, 2 in es-2 and 1 in es-3) started dissolving once the salinity was further lowered to 15 psu. one specimen showed visible signs of dissolution only when the salinity was decreased to 13 psu. all the specimens continued to dissolve till they were subjected to seawater of 10 psu salinity. the specimens were alive as evident from the collection of food but the pseudopodial activity was very limited. the dissolution started from the last chamber, however, only partial dissolution of chambers was noted. outline of almost all the dissolved chambers remained visible (pl. 1). although the response of specimens varied when the salinity was lowered, all the specimens started rebuilding the test, once the salinity was increased from 10 psu to 13 psu. the increase in salinity not only resulted in recalcification of partially dissolved chambers and addition of new chambers but also in increased pseudopodial activity. specimens completely regenerated the dissolved chambers and attained the size almost equal to or slightly lower than that of the specimens in control set (fig. 3). at the end of the experiment an average growth of ~357 µm (varying from 328-394 µm) was noted in all the six sets. the final size attained by the specimens in both the control and treatment sets was comparable. the most interesting feature was abnormalities in the regenerated chambers (chambers added after dissolution). all the 24 specimens subjected to hyposaline seawater developed abnormalities after regeneration (pl. 1), whereas only one out of the 12 specimens in the control sets developed abnormal test in the course of experiment. the abnormality in case of the specimen from the control sets was very minor with slightly bigger chamber and without any change in the plane of orientation of newly added chambers. abnormalities in the case of treatment set specimens included bigger than normal chamber, disoriented chambers and differently shaped chambers. not all the fig. 4 relationship between seawater salinity and ph as observed for the media prepared in the laboratory. the ph decreased with the lowering of salinity. plate 1 dissolution (a-f) and abnormalities (g-n) in rosalina leei specimens subjected to hyposaline seawater. in most of the specimens, hyposaline seawater resulted in partial dissolution of chambers (a-c) while in others (d-f), last few chambers got completely dissolved (arrows indicate dissolution in the specimens). almost all of the specimens regenerated the dissolved chambers, but became abnormal (g-n). abnormalities included addition of larger or smaller chambers, in planes others than the normal plane of addition of chambers. regeneration and abnormality in benthic foraminifer rosalina leei 193 194 sujata k.r., nigam r., saraswat r. & linshy v.n. specimens subjected to hyposaline conditions could recover; three specimens died during the experiment. the dissolution of tests probably took place because of decreased ph of seawater, associated with the decreased salinity. it was observed that ph decreased with the decreasing salinity, both in case of media prepared in the laboratory (fig. 4) as well as in case of observations made in the field as part of this study (fig. 2) as well as previous reports (brown et al. 1999). earlier boltovskoy & wright (1976) noted that ph lower than 7.8 induces dissolution of calcareous tests, whereas bradshaw (1961) and angell (1967) observed dissolution of the selected foraminiferal species only under acidic ph conditions. similarly, stouff et al. (1999b) also reported that dissolution in ammonia beccarii started when the seawater ph decreased below 5. however, the dissolution in case of r. leei started at ph lower than 7.5 (20 psu salinity), different than the earlier observations. the dissolution in the present study started well above the ph value (7.0) reported by le cadre et al. (2003) for ammonia beccarii. though, le cadre et al. (2003) postulated that dissolution will start below 7.5, but the specimens were only subjected to seawater ph 7.5 and 7.0. no dissolution was observed in case of specimens subjected to seawater ph 7.5 and even the specimens subjected to ph 7.0 do not show any signs of dissolution till five days. the dissolution in case of r. leei at significantly higher ph value than that for ammonia beccarii probably arises due to the comparatively thinner tests of r. leei. this study helped to refine the seawater ph value at which dissolution begins in r. leei, as against the large range proposed by earlier workers (bradshaw 1961; le cadre et al. 2003). the study shows that the dissolution of calcareous foraminiferal tests can take place at much more alkaline seawater ph than reported before. another possible reason for the dissolution of the tests might be the decrease in the concentration of the calcium carbonate at lower salinities as it has been cited as a potential cause for the dissolution of foraminiferal tests in numerous paleoclimatic reconstruction studies (de rijk 1995; murray & alve 1999; kimoto et al. 2003). the change in carbonate ion concentration might have occurred due to the preparation of hyposaline water by adding distilled water. however, unfortunately we don’t have any measured values for the calcium carbonate concentration of the differently saline media. the series of stages followed during dissolution under low salinity in the present experiment (pl. 1), starting with tests becoming slightly opaque and then dissolution starting from last chamber, has also been reported by le cadre et al. (2003), while observing the effects of low ph on ammonia beccarii. opaque tests have also been reported from field. however, the dissolution does not progress chamber-by-chamber, except a last few chambers. after near complete dissolution of the last one or two chambers, almost all the chambers of the last whorl were equally affected by the dissolution. though the differences in the degree of resistance to dissolution of individual tests were also noted, it probably shows the slight differential individual response. the recalcification of dissolved and addition of new chambers after increasing the salinity, resulted because of revival of favorable conditions. this recalcification confirms the views expressed by boltovskoy & wright (1976) that foraminifera can repair and/or regenerate their tests after damage arising out of either physical injury or chemical effects. decalcified living specimens, when cultured under favorable conditions, showed pseudopodial emissions and recalcification of chambers leading to morphological abnormalities (stouff et al. 1999b; le cadre et al. 2003). the results are significant in view of the reported presence of abnormal tests near river mouths, where the possibility of breaking and later abnormal regeneration has been expressed (vilela 2003). geslin et al. (2002) also reported abnormal tests from areas exposed to periodic salinity changes and acidification. however, the abnormal tests were reported from areas subjected to hypersaline waters, whereas in our study only those specimens which were subjected to hyposaline water developed abnormalities during recalcification after partial dissolution of the tests. the rate of regeneration of test was comparable with that of the dissolution. the thickening of wall of decalcified chambers took place contemporaneously with the addition of new chambers. angell (1967) also reported similar observations for rosalina floridana, where a hydrochloric acid decalcified specimen recovered once transferred to normal saline water. he explained this recalcification as a result of addition of calcium carbonate layer to the whole test, every time a new chamber is formed, rather than any healing mechanism. however, such regeneration of dissolved chambers by the specimens, not through any specific mechanism to recover damage induced to the chambers, rather as a result of normal process of addition of calcitic layer to the whole test in certain foraminiferal species, every time a new chamber is added, was not noticed in the present experiment. another significant finding of the present experiment was the abnormality in the chambers added during the regeneration of the dissolved tests. the added chambers were abnormally oriented away from the normal plane of orientation of the earlier chambers formed under normal conditions (pl. 1). these findings can help explain the increased abundance of abnormal specimens in areas subjected to short-term ecological variations, especially salinity variations (murray 1989). regeneration and abnormality in benthic foraminifer rosalina leei 195 this abnormality in the chambers added during regeneration of the tests probably arises because of either physiological or structural damage during dissolution of the tests, which the r. leei specimens are unable to recuperate. geslin et al. (2002) also suggested that regeneration after damage of tests may also induce high proportion of abnormalities in environments with strong hydrodynamics. interestingly, the size of specimens subjected to hyposaline condition was as big as that of control specimens, even after considerable dissolution of the test. it appears that the physiological activity of the specimens increased once they got favorable conditions after severe damage under hyposaline seawater. based on the laboratory culture experiment carried out to understand the response of inner shelf benthic foraminifer rosalina leei to short-term decrease in salinity we conclude that lower than normal saline water leads to partial dissolution of the tests. we further conclude that although r. leei is capable of recovering from short-term low salinity changes, the signatures of hyposaline conditions are retained by the test in the form of visible morphological abnormalities. the results can help in understanding the cause of anomalously high abundance of abnormal specimens as well as in changes in abundance of certain species in the samples collected from marginal marine areas in the field. acknowledgements. the authors are thankful to the director, national institute of oceanography for permission and support. authors are thankful to prof. maurizio gaetani and prof. emmaneul le geslin for comments and suggestions to improve the manuscript. skr and vnl are thankful to the council of scientific and industrial research, new delhi, for awarding the senior research fellowships. the work was carried out as part of department of science and technology, government of india funded project on laboratory culturing of foraminifera for paleoclimatic studies. references angell r.w. 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"� �-��#�% )��/��" %�"�� �#����� �"� %5 ����������& �%%#��"$ ���� �( �!��� �"�������"� �� �!� �"$���"� �����"� ��-��#� �!�� �!�� (�� /�## /��! �!� ����"� %�"�� �#����� +��)& 5,& �� ���*�!� -���f + @1<), ������)�� �!� �2��"�� %�"�� ����!��� (��� �!� ����#� �#�������"� �( ?��"�� �� �!� %��)�)#� �"������ �( )��! %5 �#����� �"� %5 ���( ����"�� �� �� ���)�"�� �!������������� �( �!� �/� �%�����4 %�"�� ����!��� �� �!����������� )� �!� ������������� �� ������� �5 6 &���� ,5:@� %����" ��� ������������ � �#� *��� &"��������� 7������� ���� ������� :@5 total length proximal width proximal depth distal width distal depth width in the middle humerus falco cherrug ing. a 72 84.6 16.2 9.7 7.6 falco cherrug ing. a 76 99.7 7.9 18.9 11.1 9.2 falco cherrug ing. a. 1129 82.8 15.7 10.1 7.5 falco cherrug ing. a. 1133 18.5 10.9 falco cherrug ing. a. 1134 83.1 20.3 17.2 10.4 7.0 falco cherrug ing. a. 1135 84.9 20.7 10.0 16.6 9.3 7.2 falco cherrug ing. a. 1182 16.0 falco cherrug ing. a. 1221 87.3 19.5 11.0 17.1 10.2 8.3 falco cherrug – recent 87.1 99.3 (n = 9) 19.3 22.5 (n = 7) 10.0 11.1 (n = 3) 16.5 18.9 (n = 9) 9.4 10.4 (n = 3) 7.4 8.3 (n = 9) falco rusticolus – recent 96.5 110.0 (n = 7) 19.4 24.8 (n = 9) 10.1 12.6 (n = 8) 18.2 21.0 (n = 8) 10.8 12.1 (n = 6) 8.3 9.6 (n = 8) falco biarmicus – recent 74.8 84.1 (n = 6) 17.5 18.7 (n = 6) 9.1 9.2 (n = 2) 13.3 15.7 (n = 6) 8.7 9.4 (n = 2) 6.2 7.3 (n = 6) falco peregrinus – recent 66.8 91.4 (n = 21) 16.5 21.3 (n = 20) 7.2 11.6 (n = 20) 13.7 18.0 (n = 20) 8.0 10.5 (n = 17) 6.7 9.0 (n = 21) ulna falco cherrug ing. a. 1 117.0 13.5 10.7 9.7 6.8 falco cherrug ing. a. 2 11.3 10.4 falco cherrug ing. a. 3 10.9 falco cherrug ing. a. 9 10.9 8.7 falco cherrug ing. a. 11 10.3\ 8.8 falco cherrug ing. a. 12 10.0 8.0 falco cherrug ing. a. 14 9.1 7.9 falco cherrug ing. a. 1186 9.7 8.0 falco cherrug – recent 100.2 117.7 (n = 8) 11.2 13.3 (n = 8) 10.3 12.2 (n = 8) 9.6 12.6 (n = 8) 8.6 9.1 (n = 3) 5.7 7.5 (n = 8) falco rusticolus – recent 107.7 124.7 (n = 8) 12.6 14.3 (n = 8) 11.6 13.0 (n = 8) 10.4 13.7 (n = 8) 9.1 10.6 (n = 6) 6.4 7.6 (n = 6) falco biarmicus – recent 87.3 98.1 (n = 6) 9.7 10.9 (n = 6) 9.0 10.9 (n = 6) 9.5 10.8 (n = 5) 7.8 8.2 (n = 2) 5.2 6.0 (n = 6) falco peregrinus – recent 77.0 105.8 (n = 20) 9.3 12.3 (n = 18) 9.4 12.4 (n = 15) 8.0 11.2 (n = 15) 7.1 – 9.3 (n = 15) 4.8 6.7 (n = 21) carpometacarpus falco cherrug ing. a. 87 60.5 15.3 6.8 5.7 4.6 falco cherrug ing. a. 89 7.5 falco cherrug ing. a. 90 69.5 7.6 11.9 7.3 5.0 falco cherrug ing. a. 91 58.5 15.4 6.6 9.7 5.8 falco cherrug ing. a. 94 56.9 14.8 6.6 9.6 7.2 4.1 falco cherrug ing. a. 1121 15.0 6.7 falco cherrug ing. a. 1122 68.8 17.2 7.4 11.1 7.1 4.7 falco cherrug ing. a. 1123 17.2 7.5 falco cherrug ing. a. 1137 6.3 falco cherrug – recent 59.4 69.3 (n = 9) 14.8 17.7 (n = 8) 5.6 5.8 (n = 2) 9.0 11.0 (n = 8) 6.5 7.5 (n = 2) 3.7 4.4 (n = 9) falco rusticolus – recent 66.2 72.5 (n = 7) 17.4 19.6 (n = 7) 6.3 7.7 (n = 6) 10.8 12.0 (n = 8) 6.8 8.0 (n = 6) 4.2 5.1 (n = 8) falco biarmicus – recent 52.5 58.7 (n = 6) 13.5 15.5 (n = 6) 5.7 5.8 (n = 2) 9.0 9.9 (n = 6) 5.8 6.0 (n = 2) 3.3 4.2 (n = 6) falco peregrinus – recent 48.0 65.2 (n = 25) 13.4 17.3 (n = 19) 5.0 7.3 (n = 18) 8.6 11.5 (n = 19) 5.1 6.5 (n = 19) 3.5 4.7 (n = 25) ��)& * ���� ����"�� �( �!� (���#��) )�"�� �( ���� �#�������"� %�"�� �#����� (��� �"$���"�� ���%���� /��! �!��� �( ����"� %�"�� �#������ %5 �������"��� %5 ��� ���� �"� %5 ���������� +���� (��� ��#�� @;=� @; �� @; ) �"� �� ����� �& �� ���*�!� -���f,& ������� �5 6 &���� ,5:@: total length proximal width proximal depth distal width distal depth width in the middle femur falco cherrug ing. a. 68 13.4 8.1 13.8 6.5 falco cherrug ing. a. 69 76.5 15.6 11.9 7.4 falco cherrug ing. a. 1124 13.2 8.6 falco cherrug ing. a. 1131 13.8 8.4 13.7 11.9 6.2 falco cherrug ing. a. 1132 77.5 15.9 13.4 7.2 falco cherrug – recent 69.3 77.1 (n = 7) 13.3 14.5 (n = 7) 8.3 10.6 (n = 6) 13.7 15.5 (n = 7) 10.3 11.9 (n = 6) 6.2 7.4 (n = 7) falco rusticolus – recent 82.0 92.3 (n = 9) 15.3 18.3 (n = 9) 9.4 11.3 (n = 9) 15.8 19.0 (n = 9) 12.4 14.0 (n = 9) 7.2 9.1 (n = 9) falco biarmicus – recent 60.5 66.6 (n = 6) 11.2 12.6 (n = 6) 7.7 8.4 (n = 5) 11.3 13.3 (n = 6) 8.6 11.2 (n = 6) 5.4 6.5 (n = 6) falco peregrinus – recent 55.3 73.2 (n = 27) 11.2 15.0 (n = 25) 6.5 9.3 (n = 25) 11.2 14.7 (n = 25) 9.2 12.6 (n = 25) 5.3 7.4 (n = 27) tibiotarsus falco cherrug ing. a. 24 12.8 8.8 falco cherrug ing. a. 43 92.6 14.3 10.5 6.9 falco cherrug ing. a. 46 14.9 9.7 falco cherrug ing. a. 55 12.2 12.8 falco cherrug ing. a. 1125 9.1 falco cherrug ing. a. 1225 96.9 14.9 12.1 15.1 9.7 6.7 falco cherrug – recent 88.0 97.9 (n = 8) 12.0 12.2 (n = 2) 12.8 13.8 (n = 2) 12.1 15.3 (n = 8) 9.1 9.7 (n = 3) 4.3 6.3 (n = 8) falco rusticolus – recent 99.5 110.5 (n = 8) 13.2 15.0 (n = 6) 13.9 16.1 (n = 6) 13.8 17.2 (n = 8) 9.8 12.1 (n = 6) 5.3 7.2 (n = 6) falco biarmicus – recent 79.2 87.0 (n = 6) 10.3 12.1 (n = 5) 11.6 12.6 (n = 2) 11.0 12.1 (n = 4) 6.4 8.5 (n = 2) 4.0 5.4 (n = 6) falco peregrinus – recent 71.5 92.7 (n = 21) 9.5 15.0 (n = 19) 9.2 14.2 (n = 19) 10.2 14.5 (n = 19) 7.1 9.9 (n = 19) 4.1 6.7 (n = 21) tarsometatarsus falco cherrug ing. a. 21 15.2 10.1 falco cherrug ing. a. 22 13.8 9.3 falco cherrug ing. a. 29 14.7 falco cherrug ing. a. 31 13.2 10.3 falco cherrug ing. a. 1223 54.4 13.2 14.0 9.2 6.8 falco cherrug ing. a. 1124 58.3 14.6 15.8 10.2 6.2 falco cherrug ing. a. 1226 55.4 13.7 13.6 9.2 5.8 falco cherrug ing. a. 1227 52.9 13.2 10.1 14.1 9.3 6.3 falco cherrug ing. a. 1228 13.8 9.3 falco cherrug – recent 53.8 60.5 (n = 9) 13.3 15.9 (n = 9) 10.0 11.2 (n = 3) 13.0 15.8 (n = 8) 9.3 9.5 (n = 2) 5.0 6.3 (n = 9) falco rusticolus – recent 60.4 62.3 (n = 8) 14.6 18.8 (n = 8) 11.3 13.6 (n = 6) 15.1 17.3 (n = 8) 9.7 12.0 (n = 6) 5.3 7.6 (n = 8) falco biarmicus – recent 51.1 53.0 (n = 5) 11.5 13.2 (n = 6) 9.9 10.0 (n = 2) 10.7 12.9 (n = 4) 8.8 4.6 5.8 (n = 6) falco peregrinus – recent 44.4 56.0 (n = 23) 11.0 14.4 (n = 21) 8.3 11.0 (n = 21) 10.8 15.0 (n = 21) 7.4 10.2 (n = 15) 4.3 6.2 (n = 21) ��)& � * ���� ����"�� �( �!� !�"�#��) )�"�� �( ���� �#�������"� %�"�� �#����� (��� �"$���"�� ���%���� /��! 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���� +����� �,� �/� #"�� +����� �,� �"� (�� � +����� �,� �/� ��)������� +����� �,� (�-� �������������� + #���� �� : #���� �,& ����!�%$��� ��& ��"%�!�����& �!� $�" � �"��( ����� �� ��(�"���-�#� #��$�� �!�" �!� $�" � &����' �" ���! %���*���"��# �#���"� �"� ��" �#�� )� ��%������ (��� �!� ������� �5 6 &���� ,5:@a ?�$& 1 * ������� ���$���� �( �!� ���* � ����"�� �( ������������� � �( ����"� %�"�� ����������� %5 ��� ����� %5 �������"��� %5 �#����� �"� �!� (����# %5 �#��( ��� (��� �!� ���� �#�����* ��"� �( �"$���"� +���� (��� ��#�� @;=� @; �� @; ) �"� �� ����� �& �� ���*�!� * -���f,& falco cherrug ingarano falco cherrug falco peregrinus min. max. x min. max. x min. max. x humerus total length / distal width 5.10 5.43 5.23 5.12 5.34 5.25 4.86 5.08 4.96 carpometacarpus total length / proximal width 3.80 4.00 3.89 3.82 4.13 3.94 3.61 3.376 3.69 tibiotarsus total length / distal width 6.41 6.47 6.44 6.31 6.78 6.48 6.39 7.15 6.69 tarsometatarsus total length / proximal width 3.99 4.12 4.04 3.62 4.04 3.86 3.71 3.90 3.78 ��)& 5 * ������ )��/��" ��((���"� ���� ����"�� �( -���� � #�"$ )�"�� �( ���� �#�����* ��"� %�"�� �#����� (��� �"* $���"�� ���%���� /��! 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spinosum kozur & mostler x x x x spinotriassocampe carnica kozur & mostler x x hinedorcus koeseyahyaensis tekin n. sp. x x x hinedorcus sp. a x x hinedorcus ? sp. b x picapora robusta kozur and mostler x x x x picapora sp. a x silicarmiger curvatus (kozur & mostler) x x x katroma ? ornata tekin n. sp. x x katroma ? proba tekin n. sp. x x x katroma ? tunoglui tekin n. sp. x x x syringocapsa firma tekin n. sp. x x x x syringocapsa nuda tekin n. sp. x x x praeprotunuma sp. a x nabolella parvispinosa (kozur & mock) x x x x nabolella sp. a x x annulopoulpus parviaperturus (kozur & mostler) x x x parapoulpus sp. a x poulpus piabyx de wever x x x x x veghia goestlingensis kozur & mostler x x x x veghia ? sp. a x pseudosaturniforma carnica kozur & mostler x x x x x sanfilippoella carterae tekin n. sp. x x x sanfilippoella tortilis kozur & mostler x x x x sanfilippoella sp. a x sanfilippoella sp. b x castrum sp. aff. c. perornatum blome x x castrum sp. a x castrum sp. b x x x trialatus ? panus (de wever) x x x triassocyrtium sp. aff. t. hamatum kozur & mostler x nassellaria gen. and sp. indet. a x nassellaria gen. and sp. indet. b x x d�1# ? 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/qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /colorimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000coloracsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000colorimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasgrayimages false /downsamplegrayimages true /grayimagedownsampletype /bicubic /grayimageresolution 300 /grayimagedepth -1 /grayimagedownsamplethreshold 1.50000 /encodegrayimages true /grayimagefilter /dctencode /autofiltergrayimages true /grayimageautofilterstrategy /jpeg /grayacsimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /grayimagedict << /qfactor 0.15 /hsamples [1 1 1 1] /vsamples [1 1 1 1] >> /jpeg2000grayacsimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /jpeg2000grayimagedict << /tilewidth 256 /tileheight 256 /quality 30 >> /antialiasmonoimages false /downsamplemonoimages true /monoimagedownsampletype /bicubic /monoimageresolution 1200 /monoimagedepth -1 /monoimagedownsamplethreshold 1.50000 /encodemonoimages true /monoimagefilter /ccittfaxencode /monoimagedict << /k -1 >> /allowpsxobjects false /pdfx1acheck false /pdfx3check false /pdfxcompliantpdfonly false /pdfxnotrimboxerror true /pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /fra /enu (use these settings to create pdf documents with higher image resolution for improved printing quality. the pdf documents can be opened with acrobat and reader 5.0 and later.) /jpn /deu /ptb /dan /nld /esp /suo /ita /nor /sve /kor /chs /cht >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [595.000 842.000] >> setpagedevice rivista italiana di paleontologia e stratigrafia volume 117 no. 1 pp. 149-160 april 2011 new pterosaur specimens from the kem kem beds (upper cretaceous, cenomanian) of morocco taissa rodrigues 1, alexander w. a. kellner 1, bryn j. mader 2 & dale a. russell 3 received: october 11, 2010; accepted: february 1st, 2011 1 setor de paleovertebrados, dept. geologia e paleontologia, museu nacional/ufrj. quinta da boa vista s/n, são cristóvão, 20940-040, rio de janeiro, brazil. e-mail: taissa.rodrigues@gmail.com, kellner@mn.ufrj.br 2 queensborough community college, 222-05 56th avenue bayside, ny 11364, usa. e-mail: bjmader@email.msn.com 3 north carolina state university, dept. of marine, earth and atmospheric sciences, jordan hall 5145, box 8208, raleigh, nc 27695, usa. e-mail: dale_russell@ncsu.edu key words: pterosauria, azhdarchidae, morocco, cretaceous, cenomanian, kem kem beds. abstract. although pterosaurs from africa are still rare, in recent years several specimens have been described from the kem kem beds (upper cretaceous, cenomanian) of morocco. here we describe four additional specimens from this informal lithostratigraphic unit: a jaw fragment, two mid-cervical vertebrae, and a humerus. all these specimens show three-dimensional preservation, differing much from the flat condition found in most pterosaur material. the vertebrae are particularly well preserved, and allow accurate observations on the pneumatization of the neural arch. based on comparable material, we show that at least two edentulous pterosaur species were present in this informal lithostratigraphic unit, thus adding to the growing evidence of considerable pterosaur diversity in northwestern africa during the “middle” cretaceous. so far, the kem kem beds have the most diverse pterosaur fauna in this continent, with the presence of anhanguerids, azhdarchids, pteranodontids, and tapejarids. riassunto. sebbene gli pterosauri rinvenuti in africa siano ancora rari, negli ultimi anni diversi esemplari sono stati descritti dai cosiddetti kem kem beds del marocco (cretacico superiore, cenomaniano). nell’ articolo descriviamo quattro nuovi reperti provenienti da questa unità litostratigrafica informale: un frammento di mandibola, due vertebre medio-cervicali e un omero. tutti questi esemplari offrono una conservazione tridimensionale, differendo in questo dalle condizioni di appiattimento proprie di molti reperti di pterosauri. le vertebre sono particolarmente ben conservate e consentono una accurata osservazione della pneumatizzazione dell’arco neurale. sulla base di materiale di confronto noi mostriamo che almeno due specie edentule di pterosauri sono presenti in questa unità litostratigrafica informale. ciò incrementa l’ evidenza di una considerevole diversità tra gli pterosauri che popolarono l’africa nordoccidentale durante il “cretacico medio’’. i kem kem beds contengono la fauna a pterosauri più diversificata in questo continente, con la presenza di anhangueridi, azhdarchidi, pteranodontidi e tapejaridi. introduction african pterosaurs are still poorly known, limited to relatively few incomplete skeletons and isolated remains (swinton 1948; monteillet et al. 1982; unwin & heinrich 1999; benton et al. 2000; dalla vecchia et al. 2001; blackburn 2002; dal sasso & pasini 2003; ntamak-nida et al. 2006; blackbeard & yates 2007; kellner et al. 2007; costa & kellner 2009; ibrahim et al. 2010). morocco represents several of the recent finds of pterosaur fossils. sigogneau-russell et al. (1998) first reported on some pterosaur material from a lower cretaceous calcareous lens in the locality of anoual. these specimens, nearly 300 isolated teeth, were described by knoll (2000), who identified them as belonging to ornithocheirids and gnathosaurines, the latter questioned by rodrigues & kellner (2010) who pointed out that at least some of the gnathosaurin teeth could actually belong to the anhangueridae or closely related taxa. another moroccan pterosaur occurrence is the sequence of cervical vertebrae from the maastrichtian of the oulad abdoun phosphatic basin that represents the azhdarchid phosphatodraco mauritanicus pereda-suberbiola, bardet, jouve, iarochène, bouya & amaghzaz, 2003 (see pereda-suberbiola et al. 2003; kellner 2010). notwithstanding these occurrences, most pterosaur material comes from the cenomanian ifezouane and aoufous formations (informally known as the kem kem beds) (sereno et al. 1996; cavin et al. 2010). 150 rodrigues t., kellner a.w.a., mader b.j. & russell d.a. so far, these deposits have yielded isolated anhanguerid teeth (kellner & mader 1997; wellnhofer & buffetaut 1999), isolated azhdarchid vertebrae (kellner & mader 1996; rodrigues et al. 2006; ibrahim et al. 2010) and fragmentary jaws referred to the anhangueridae, azhdarchidae, pteranodontidae, and tapejaridae (mader & kellner 1999; wellnhofer & buffetaut 1999; kellner et al. 2007; ibrahim et al. 2010), including two described species: the anhanguerid siroccopteryx moroccensis mader & kellner, 1999 (see mader & kellner 1999; rodrigues & kellner 2008; but see ibrahim et al. 2010 for a different classification of this species) and the azhdarchid alanqa saharica ibrahim, unwin, martill, baidder & zouhri, 2010 (see ibrahim et al. 2010). the pterosaur specimens, although relatively common, tend to be fragmentary due to the high energy depositional environment of both these units (cavin et al. 2010). here we describe four specimens that have been only briefly mentioned in the literature (kellner & mader 1996; rodrigues et al. 2006), adding to the better understanding of the moroccan pterosaur fauna. institutional abbreviations: bsp, bayerische staatssammlung für paläontologie und geologie, munich, germany; ccmge, museum of central scientific research institute for geological exploration, university of st. petersburg, saint petersburg, russia; cmn, canadian museum of nature, ottawa, canada; fsac, faculté des sciences ain chock, université hassan ii, casablanca, morocco; linhm, long island natural history museum, new york city, usa; mdm, mifune dinosaur museum, mifune, japan; mguv, museo del departamento de geología / universidad de valencia, valencia, spain; mn, museu nacional / universidade federal do rio de janeiro, rio de janeiro, brazil; mtm, magyar természettudományi múzeum, budapest, hungary; nhmuk, natural history museum, london, england; smnk, staatliches museum für naturkunde karlsruhe, karlsruhe, germany; and tmm, texas memorial museum, austin, usa. anatomical abbreviations: cap, capitulum or radial condyle of the humerus; co, cotyle; con, condyle; dpc, deltopectoral crest of the humerus; ect, ectepicondyle of the humerus; ent, entepicondyle of the humerus; fo, foramen; fopn, foramen pneumaticum; hyp, hypapophysis; mus, muscle scar; nc, neural canal; ns, neural spine; poex, postexapophysis; poz, postzygapophysis; prz, prezygapophysis; ?ri, possible vestigial cervical rib; rid, ridge; sul, sulcus; tro, trochlea or ulnar condyle of the humerus; tub, tubercle; and ulc, ulnar crest of the humerus. geological settings the so-called ‘continental intercalaire’ comprises extensive deltaic and brackish deposits in the lower cretaceous of north africa (kilian 1931; lapparent 1960; cavin et al. 2010). in morocco, it is known informally as the kem kem beds (sereno et al. 1996), whose fauna, paleoenvironment and paleogeography were recently reviewed by cavin et al. (2010). the age of the kem kem beds is not easy to be established due to the lack of marine intercalations, but by the constraint of the overlying akrabou formation, whose lower part contains ammonoids characteristic of the lower part of the upper cenomanian, and by comparison to other north african faunas of the ‘continental intercalaire’, it has been concluded that they were deposited during the cenomanian (sereno et al. 1996; cavin et al. 2010). cavin et al. (2010) emphasized that the kem kem beds actually include two formations: the ifezouane formation at the bottom and the aoufous formation at the top, as pointed out before (e.g., ettachfini & andreu 2004). the ifezouane formation is composed mostly of sandstone, while the aoufous formation presents marls with intercalations of sandstone and microconglomerates (cavin et al. 2010). as most of the fossils from the kem kem beds were collected by commercial dealers, it is not possible to determine their provenience, both in terms of locality and lithostratigraphical unit (e.g., mcgowan & dyke 2009). thus, the kem kem beds paleovertebrate fauna is, in fact, a compound assemblage, including terrestrial, freshwater and brackish vertebrates (cavin et al. 2010). systematic paleontology pterosauria kaup, 1834 pterodactyloidea plieninger, 1901 dsungaripteroidea sensu kellner 2003 dsungaripteroidea indet. material: ?lower jaw (cmn 50859) the specimen cmn 50859 consists of the anterior fragment (preserved length is 43 mm) from an edentulous jaw, possibly a mandible (fig. 1). it tapers anteriorly and shows the margin opposite to the occlusal surface rounded. in cross-section, the dorsal margin is concave and the lateral edges are elevated. the lateral sides are slightly convex, and a central oval lumen can also be observed. there is no median ridge or groove on the occlusal surface. small slit-like foramina are also present on the occlusal and lateral surfaces, suggesting that this portion of the jaw was well vascularized or innervated. the ventral margin lacks any evidence of a sagittal crest or keel. proper identification of such a small fragment can only be made tentatively. the dsungaripteridae, a clade which comprises pterosaurs that lack teeth on the tip of the jaws (young 1964, 1973), can be ruled out since the cross-section is not circular in cmn 50859. among tapejarids sensu kellner (2003), all species of the tapejarinae possess dentary crests (wellnhofer & kellner 1991; li et al. 2003; wang & zhou 2003; lü & yuan 2005; lü et al. 2006; kellner & campos 2007), a new pterosaur specimens from the kem kem beds (upper cretaceous, cenomanian) of morocco 151 feature absent in the preserved portion of cmn 50859. the new material also differs from the thalassodromin thalassodromeus sethi kellner & campos, 2002 by the lack of the distinctive sharp dorsal edge (kellner & campos 2002). the tips of the jaws in the tapejarid genus tupuxuara, in the pteranodontidae sensu kellner (2003), nyctosauridae, chaoyangopteridae, and in most of the azhdarchids, are similar to cmn 50859 and we cannot properly assign the material to any of these. compared with other moroccan toothless jaws, the preserved portion of cmn 50859 shares the concave dorsal surface with bsp 1993 ix 338 (wellnhofer & buffetaut 1999), here regarded as a lower jaw (contra ibrahim et al. 2010), and mn 7054-v (kellner et al. 2007), tentatively referred to the pteranodontoidea (sensu kellner 2003) (wellnhofer & buffetaut 1999; kellner et al. 2007) or to the azhdarchidae (bsp 1993 ix 338; ibrahim et al. 2010). cmn 50859 lacks the low sagittal crest and has a quite distinct cross-section from the recently described azhdarchid alanqa saharica (see ibrahim et al. 2010), indicating that a second edentulous species was present in the kem kem beds compound assemblage. as comparisons are limited and cmn 50859 also can not be undoubtedly identified as a lower jaw, it is here referred in the dsungaripteroidea, a clade which includes all toothless pterosaurs besides the anhangueridae, the istiodactylidae and the dsungaripteridae (kellner 2003, wang et al. 2009; but see unwin 2003 for a different hypothesis of pterosaur phylogeny). azhdarchidae nessov, 1984 azhdarchidae indet. material: mid-cervical vertebrae (cmn 50801 and linhm 014) the two cervical vertebrae described here show similar basic morphology to one another. both are elongated and clearly belong to the middle series (between cervical vertebrae 4 and 7). cmn 50801 (figs 2, 3) is well preserved, with no major signs of compression. the centrum is elongate (119 mm long, 18 mm minimum width) and somewhat flattened dorsoventrally. no lateral pneumatic foramina are present. two well-defined longitudinal sulci are observed on the anterior part, at the ventrolateral surfaces. the anterior cotyle is saddle-shaped, bearing a short and blunt hypapophysis on the ventral surface. the posterior condyle is expanded, sub-oval in shape, somewhat flattened dorsoventrally, and has a shallow depression underneath. the neural spine is low and elongated anteroposteriorly, extending along the whole length of the neural arch. in anterior and posterior views, two proportionally large, oval pneumatic foramina are present lateral to the neural canal. a third pneumatic foramen is also present on the anterior side, located dorsally to the neural canal, but is absent posteriorly. the present specimen does not have the right prezygapophysis preserved. the articular facet of the left prezygapophysis is broad, dorsally flattened, and oriented anteroventrally. a small ossification is present ventral to the prezygapophysis and lateral to the centrum, which appears to be the vestige of the cervical rib (either a fused cervical rib or an apophysis for the articulation of an unfused one), including a foramen (foramen transversarium). a shallow sulcus starts at this foramen and ends at the middle part of the centrum. the postexapophyses are horn-like and slightly divergent from each other. each bears small dorsolateral tubercles. the articular surfaces of the postzygapophyses are wide, flat and directed lateroposteriorly, somewhat inclined downwards. the vertebra linhm 014 (fig. 4) is also wellpreserved, with a very elongate centrum (182 mm long, 32 mm minimum width) that lacks lateral pneumatic foramina. longitudinal, ventrolateral sulci are also clearly visible in this specimen, as is the ossification regarded as the vestige of the cervical ribs. on the anterior side, a blunt hypapophysis is present. ventrally, there are two large, round pneumatic openings, each one located laterally to the hypapophysis. being paired structures, they can be considered original morphological features and not artifacts derived from preparation, pathological or taphonomic processes. the neural spine is low and the condyle laterally expanded. two large pneumatic foramina are also observed lateral to the neural canal, in both anterior and posterior views. the articular region of the left prezygapophysis is missing. a prominent foramen is present beneath each prezygapophysis and continued as a ventrolateral sulcus, as seen in cmn 50801. the postexapophyses are fig. 1 ?mandible, cmn 50859. a) cmn 50859 in dorsal view; b) cmn 50859 in ventral view; c) cmn 50859 in left lateral view; d) cmn 50859 in proximal view. scale bar = 10mm. 152 rodrigues t., kellner a.w.a., mader b.j. & russell d.a. fig. 2 mid-cervical vertebra, cmn 50801. a) cmn 50801 in dorsal view; b) cmn 50801 in left lateral view; c) cmn 50801 in right lateral view; d) cmn 50801 in ventral view. abbreviations in the text. scale bar = 50 mm. fig. 3 mid-cervical vertebra, cmn 50801. a), cmn 50801 in anterior view; b) cmn 50801 in posterior view. abbreviations in the text. scale bar = 10 mm. new pterosaur specimens from the kem kem beds (upper cretaceous, cenomanian) of morocco 153 horn-like and also bear dorsolateral tubercles. the very elongate centrum, low neural spine, lack of lateral pneumatic foramina on the centrum, and a neural arch fully integrated into a tubular vertebral body allow the placement of both vertebrae in the azhdarchidae (kellner 2003, 2004; unwin 2003; andres & ji 2008). among azhdarchids, ventrolateral sulci and canals are observed in azhdarcho lancicollis nessov, 1984 (see nessov 1984; averianov 2010), phosphatodraco mauritanicus (see pereda-suberbiola et al. 2003), and some isolated specimens (buffetaut 1999; ősi et al. 2005; henderson & peterson 2006). the saddle-shaped anterior cotyle is similar to that of other azhdarchid vertebrae but differs from arambourgiania philadelphiae (arambourg, 1959), where the cotyle is as high as, or higher than, wide (martill et al. 1998). a pneumatic foramen located dorsally by to the neural canal, as seen in cmn 50801, is also present in azhdarcho lancicollis (including the same proportions seen in the holotype specimen, ccmge 1/11915, nessov 1984; averianov 2010), but absent in arambourgiania philadelphiae (see martill et al. 1998) and other isolated azhdarchid cervical vertebrae (company et al. 1999; buffetaut 2001; ősi et al. 2005; henderson & peterson 2006). the presence of this third pneumatic opening has been regarded as a possible generic character for azhdarcho (bennett 1989); however, it was later found to be also present in pteranodon (bennett 2001). it is also noteworthy that the pneumatic foramina present in the neural arch of azhdarcho lancicollis vary in number, size and conspicuity, depending on the pofig. 4 mid-cervical vertebra, linhm 014. a) linhm 014 in dorsal view; b) linhm 014 in right lateral view; c) linhm 014 in left lateral view; d) linhm 014 in ventral view. abbreviations in the text. scale bar = 50 mm. 154 rodrigues t., kellner a.w.a., mader b.j. & russell d.a. sition of the vertebra in the neck (nessov 1984; averianov 2010). detailed studies are still needed to evaluate how this feature evolved within the pterodactyloidea and which are its systematic implications. in anterior view, the anterior margin of the dorsal surface of cmn 50801 presents two concave areas separated by the median elevation of the neural canal, similar to the condition seen in azhdarcho lancicollis (ccmge 1/11915, nessov 1984; averianov 2010). at the posterior end, the dorsal surface forms a distinctive convex roof, thus differing from the hungarian vertebra mtm gyn/450 (ősi et al. 2005), the spanish vertebra mguv 2271 (company et al. 1999), and also from azhdarcho (cast bsp 1992 i 19), all of which present two small concave areas, separated by the low neural spine. furthermore, both moroccan specimens do not show the ridge parallel to the neural spine observed on the right side of the neural arch of an azhdarchid vertebra from japan (ikegami et al. 2000). the prezygapophyses in linhm 014 are more divergent and shorter than in cmn 50801, the hungarian azhdarchid vertebra mtm gyn/448 (ősi et al. 2005), arambourgiania philadelphiae, quetzalcoatlus sp. and azhdarcho lancicollis (see nessov 1984; martill et al. 1998, ősi et al. 2005; averianov 2010). in linhm 014, the postzygapophyses are also shorter and less divergent than in the hungarian vertebra mtm gyn/450 (ősi et al. 2005), and broader than in cmn 50801. however, linhm 014 has a concave surface between the postexapophyses in ventral view, which is absent in fsac-kk 34, another azhdarchid cervical vertebra from the kem kem beds of morocco (ibrahim et al. 2010). furthermore, linhm 014 has postexapophyses larger than the postzygapophyses, as also reported in other specimens (buffetaut 1999; company et al. 1999), and opposed to cmn 50801. azhdarchoidea (sensu kellner 2003) azhdarchoidea indet. material: humerus (cmn 50814) cmn 50814 is a right humerus consisting of proximal and distal portions separated by a small gap. the preserved proximal part (fig. 6) is 134 mm long and the widest preserved portion is 78 mm. the humeral head is wide (65 mm) and shows the saddle-shaped fig. 5 azhdarchidae cervical vertebrae in dorsal view. a) linhm 014; b) cmn 50801, inverted; c) mtm gyn/448; d) quetzalcoatlus sp. (tmm 41544-16, cast nhmuk pv r 9323), inverted; e) azhdarcho lancicollis (ccmge 1/11915, cast bsp 1992 i 17), inverted ; f) arambourgiania philadelphiae (cast at the smnk); g) mdm 349; h) mtm gyn/450; i) fsackk 34. scale bars = 50 mm. new pterosaur specimens from the kem kem beds (upper cretaceous, cenomanian) of morocco 155 condition typical of pterosaurs. breakages show a very thin cortical bone supported internally by a system of trabeculae. on the ventral surface there is a small pneumatic foramen (10.5 mm diameter) located medially at the base of the deltopectoral crest. an internal mould of this foramen is observed. both the ulnar and deltopectoral crests are broken off. a well developed muscle scar (probably for m. triceps; see bennett 2003), with a rugose surface, is observed on the dorsal surface of the shaft, close to the medial side. it starts about 80 mm from the humeral head, shortly below the level of the base of the ulnar crest, running for 44 mm until reaching the end of the preserved portion of the humerus, suggesting that it was longer. the distal preserved portion of the humerus (fig. 7) is 128 mm long. ventrally, the capitulum (radial condyle) and the trochlea (ulnar condyle) are well delimited. the intertrochlear sulcus is deep and somewhat wide. there is a pneumatic opening of about 5 mm in diameter on the lateral side of the capitulum, entering beneath it (fig. 7c). a second, larger pneumatic foramen (30 mm long and 11 mm wide; fig. 7a) and a strong muscle scar are present proximal to the capitulum. the ectepicondyle is well marked, and the entepicondyle is very robust and prominent. the distal end is complete and has a maximum width of 100 mm. in distal view, it is rectangular, with a depressed middle part. a pneumatic foramen located on the ventral side of the proximal part of the humerus is present in nyctosaurus, pteranodon and the azhdarchoidea, while a rectangular distal end in distal view is similar to the condition observed only in the azhdarchoidea, differing from the sub-triangular shape present in the pteranodontoidea (bennett 2001; kellner 2003). thus, this specimen can be confidently referred as an azhdarchoid, although a more restricted identification to a particular taxon is not possible at the time. among azhdarchoids, there are some humeri reffig. 6 proximal part of the right humerus, cmn 50814. a) cmn 50814 in ventral view; b) cmn 50814 in dorsal view. abbreviations in the text. scale bar = 50 mm. fig. 7 distal part of the right humerus, cmn 50814. a) cmn 50814 in ventral view; b) cmn 50814 in dorsal view; c) inset showing the pneumatic foramen on the lateral side of the capitulum. abbreviations in the text. scale bar equals 50 mm in a and b, and 10 mm in c. 156 rodrigues t., kellner a.w.a., mader b.j. & russell d.a. erable to the azhdarchidae preserved in three dimensions. cmn 50814 is somewhat similar in shape to montanazhdarcho minor padian, de ricqlès & horner, 1995 (mcgowen et al. 2002), although substantially larger. the humeri of quetzalcoatlus sp. (tmm 415449, cast mn 4715-v), quetzalcoatlus northropi lawson, 1975 (tmm 41450-3, cast mn 6952-v), montanazhdarcho minor (mcgowen et al. 2002), and a yet undescribed tapejarid specimen from brazil (mn 6505-v) have a pneumatic opening beneath the capitulum. a pneumatic foramen proximal to the capitulum, as clearly observed in cmn 50814, also occurs in pteranodon (bennett 2001) and mn 6505-v. quetzalcoatlus sp. has a shallow depression located on the same place, but its association with a pneumatic diverticulum is unclear, and quetzalcoatlus northropi seems to lack this character. furthermore, the distal articulation of cmn 50814 lacks the bony ridge present in quetzalcoatlus northopi (fig. 8c), quetzalcoatlus sp. (fig. 8d), and in azhdarcho lancicollis (see averianov 2010), and therefore the moroccan specimen can not be referred to quetzalcoatlus or azhdarcho. the proximally located scar found in the moroccan specimen is much stronger than in other pterodactyloid pterosaurs, including quetzalcoatlus northopi and quetzalcoatlus sp., where it is only slightly marked. in both quetzalcoatlus northopi and quetzalcoatlus sp. the entepicondyle is comparatively smaller than in cmn 50814 (fig. 7, 8). discussion three edentulous pterosaur jaws from the kem kem beds were described by wellnhofer & buffetaut (1999), who tentatively regarded them as one pteranodontid upper jaw (bsp 1993 ix 338), one azhdarchid upper jaw (bsp 1996 i 36) and one tapejarid lower jaw (bsp 1997 i 67). recently, averianov et al. (2008) considered all of them as belonging to the same azhdarchid taxon, from different parts of the jaw. as ibrahim et al. (2010), we agree with the original description that bsp 1997 i 67 is a partial lower jaw and best referable to the tapejaridae (wellnhofer & buffetaut 1999; kellner 2004). as in all tapejarines, it has a concave dorsal surface and a deep crest (wellnhofer & kellner 1991; kellner & campos 2007). in fact, this fragment is very similar to the correspondent portion of the mandible of sinopterus dongi wang & zhou, 2003 from the aptian of china (wang & zhou 2003), although being larger. ibrahim et al. (2010) identified bsp 1996 i 36 as a lower jaw fragment, a view with which we agree, and referred it in a new species, alanqa saharica. we agree that this mandible is diagnostic but we do not support the identification of the third jaw fragment bsp 1993 ix 338 as the upper jaw of the same taxon (ibrahim et al. 2010), because it is based only on overall shape and size similarities. as it is shown above, the new lower jaw fragment described here (cmn 50859) has a much different cross-section with respect to the tip of the mandible of alanqa saharica, suggesting the presence of another edentulous pterosaur from the kem kem beds compound assemblage. therefore, we suggest that the material referred to alanqa saharica should be constrained to the holotype and two other lower jaws (bsp 1996 i 36 and fsac-kk 31), at least until more complete or associated material comes to light and confidently solves this ambiguity. furthermore, bsp 1993 ix 338 is very low and therefore can be regarded as a fig. 8 azhdarchoidea humeri in distal view. a) cmn 50814; b) tapejaridae (mn 6505-v); c) quetzalcoatlus northropi (tmm 41450-3, cast mn 6952-v); d) quetzalcoatlus sp. (tmm 41544-9, cast mn 4715-v). abbreviations in the text. scale bar equals 10 mm in a, b and d, 50 mm in c. new pterosaur specimens from the kem kem beds (upper cretaceous, cenomanian) of morocco 157 lower jaw, sharing the concave dorsal surface and lack of a sagittal crest with the lower jaw of pteranodon and dawndraco (e.g. bennett 2001; kellner 2010). for the time being, we agree with the original attribution by wellnhofer & buffetaut (1999) that bsp 1993 ix 338 is most probably a pteranodontidae or closely related taxon (as also pointed out by cavin et al. 2010). both vertebrae described here are referred to the azhdarchidae. apart from their overall resemblance, linhm 014 is larger and comparatively less elongate than cmn 50801. besides, the paired ventral pneumatic openings present only in linhm 014 are unknown in any other pterosaur specimen. as both vertebrae were found in isolation, it is not possible to know if they pertained to different positions on the neck (such that the presence of these foramina would be typical of one of the cervical vertebrae but absent in others) or if these features suggest that two different azhdarchid species coexisted in this region. cmn 50801 shares an overall similar morphology with the specimen ccmge 1/11915, holotype of azhdarcho lancicollis, which was referred as the ?fifth cervical (bakhurina & unwin 1995). comparisons to the published proportions of phosphatodraco mauritanicus (see pereda-suberbiola et al. 2003) indicate that both moroccan vertebrae compare well with the putative sixth cervical; however, the described fifth cervical actually consists of two separate, closely articulated third and fourth cervical vertebrae (kellner 2010). therefore, both moroccan vertebrae here described show proportions similar to what is actually the fifth cervical of phosphatodraco mauritanicus. if these suppositions are correct, then both specimens are fifth cervicals and therefore representatives of two different azhdarchid taxa. both moroccan vertebrae described here differ from the fifth cervical of quetzalcoatlus sp. (tmm 41544-15, cast nhmuk pv r 9325), which is extremely elongate, demonstrating that they concern relatively small individuals when in comparison with derived azhdarchids. the humerus cmn 50814 allows a better estimative of the wingspan of this individual. it can be estimated that at least between 35 mm and 50 mm of the shaft is missing, rendering a minimum total length of the humerus of about 300 mm. comparisons with the estimated wingspans of quetzalcoatlus northropi and montanazhdarcho minor (whose holotypes possess complete humeri) rendered an estimative of ca. 5.5-6 m for this individual. even if different growth rates in wing bones of the same individual are taken into consideration (as shown by sayão 2003), such estimative indicates that cmn 50814 concerns a large but not gigantic pterosaur. as both distal epiphyses are well fused to the humeral shaft, cmn 50814 can be considered at least a sub-adult individual (bennett 1993). conclusions despite the large number of pterosaurs known to date, the attribution of isolated edentulous jaws is still not well understood, as illustrated by the scarce number of characters in phylogenetic matrices that could be used in this regard (e.g., kellner 2003; unwin 2003; andres & ji 2008; lü et al. 2009; wang et al. 2009). clades such as the azhdarchoidea might comprise a relatively high morphological diversity, and more complete material is still needed to better understand this. perhaps somewhat surprisingly, the identification of isolated postcranial material of edentulous taxa is less of a challenge. certain features (e.g., the rectangular distal end of the humeri) provide clades good support and can be used in referrals to more inclusive clades with confidence. furthermore, the vertebrae here described provide a hint on azhdarchid evolution, showing a low but not vestigial neural spine, suggesting that the height of the neural spine reduced in at least some more derived and younger azhdarchid forms, such as the maastrichtian species quetzalcoatlus northropi and arambourgiania philadelphiae, which also possess an extreme elongation of the cervical vertebrae (kellner & langston 1996; martill et al. 1998; henderson & peterson 2006). pterosaur specimens from the ifezouane and the aoufous formations of morocco, even though being incomplete and found isolated, can be extremely well preserved, as it is the case of the two cervical vertebrae here described. they comprise a rather wide range of dsungaripteroid clades, and represent so far the most diverse pterosaur fauna from africa (kellner et al. 2007; ibrahim et al. 2010). whether such a diverse fauna coexisted temporally or is an artifact of the lack of detailed stratigraphic information still remains to be accessed. in any case, the existence of such high diversity of clades shows the potential of new finds that these formations could yield. acknowledgments. the authors would like to thank oliver rauhut (bayerische staatssammlung für paläontologie und geologie), lorna steel (natural history museum) and eberhard frey (staatliches museum für naturkunde karlsruhe) for access to the pterosaur collections, eric buffetaut (cnrs, laboratoire de géologie de l’ecole normale supérieure) for providing information and bibliography on the kem kem beds, julio company (universidad politécnica de valencia) for the articles, photos and replica sent, attila ősi (hungarian academy of sciences) for photos of mtm gyn/448 and mtm gyn/450, and naoki ikegami (mifune dinosaur museum) for the photo of mdm 349. the article was greatly improved by the many suggestions of the reviewers fabio dalla vecchia (institut català de paleontologia) and kevin padian (university of california). this study was partially funded by the conselho nacional de desenvolvimento científico e tecnológico (cnpq – grants 140407/2007-3 and 290019/2008-7 to t.r. and 307276/2009-0 to a.w.a.k.) and fundação carlos chagas filho de amparo à pesquisa do rio de janeiro (faperj – grant e-26/102.779/2008 to a.w.a.k.), with support from the deutscher akademischer austauschdienst (daad – to t.r.). 158 rodrigues t., kellner a.w.a., mader b.j. & russell d.a. andres b. & ji q. 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si s a . s o ld a d o e n s is c re ta ce o u s ta xa f o ra m in if e ra l b io zo n a ti o n 6 0 5 5 5 0 4 5 4 0 38 .5 paleoceneeocene danianselan.thanet.ypresianlutetianbart. p r ia b . p 3 p 4p 5 e 1 e 2 e 3 e 4e 6 e 5 e 7 e 8 e 9 e 1 0 e 1 2 e 11 if p 1 6 (= e 1 3 ) if p 1 7 p0-p2a ab bc 6 5 c r e t a c e o u s c . te n u is c re ta ce o u s ta xa c . d a n ic u s s p h e n o lit h u s . sp p f a s c ic u li th u s s p p . h . k n e in p e ll ii h . ri e d e lii d . m o h le ri d . m u lt ir a d ia tu s d . d ia s ty p u s f. ty m p a n if o rm is d . lo d o e n s is d .b is e c tu s c . o a m a ru e n si s f c o c . re ti c u la tu m d . s u b lo d o e n s is n a n n o te tr in a s p p . n a n n o te tr in a s p p . t. o rt h o st yl u s t. o rt h o st yl u s c . g ig a s c . g ig a s r . u m b ili cu s > 1 4 m ic ro n n a n n o fo s s il b io zo n a ti o n n p 1 5 * n p 1 6 * n p 1 7 * n p 1 8 * paleoceneeocene danianselan.thanet.ypresianlutetianbart. p r ia b . n p 1 n p 2 n p 3 * n p 4 * n p 5 * n p 6 n p 7 n p 8 n p 9 n p 1 0 n p 11 * n p 1 2 n p 1 3 n p 1 4 * 6 0 6 5 5 5 5 0 4 5 4 0 3 8 .5 c r e t a c e o u s s e d ic o 1 0m bellunoflysch venad’oromarl s ca g lia 12 50 s h s s c g s h s s c g gallaregroupjesoloflysch 15 00 24 69 scaglia e r a c l e a 1 � �� � � � � � � � � � � �� �� �� � � � � � �� �� � � � �� �� � � �� � � � � � � � �� �� �� �� �� � � � �� � � �� �� � �� � �� �� � � � � �� � � �� �� � � �� � �� �� � � �� � � � � � �� �� � � � �� �� � � � �� � � � � � �� �� � � �� � � �� � �� � � � �� �� �� �� � � �� �� �� � � � � �� � � �� ! � � � � � �� � � �� � � �� � � � � � � �� �� �� ��� � ��� ��� ��� � !���� "� # ������� � ���" 0 m 1 0 0 m 2 0 0 m t h ic kb e d d e d tu rb id ite sa n d st o n e c a rb o n a te h e m ip e la g ic m u d st o n e m a rl y h e m ip e la g ic m u d st o n e l o w e st o cc u rr e n ce (l o ) h ig h e st o cc u rr e n ce (h o ) u n co n fo rm ity f o ra m in if e ra l b io zo n a ti o n 6 0 5 5 5 0 4 5 4 0 38 .5 paleoceneeocene danianselandianthanetianypresianlutetianbart. p r ia b o n ia n p 3 p 4p 5 e 1 e 2 e 4 e 6 e 5 e 7 e 8 e 9 e 1 0 e 11 e 1 2 if p 1 6 (= e 1 3 ) if p 1 7 p0-p2 e 3 a ab bc m . v e la s c o e n s is m . fo rm o sa m . s u b b o ti n a e m . a ra g o n e n si s m . a ra g o n e n s is o . b e ck m a n n i m u ri c a te s p e c ie s o . b e ck m a n n i t. c e rr o a zu le n s is fr o n to s a a . cu n e ic a m e ra ta g . n u tt a lli g . n u tt a lli g . m e x ic a n a k u g le ri p. w ilc o xe n si s m . a n g u la ta i. a lb e a ri g . p se u d o m e n a rd ii g . p s e u d o m e n a rd ii p. va ri o sp ir a a . so ld a d o e n si s a . si b a iy a e n si s a . s o ld a d o e n s is 6 5 c r e t a c e o u s c re ta c e o u s ta x a n a n n o fo s s il b io zo n a ti o n n p 1 5 * n p 1 6 * n p 1 7 * n p 1 8 * paleoceneeocene danianselandianthanetianypresianlutetianbartonian p r ia b o n ia n n p 1 n p 2 n p 3 * n p 4 * n p 5 * n p 6 n p 7 n p 8 n p 9 n p 1 0 n p 11 * n p 1 2 n p 1 3 n p 1 4 * 6 0 6 5 5 5 5 0 4 5 4 0 3 8 .5 c . te n u is c re ta c e o u s ta x a c . d a n ic u s s p h e n o lit h u s sp p . f a s c ic u li th u s s p p . h . kn e in p e lli i h . ri e d e lii d . m o h le ri d . m u lt ir a d ia tu s d . d ia st yp u s d .l o d o e n s is d .b is e c tu s c . o a m a ru e n si s f c o c . re ti c u la tu m d .s u b lo d o e n s is n a n n o te tr in a s p p . n a n n o te tr in a sp p . t. o rt h o st yl u s t. o rt h o st yl u s c . g ig a s c . g ig a s r . u m b ili cu s > 1 4 m ic ro n c r e t a c e o u s a m ir a 1 15 00 a d a 1 a s s u n ta 1 s h s s c g s h s s c g s h s s c g gallaregroup gallaregroup gallaregroup scagliafm. scagliafm. scagliafm. 22 00 17 00 30 32 26 00 24 70 21 00 1 2 3 4 1 2 3 4 � �� � # � � � � � � � � �� �� �� � � � � � �� �� � � � �� �� � � �� � � � � � � � �� �� �� �� �� � � � � � �� �� � �� $ �� � � $ � � $ � �� � �� �� � � �� � �� �� � � �� � � � � � �� �� � � � �� �� � � � �� � � � � � �� �� � � �� � � 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l io c . p le . s e l a n . danian t h a n . y p r e s ia n l u t e t ia n b a r t. p r ia b . r u p e l ia n c h a t t ia n aquitan. burdigal. langhian serraval. tortonian messinian zanclean piacenzian gelasian calabrian ion-tyr p3 p2 p1c e6 p4 p5 e4 e2 e1 e5 e7 e8 e9 e10 e11 e12 (=e13) ifp17 ifp16 ifp21 np5* np4* np6 np7 np8 np9 np10 np11* np12 np13 np14* np15* np16* np17* np18* mnp19 mnp20 mnp21a mnp21b mnp22 mnp23 ifp20 ifp19 ifp18 ifp22 ifn1 ifn2 ifn3 praeorbulina spp. mpl1 mpl 2 mpl 3 mpl 4 mpl 5 mpl 6 und. a a a e3 a a b b b b b c ifn4 mnp24 mnp25a mnn2a mnn2b mnn3a mnn3b mnn4a mnn4b mnn5 5a 5b mnn6a mnn6b mnn7 mnn8 mnn9 nn10 nn11 n o t co n si d e re d 11a 11b 11c 11d mnn1 1a 1b 1c 1d mnp25b g. peripher. p. mayeri t. conomiozea g. menardii / n. acostaensis 60 55 50 45 40 35 30 25 20 15 10 5 ���� 7 � =� �!��� � � �� �������� �� � � ��� ���� �������� ;��� � � � ����� ������ � � ���� �� �� � �� �� �� �� � �� +"--� �� � ����.� ;��!� �� ��> )/� 9)/ � 9)) ��� � �� ����� ��������? /� 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sandstone thin-bedded turbidite sandstone all. era. san. gs b v.v. s. donà gall. cav. poss. jesolo sc. : alluvial deposit : eraclea sandstone : santerno group : gessoso solfifera : boreca conglomerate : vittorio veneto sandstone : san donà marl : gallare group : cavanella group : possagno marl : jesolo flysch : scaglia rossa60 65 55 50 45 40 35 30 25 20 15 10 5 0 p a l e o c e n e cretaceous e o c e n e m io c e n e o l ig o c e n e p l io c e n e p l e i. d a n ia n s e l a n d . t h a n e t. y p r e s ia n l u t e t ia n b a r t. p r ia b o n ia n r u p e l ia n c h a t t ia n a q u it a n . b u r d ig . l a n g . s e r r . t o r t o n . mess. zanc. piac. gelas. calab. ion-tyr p4 p5 e1-e2 e 3 -e 5 e 8 -e 9 e 1 0 -e 1 2 e6-e7 ifp16= e13 ifp17 p 0 p 3 ifp20 ifp21 ifp19 ifp18 ifp22 praeorbul. if n 1 -i f n 3 1-2 4-5 3 6 u n d . a b if n 4 m p l t. conomiozea g . p e ri p h e r. p. m a ye ry g .m en ar di i g . a co st ae ns is s c. s c. ���� � � ��� �� � ������ ������ ��������������� 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"''�' #$ &���� ! �� �� $ �( -� � �"'-#.*�"� �"�*$ #$ �#�� "'!� �"$$ ! �#& /�$$" #$ �*&� �� �' �&&�*$"� (# ,� !� !��!� ��, #$ )%$�"� "$ "�� "�$ #' !� ���' ,��( $� ������� ���� ���� ������ ����"�#$�' ,#!� /�$$#� $� &#� '$ ��� !� *�� � ! !�� !� "'"��$#$ ,"$ �#�#! !� !� g*�"� ! !�� � 5����� "'"��$#$ $��, !�"! !� -#// � '& $ . !, ' � "' "� $#�'#/#&"'! 2d#�1e$ �"�.% -"� >�� ::3 �� ���8:�%>84� 5����� � ( "� !�"! !� �( !�"'" ,��( $ -#// � /��� !��$ �/ ?��!!" ���"' ��# 2�%("�* h >�>>�04 "'5 ��#�'"'� 2�%("�* h >�>>>:4 .*! '�! /��� !��$ �/ �'�"�"'� 2�%("�* h >�>9004 "')"' )#�� 2�%("�* h >�8>9;4 2 ".� ;4� � -#$&�#�#'"'! "'"��$#$� � �/��� !� ! $! !� -#// � '& $ . !, ' !� �( !�"'" $� &#� '$ "'!� �!� � /�$$#� ,��( $� #$ $#�% '#/#&"'! 2�h >�>> 904 "'!� $� &#� '$ &��� &!�� &�"$% $#/# "� 0>�9i� � ��� � �/��� �' !� *�� � ! !� $��,$ $#�#�"� -#// � '& $ 2�#�� @3 ".� 94� � /#�$! &����' '! ��� 2�' !� ���#7�'!"� "6#$4 "&&�*'!$ /�� !� 90�;�i �/ !� !�!"� ("�#"'& � "'�"$ #$ ��"-#'�$ /�� !� � '�!� �/ �;� � $ &�'&����' '! 2( �!#&"� "6#$4 6��"#' �9�@:i �/ !� ("�#"'& "'#$ �"#'�� #'/�* '& 0����1 /"," 2 %�����.� )"�: area site age mis references italy melpignano and san sidero late pleistocene bertè 2013; sardella et al. 2014; this work italy grotta romanelli late pleistocene cassoli et al. 1994; sardella et al. 2014; this work italy avetrana late pleistocene petronio et al. 2008; pandolfi et al. 2013a; this work italy veja grotta a late pleistocene 3 this work italy pocala late pleistocene 3 this work italy buco del frate late pleistocene 3 bona pers. com.; this work italy ingarano late pleistocene 3 bertè 2013; sardella et al. 2014; this work italy grotta sant'agostino late pleistocene 3 tozzi 1970; this work italy grotta di ladrenizza late pleistocene 3 this work italy grotta mora cavorso late pleistocene 2 salari et al. 2011; this work italy covoli di velo late pleistocene 2 this work italy grotta paglicci late pleistocene 2 bartolomei 1975; this work italy grotta bella holocene, ancient 1 guerreschi et al. 1992; this work italy colle cappuccini holocene, ancient 1 wilkens 1990; this work italy central italy, apennine area holocene, recent 1 bertè 2013, this work france aven de l'arquet late pleistocene 3 boudadi-maligne 2010, 2012; brugal & boudadi-maligne 2011 france jaurens late pleistocene 3 boudadi-maligne 2010, 2012; brugal & boudadi-maligne 2011 france maldidier late pleistocene 3 boudadi-maligne 2010, 2012; brugal & boudadi-maligne 2011 france igue du gral late pleistocene 2 boudadi-maligne 2010, 2012; brugal & boudadi-maligne 2011 4 3 4 3 ".� � % �#$! �/ !� /�$$#�#/ ��*$ ��&"�#!# $ �# �� �"#'$ �/ ���� ���� #'&�*#' !�#$ $!*-�� )��� %��� ������ �"'#$ �*�*$ .�� �������� 3-���#��� 4����5 �:� site p4 l p4 b gb p4 m1 l m1 b m2 l m2 b melpignano/san sidero mean 21.9 8.7 11.1 13.1 16.7 8.3 12.1 n 9 9 8 5 5 3 3 grotta romanelli mean 22.1 8.9 10.8 13.9 17.0 8.3 12.0 n 5 5 3 6 6 6 6 veja grotta a mean 27.0 10.7 13.0 16.4 19.0 10.0 13.6 n 1 1 1 1 1 1 1 buco del frate mean 26.0 10.6 13.9 16.0 19.3 8.9 13.1 n 13 13 13 11 11 6 6 ingarano mean 23.5 9.0 12.2 14.7 18.1 8.7 12.0 n 5 5 5 7 7 4 4 grotta sant'agostino mean 24.9 13.2 17.3 22.0 n 3 2 3 3 grotta ladrenizza mean 23.9 9.5 14.4 15.2 18.2 8.8 12.1 n 2 2 2 2 2 2 2 avetrana mean 23.7 10.3 12.9 16.0 19.5 9.2 12.2 n 12 10 10 13 14 3 3 grotta mora cavorso mean 24.87 15.1 17.5 23.4 n 2 2 2 2 covoli di velo mean 25.4 10.2 15.0 17.0 19.5 9.3 13.4 n 2 2 2 1 1 2 2 grotta paglicci mean 22.7 8.9 11.9 n 1 1 1 appennine area mean 24.2 9.7 12.8 15.3 18.4 8.3 12.8 n 250 250 250 252 253 253 252 aven de l'arquet mean 25.7 10.3 13.6 16.3 21.5 9.0 13.5 n 27 26 27 16 16 15 15 jaurens mean 26.6 10.7 14.5 16.6 20.0 8.8 14.0 n 8 8 7 5 5 5 5 maldidier mean 27 11.07 14.8 16.2 21.95 9.2 14.5 n 3 3 3 2 2 4 4 igue du gral mean 25.47 10.36 13.9 n 10 10 10 ".� ; % ����"�"!#( -#� '$#�'$ 2��4 �/ !� *�� � ! !� �/ ���� ���� �/ �( !�"'" "'!��$ �/ $ ( �"� )�*!�% �' �!"�#"' ��&"�#!# $ "'�� ''#' "� "� �#�� @ % ��#'&#�"� �����' '! �'"��% $#$ � �/��� �' !� *�� � ! !�� ���$$� /*�� �#' � �( % !�"'"3 &#�&� � .#� ��#'! �#' � 5 ��#�'"'�j)"' )#��3 !�#% "'�� � ��#'! 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'�! /��� !��$ �/ �'�"�"'� 2�%("�* h >��;;�; 4� 5 ��#�'"'� 2�%("�* h >�>:�0;::4 2 ".� @4� � ��� � �/��� �' !� ��, � ! !� $��,$ $#�#�"� -#// � '& $ 2�#�� 93 ".� 94� � /#�$! &����' '! ��� 2�' !� ���#7�'!"� "6#$4 "&&�*'!$ /�� !� @@�@i �/ !� !�!"� ("�#"'& � "'�"$ #$ ��"-#'�$ /�� !� � '�!� �/ 5�� � $ &�'&����' '! 2( �!#&"� "6#$4 6��"#' �9�>i �/ !� ("�#"'& "'#$ �"#'�� #'% /�* '& .� !� .� "-!� �/ � � ����� "'"��$#$ ,"$ � �/��� �' ��� "'�� 2 *1 � �"#�,"�$ ! $! �h 0�:9��%>94� � -#� '$#�'"� �"'� �/ !� � '�!� �/ 5� #$ $#�#�"� !� !�"! �/ !� �� '&� ��&"�#!# $ �/ a"*� '$ "'�( ' �e��c* !� #' "--#!#�' #! #$ $#�#�"� !� !�"! �/ �*&� � ��"! � ��&"�" "'�!� � ���!� �' "'� '% !�"� �!"�#"' ��&"�#!# $� &���'����#&"��� � �"! !� 5�) � "'5�) 2�#�� :4� � � "' ("�* �/ !� �"!#� . % !, ' !� !�!"� � '�!� "'!� !"��'#� '�!� �/ 5� $��,$ $#�#�"� ("�* $ !� !� $"��� $ *'� #'( $!#�"!#�' 2 ".$ ;� @4� � $� &#� '$ �/ �( !�"'" "� '�! $#�'#/#&"'!�� -#// � '!� .�!� /�� *�� � "'��, � ! !�� /��� !� ��% �*�"!#�'$ �/ �*&� � ��"! � "'a"*� '$� 0����1 /"," 2 %�����.� )"�:; site p1 l p1 w p2 l p2 w p3 l p3 w p4 l p4 w m1 l m1 w ltal m2 l m2 w m3 l m3 w melpignano/san sidero mean 5.3 4.2 11.7 5.4 13.0 6.1 14.8 7.0 24.2 10.0 7.4 11.1 7.8 5.5 4.8 n 4 4 8 8 10 10 10 10 7 7 7 6 6 3 3 grotta romanelli mean 5.4 4.0 10.5 5.0 12.0 5.1 13.7 6.2 23.9 9.5 7.1 10.8 7.5 4.3 4.0 n 2 2 3 3 2 2 5 5 4 4 4 4 4 2 2 veja grotta a mean 23.0 9.2 6.8 10.2 7.2 n 1 1 1 1 1 pocala mean 5.7 4.4 13.2 6.2 13.0 6.3 15.6 7.8 27.8 11.2 8.5 11.7 8.4 n 1 1 1 1 2 2 3 3 7 7 7 3 3 buco del frate mean 5.9 5.0 12.3 7.7 14.1 7.5 16.2 8.2 28.4 11.8 7.7 11.8 8.7 n 1 1 11 11 10 10 17 17 27 27 27 14 14 ingarano mean 10.9 5.3 12.7 6.0 14.0 6.6 25.0 10.0 7.0 11.6 7.9 n 3 3 2 2 2 2 5 5 5 3 3 grotta sant'agostino mean 28.2 11.3 n 5 5 avetrana mean 5.5 4.9 12.5 6.0 14.2 6.7 15.5 8.0 28.5 11.7 8.8 12.2 8.8 6.0 5.1 n 1 1 12 12 9 8 11 9 19 17 16 11 11 1 1 grotta mora cavorso mean 29.1 12.1 n 3 3 grotta paglicci mean 11.9 5.3 12.7 5.8 14.6 6.8 26.3 10.8 6.3 n 1 1 1 1 3 3 1 1 1 grotta bella mean 28.3 n 4 grotta cappuccini mean 28.6 n 1 appennine area mean 5.7 4.3 11.6 5.7 13.1 6.2 14.9 7.1 27.2 10.8 7.6 10.9 8.0 5.2 4.8 n 167 167 248 248 247 247 254 254 251 251 249 247 247 198 198 aven de l'arquet mean 6.2 4.9 12.6 6.1 13.2 6.8 16.1 8.0 28.2 11.4 10.2 11.8 8.8 5.9 6.0 n 7 7 17 17 15 14 18 18 28 28 28 21 21 1 1 jaurens mean 6.6 5.0 12.7 6.2 14.6 7.2 16.5 8.5 29.5 12.0 10.2 12.0 8.9 n 4 4 10 10 13 13 15 15 19 19 19 9 9 maldidier mean 6.2 5.0 13.0 6.7 15.0 7.7 17.2 8.9 30.3 12.4 10.6 10.6 8.5 5.5 5.3 n 4 4 4 4 4 4 5 5 3 3 3 2 2 3 3 igue du gral mean 6.1 5.0 12.7 6.4 14.5 7.3 16.7 8.5 29.3 11.9 11.0 12.0 9.2 6.0 5.5 n 6 6 10 10 10 10 12 12 10 10 10 7 7 7 7 ".� @ % ����"�"!#( -#� '$#�'$ 2��4 �/ !� ��, � ! !� �/ ���� ���� �/ �( !�"'" "'!��$ �/ $ ( �"� )�*!� �' �!"�#"' ��&"�#!# $ "'�� ''#' "� "� ����"�#$�' ,#!� 6!"'! ,��( $ � 5����� "'"��$#$ $��, !�"! !� -#// � '& $ . !, ' � "' "� $#�'#/#&"'! 2d#�1$ �"�.-"� >�>;0@3 �h � ;>>��%>94� � $"��� /��� �( !�"'" "'!� 6!"'! ,��( $ "� $#�'#/#&"'! -#// � '! 2�%("�* � � ;>>��%>94 "'!� �%("�* � �"#'$ !� $"� "/! � !� ��'/ ���'# &��� &!#�'� � -#$&�#�#'"'! 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6�@<@8 �$, %�� ,9��" ����%(� �����%��* "%$$� ��$�%(,�� f���&$%��((% �� ��$"�&$%��((% �� ,� g�( &%�����%��* � �& 3�9 �)&# ��������& �3& ����� ���&� ���&#�����3��& ������ �� ;� ��@* << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /all /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /warning /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true /embedjoboptions true /dscreportinglevel 0 /syntheticboldness 1.00 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true 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/pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /fra /enu (use these settings to create pdf documents with higher image resolution for improved printing quality. the pdf documents can be opened with acrobat and reader 5.0 and later.) /jpn /deu /ptb /dan /nld /esp /suo /ita /nor /sve /kor /chs /cht >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [595.000 842.000] >> setpagedevice rivista italiana di paleontologia e stratigrafia volume 117 no. 1 3 pls. pp. 39-50 april 2011 discovery of the lower murgabian (middle permian) based on neoschwagerinids and verbeekinids in the taurides, southern turkey fumio kobayashi 1 & demir altiner 2 received: september 9, 2010; accepted: december 1st, 2010 1 corresponding author. institute of natural and environmental sciences, university of hyogo, sanda, hyogo 669-1546, japan. e-mail: kobayasi@hitohaku.jp 2 department of geological engineering, middle east technical university, ankara 06531, turkey. e-mail: demir@metu.edu.tr key words: fusulines, early murgabian, taurides, turkey. abstract. lower murgabian (roadian) beds have been discovered for the first time in a thick carbonate sequence ranging from devonian to triassic in the hadim area, central taurides, southern turkey. the roadian limestone consists of black algal fusuline packstone and black bioclastic packstone, and contains presumatrina ciryi n. sp., an evolved form of the genus, verbeekina erki n. sp., an earliest species of verbeekina, dunbarula protomathieui n. sp., an ancestral form of dunbarula mathieui, and several smaller foraminifera. riassunto. sono stati individuati per la prima volta livelli fossiliferi di età murgabiano inferiore (roadiano) in una potente successione carbonatica che si estende dal devoniano al triassico nell’area di hadim, tauridi centrali, turchia meridionale. i calcari di età roadiana sono costituiti da packstone scuri con alghe e fusuline e da packstone bioclastici scuri. essi contengono presumatrina ciryi n. sp., una forma evoluta del genere; verbeekina erki n. sp., una specie primitiva di verbeekina, ed infine dunbarula protomathieui n. sp., forma ancestrale di dunbarula mathieui, oltre a numerosi piccoli foraminiferi. introduction more than 5.500 m thick carbonate rocks of the aladag unit ranging from the devonian to triassic are widely exposed in the hadim-taskent area, central taurides, southern turkey. in altiner & özgül (2001), the carboniferous sequence measuring 910 m in thickness was divided into 14 foraminiferal zones within the interval from visean to moscovian and the permian consisting of a 1.090 m thick succession was mostly assigned to the upper middle permian and upper permian (a questionable interval of midian, midian, dzhulfian and dorashamian). the upper carboniferous to lower permian interval had not been zoned in altiner & özgül (2001) and its biostratigraphy had been left for a further study. we recognized 11 fusuline zones in the upper carboniferous (kasimovian and gzhelian) and lower permian (asselian to artinskian?) of the hadim area, and compared these fusuline faunas with contemporaneous ones of other west tethyan, southern ural, and russian platform regions (kobayashi & altiner 2008a; 2008b). on the other hand, the presence of upper lower permian to the middle part of middle permian (artinskian to wordian or yakhtashian to murgabian) is still uncertain in the hadim area. in other permian localities of the tauride block, the “assemblage” characterized by four species of eopolydiexodina (altiner, 1984) and the eopolydiexodina zone (köylüoglu & altiner 1989) reported from the hakkari area of southeast turkey were thought to be murgabian based on leven (1981) who reported at that time that eopolydiexondina is frequently present in the neoschwagerina craticulifera and neoschwagerina margaritae zones corresponding to the middle and upper murgabian. in the absence of neoschwagerinids and verbeekinids in the hakkari faunas it was not possible to assign a more detailed age to those levels. on the other hand, more prolific and variable fusulines of murgabian or midian age were reported from the blocks of the karakaya orogen in northern turkey (ciry 1938; erk 1942; skinner 1969; leven & okay 1996; turhan et al. 2004; altiner & özkan-altiner 2010). 40 kobayashi f. & altiner d. the difference between the permian faunas of turkey were analyzed in altiner et al. (2000) who defined two biofacies belts in the middle to upper permian. the northern biofacies belt comprises the permian exposures of the karakaya orogen and the outer platform or platform margin deposits of the tauride belt recognized in the bolkar dag unit. the biostratigraphy in the northern biofacies belt is based on a more diverse foraminiferal fauna and the kubergandian and murgabian stages are present. the hakkari area with eopolydiexodina belongs to the southern biofacies belt very poor or absent in neoschwagerinids and verbeekinids. an advanced form of presumatrina and a primitive form of verbeekina reported in this paper are recognized in the aladag unit of the hadim area. throughout the taurides, the aladag unit assignable to the southern biofacies belt is tectonically overlain by the bolkar dag unit belonging to the northern biofacies belt. these two units were tectonically and paleogeographically separated until their amalgamation during the late cretaceous to late eocene (e.g., sengör & yilmaz 1981). the limestone beds with murgabian and midian neoschwagerinids reported from the taskent area (altiner & özgül 2001) belong to the bolkar dagi unit. the purpose of this paper is to report the discovery of the lower murgabian (roadian) strata from the aladag unit of the hadim area and to describe the murgabian foraminiferal faunas including three new species of presumatrina ciryi, verbeekina erki, and dunbarula protomathieui. two limestone samples used in this paper were collected on the occasion of our field work of carboniferous-permian boundary sequences in the hadim area in 2004. limestone thin sections of the hadim area used in this paper are stored in the collection of the museum of nature and human activities, hyogo, japan (fumio kobayashi collection, mnhah). material the permian çekiç dagi formation of the hadim area is subdivided into four members in ascending order: keltas, çamalani, kizilgeris, and yellice. the keltas member is of asselian to sakmarian age (kobayashi and altiner 2008a). the çamalani member is unknown in age whereas the kizilgeris member is questionably assigned to the midian. the thick carbonate succession of the yellice member belongs to the midian to dorashamian interval (altiner & özgül 2001). two samples were collected from the 17-m-thick, thin bedded limestone succession intercalated within the quartz arenitic sandstone of the kizilgeris member (figs 1, 2). they are located at about 12 km sw of the town of hadim, southern turkey (36°54'32˝n, 32°23'08˝e). sample a is a black algal fusuline packstone poor in fossil diversity. in addition to a dominant fusuline fauna, algae referable to mizzia and other algal bioclasts (mostly replaced by calcite) are packed within a lime mud matrix (pl. 1, fig. 1). sample b is different in microfacies from sample a and is classified as black, bituminous, bioclastic packstone (pl. 1, fig. 2). foraminifers (pl. 1, fig. 6) and a microproblematicum referable to pseudovermiporella sodalica elliott (pl. 1, figs. 3-5) are the principal constituents in the sample. fusulines, however, are completely absent in this bioclastic packstone except some rare staffellids (sphaerulina and nankinella). fauna and age foraminifers contained in sample a are mostly composed of presumatrina ciryi n. sp. and other species consisting of verbeekina erki n. sp., dunbarula protomathieui n. sp., dunbarula sp., rauserella? sp. schubertellidae gen. and sp. indet. a, schubertellidae gen. and sp. indet. b, nankinella sp., endothyra sp., climacammina sp., palaeotextularia sp., globivalvulina sp., geinitzina? sp., pachyphloia spp., hemigordius discoides lin, li & sun, hemigordius sp., multidiscus sp., and neodiscus spp. fusulines referable to schwagerinidae are absent in this fauna. fig. 1 upper: map showing the distribution of the tauride block and the hadim area, southern turkey. lower: geologic map of upper paleozoic strata in the hadim area, about 12 km sw of the town of hadim and the limestone sample locality of early murgabian foraminifers. the bold line in right of the sample locality shows the location of the columnar section shown in fig. 2 (modified from altiner & özgül 2001). discovery of lower murgabian (middle permian) in the taurides, southern turkey 41 faunal composition of sample b is largely different from that of sample a. fusulines are very poor and confined to unidentified species of sphaerulina and nankinella in 39 slides of thin sections. non-fusulinoidean foraminifers distinguished are neodiscus spp., pachyphloia robusta miklukho-maklay, pachyphloia sp. geinitzina spp., nodosinellodes sp., and an indeterminate milioline species. age-diagnostic species are not found in sample b. the species composition in smaller foraminifers is highly different from that of the sample a and coeval ones previously described. palaeotextulariidae, biseriamminidae, and endothyridae are totally absent in sample b. presumatrina ciryi n. sp. is thought to be one of the evolved species of the genus along with presumatrina ozawai (hanzawa) described from the lower murgabian akiyoshi limestone of japan (hanzawa 1954), presumatrina grandis leven from the lower murgabian of the southeast pamir (leven 1967), and presumatrina uruzganensis leven from the lower murgabian of afghanistan (leven 1997). these three species of presumatrina have larger tests and more whorls than other described species of the genus. secondary transverse septula first appear in the fourth whorl in this new species. they appear ontogenetically earlier than those of other species such as presumatrina schellwieni (deprat) and p. neoschwagerinoides (deprat). secondary transverse septula are less developed in outer whorls in comparison with those of afghanella. presumatrina is almost exclusively known from the early murgabian (neoschwagerina simplex zone and its equivalents) in the tethyan regions (e.g., leven 1967, 1993, 1997, ozawa 1970; ozawa & kobayashi 1990; ueno 1992). accordingly, sample a is thought to be early murgabian in age. on the other hand, the correlation of early murgabian with the global middle permian time scale is a subject of dispute due to more or less different correlation schemes among authors based on zonations of fusulines, conodonts, and ammonoids. its correlation to late roadian (early guadalupian) is adopted by menning et al. (2004) and to early roadian by wardlaw et al. (2004). early murgabian age of sample a suggests that verbeekina erki n. sp. is the earliest among the known species of the genus. because almost all species of verbeekina are restricted to the middle and upper part of the murgabian except for a few species ranging up to the midian. occurrence of dunbarula and rauserella?, rarely present in sample a is not conflict with this age determination of the sample. because both genera first appear in the cancellina nipponica zone (upper kubergandian) in the akasaka limestone of japan (kobayashi’s unpublished data) and in the parafusulina yabei zone (upper kubergandian to lower murgabian) in the nabeyama formation of japan (kobayashi 2006a; 2006b), and range up to the midian (e.g., kobayashi 2006c). sample b is also thought to be coeval with sample a because of their nearly the same stratigraphic level in spite of different faunal composition between them. these lines of evidence lead to the conclusion that the questionable midian age assigned to the kizilgeris member of the çekiç dagi formation by altiner & özgül (2001) should be revised to early murgabian. this revision suggests that the undoubted lower fig. 2 the columnar section of the çekiç daji formation along the bold line shown in fig. 1. the stratigraphic level of the sample corresponds to the lower part of the kizilgeris member (modified from altiner & özgül 2001). 42 kobayashi f. & altiner d. middle permian is present in the hadim area of southern turkey. systematic paleontology suborder fusulinina wedekind, 1937 superfamily fusulinoidea von möller, 1878 family schubertellidae skinner, 1931 subfamily boultoniinae skinner and wilde, 1954 genus dunbarula ciry, 1948 dunbarula protomathieui n. sp. pl. 2, figs 40-43 origin of the name: ancestral form of dunbarula mathieui ciry. type series: holotype d2-032813 (pl. 2, fig. 40; tangential section). paratypes d2-032784 (pl. 2, fig. 41; tangential section), d2032773 (pl. 2, fig. 42; oblique section), d2-032780 (pl. 2, fig. 43; parallel section). material: two tangential, one oblique, and one parallel sections illustrated. diagnosis: ellipsoidal to short fusiform dunbarula having a minute proloculus, a few lenticular juvenile whorls succeeded by rapidly expanding fusiform whorls, closely spaced septa folded weakly in tunnel region and more intensely in axial and polar regions of outer whorls, and indistinct mural pores. description. test ellipsoidal to short fusiform with broadly rounded periphery, slightly convex to straight lateral slopes, rounded to bluntly pointed poles, and straight axis of coiling. mature test with five to six whorls, about 1.1 mm in length and 0.8 mm in width. proloculus minute and inner few whorls discoidal to lenticular and tightly coiled. with a sharp change of axis of coiling, the following ellipsoidal to fusiform whorls rapidly increasing their length and width. wall consists of tectum and translucent layer with indistinct mural pores in outer fusiform whorls variably coated by secondary deposits. septa closely spaced, partly very finely perforated, also coated by secondary deposits, and folded weakly in the tunnel region and more intensely in axial and polar regions. chomata one-third to half as high as chambers in inner fusiform whorls, but indistinct in outer ones due to secondary deposits on chamber floor in tunnel region. discussion. in the absence of axial sections, morphologic variation of this new species is not totally clarified. however, this species is distinguished from the known species of dunbarula in having diagnostic characters such as finely perforate wall and septa rather thick for the genus. in addition, smaller test and more weakly folded and smaller number of septa suggest that this new species represents the probable ancestral form of dunbarula mathieui, originally described by ciry (1948) and subsequently by thompson (1954) and skinner & wilde (1967) from tunisia (fig. 3.1-3.4). similarly, this new species has more primitive test characters than the specimens of d. mathieui recovered from the midian yellice member in the hadim area (figs 3.5, 3.6) in the test size, and the number and folding of septa. septal pores and mural pores are not prominent in this new species as in d. mathieui. stratigraphic evidence in the hadim area agrees with the phyletic relation from dunbarula protomathieui to more advanced dunbarula mathieui. this new species differs from dunbarula nana described by kochansky-devidé & ramovš (1955) from the murgabian to midian of slovenia in having larger and more rapidly expanding test with more rounded poles, and more intensely fluted septa. primitive forms of dunbarula are common in the lower murgabian limestone of japan (e.g. kobayashi 2006b). this new species is easily distinguished from these forms by its thicker wall and septa, larger test with more rounded poles, and more intensely fluted septa. occurrence. rare in sample a in association with presumatrina ciryi n. sp., verbeekina erki n. sp., and others. plate 1 fig. 1 algal fusuline packstone, sample a, × 8.5. fig. 2 bioclastic packstone containing many foraminifers and fossils of unknown affinity (light part in the upper and middle parts), sample b, × 6. figs 3-5 microproblematica referable to pseudovermiporella sodalica, sample b, all × 30. fig. 6 indeterminate miliolina and pseudovermiporella sodalica contained in the bioclastic packstone, sample b, × 15. figs. 7, 8, 9(?), 10-14 hemigordius discoides lin, li & sun. 7: d2032800; 8: d2-047880; 9: d2-047879; 10: d2-047876; 11: d2-032813; 12: d2-032816; 13: d2-032798; 14: d2032797, all sample a, 7 and 11: × 60; others: × 50. fig. 15 hemigordius sp. d2-032802, sample a, × 40. figs. 16, 18-21 multidiscus sp. 16: d2-032792; 18: d2-032792; 19: d2-032805; 20: d2-047880; 21: d2-032805, all sample a, × 40. fig. 22 pachyphloia robusta k. m. miklukho-maklay. d2032765, sample b, × 60. fig. 23 langella ? sp. d2-047875, sample a × 50. fig. 24 pachyphloia schwageri sellier de civrieux & dessauvagie, d2-032789, sample a, × 50. fig. 25 pachyphloia sp. d2-032814, sample a, × 40. figs. 26, 31, 33, 34 nodosinelloides sp. 26: d2-047880; 31: d2047854; 33: d2-047878; 34: d2-032827, 31: sample b; others: sample a, all × 50. fig. 32 geinitzina? sp. d2-032800, sample a, × 50. figs. 27-30 indeterminate miliolina 27: d2-047844; 28: d2-032766; 29: d2-047862; 30: d2-047863, all sample b, × 30. discovery of lower murgabian (middle permian) in the taurides, southern turkey 43 44 kobayashi f. & altiner d. family verbeekinidae staff & wedekind, 1910 genus verbeekina staff, 1909 verbeekina erki n. sp. pl. 3, figs 23-28 origin of the name: in honor of the fusuline paleontologist who established the genus reichelina based on materials from bursa, northwestern turkey. type series: holotype d2-032775 (pl. 3, fig. 23; axial section). paratypes d2-032799 (pl. 3, fig. 24; tangential section), d2-032774 (pl. 3, fig. 25; axial section), d2-032775 (pl. 3, fig. 26; tangential section), d2-032825 (pl. 3, fig. 27; parallel section), d2-032812 (pl. 3, fig. 28; oblique section). material: two axial, two tangential, and one parallel and one oblique sections illustrated. diagnosis: a primitive form of verbeekina having a small test and less number of juvenile whorls for the genus, and low and welldeveloped parachomata in outer whorls. description. test nearly spherical with shallow umbilical depressions and straight axis of coiling. mature test with 12 to 13 whorls, about 4.7 to 5.1 mm in length, about 4.6 to 4.7 mm in width, and about 0.97 to 1.0 in form ratio. proloculus spherical, minute, 0.03 to 0.04 mm in diameter, and 0.036 mm in the holotype. inner two whorls lenticular and very tightly coiled. the next one to one and half whorls become thick lenticular to subspherical with slight change of axis of coiling. beyond fourth whorl, they become nearly spherical with shallowly umbilicated poles. length from the 1st to 12th whorls 0.04, 0.10, 0.17, 0.44, 0.83, 1.24, 1.75, 2.46, 2.94, 3.50, 4.07, and 4.61? mm; width from the 1st to 12th plate 2 figs 1-3 endothyra sp. 1: d2-032810, × 40; 2: d2-032785, × 40; 3: d2-032778, × 50; all sample a. figs 4, 5 palaeotextularia sp. 4: d2-047869, × 40; 5: d2-032788, × 30; both sample a. figs 6-10 climacammina sp. 6: d2-032804; 7: d2-032787; 8: d2047879; 9: d2-032793; 10: d2-047875, all sample a, × 20. figs 11-17 globivalvulina sp. 11: d2-047874; 12: d2-047874; 13: d2-032770; 14: 47880; 15: d2-032798, × 50; 16: d2047868, × 50; 17: d2-032783, all sample a, 13 and 17: × 40, others: × 50. figs 18-20, 23-26 sphaerulina sp. a 18: d2-047858; 19: d2-047866; 20: d2-032763; 23: d2-047863; 24: d2-047862; 25: d2047859; 26: d2-047863, all sample b, 20: × 60, others: × 40. fig. 21 nankinella sp. b d2-032825, sample a, × 40. fig. 22 nankinella sp. a d2-032764, sample b, × 15. figs. 27, 28 sphaerulina sp. b 27: d2-047863, × 40; 28: d2-032769, × 30; both sample b. figs. 29-31 schubertellidae gen. & sp. indet. a 29: d2-032778, × 40; 30: d2-032781, × 50; 31: d2-032779, × 40; all sample a. figs. 32, 34-36 schubertellidae gen. & sp. indet. b 32: d2-032800, × 40; 34: d2-047876, × 50; 35: d2-032796, × 40; 36: d2032788, × 50; all sample a. fig. 33 rauserella? sp. d2-032814, sample a, × 40. fig. 37 dunbarula sp. d2-032806, sample a, × 40. figs. 38, 39, 44-51 neodiscus spp. 38: d2-047859; 39: d2-032794; 44: d2-032777; 45: d2-047863; 46: d2-032776; 47: d2047851; 48: d2-032774; 49: d2-047843; 50: d2-047851; 51: d2-032780; 38, 39, 45, 47, 49, 50: sample b; 44, 46, 48, 51: sample a; 38, 39, 45, 47, 50: × 50; 44, 46, 48, 51: × 60; 49: × 40. figs. 40-43 dunbarula protomathieui n. sp. 40: tangential section of the holotype, 41: tangential section of the paratype, 42: oblique section of the paratype, 43: parallel section of the paratype. all sample a, × 40. registered numbers are shown in the description part. fig. 3 six specimens of dunbarula mathieui ciry for comparison with dunbarula protomathieui n. sp. 1, 2: types from tunisia (after ciry 1948); 3: topotype (after skinner & wilde 1967); 4: topotype (after thompson 1954); 5, 6: specimens from the yellice member of the çekiç dagi formation in the hadim area. all × 30. discovery of lower murgabian (middle permian) in the taurides, southern turkey 45 46 kobayashi f. & altiner d. whorls 0.08, 0.15, 0.17, 0.40, 0.65, 1.10, 1.63, 2.19, 2.78, 3.36, 3.96, and 4.42? mm; form ratio from the 1st to 12th whorls 0.50, 0.67, 1.00, 1.10, 1.28, 1.13, 1.07, 1.12, 1.06, 1.04, 1.03, and 1.04?, respectively, in the holotype. septa gently inclined anteriorly, long, and partly in contact with parachomata. wall structureless in inner three whorls, consisting of tectum and protheca in the 4th and 5th, and of tectum and very finely alveolar keriotheca beyond the 6th. thickness of wall from the 1st to 12th whorls 5, 8, 8, 16, 18, 17, 20, 22, 33, 37, 43, and 45 microns in the holotype. parachomata low, well developed in the 3rd or 3rd to 5th whorls, rudimentary or absent in the succeeding ones, and well developed in further outer whorls. discussion. test characters such as the size and expansion of the test, and the development of parachomata suggest that the new species is close to a transitional form between armenina and verbeekina. it is better assignable to verbeekina than to armenina in its thinner wall and lesser developed parachomata. among described species, verbeekina erki n. sp. is most similar to v. furnishi skinner & wilde originally described from the upper murgabian of sicily (skinner & wilde 1966), subsequently reported from the murgabian to midian of the central afghanistan (leven 1997) and from the middle to upper murgabian of the abadeh formation of iran (kobayashi & ishii 2003). however, this new species has a smaller test, fewer juvenile whorls and more developed parachomata. verbeekina grabaui thompson & foster described from the middle murgabian of sichuan in south china (thompson & foster 1937) resembles this new species, but the former has more whorls, a thinner wall, and lesser developed parachomata. occurrence. rare in sample a and found in association with presumatrina ciryi n. sp. and others. family neoschwagerinidae dunbar & condra, 1927 subfamily sumatrininae silvestri, 1933 genus presumatrina tumanskaya, 1950 presumatrina ciryi n. sp. pl. 3, figs 1-22 origin of the name: in honor of the fusuline paleontologist who established the genus dunbarula. type series: holotype d2-032818 (pl. 3, fig. 2; axial section). paratypes d2-032785 (pl. 3, fig. 1; axial section), d2-032797 (pl. 3, fig. 3; axial section), d2-032815 (pl. 3, fig. 4; axial section), d2-032781 (pl. 3, fig. 5; axial section), d2-032800 (pl. 3, fig. 6; axial section), d2-032783 (pl. 3, fig. 7; axial section of the microspheric form), d2032796 (pl. 3, fig. 8; axial section), d2-032793 (pl. 3, fig. 9; axial section), d2-032802 (pl. 3, fig. 10; axial section), d2-032814 (pl. 3, fig. 11; axial section), d2-032794 (pl. 3, fig. 12; axial section), d2-032784 (pl. 3, fig. 13; axial section), d2-032777 (pl. 3, fig. 14; sagittal section), d2-032801 (pl. 3, fig. 15; axial section), d2-032810 (pl. 3, fig. 16; axial section), d2-032776 (pl. 3, fig. 17; axial section), d2-032808 (pl. 3, fig. 18; sagittal section), d2-032778 (pl. 3, fig. 19; sagittal section), d2032820 (pl. 3, fig. 20; sagittal section), d2-032795 (pl. 3, fig. 21; sagittal section), d2-032791 (pl. 3, fig. 22; sagittal section). material: fifteen axial and six sagittal sections of megalospheric forms and one axial section of microspheric form illustrated. diagnosis: a large-sized species of presumatrina with broadly rounded poles and large proloculus in megalospheric forms, well developed secondary transverse septula that first appear in the fourth whorl, thin wall and septa, and low and massive parachomata. description. test ellipsoidal, with broadly rounded poles and straight axis of coiling. mature specimens with 8 to 9.5 whorls, about 3.5 to 4.4 mm in length, about 2.1 to 2.8 mm in width, and about 1.5 to 1.9 in form ratio in megalospheric form; with 12 whorls, about 5.5 mm in length, about 2.7 mm, and about 2.0 in form ratio in the microspheric form (tab. 1). proloculus spherical to subspherical and 0.18 to 0.27 mm in megalospheric form, and 0.02 mm in microspheric form. the first whorl subspherical to short fusiform, then becoming short fusiform to ellipsoidal. test expands gradually decreasing their form ratio and increasing roundness of poles outwards. length from the first to 9th whorls 0.32 to 0.45, 0.64 to 0.86, 1.12 to 1.42, 1.60 to 2.03, 2.04 to 2.61, 2.57 to 3.13, 3.02 to 3.70, 3.51 to 4.14?, and 3.98 or 4.1? mm in illustrated 13 specimens of megalospheric form. length from the first to 12th whorls 0.03, 0.11, 0.25, 0.44, 0.80, 1.29, 1.94, 2.75, 3.36, 4.12, 4.87, and 5.5 mm in microspheric form. width from the first to 9th whorls 0.25 to 0.37, 0.34 to 0.55, 0.51 to 0.74, 0.70 to 1.02, 0.93 to 1.35, 1.20 to 1.71, 1.49 to 2.07, 1.80 to 2.25, and 2.18 to 2.62 mm in illustrated 19 specimens. width of the first to 12th whorls 0.06, 0.12, 0.20, 0.26, 0.37, 0.54, 0.78, 1.07, 1.42, 1.83, 2.24, and 2.7? mm in microspheric form. wall thin for the test size and less than 0.05 mm in the thickest part of outer whorls, mostly structureless in the first and second whorl, and consists of tecplate 3 figs. 1-22 presumatrina ciryi n. sp. 2: axial section of the holotype; 1, 3-13, 15-17: axial sections of paratypes; 14, 19-22: sagittal sections of paratypes. 7: microspheric form; others: megalospheric forms, all sample a, 7b: × 30; others: × 10. registered numbers are shown in the description part. figs. 23-28 verbeekina erki n. sp. 23: axial section of the holotype; 24, 26: tangential sections of paratypes; 25: axial section of the paratype; 27: parallel section of the paratype; 28: oblique section of the paratype, all sample a, 23b: × 30; others: × 10. registered numbers are shown in the description part. discovery of lower murgabian (middle permian) in the taurides, southern turkey 47 48 kobayashi f. & altiner d. tum and very finely alveolar keriotheca from the third whorl. septa thin and closely spaced. septal counts from the first to 9th whorls 5 to 7, 11 to 13, 14 to 16, 15 to 17, 17 to 21, 20 to 23, 23 or 24, 24 to 27, and 29 in six megalospheric sagittal sections illustrated. primary transverse septula long, slender, and well developed and present in all whorls. secondary transverse septula short and slender, and first appear in the fourth whorl in most specimens. one secondary transverse septulum, rarely two secondary transverse septula, inserted between adjacent primary transverse septula. axial septula absent in inner few whorls, poorly developed in the succeeding one or two whorls, and relatively well developed, but short for the width of outer whorls. one or two axial septula present between adjacent septa in outer whorls. parachomata, partly in contact with septa in middle and outer whorls, low, massive, node-like, and well developed from the second whorl and also present on the proloculus in some specimens. discussion. this new species is easily distinguished from the previously described species of presumatrina in its shape and size of the test, proloculus size, and development of secondary transverse septula and parachomata. it is different from presumatrina schellwieni (deprat) and p. neoschwagerinoides (deprat) in having a larger test with much more rounded poles, a larger proloculus, and better developed secondary transverse septula. presumatrina grandis leven described from the lower murgabian of the southeast pamir (leven 1967) seems to be the closest taxon to presumatrina ciryi n. sp. among the described species. however, the former has more whorls and lesser developed secondary transverse septula than the latter. further comparison is impossible since the morphologic variation of p. grandis is uncertain from two specimens illustrated by leven (1967). presumatrina uruzganensis leven described from the lower murgabian of the bamian zone of northern afghanistan (leven 1997) has a smaller test and proloculus, more whorls, smaller proloculus, better developed secondary transverse septula, and higher and more massive parachomata. with respect to lesser developed secondary transverse septula and axial septula, this new species is distinguished from presumatrina ozawai, originally described by hanzawa (1954) from the akiyoshi limestone and the index species of the upper part of the parafusulina kaerimizensis zone (lower murgabian) of the akiyoshi terrane (kobayashi 1988; ozawa & kobayashi 1990; ueno 1992). occurrence. common in sample a and found in association with verbeekina erki n. sp. and others. acknowledgements. we are grateful to drs. maurizio gaetani, katsumi ueno and ernst ja. leven for their critical review of the manuscript, and to mrs. atsuko ujimaru for her help drawing figures. funds for travel expenses in turkey for the senior author were provided through the grant-in aid for overseas scientific research from the hyogo science and technology association in 2004. tab. 1 measurement of presumatrina ciryi, n. sp. pl. 1, fig. 7: microspheric form; others: megalospheric forms. discovery of lower murgabian (middle permian) in the taurides, southern turkey 49 altiner d. 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(���"�� %��� c���. >?2�3� > a�> ? �$��. � ''� �. ��1��"���� �. �::�' �� �. 4�!!�'���� �. ��,� �. ���� "� �. � ��,'� �. 2 cc�3 � ���� *�,"���! !�� ��1'��� " # �(� ���� &����#'�"!(��" *�**�� +"��$ #' * �(� �--�' &����'" !�"�'�� ����%. �� ��� %��� 0 . �c� ��a���� � *�. � ''� �. ��1��"���� �. �'��"� �. 4�!!�'���� �. ��$�"� 4. � ���� "� �. 2 cc=3 � ���� *�,"���! !���1'��� "$ # ��� �����$� !�"� *�**�� � !������$ �" !�"�'�� ����%. �� � @��� 5�����. �c2 3� aac�a>@ l'�6 ). � '� "�$� �. 2 c>@3 � �$���!(�(%�$ 2��$!� $$��3. ��'�� -'�� *�. �"� 7����'�"� 2��.3 � 4�+"� �9������ @� �a=a � ,"�. � '� "�$� �. 2 c>a3 � �$���!(�(%�$ 2��$!� $$��3. ��'�� $�� ! "��. �"� 7����'�"� 2��.3 � 4�+"� �9������ � �=�= � ,"�. �+"1'��,� �. �. 2 c?�3 � 4�!��$ * ��� # ' � $�"�% �-(�*�'�� $�'��* �"� !��% -��%� ! *-��ed �(� ������ ��/ "��" �'�"��$( � 4 '*��� " # � '�( ��/ " �.l.. ����$ �� ���"� � � =c>�> a �e# '�. &�" ��' ��+��" �. �. 2 c>�3 � ������ ����$� !�"� �*����' ��**��$ #' * �(� � "�� ��,��� 2�'/��� ����%3 )��( �-�!��� ��#�'�"!� � �(� �(%� ,�"% # ����� �� 2�'� /�! ����� � ��"���3. ?�� ������� >� � �� �*$��'� ��*. &�" l �#$!( ��" �(. /�" ��' ��+��" �.�. �"�,6 g. 2 cc?3 � �(� ���� ��� !�"� � ��"�$ 2��**����3 #' * 4'�!(�" 2� )�' �(�"� �$�" ��'*�"%3. �"� &�" l �#$!( ��" �( � ��11�'� �.�. 2��$.3 �(� ��)" # �(� 0+���'"�'%. ������ :����� 0�� 1��" d� 1:, =@� = � > �����". &�'� � �. 7(���"� �. � �!(��# �. 2 cc@3 � 0+�"��#�!��� " �+ ,'��+���$*� -(%��i��5+� �� �� � �� ���� ���� �8 �� � �� �� �� ������� 2�'/�! ����� � ��"���3 �8 �9���� �� �9�"��%$� �9�*�,�$. 5 � ���� ��� 3���� � @ ��i'�� ��� >aa� >=@ ��'�$. ��,)�b" g. �. 2 c>�3 � �(� ��� !�"������$� !�"� 1 +"��'% �" )�$��'" �"� $ +�(�'" �+' -�. /����� �� >a�c> �p '� (+$ 7. rivista italiana di paleontologia e stratigrafia volume 104 numero i tavole 1 paglne r t)-i_l aprile 1998 noa bre'ry a phytosaur skull from ti1e*ry_9rian (late triassic) of lombardy (northern italy) slvio renestox s. anna paganoni** receitled june 17, 1997; dccepted noaember 12, 1997 key-uords: phytosauria (reptilia, archosauria); isolated skull; description; norian (late triassic), lombardy, nonhern italy. riassunto. viene descritto per la prima volta un cranio isolato di un rettile fitosauro, rinvenuto nel calcare dt zorzino, di età norica (triassìco superiore) nella località di endenna nelle prealpi lombarde. il cranio, per il resto completo, è privo della mandibola ed è stato notevolmente deformato nella sua pane posteriore. è ,t"t" trrtt"_ via possibile una descrizione che ha consentito di attribuirlo con ra_ gionevole cefîezz^ al genere mystriosuchus, probabilmente à4. planiro. srrls, già noto da eccellenti esemplari della formazione tedesca dello stubensandstein. questo cranio rappresenta uno dei rari retrili di grosse dimensioni rinvenuti nelle località fossilifere del norico lombardo, essendo stati rinvenuti solamente un esemplare completo di fitosauro lungo 4 metri, ancora in preparazione e un esemplare del rettile placodonte pseboderma alpinum lungo circa due metri. abstract. an isolated phjtosaur skull found in the calcare di zorzino (norian, late triassic), near rhe locality of endenna (berga_ mo prealps, lombardy, nonhern italy), is described. the skull lacks the mandible and is severely compressed and distoned in its posterior portion. nevertheless it is possible to ascribe it to the gews mystrio. sucbus, likely to mystriosucbus planirostris, already known on the basis of excellent specimens from the stubensandsrein formatron in germany. this finding represents one of the rare large reptile specimens found in the norian fossil-bearing localities of lombardy, along with a new phytosaur yet to be prepared and a nearly two meters long specimen of the placodont psephoderrna alpinum. introduction. the norian (late triassic) formations of calcare di zorzino (zorzino limestone) and argillite di riva di solto (riva di soito shale) crop our exrensiveiy in northern italy. some localities are fossiliferous and have yielded a rich vertebrate fauna of great scientific interest. the richest fossiliferous unit is the calcare di zorzino, deposited in intraplatform basins some hundreds of meters deep and several kilomerres wide, surrounded by the huge dolomia principale (hauptdoiomit) carbonate platform. the centre of these basins was anoxic, while the margins were in the oxic environment (tadoul et. al., 1992) and sustained a rich vertebrare and invertebrate life. fossil findings consist mainly of fishes and are characterised by thousands of superbly preserved specimens, representing a well differentiated communiry, (tintori, 1992), with also some previously unknown genera (tintori & lombardo, 1996). reptiles are much rarer, albeir of great interesr. aquatic reptiies include the first complete specimens of the placo dont psephoderma alpinum meyer (pinna, 1979, pinna & nosotti, 19g9, renesto & tintori, 1995) and the thalattos aur endenna_ saurus (renesto, 1984, 1992). despire rhe depositional environment, terrestrial reptiles are more common than marine forms, offering an insight into rhe complex faunal community living on the emerged areas surrounding the basins, possibly islands with freshwarer reservoirs (tintori et. a1., 1985). the oldest prerosaurs described so far (vild, 1928; renesto, 1993) .were found associated with fragments of the armour of the archo sallr aetosdurus (wil.d, 1991). other finds are rare diapsids, often unknown from other localities, like rhe arboreal genera megalancosaurus (calzavara ef ai, 1980; renesto, i994a) and drepanosaurus (pinna, 1980, 1984; renesto, ig94b). two small prolacertiform reptiles related to the ranystropheids and belonging ro rhe new genus langobard.isau',4s (renesto, 1994c) were found along with a tiny specimen of the sphenodontid diphydontosaurus (renesto, 1995a). most of these reptiles are of small or medium size, excluding the large specimen of psephoderma (renesto & tintorl, 1995), nearly t.wo meters long, and a newly found complere phytosaur, currently under preparation, that reaches nearly four merers in length (tintori et al., t996). the isolated skull described here be_ iongs aiso to a large phytosaur, as characterised by the elongate, crocodile-like snour, with a long rostrum formed principally by the premaxillae, and by the partern 'r dipartimento di scienze della terra dell'università degli studi di milano, via mangiagalli 31, 20133 mrlano, italy; e-mail: renesro@imiuccr.csi.unimi.it 'r'r museo civico di scienze naturali "e. caffi", piazza cittadella, bergamo, italy. 116 s. renesto & '4. paganoni of other skull bones. phytosaurs are archosaurs well known from several triassic fossiliferous localities, mainiy in the nofthern hemisphere, europe, usa and india (i4cgregor, 1906; gregory 1962; chatterjee, 1978, ballew; 1989; hunt e lucas, 1989; doyle & sues, 1995) but known also from fragmentary remains from north africa (chatterjee, 1978) and turkey (buffetaut et. a1., 1988). the phytosaurs were carnivorous reptiles of late triassic age, which resembled crocodiles both in general shape and possibly in life habits. they are usually consi dered to be associated with freshwater environments, but it cannot be excluded, at least in the case of our finding, that occasionally they might have visited the seashore, as do some modern crocodiles. alternatively, it may be hypothesised that the isolated skull belongs to a floating carcass that was swept away by water currents (renesto 1995b).it is worth mentioning, though, that other findings of phytosaur remains have been reported from brackish environments (\íestphal, i976; buffetaut, 1ee3). the skull described here was found several years ago in the fossiliferous quarry of endenna, near zogno (bergamo, lombardy, northern italy), but it was only mentioned incidentally by pinna (1987) as possibly belonging to the genus mystriosuchws and no description was provided. buffetaut (1993) also mentioned the specimen under the name mystriosuchus planirostris, but did not provide any evidence for this identification. at the time the fossil was exposed only on its ventral side. further preparation by mr. m. pandolfi, on staff at the museum of natural history of bergamo, revealed the dorsal side, allowing a complete description and a positive generic attribution, despite the fact that many of the bones of the postnarial region were flattened and crushed. systematic palaeontology class reptilia superdivision neodiapsida benton, l985 division archosauromorpha fiuene, 1946 subdivision archosauria cope, 1869 order phytosauria meyer, 1861 family pbytosauridae meyer, 1861 genus mystriosuchus fraas, 1896 mystriosuchus planirostris (meyer), 1863 pl. 1 1.863 belod.on planirostris von meyer, p. 244 7896 mystriosuchus planirostris-f raas, p.75-17 19a6 mystriosuchus planirostis -mcgregor, p. 36 material. an isolated sku1l, 21.5 cm long from the tip of the snout to the occipital condyle. the skull is transversely broken into three pieces, lacks the mandible and is flattened dorsoventrally so that many bones of the postnarial region are crushed or overlapped. funhermore, the antorbital fenestrae are obliterated. the left ponion is the best preserued and protides the basis for most of the description. repository. museo civico di scienze naturali "e.caffi", bergamo (lombardy, nonhern italy). curator dr. anna paganoni. catalogue number mbsn 2. provenance. ijppermost level of the calcare di zorzino (zorzino limestone), norian age (upper triassìc), in the small quarry of endenna, near zogno (bergamo prealps, bmbardy, nonhern ltaly). measurements (in cm). length of the skull from the tip of the snout to the occioital condvle lenght of the rostrum number of alveola on the left premaxilla number of alveola on the left maxilla 7t.5 54 21 20 description. the premaxillae (pl. 1) are long, slender and narrow, only slightly expanded at the anterior tip, which is slightly bent ventrally. the entire outline of the skull bears a gavialjike appearance. the dorsal surface of the premaxillae is smooth and the snout lacks crests or ridges. both show two medial rounded ridges on the ventral side, separated by a trough ìn the middle. according to mcgregor (1906, p. 38) these features "prevent the close approximation of upper and lower alveolar region and thus serve to prevent the breaking of the teeth when the jaws are forcibly closed". a cavity is visible in cross section along the greàfer part of the rostrum. the left premaxilla bears 21 rounded alveoli which are spaced, suggesting that the dentition was unserrated. in some cases the margin of the alveolus has a 1ow lateral crest. the teeth are aimost completely lost, apart for some elements on the left premaxilla. fragments of the first tooth, which had to be among the largest, are visible, but its size, judging from that of the alveolus, was not too different from that of the other teeth. the 18th tooth, a very small replacement tooth, which bears a faint posterior carina and delicate fluting, is preserved. the last preserved tooth, the 21st, is crushed, so that only its rounded cross section may be observed. the median margins of the dorsal processes of the premaxiliae are very extended posteriorly, nearly reaching the anterior border of the external nares. the maxillae are extremely difficult to reconstruct, because their dorsai region is crushed and split, and the dorsal outline m y not be observed; the antorbital fenestra also is obliterated. about 20 rounded, spaced alveoli are visible on the left maxilla, and their size decreases gradually toward the posterior end of the bone. no teeth are preserved. pl. 1 a plrytosaur skull from the norian of lombardy plate 1 fig. 1 mys*iosucbus planirostk, isolated skull; a) dorsal, b) ventral views. the entire length of the sku1l is 21.5 cm. 118 s. renesto & a. paganoni mystriosuchus planirostris, schematic restoration of the pattern of the post rosrral region of the skull, from dorsal view. scale bar equals 10 cm. fractures were omitted for clarity. abbreviations are: bo) basioccipital; f) frontal; .j) juga1, l) lacrimal; mx) maxilla; na) nasal; opo) opisthotic; pa) parietal; pmx) premaxilla; po) postorbital; pof) postorbitofrontal; prf prefrontal; qj) quadratojugal; so) supraoccipital; sq) squamosal. the septomaxillary is difficult ro see, due to the shifting and crushing of the bones. the nasals, on the contrary, seem to form most of the margin of the narial openings and are sculptured with grooves and pits in their posterior portion. their posterior margin meets the prefrontals, the frontals and the wide lacrimals. the narial openings are anteroposteriorly elongate, open anterodorsally and raised with respect to the dorsal profile of the snout, but they do not raise over the level of the skull roof. the frontals are heavily sculptured and seem to be longer than wide, but their suture with the parietals is not visible, while the sagittal suture berween the two bones is clearly detectable. a rounded postfrontal forms, along with the frontal, the posterodorsal margin of the orbit. a iarge postorbital is presenr ventrally to this bone. the parietals seem to be small and rugose in their dorsal portion. the interparietal suture is persistent and a pineal foramen is absent. posteriorly they contribute, along with the squamosal, to the greatly depressed and reduced upper temporal arcade that is distinctive for the genus (mcgregor 1906; gregory, 7962; ballew, 1989; hunt & lucas, 1989). the squamosals show a strongly scupltured dorsal portion and form the outer part of the highly modified post-temporal arcade. the supratemporal bar is formed by postorbital and squamosal that becomes flattened dorsoventrally and moderately stout and wide mediolaterally. a ridge divides rhe squamosal into lateral and dorsal portions. the posterior process of the squamosal is short and not pointed as in pseudopalatus (hungerbùhler, pers. comm.). a wide lacrimal meers rhe large jugal anteriorly. the latter bone bears a posteroventral process that forms most of the ventral margin of the large, lower temporal fenestra and is in contact with the quadratojugal. the dorsal process of the jugal separates the orbit from the lower temporai fenesrra and contributes to the formation of the ventral margin of the orbit. the quadratojugal is subtriangular, ir conracrs rhe quadrate and overlaps part of the squamosal. its surface is smooth. the ventral surface of the skull is badly crushed and many bones are reduced to splinters, making it very difficult to reconstruct their outlines. for this reason this surface is only superficially described and it is not illustrated. palatine bones" the anterior portion of the narrow; elongate vomers is visible, while the choanae are obliterated by bone fragments and are not detectable. the paiatines are wide and smooth, and are in contact a plryrosaur skull from the norian of lombarch medially with the large pterygoids. part of the left ectopterygoid is preserved. the basisphenoid is heavy and sturdy strong in its distal portion; the junction vzith the basioccipital is strong and no surure line is visible. occipital bones. the foramen magnum is oblirerated but it can be reconstructed from the pattern of the surrounding bones. a small triangular supraoccipital is surrounded dorsaily and laterally by the parietals, forming an inverted ij strucrure (ballew; 1989; hunt & lucas, 1989). its surface is slightly concave and rugose for the attachment of muscles. the two exooccipitals are strong and contribute also to the formation of the occipital condyle. they continue lareroposreriorly into the long and stour paroccipital processes of the opisthotic, but no suture is visible between these bones. the paroccipital processes of the opisthotic are surrounded distally by the squamosals and support dorsally the greatly depressed parieto-squamosal arcade. the posttemporal fenestra is reduced to a smail slit. the balllike basioccipital forms most of the condyle. discussion. the characteristics observed in the specimen show that mbsn2 belongs to the phytosauridae (intended as the group including the last common ancestor of angistorhinus, mystriosucbus, pseudopalatus, nicrosaurus and rutiodon) as diagnosed by ballew (1989) and doyle & sues (1995). the diagnostic characteristics visible in mbsn2 are the exrernal nares facing dorsally and the squamosal with posterior and hooklike ventral process. according to gregory (962) and hunt & lucas (1989) the following characteristics suggesr that mbsn2 belongs to the genus mystriosucbu.s: unserrated and probably homodonr teeth that decrease in size posteriorly, heavily sculptured skull roof region and greatly depressed, compressed, supratemporal fenestra, inverted u shaped parietal-supraoccipital complex, short and not pointed posterior process of the squamosal, supratemporal openings that are deeply incised inro the sku1l roof and nares which are not raised above the level of the skull roof. poor preservation of the region posterior to the nares prevents a detailed comparison with other mystriosuchus skulls, resulting in a tenrative specific assignment to mystriosuchus planirostris. "m. plieningeri", specimen gpit stored at the institur und museum fùr geologie und palàontologie, university of tùbingen, number 264/001 (f{uene, 7922, pls. i2-i4) may well represenr a different species, blrt according to fiunt & lucas (1989) it shows, among other differences with m. planirostris, external nares depressed well below the skull table, a characteristic not observed in mbsn2. on the other hand this trait is questioned by iong u murry (1995) who state that the nares of specimen gpit are not significantly more depressed than in m. planiro.srds. f{owever, specimen gpit has also a much more massive snout (hunt & lucas, 1989), iarge posterior maxillary tooth sockets (huene, 1911), a shorter prenarial snout and a ventrally bulging rim of the maxillary (long & murry, 1995), all characters that are different from those of mbsn 2. the classification of mbsn 2 as mystriosuchus planirostris, typical of the middle norian stubensandstein fauna, confirms rhe age determination of the locality, based on fragments of aetosaurus (wild, 1991). palaeoecology. the elongate, cylindrical snout with conical teeth indicates a piscivorous diet as already pointed our by hunt (1989), even if an occasional reptile prey cannor be exciuded, as reported by chatterje e (i97g) for the long-snouted species parasucbus hislopi. mystriosucbus is considered linked, like other phyrosaurs, ro a conrinental, freshwater environment, and therefore the finding of mbsn2 amid a marine basin has to be explained. the finding of such isolared remains of animals, common in coeval continental deposits such as the stubensandstein, like the aetosaurus scutes and mystriosuchu-s skull can be explained as represenring sunken parts of decaying carcasses coming from distant life environmenrs, as hypothesised by renesto (1995b). fiowever, the recent finding of a new, iarge (4 merer in length) phytosaur (tintori et al., 1996), possibly another mystriosucbus, requires changing this scenario. the skeleton of this new specimen is virtually complete and articulate and suggests thar rhe life environment of these animals might have been closer to the basin, and perhaps might have corresponded to the islands (with freshwater reservoirs, tintori et al., 1985) which surrounded the basin and hosted many endemic reptiles. if this is rhe case, ir cannor be excluded that these phytosaurs became acquainted with a marine environment, preying on the abundant fish fauna of these basins. further strength to this hypothesis is provided by the discovery of phytosaur remains (possibly mystriosuchus, buffetaut, 1993) rn the southern part of the totes gebirge in styria (austria). the specimens were found in the norian dachsteinkalk, that is considered as deposited in a shallow marine environmenr. among phytosaurs, mystriosucbu.s is the most adapted to a piscivorous diet, and thus may have been a mosrly aquatic form, which lived in both freshwater and marine environments, like some modern crocodilians. actually, crocodylws porosus inhabits both freshwater and marine environments and can swim for several kilometers in the open sea (ross & magnusson, 1989); mystriosuchus may have done the same. 1,24 s. renesto g a. paganoni references acknouledgements. \le must thank mr. mario pandolfi for the skillful preparation of the dorsal side of the skull, dr. r. \fild, stuttgan and dr. h.-d. sues, toronto, for useful advice and bibliographic help. we are deeply indebted to prof. m. j. benton and dr. a. hungerhbùhler (both of bristol) for critically reading the manuscript and language corrections; our sincere thanks especially to a. hungerbùhler for useful advice that greatly improved the manuscript. thanks are due also to the staff of the museo brembano di scienze naturali imbsn) s. pellegrino (bergamo), where the fossil was formerly stored: p. geruasoni, who found the fossil, the late a. torriani, who directed the museum and f. galizzi, who made the first preparation of the ventral side of the skull. financial support from the museo civico di scienze naturali "e. caffi", bergamo. ballew k. 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�" ������� ��� ����� �%� ���*��"��( �����" b����""��� ��f�� ���f��� ������ �$ ��$ >=-@dm.c�c 0��1� � ����� �� ���)$ ��f�� ����% �%��!" ��� ��� �� �%� � � ��*�� ��"� :�$ ��(���� 5$ 2����� ��� :�$ ������ ���!)���� %�"� �������" ��� %���)�� �� �����*��+ �%� �����$ ����� �( * &��� �(d. p. theobaldi present paper p. palaeindicus colbert 1935 p. porcus colbert 1935 p. magnus arribas & garrido 2008 p. afarensis bishop 2011 p. chinhsienensis li 1963 measurement pua/sk07/94 a.m. 19878 a.m. 53727 fp-2001-0251 ep1058/04 p 4 max. mesio-distal diameter (l) 13.2 15.0 13.0 15.0 max. bucco-lingual diameter (b) 16.2 17.0 15.0 18.1 index (b/l x 100) 123.0 113.3 115.4 120.5 m 1 max. mesio-distal diameter (l) 16.5 19.0 19.0 18.2 16.4 16.0 max. bucco-lingual diameter (b) 15.4 17.0 17.0 19.1 12.6 16.2 index (b/l x 100) 94.0 89.5 89.5 105.1 76.8 101.2 m 2 max. mesio-distal diameter (l) 21.6 26.0 23.0 28.5 27.5 max. bucco-lingual diameter (b) 19.7 23.0 22.0 24.1 21.5 index (b/l x 100) 91.2 88.5 95.6 84.7 78.2 ��&$ � b��������*� ���"�������" 0��1 �) ��'�����( ���������" �) "��� "�����" �) �������������$ � � � � � � � b � � ����&�" �$ ������� �$ 0.cc;1 � �� ��� &��� &����+��+ �� �%� +���" ������������� 0������� ���������(��� ��������1 )��� �%� ����"��� 5��� ����� �������� 0������" �-�� b����� �� �����'� �������1$ %���( 2��( &���� ���� �c� >>@->d,$ �66����� �$ ��������� �$ 0�=;.1 ��+����"�����+���%�� ��*�"��+�����" �) �%� ����� �� ���! 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' ��� +����-�%� +��,)�� -�" )"�* +�� $�,# ����"�%"� �d� i �= 0.#�5h�� -���� 77.88 �" !�� ,��")��# ,�%! �+ � "��$�,�% !� $�,���� 77�88 �" !�� ,��% �+ !�� ,��")��,�%! �+ !�� "!�%*��* "�,��� 0!�� "!�%*��* "�,# ��� �" �% !��" $�"� !�� ���)��!��% +��, ��"��$� �5 77��88 �" !�� "!�%*��* *�(��!��% �+ !�� ,��")��,�%! �+ !�� "!�%*��* "�,���' ��� (��)�" ��!��%�* -��� ���!!�* �% ��"!����," ���*)�!�* �% "�(���� "!�%*��* *�(��!��%" +��, !�� "!�%*��*' ��� "!�%*��* *�,�%"��% �" "�! �! /��� �%* � ,��")��,�%! �%� "!�%*��* *�(��!��% ������ !��% !�� "!�%*��* *�,�%"��% �" ���!!�* �! �= �%� "!�%*��* *�(��!��% ",����� �! #�= 0���*��19<5 0���' �5' ��� ��"!$��%��� ,�!����� �+ ����� ���� � �����# ���� +��, ��"��$� � 0 ���%*�.2 �� �!�+�%� � ��# !��%�� �11�5 -�" )"�* �" "!�%*��* "�,���' ���" $���$� ��" ���% ,�*� ��$�)"� !��! ���)��!��% �" ����� ��,�# ��%��)" �%* +�� +��, !�� �!��� ��,��%" $�%"�*���* +��, !�� ���������$�� �%* $���%�����$�� ���%! �+ (��-' �� ��!!�� ��"��(� !�� "�/� (����!��%" �! � �����%�� ��(�� -� *�(�*�* !�� ��$���!��" �%!� !���� ���)�" �����"�%!�%� !���� *�++���%! ���������$�� ����"� %��!���% �!�� 0��$��# %� &��'��� ("� )�����'� *"� )������� �" + &����� *"�4< site region geographic area age reference isorella lombardy northern italy holocene bon et al. 2005 santorso venetian northern italy holocene cassoli & tagliacozzo 1989 fimon venetian northern italy holocene riedel 1948 val liona venetian northern italy holocene riedel 1948 isolone della prevaldesca lombardy northern italy holocene riedel 1975 ledro trentino alto adige northern italy holocene riedel 1976a barche di solferino lombardy northern italy holocene riedel 1976b colombare di negrar venetian northern italy holocene riedel 1976c pozzuolo friuli friuli venezia giulia northern italy holocene riedel 1983 colognola ai colli venetian northern italy holocene riedel 1984a cladrecis friuli venezia giulia northern italy holocene riedel 1984b appiano trentino alto adige northern italy holocene riedel 1985 grotta d’ernesto trentino alto adige northern italy holocene riedel 1991 canàr polesine venetian northern italy holocene riedel 1998 vadena trentino alto adige northern italy holocene riedel 2002 ancona cappuccini marche central italy holocene wilkens 1989 bachero marche central italy holocene wilkens 1989 capo d'acqua abruzzi central italy holocene wilkens 1989 conelle marche central italy holocene wilkens 1989 grotta continenza abruzzi central italy holocene wilkens 1989 grotta dei piccioni abruzzi central italy holocene wilkens 1989 ortucchio abruzzi central italy holocene wilkens 1989 ripoli abruzzi central italy holocene wilkens 1989 s. maria in selva marche central italy holocene wilkens 1989 grotta s. angelo abruzzi central italy holocene wilkens 1996 osimo marche central italy holocene wilkens 1997 coppa nevigata apulia southern italy holocene siracusano 1995 torre mordillo calabria southern italy holocene tagliacozzo & curci 2001 latronico basilicata southern italy holocene wilkens 1989 punta delle terrare apulia southern italy holocene wilkens 1989 bari s. maria apulia southern italy holocene wilkens 1991 arene candide liguria northern italy mis 2 cassoli & tagliacozzo 1994 riparo tagliente venetian northern italy mis 2 rocci ris et al. 2005 palidoro latium central italy mis 2 this work grotta delle mura apulia southern italy mis 2 bon & boscato 1993 grotta la cala campania southern italy mis 2 boscato et al. 1997 grotta della serratura campania southern italy mis 2 boscato et al. 2005 grotta romanelli apulia southern italy mis 2 tagliacozzo 2003, and this work riparo di fumane venetian northern italy mis 3/2 cassoli & tagliacozzo 1991, and this work grotta del fossellone latium central italy mis 3/2 this work botro ai marmi tuscany central italy mis 3 boscato 1995 grotta di parignana tuscany central italy mis 3 caterini 1921; farina 2009 grotta s.agostino latium central italy mis 4/3 tozzi 1970 ingarano apulia southern italy mis 4/3 this work sora -valle radice latium central italy mis 5 this work avetrana apulia southern italy mis 5 this work ���' � # ��!� ����"!�$�%� �%* ��*�� &���$�%� ��$���!��" �+ �!�� $�%"�*���* �% !��" -��6' �!��" %��!���% !��% !�� �,����%#�)"$�% ��%%�%�"5 $�%!��� �!�� 0��$���!��" +��, !�� �,����%#�)"$�% ��%# %�%�" !� !�� "�)!���% ��!�),5 �%* "�)!���% �!�� 0��$��# �!��" �% �)��� b�"���$�!� �%* ��������5 0���' �5' ��� (����!��% �+ �d� ��" ���% �����"�%!�* +�� �(�� ���� +�� +�)� $���%�����$�� �%!��(��" $����"��%*# �%� !� ��� = ��� ah: 0��� a �%* ��� :5 ��� � �%* !� ��*�� &���$�%�' +�*���"�!" $���%�����$��� ��# +����* ��!-��% !�� �%* �+ ��� : �%* !�� ����%%�%� �+ !�� ��� � ��� �%$�)*�* �%!� !�� ���)� �+ ��� �' ��� ��� �+ !�� ��$���!��" ��" ���% !�6�% +��, ��!���!)�� "�,�# !�,�" ��(��-�* � !�� �)!���" �% !�� ��"�" �+ !�� �%# +��,�!��% �(������� +��, !�� +�)%�� ��"! 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# �� ���!!� ��%!8 %���� ")��� ��%!��%� *�� ����� 0����,�5� �33#�1: ���� ��"�# ��,�' ;��6�%" b' 0�1135 # � ��"!� +�)%�"!�$� *�� ��(���� ��$�%� *� �"�,�' �� " ��� " �a� �:3#�=�' %� &��'��� ("� )�����'� *"� )������� �" + &����� *"��a �������� ���")��,�%!" �+ !�� ��"!#$��%��� ��,��%" �+ ��* *��� +��, !�� ��**�� ����"!�$�%� �+ ��"��$� � 0���,�% 5 "!���* �! !�� �!)���"# !���"$��" �)"�), �+ ���%!/ �%* +��, !�� ��!� ����"!�$�%� �!����% "�!�" �+ �%����%� �%* (�!��%� 0"!���* �! !�� �����!,�%! �+ ���!� �$��%$� �� �����%/� �%�(��"�! �+ ��,�5 ���� ��""����%� ����*��� ���!!� ��,�%���� �%* ������ �),�%� 0"!���* �! !�� �)"�� �/��%��� ����"# !���$� �!%����+�$� �)��� ������%� ��,�5' � i ,�.�,), ��%�!�2 ��� i ���.�,�� !��%"(��"�� *��,�!��2 � �� i ���.�,�� �%!���#��"!����� *��,�!��2 ��� i !��%"(��"�� *��# ,�!�� �+ !�� "��+!2 � �� i �%!���#��"!����� *��,�!�� �+ !�� "��+!2 ��� i *�"!�� !��%"(��"�� *��,�!��2 � �� i *�"!�� �%!���#��"!����� *��,�!��2 &� i ��!���� �����!2 &, i ,�*��� �����!2 �� i ,�.�,), !��%"(��"�� *��,�!��' radius site specimen l ptd papd mtd mapd dtd dapd mosbach mm 1949/61 281.0 53.0 30.0 38.0 21.0 50.0 31.0 mosbach mm 1949/248 288.0 56.0 29.0 34.0 21.0 48.0 33.0 mosbach mm 1956/358 281.0 55.0 29.0 34.0 19.0 46.0 30.0 mosbach mm 1964/529 308.0 60.0 33.0 39.0 25.0 56.0 39.0 mosbach mm 1955/1136 319.0 60.0 31.0 37.0 22.0 54.0 40.0 mosbach mm 1958/42 335.0 65.0 36.0 41.0 23.0 60.0 43.0 mosbach mm 1973/299 302.0 57.0 30.0 51.0 32.0 mosbach mm 1961/69 66.0 35.0 34.0 24.0 mosbach mm 1966/192 323.0 65.0 35.0 39.0 22.0 58.0 41.0 mosbach mm 1959/217 62.0 33.0 41.0 23.0 mosbach mm 1958/355 54.0 29.0 32.0 18.0 mosbach mm 1962/1233 69.0 36.0 39.0 25.0 mosbach mm 1961/292 324.0 69.0 35.5 39.0 27.0 61.0 40.0 ingarano i1 268.5 56.0 32.0 32.0 24.0 51.0 37.5 ingarano i2 53.5 29.0 ingarano i3 56.0 34.0 ingarano i4 57.0 32.5 34.0 21.0 avetrana a1 46.0 24.0 27.5 16.0 avetrana a2 52.0 30.0 avetrana a3 55.0 31.0 avetrana a4 53.0 29.0 avetrana a5 30.0 20.0 sora valle radice p4215 49.0 25.5 sora valle radice p4217 45.2 33.0 sora valle radice sn 48.4 25.5 sora valle radice sn 45.8 27.5 sora valle radice sn 53.1 27.7 sora valle radice sn 49.0 25.4 sora valle radice sn 44.4 22.7 sora valle radice sn 42.5 31.0 sora valle radice sn 44.9 31.9 sora valle radice sn 45.3 31.3 sora valle radice sn 39.7 29.2 grotta romanelli p1691 54.0 30.0 grotta romanelli p1692 50.0 27.0 grotta romanelli p1696 45.0 32.0 grotta romanelli p1698 45.0 33.0 grotta romanelli p1690 50.0 28.5 grotta romanelli p1694 52.0 29.5 grotta romanelli p1699 46.0 32.0 grotta romanelli p1700 46.5 31.0 grotta romanelli p1697 46.0 30.0 grotta romanelli r54v 54.2 30.6 grotta romanelli sn 45.5 31.0 grotta romanelli sn 44.4 30.9 grotta romanelli sn 45.9 32.4 ���()" �����)" '��, )��� ������ �' -������ )���� ��� ��= metacarpus site specimen l ptd papd mtd mapd dtd dapd mosbach mm 1954/441 294.0 47.0 35.0 28.0 27.5 48.0 27.0 mosbach mm 1954/442 286.0 41.5 32.0 27.0 26.5 46.5 27.5 mosbach mm 1961/534 277.0 45.0 31.0 25.5 26.0 42.5 26.0 mosbach mm 1961/1308 278.0 47.0 35.5 29.0 28.5 48.5 28.0 mosbach mm 1961/1059 279.0 45.0 31.0 27.0 26.0 mosbach mm 1962/684 46.0 34.0 28.0 24.0 mosbach mm 1956/559 45.5 32.0 29.0 28.0 mosbach mm 1955/306 268.0 38.5 29.5 25.0 24.0 43.0 25.0 mosbach mm 1957/528 271.0 39.0 30.5 22.5 25.0 40.0 26.0 mosbach mm 1955/968 290.0 47.0 33.5 27.5 28.0 47.0 29.0 mosbach mm 1951/103 277.0 43.0 31.0 25.5 25.0 45.0 26.0 mosbach mm 1954/324 298.0 47.0 34.5 29.0 26.0 mosbach mm 1957/956 25.5 28.0 46.5 27.5 mosbach mm 1958/354 52.0 36.0 35.0 25.0 mosbach mm 1958/290 29.0 27.0 46.0 28.0 mosbach mm 1965/281 45.5 33.0 29.5 24.0 mosbach mm 1959/214 288.0 46.0 33.0 30.0 27.0 47.0 28.0 mosbach mm 1956/359 278.0 47.0 32.0 27.0 27.0 45.0 27.0 mosbach mm 1958/353 264.0 40.0 29.0 24.5 24.0 mosbach mm 1959/581 43.0 32.0 25.0 24.0 mosbach mm 1959/580 46.0 32.0 27.0 25.0 47.0 27.0 mosbach mm 1952/360 52.0 36.0 mosbach mm 1955/1359 49.0 36.0 28.0 29.0 mosbach mm 1963/638 45.0 33.5 27.0 27.0 mosbach mm 1959/132 45.0 33.0 26.5 27.0 mosbach mm 1956/706 45.0 32.0 26.0 25.5 mosbach mm 1962/971 44.0 33.0 mosbach mm 1952/352 294.0 49.0 35.0 29.0 27.0 48.0 30.0 mosbach md mb 665 272.0 46.0 33.0 26.5 27.0 48.0 28.0 mosbach md mb 389 282.0 44.0 31.0 27.0 27.0 46.5 27.0 ingarano i5 245.0 40.5 31.0 24.0 23.5 40.5 27.5 ingarano i6 247.0 40.5 33.0 23.5 23.5 41.0 27.5 ingarano i7 37.5 27.0 ingarano i8 29.0 ingarano i9 42.0 27.0 ingarano i10 23.0 22.0 avetrana a6 39.0 28.0 avetrana a7 31.0 avetrana a8 36.0 25.5 avetrana a9 40.5 29.0 avetrana a10 36.0 26.0 21.5 21.0 avetrana a11 25.0 28.0 44.0 32.0 avetrana a12 40.0 28.0 grotta del fossellone p90.1 265.0 44.0 34.0 30.0 30.0 46.0 28.5 grotta del fossellone p90.1 256.0 42.0 31.0 24.0 26.0 43.0 29.0 grotta del fossellone p10.58 39.0 26.0 sora valle radice p4230 244.0 36.0 27.0 20.5 22.0 38.5 26.0 sora valle radice sn 36.7 25.8 sora valle radice sn 41.2 29.6 sora valle radice sn 35.7 27.0 sora valle radice sn 36.4 27.5 sora valle radice sn 40.0 28.6 sora valle radice sn 39.8 28.2 sora valle radice sn 41.0 26.5 sora valle radice sn 41.5 27.8 grotta romanelli p1724 244.5 38.0 28.0 22.0 22.5 38.5 27.0 ���������� %� &��'��� ("� )�����'� *"� )������� �" + &����� *"��< metacarpus site specimen l ptd papd mtd mapd dtd dapd grotta romanelli p1732 46.0 29.0 grotta romanelli p1725 44.0 30.0 grotta romanelli p1737 38.5 27.0 grotta romanelli p1735 37.0 25.2 grotta romanelli p1733 45.8 31.0 grotta romanelli sn 37.5 25.4 grotta romanelli sn 39.2 27.3 grotta romanelli sn 42.8 28.7 grotta romanelli sn 41.1 28.0 grotta romanelli sn 44.5 29.8 grotta romanelli sn 38.4 25.6 grotta romanelli sn 41.2 27.7 grotta romanelli sn 46.7 27.6 grotta romanelli sn 39.7 27.9 grotta romanelli sn 45.4 28.5 grotta romanelli sn 43.4 26.7 grotta romanelli sn 40.3 26.1 grotta romanelli sn 42.7 27.5 grotta romanelli sn 36.9 25.0 grotta romanelli sn 40.1 26.4 grotta romanelli sn 36.6 25.4 grotta romanelli sn 42.5 26.8 tibia site specimen l ptd papd mtd mapd dtd dapd mosbach mm 1972/135 409.0 95.0 40.0 35.0 55.0 mosbach mm 1957/587 48.0 35.5 mosbach mm 1954/68 49.0 38.0 mosbach mm 1959/137 48.5 38.0 mosbach mm 1961/1118 60.0 42.0 mosbach mm 1962/1772 50.5 37.0 mosbach mm 1961/201 37.0 30.5 55.0 42.5 mosbach mm 1956/186 30.0 28.0 50.0 35.0 mosbach mm 1957/357 33.0 29.0 45.0 34.0 mosbach mm 1957/181 43.0 37.0 61.0 47.0 mosbach mm 1954/519 37.0 28.0 55.0 43.0 mosbach mm 1961/76 55.0 43.0 mosbach mm 1955/377 411.0 92.0 36.0 31.0 58.0 42.0 ingarano i11 336.5 76.0 71.5 32.5 32.0 50.5 40.0 ingarano i12 31.5 27.0 51.0 42.0 ingarano i13 57.0 42.0 ingarano i14 375.0 85.0 78.0 25.0 30.0 51.0 41.0 avetrana a13 32.0 27.0 51.5 41.0 avetrana a14 50.0 40.0 grotta del fossellone p1065 50.0 40.0 riparo di fumane a10v75 56.0 44.5 sora valle radice p4235 325.0 64ca 68.0 25.0 27.0 48.0 36.0 sora valle radice sn 63.2 67.0 sora valle radice sn 41.7 34.0 sora valle radice sn 41.7 32.1 sora valle radice sn 40.0 32.6 sora valle radice sn 53.6 42.3 sora valle radice sn 42.4 35.0 sora valle radice sn 41.8 33.8 sora valle radice sn 43.1 33.5 ���������� ���()" �����)" '��, )��� ������ �' -������ )���� ��� ��3 tibia site specimen l ptd papd mtd mapd dtd dapd sora valle radice sn 40.5 30.2 sora valle radice sn 42.6 33.5 grotta romanelli p1755 71.0 62.5 grotta romanelli p1765 45.0 34.0 grotta romanelli p1759 47.0 39.0 grotta romanelli p1754 78.2 75.0 grotta romanelli p1756 62.0 61.0 grotta romanelli p1766 50.0 41.0 grotta romanelli p1760 49.0 37.0 grotta romanelli p1764 47.0 37.0 grotta romanelli p1758 47.0 36.0 grotta romanelli p1762 47.2 37.5 grotta romanelli sn 45.8 32.7 grotta romanelli sn 46.4 35.4 grotta romanelli sn 45.2 36.4 metatarsus site specimen l ptd papd mtd mapd dtd dapd mosbach mm 1961/324 316.0 41.0 46.0 29.0 30.0 45.0 30.0 mosbach mm 1956/667 312.0 26.0 30.0 44.5 mosbach mm 1955/459 40.0 41.0 25.0 30.0 mosbach mm 1962/847 329.0 42.0 45.5 28.0 31.5 49.0 30.0 mosbach mm 1957/324 302.0 34.0 40.0 25.0 30.5 41.5 25.5 mosbach mm 1961/535 35.0 40.0 24.0 27.0 mosbach mm 1951/355 303.0 41.0 24.0 28.0 44.5 29.0 mosbach mm 1955/1021 40.0 42.0 26.0 29.0 mosbach mm 1961/126 316.0 42.0 42.0 26.0 30.0 mosbach mm 1955/1134 40.0 43.0 26.5 33.0 mosbach mm 1957/703 302.0 38.0 43.0 24.0 32.0 42.0 28.0 mosbach mm 1957/806 318.0 44.0 48.0 27.5 34.0 49.0 29.0 mosbach mm 1957/569 39.0 43.5 26.5 28.0 mosbach mm 1957/991 47.5 50.0 29.0 35.0 mosbach mm 1958/807 44.0 47.0 mosbach mm 1971/34 36.5 40.0 24.5 26.0 mosbach mm 1957/759 37.5 43.0 25.0 27.0 mosbach mm 1967/50 43.0 47.0 26.0 29.5 mosbach mm 1957/758 36.5 41.0 24.0 29.0 mosbach mm 1958/122 312.0 44.0 47.5 26.5 29.0 48.5 30.0 mosbach mm 1958/502 312.0 41.0 44.5 25.0 30.0 44.0 30.0 mosbach mm 1958/121 335.0 48.0 51.0 32.0 36.0 58.0 34.5 mosbach mm 1959/353 321.0 42.0 45.5 28.0 29.5 28.0 mosbach mm 1954/222 48.0 29.0 mosbach mm 1962/571 39.5 42.0 mosbach mm s. n. 41.0 44.0 mosbach mm 1970/123 27.0 30.5 47.0 30.0 mosbach mm 1969/130 27.0 31.5 47.0 31.0 mosbach mm 1961/824 48.0 29.0 mosbach mm 1962/970 46.0 28.5 mosbach mm 1961/127 29.0 31.5 49.5 30.0 mosbach mm 1959/756 27.0 28.0 53.0 29.0 mosbach mm 1962/419 49.5 32.5 mosbach mm 1961/718 48.0 30.0 mosbach mm 1970/122 47.5 30.5 mosbach mm 1957/244 36.0 38.0 25.5 28.0 mosbach mm 1956/764 40.0 44.0 25.5 27.5 ���������� %� &��'��� ("� )�����'� *"� )������� �" + &����� *"��9 metatarsus site specimen l ptd papd mtd mapd dtd dapd mosbach mm 1953/393 39.0 41.0 26.0 27.0 mosbach mm 1961/604 39.0 44.5 mosbach mm 1962/1376 37.0 39.5 25.0 25.5 mosbach mm 1961/1057 38.0 41.5 25.5 27.0 mosbach mm 1957/581 38.0 40.0 mosbach mm 1968/76 40.0 43.0 24.0 27.0 mosbach mm 1949/379 321.0 44.0 47.0 28.0 31.0 52.0 32.0 mosbach md mb 372 317.0 47.0 29.0 32.0 52.0 33.0 mosbach md mb 371 47.0 46.0 28.0 29.0 ingarano i15 276.0 39.5 42.0 28.0 27.0 40.5 30.0 ingarano i16 35.5 36.0 18.5 25.5 ingarano i17 36.5 36.5 ingarano i18 40.0 46.0 28.0 33.0 avetrana a15 29.0 31.0 avetrana a16 34.0 37.0 avetrana a17 32.0 33.0 20.5 23.5 avetrana a18 21.0 24.0 avetrana a19 34.0 37.0 21.0 avetrana a20 35.0 38.0 22.0 28.0 grotta del fossellone p1061 42.0 27.5 grotta del fossellone p90.3 279.2 39.0 41.0 24.5 28.5 45.0 29.5 grotta del fossellone p1059 34.0 37.5 grotta del fossellone p1062 46.0 29.0 sora valle radice p4246 282.0 39.0 39.0 23.5 28.0 42.0 27.0 sora valle radice sn 36.7 38.3 sora valle radice sn 36.3 37.2 sora valle radice sn 33.4 34.6 sora valle radice sn 36.6 40.2 sora valle radice sn 39.8 25.6 sora valle radice sn 39.2 25.1 sora valle radice sn 34.7 24.7 sora valle radice sn 38.6 26.5 grotta romanelli p1796 40.2 28.0 grotta romanelli p1788 35.0 37.0 grotta romanelli p1795 42.0 27.0 grotta romanelli sn 33.7 35.1 grotta romanelli sn 39.0 26.8 astragalus site specimen hl hm td mosbach mm 1957/518 62.0 57.0 40.0 mosbach mm 1955/318 62.0 57.0 38.0 mosbach mm 1950/585 69.0 63.0 46.0 mosbach mm 1955/1150 65.0 62.0 40.0 mosbach mm 1954/642 64.0 60.0 42.0 mosbach mm 1957/48 63.0 59.5 40.0 mosbach mm 1954/770 60.0 57.0 39.0 mosbach mm 1956/587 58.0 55.0 38.0 mosbach mm 1959/799 60.0 55.0 38.0 mosbach mm 1954/619 64.0 60.0 40.0 mosbach mm 1955/978 60.0 55.0 mosbach mm 1955/975 65.0 61.0 42.0 mosbach mm 1959/856 63.0 57.0 41.0 mosbach mm 1955/1250 61.0 57.0 37.0 ���������� ���()" �����)" '��, )��� ������ �' -������ )���� ��� ��1 astragalus site specimen hl hm td mosbach mm 1961/512 63.0 57.0 40.0 mosbach mm 1961/901 65.0 61.0 40.0 mosbach mm 1960/61 64.0 59.5 41.0 mosbach mm 1959/745 69.0 65.0 45.0 mosbach mm 1960/51 65.0 63.0 42.0 mosbach mm 1961/598 66.0 61.0 44.0 mosbach mm 1961/58 62.0 57.0 40.0 mosbach mm 1957/533 64.0 59.0 42.0 mosbach mm 1968/253 63.0 60.0 41.5 mosbach mm 1958/575 63.0 57.0 41.0 mosbach mm 1961/131 62.0 58.0 38.0 mosbach mm 1959/668 63.0 59.0 38.0 mosbach mm 1957/590 80.0 76.0 47.0 ingarano i19 55.0 52.0 37.0 ingarano i20 49.0 47.0 32.0 ingarano i21 54.0 49.5 37.0 ingarano i22 54.0 48.0 38.0 ingarano i23 35.0 ingarano i24 53.0 49.0 34.0 ingarano i25 53.0 49.5 36.0 ingarano i26 46.0 ingarano i27 47.0 ingarano i28 49.0 34.0 ingarano i29 51.5 ingarano i30 45.0 ingarano i31 50.0 44.5 32.0 ingarano i32 49.5 45.0 32.0 avetrana a21 49.0 avetrana a22 53.5 51.0 35.0 avetrana a23 52.0 47.5 34.0 avetrana a24 57.5 54.0 38.0 grotta del fossellone p1069-1 53.5 49.0 35.0 grotta del fossellone p1068 49.0 45.0 31.1 grotta del fossellone p1071 57.0 53.0 36.0 grotta del fossellone p1070 54.0 51.0 37.8 sora valle radice p4242 48.7 45.2 30.5 sora valle radice sn 48.0 42.8 31.3 sora valle radice sn 51.9 47.2 33.5 sora valle radice sn 53.2 49.5 34.3 sora valle radice sn 49.0 33.0 sora valle radice sn 47.5 45.9 28.6 palidoro pal59 55.3 50.7 34.0 palidoro pal59 53.5 49.3 34.7 palidoro pal59 55.1 50.7 34.6 palidoro pal59 55.4 52.5 35.6 palidoro pal59 50.0 47.0 30.3 grotta romanelli p1774 51.0 48.0 33.0 grotta romanelli p1769-2 52.5 49.5 33.0 grotta romanelli p1769-1 54.0 50.0 33.0 grotta romanelli p1770-1 48.0 46.0 29.0 grotta romanelli p1772-2 54.0 52.0 34.0 grotta romanelli p1772-1 55.2 51.5 33.0 grotta romanelli p1771-1 51.2 49.0 28.0 grotta romanelli p1771-2 52.0 48.5 31.0 grotta romanelli p1770-2 52.0 48.5 32.0 grotta romanelli p1773 45.0 42.0 28.0 %� &��'��� ("� )�����'� *"� )������� �" + &����� *"��4 astragalus site specimen hl hm td grotta romanelli sn 54.8 53.3 33.0 grotta romanelli sn 49.9 47.5 30.3 grotta romanelli sn 52.1 48.7 32.5 grotta romanelli sn 53.0 37.5 grotta romanelli sn 58.3 52.4 32.3 grotta romanelli sn 55.0 50.6 35.0 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /all /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /warning /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy 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("* "& ��?2 �+! #**! ."$( $�.*�! '%3!$(')�(!0 '% (+'$ $(#0, 5�� <�6 !�" 0$ (+! �� "& ���������� ���������� 9+'�+ 0!&'%!$ (+! /�$! "& ���7 "& �� �, �"�!%! �)!2 �+! ��?1���7 4"%�� /"#%0� , ��%%"( /! * !�'$!�, .� :!0 $'%�! � �"9!$"�#('"% $�.*�'%) 0#! (" *"" !=*"$# ! "& (+! "#(� "* 5�')2 �62 �+#$� '( '$ #%��!� '& (+! ���!"�!%!-�"�!%! �+! .�� �='.#. '%(! 3�� '$ �/$!%( �%0 (+! ��?���7 ( �%$'('"% '$ �+� ��(! '4!0 /, .�8" �+�%)!$ '% $!0'.!%(�('"% �(! " $( �(') �*+'� )�*$2 ����������� �!3! �� '.*" (�%( %�%%"&"$$'� !3!%($ +�3! /!!% !�")%'4!0 �%0 ."$( 4"%�� .� :! $ 0'$*��, � �"%('%#"#$ "��# !%�! '% (+! ��/'�)" $!�('"%2 �+! '%3!$(')�(!0 $!�('"% $*�%$� ��."$( �"%('%#"#$�,� (+! !� �, ��$( '�+('�% ���� (" (+! �� �, �"�!%! ���7 %�%%"&"$$'� 4"%!2 '.'(!0 (!�("%'� 0'$*���!.!%($� $#�+ �$ &�#�($ �%0 &"�0$� "/$! 3!0 '% "#(� "*$ � ! /!�"9 /'"$( �(') �*+'� !$"�#('"%2 �+! � !(��!"#$ �%0 �! ('� , '%(! 3��$ '%3!$(')�(!0 $+"9 * ".'%!%( 3� '�('"%$ '% $!0'.!%(�('"% �(!2 �+! � !%%" �" .�('"%$ '$ �+� ��(! '4!0 /, � $!0'.!%(�('"% �(! "& ;�7 .1.2,2� 9+'�$( (+! ��/'�)" �" .�('"% '$ .� :!0 /, 0 �.�('� 0!� !�$! (" ;�7 .1.2,2 �9" .�8" $( �(') �*+'� )�*$ $!!. (" /! �$$"�'�(!0 9'(+ (+! � !(��!"#$-�! ('� , ( �%$'('"%2 �+! &' $( *"$$'/�! +'�(#$ +�$ /!!% "/$! 3!0 '% (+! ��(!$( ��$( '�+('�%� 9+! ! (+! �/$!%�! "& (+! 4"%�� .� :! � ������� .�, '.*�, (+�( (+! 9+"�! "& (+! ���< 4"%! �%0 *"$$'/�, � *� ( "& (+! ���>� $#/4"%! .�, /! .'$$'%) " !=( !.!�, �"%0!%$!02 �'%�! "%�, "%! $�.*�! 9�$ �"��!�(!0 /!(9!!% �� �� �%0 �� ��/"((". 0#! (" *"" !=*"$# ! "& (+! "#(� "*� '( '$ #%��!� '& (+! ���: "& � ������� !* !$!%($ � * '.� , $')%��� (+! !$#�( "& 0'�)!%!('� ��(! �('"% " '%$#&&'�'!%( $�.*�'%)2 �+! $�.*�! �� ��/"((". �"%(�'%$ � %�%%"&�" �� �$$!./��)! 0".'%�(!0 /, *!%(��'(+$ 5!2)2 '���������-���� $**26 �%0 +-�������-���� $**2 '%0'��('%) (+! !� �, ���!"�!%! ���� 9+! !�$ $�.*�! �� ��("* $+"9$ (+! �� "& ( ������� 9+'�+ "��# $ �( (+! /�$! "& 4"%! ���2 �+'%-$!�('"% '%3!$(')�('"%$ *! &" .!0 "% *��%:("%'� &" �.'%'&! $ !3!�� (+! �/$!%�! "& (+! &" �.'%'&! �� �"%! �� �%0 �#/4"%! ���2 �+'$ '%0'��(!$ (+�( (+! ��� 4"%! '$ .� :!0�, !0#�!0 �$ � !$#�( "& � �"9 $!0'.!%(�('"% �(! " � $( �(') �*+'� +'�(#$2 �%$#&&'�'!%( $�.*�'%) 0#! (" *"" !=*"$# ! "& (+! "#(� "* * !3!%($ (" ��� '&, '& (+! ( �%$'('"%$ /!(9!!% ��� �%0 ��> �%0 /!(9!!% ��? �%0 ���7 !�" 0 $".! #%�"%&" .'('!$ " !0#�!0 $!0'.!%(�('"% �(!2 ��5������# ���� i! � ! ) �(!&#� (" g2 �"% ���'$� �2 �� 0'% �%0 �2 "%!�+' &" !3'!9'%) (+! .�%#$� '*( �%0 * "3'0'%) +!�*&#� �"..!%($2 2 ��!(�%' '$ 9� .�, (+�%:!0 &" (+! !0'(" '�� !3'!92 �+'% $!�('"%$ 9! ! * !*� !0 /, �2 ��'%3! %" �%0 *+"(".'� ") �*+$ /, �2 �+'"0'2 �� �%0 �� 9! ! $#**" (!0 /, ���� !$!� �+ &#%0$� �+!!( 7?< ee�! !)%"aa2 �� 9�$ ��$" $#**" (!0 /, ��� �%! ),2 ��������-���� �� 0����� ���������� �����1����� ��������#���-� 1�� �-� +� ��#� ������� 2� )����3 �> +�� ����� , � "�������� � "������ & . /�� ��� "���� ) �< nannofossil zones (sissingh 1977) sample etching overgrowth abundance watznaueria spp. tegumentum stradneri biscutum constans biscutum magnum zeugrhabdotus elegans zeugrhabdotus erectus zeugrhabdotus diplogrammus zeugrhabdotus trivectis prediscosphaera cretacea prediscosphaera spinosa prediscosphaera ponticula prediscosphaera grandis chiastozygus litterarius chiastozygus platyrhethus ahmuellerella octoradiata arkhangelskiella cymbiformis arkhangelskiella cymbiformis >10µm arkhangelskiella specillata arkhangelskiella maastrichtiana cribrosphaerella ehrenbergii microrhabdulus decoratus ceratolithoides aculeus ceratolithoides kamptneri rhagodiscus angustus rhagodiscus splendens placozygus fibuliformis placozygus sigmoides kamptnerius magnificus gartnerago obliquum staurolithites imbricatus staurolithites ellipticus staurolithites zoensis manivitella pemmatoidea cretarhabdus crenulatus cretarhabdus conicus cretarhabdus striatus eiffellithus turriseiffelii eiffellithus gorkae tranolithus minimus tranolithus orionatus quadrum gartneri uniplanarius sissinghii uniplanarius trifidus reinhardtites spp. reinhardtites levis lithraphidites carniolensis lithraphidites praequadratus lithraphidites quadratus micula decussata micula concava micula murus thoracosphaera spp. braarudosphaera spp. n p 1 t t 33bottom e 3 o 3 c r a a h iatus? c c 25c t t 32 e 3 o 3 f f r f f r c r t t 31 e 2 o 2 f f r r r r f r r f t t 30 e 1 o 1 f c r r r r r r r f r r r r r r f t t 29 e 1 o 1 f f r r r r r r r r c t t 28 e 2 o 2 a a r r r r r r r r r r r r r r c t t 27 e 1 o 1 a a r r r r r f r r r r r r r f t t 26 e 2 o 1 a c r r r r r f r f r r r f t t 25 e 2 o 2 a c r r r r r r f r r r r r r r f t t 24 e 2 o 2 c c r r r r r r r f r r r r r r r f f t t 23 e 1 o 2 a a r r r r r f r r r r r r f r r f t t 22 e 2 o 1 f f r r f f r r r r f r r r r f r t t 21 e 1 o 1 a a r f f r r r r r f t t 20 e 2 o 2 c c r r r r r r c r r r r r r t t 19 e 1 o 1 c c r r r r r r f r r r r r f r f t t 18 e 2 o 1 c c r r f f r f c r r r r r r r r r f f t t 17 e 1 o 1 f f f r r r r f r f f r r r r f f t t 16 e 1 o 1 a f r r r f r f r c r r r r r r r r f t t 15 e 2 o 2 f f r r r r f f r r r f r r r r f f t t 14 e 1 o 1 a c r r r r r r r r r r r c r r r r r r f r r f f t t 13 e 2 o 2 a c r r r c r r r r r r f r t t 12 e 1 o 1 a c f r r f f r r r r r r r r r r f r r r r r r f f t t 11 e 2 o 2 c f r f f r r r f r r r r r r r f r f r r r r f f t t 10 e 2 o 2 a c r f r r c f r c r r r r f r f r r r r r f r t t 9 e 2 o 2 c c r f r f r r f c r r r r f r r r r r r r f f t t 8 e 1 o 1 a c r r r r r f r r r r r r r c r r r r r f r r r r r f t t 7 e 1 o 1 a a r f r c r r r r r r f r r r r f r f r r r t t 6 e 2 o 2 a a r f f f r r r f c r r r f c r r r f r r r c t t 5 e 2 o 2 a a r f r r f r r r r r f c r r r r f f r r r r r r f f t t 4 e 2 o 2 a a f c f r r r f c r f f r r r r f f t t 3 e 1 o 2 c c r r r f r f r r f r f r r r r r f c r r r r r r r r f f t t 2 e 1 o 1 a a r r f f r r r r f r r r f r f r r r r f f t t 1 e 1 o 1 a a r r f r f c f r r f r r r r r cc25bcc24 cc25acc23b � * * !% 0 '= � � '$( '/ # ('" % �+ � ( " & ����� !" # $ % �% % " &" $$'�$ " / $! 3 !0 '% (+ ! � !(��!" # $ * " ('" % " & (+ ! � �/ '�) " $!�('" % 2 � + ! ) �, * �((! % '% 0 '��(!$ (+ ! !3 !% ( 0 !&'% '% ) ����� !" # $ % �% % " &" $$'� 4 " % !$2 � " (�� ����� !" # $ % �% % " &" $$'� �/ # % 0 �% �!� � j � / # % 0 �% (j m > 7 $* !�'. !% $1 '� " $�" * ! &'!�0 " & 3 '!9 5� � � 6c � j � " . . " % j � 7 -> 7 $* !�'. !% $1� � � c � j � !9 j � -� 7 $* !�'. !% $1� � � c � j � � !j n � $* !�'. !% 1� � � 2 � '% ) �! (�= " % �/ # % 0 �% �!� � j � / # % 0 �% (j m � 7 $* !�'. !% $1� � � c � j � " . . " % j � -� 7 $* !�'. !% $1� � � c � j � !9 j � $* !�'. !% 1� -> 7 � � � c � j � � !j � $* !�'. !% 1m > 7 � � � 2 ��������-���� �� 0����� ���������� �����1����� ��������#���-� 1�� �-� +� ��#� ������� 2� )����3 �b nannofossil zones (martini 1971) sample etching overgrowth abundance thoracosphaera spp. braarudosphaera spp. cruciplacolithus tenuis cruciplacolithus primus coccolithus pelagicus markalius inversus praeprinsius dimorphosus placozygus sigmoides prinsius petalosus/tenuiculum/africanus chiasmolithuscf. edwarsii chiasmolithus bidens chiasmolithus danicus prinsius bisulcus prinsius martinii neochiastozygus saepes neochiastozygus perfectus neochiastozygus junctus ellipsolithus macellus ellipsolithus distichus ericsonia subpertusa ericsonia robusta fasciculithus tympaniformis fasciculithus spp. f. ulii/pileatus/janii/magnus/bitectus f. schaubii/richardii sphenolithusprimus sphenolithusanarrophus heliolithus kleinpellii bomolithus elegans toweiuscf. pertusus toweius pertusus toweius eminens discoasterspp. (unidentifiable forms) discoaster mohleri heliolithus reidelii discoaster nobilis discoaster multiradiatus zyghrablithus bijugatus discoaster diastypus n p 10 t t 6 3 e 3 o 3 f f r r f f r f f f r t t 6 2 e 2 o 2 c c r f r c f f r c r f f f t t 6 1 e 3 o 3 c c r r r r f r c f f f r c r f c f t t 6 0 e 2 o 2 c r f r r r f r f f f f r f f f r f f t t 5 9 e 2 o 3 c f r r r r f f f f r f f r f f t t 5 8 e 2 o 2 c f r r r r f f r r r c r r r f t t 5 7 e 3 o 3 c f r r r r r f r f f r f r f t t 5 6 e 2 o 2 c r f r f r f r f f r r n p 8 t t 5 5 e 3 o 3 a f r r r f c f r r r c r r r r t t 5 4 e 3 o 3 a r c r r r r r f f f r r f r r t t 5 3 e 3 o 3 c r f r r r f r r f f t t 5 2 e 3 o 3 f f r r r r f r f f f r t t 5 1 e 2 o 2 f r f r f f f r r f t t 5 0 e 3 o 3 f r f r r r f f f r f r f n p 5 t t 4 9 e 3 o 3 c c r r r r r r r t t 4 8 e 2 o 2 r f r r r r r t t 4 7 e 3 o 3 a r f a r t t 4 6 e 2 o 2 a r f r r r a r r t t 4 5 e 3 o 3 a f r r a t t 4 4 e 3 o 3 a r r r r a t t 4 3 e 3 o 3 a r r r r r a t t 4 2 e 2 o 3 a r f f f r a t t 4 1 e 3 o 3 a r f c r c r a t t 4 0 e 3 o 3 a r r r f c a a t t 3 9 e 3 o 3 a r c r a r a t t 3 8 e 3 o 2 a r r f r c c c t t 3 7 e 3 o 3 a r r f r a r r t t 3 6 e 3 o 3 a f r r r a r r t t 3 5 e 3 o 3 a r r f r a r t t 3 4 e 3 o 3 c f r f r c r r t t 3 3 to p e 3 o 3 f r r r f r f r h ia tu s n p 1 t t 3 3 b o tt o m e 3 o 3 c a a np4 np3np9 np2 n p 6np7 � * * !% 0 '= / � '$ ( '/ # (' " % �+ � ( " & �� �� � !" # $ % �% % " &" $$ '� 4 " % �� . � : ! $ �% 0 $! �! �( !0 (� = � " / $! 3 !0 '% (+ ! � �� !" ) !% ! * " ( '" % " & (+ ! � �/ '� ) " $! �( '" % 2 � / # % 0 �% �! $ " & �� �� � !" # $ % �% % " &" $$ '� $ �$ '% � * * !% 0 '= ee� aa2 � � � � � � � � � � �#/ , 2-�2 5�?d�6 ��%0/"": "& �!%"4"'� ����� !"#$ %�%%"*��%:("%2 �"": �� � (+"�'(+�! 5�'$�"�$(! $62 '� "*��!"%("�"), � !$$� �.! 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/pdfxtrimboxtomediaboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxsetbleedboxtomediabox true /pdfxbleedboxtotrimboxoffset [ 0.00000 0.00000 0.00000 0.00000 ] /pdfxoutputintentprofile () /pdfxoutputcondition () /pdfxregistryname (http://www.color.org) /pdfxtrapped /unknown /description << /fra /enu (use these settings to create pdf documents with higher image resolution for improved printing quality. the pdf documents can be opened with acrobat and reader 5.0 and later.) /jpn /deu /ptb /dan /nld /esp /suo /ita /nor /sve /kor /chs /cht >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [595.000 842.000] >> setpagedevice the southernmost occurrence of brach ycarcharias (lamniformes, odontaspididae) from the eocene of antarctica provides new information about the paleobiogeography and paleobiology of paleogene sand tiger sharks giuseppe marramà1*, andrea engelbrecht1, thomas mörs2, marcelo a. reguero3 & jürgen kriwet1 1*corresponding author. department of paleontology, university of vienna, althanstrasse 14, 1090 vienna, austria. e-mail: giuseppe.marrama@univie.ac.at, andrea.engelbrecht@univie.ac.at, juergen.kriwet@univie.ac.at 2 department of paleozoology, swedish museum of natural history, p.o, box 50007, se-104 05 stockholm, sweden. e-mail: thomas.moers@nrm.se 3 division paleontologia de vertebrados, museo de la plata, paseo del bosque s/n, 81900 fwa la plata, argentina, conicet. e-mail: egui@fcnym.unlp.edu.ar arku to cite this article: marramà g., engelbrecht a., mörs t., reguero m.a. & kriwet j. (2018) the southernmost occurrence of brachycarcharias (lamniformes, odontaspididae) from the eocene of antarctica provides new information about the paleobiogeography and paleobiology of paleogene sand tiger sharks. riv. it. paleontol. strat., 124(2): 283-298. rivista italiana di paleontologia e stratigrafia (research in paleontology and stratigraphy) vol. 124(2): 283-298. july 2018 abstract. the first record of one of the most common and widespread paleogene selachians, the sand tiger shark brachycarcharias, in the ypresian strata of the la meseta formation, seymour island, antarctica, is provided herein. selachians from the early eocene horizons of this deposit represent the southernmost paleogene occurrences in the fossil record, and are represented by isolated teeth belonging to orectolobiforms, lamniforms, carcharhiniforms, squatiniforms and pristiophoriforms. the combination of dental characters of the 49 isolated teeth collected from the horizons telms 2, 4 and 5 supports their assignment to the odontaspidid brachycarcharias lerichei (casier, 1946), a lamniform species widely spread across the northern hemisphere during the early paleogene. the unambiguous first report of this lamniform shark in the southern hemisphere in the eocene of the la meseta formation improves our knowledge concerning the diversity and paleobiology of the cartilaginous fishes of this deposit, and provides new insights about the biotic turnovers that involved the high trophic levels of the marine settings after the end-cretaceous extinction and before the establishment of the modern marine ecosystems. received: october 05, 2017; accepted: february 15, 2018 keywords: chondrichthyes; elasmobranchii; ypresian; la meseta formation; biotic turnovers. introduction living sand tiger sharks of the possibly paraphyletic family odontaspididae include three large sized species within the order lamniformes, today inhabiting marine tropical outer shelf and mesopelagic habitats (carcharias taurus) and cool-water inner shelf habitats (odontaspis ferox and o. noronhai) of the atlantic, indian, and pacific oceans (compagno 1984, 2002; cappetta 2012; nelson et al. 2016). although recent morphological and molecular analyses suggested that odontaspididae might not be monophyletic (e.g., shimada 2005; martin et al. 2002; naylor et al. 2012), several authors have recognized a set of plesiomorphic characters that are traditionally used to distinguish sand tiger sharks from all other lamniforms, including a short to moderately long, conical or slightly depressed snout, weakly protrusible jaws, first dorsal fin in front of the pelvic origin, last gill-slit in front of the pectoral origin, 156 to 183 vertebral centra, monognathic tooth heterodonty, tearing-type dentition, teeth arranged in less than 60 rows in each jaw having a tall and slender main cusp, one to three pairs of lateral cusplets, and a root with well-separated lobes and marked by a strong nutritive furrow (compagno 1999, 2002; cappetta 2012; nelson et al. 2016). although carcharias and odontaspis are the only genera having modern representatives within marramà g., engelbrecht a., mörs t., reguero m.a. & kriwet j.284 this family (compagno 1999; nelson et al. 2016), the odontaspidid fossil record has a wide temporal and geographic distribution, being indeed comprised of at least 20 extinct genera with more than 50 species dating back to the lower cretaceous (cappetta 2012; shimada et al. 2015; cappetta & case 2016). selachians from paleogene sites, such as the la meseta formation on seymour island (ne of the antarctic peninsula), are of upmost importance for understanding of the general patterns of abiotic disruptions during the paleogene. in fact, these predators can document profound changes and turnovers in marine ecosystems in concomitance to the paleocene – eocene thermal maximum (petm; ca. 55.5 ma) and subsequent cooling phase towards the establishment of the antarctic ice sheet. petm was followed by a middle to late eocene transition from the greenhouse world to icehouse conditions (ca. 49–34 ma) with marked deep-sea cooling of about 7°c (e.g., zachos et al. 2001, 2008; miller et al. 2005). the final cooling phase across the eocene – oligocene (e–o) boundary (ca. 33.7 ma) resulted in the disappearance of chondrichthyans from antarctica (e.g., kriwet et al. 2016). the vertebrate fossils of the la meseta formation are mainly represented by cartilaginous fishes, which seemingly represents the major faunal components of the pre-oligocene antarctic fish faunas (kriwet et al. 2016). fossil chondrichthyans include more than 35 species within 22 families of selachians, batoids, and chimaeroids, which have been reported from different levels within the la meseta formation and at different localities on seymour island (see welton & zinsmeister 1980; jerzmanska 1988, 1991; eastman & grande 1989, 1991; long 1992a, b, 1994; balushkin 1994; cione & reguero 1995, 1998; doktor et al. 1996; long & stilwell 2000; kriwet 2005; engelbrecht et al. 2016, 2017; kriwet et al. 2016). the ypresian strata from which the elasmobranch material of this study originated, date back about 15 ma after the end-cretaceous extinction, coinciding with a period of maximum morphological diversification of major bony and cartilaginous fish lineages (e.g., kriwet & benton 2004; friedman 2009, 2010; sorenson et al. 2014; frédérich et al. 2016; marramà & carnevale 2017; marramà et al. 2016a, b, 2017a) and correspond to the latest phase of the early eocene climatic optimum (e.g., reguero et al. 2012; kriwet et al. 2016). for this reason, these strata are crucial to reconstruct evolutionary dynamics of chondrichthyans across the paleogene of antarctica. the aim of this paper is to provide the first unambiguous record of one of the most common and widespread early paleogene marine top predators, brachycarcharias cappetta & nolf, 2005, in the early eocene la meseta formation of seymour island. paleobiogeographic, paleobiological and paleoecological implications based on this occurrence provide new insights into the biotic turnovers that occurred at a high trophic level of the marine food chain during the paleogene. geological setting the eocene la meseta formation is exposed on seymour island, which is situated approximately 100 kilometers se of the northern tip of the antarctic peninsula (fig. 1). the highly fossiliferous sediments of the 720-m-thick la meseta formation represent the uppermost part of the infill of the james ross basin, a back-arc basin developed on the eastern flank of the antarctic peninsula (elliot 1988; del valle et al. 1992; hathway 2000; marenssi et al. 2006). this formation comprises mostly poorly consolidated clastic fine-grained sediments, which were deposited in a deltaic, estuarine and shallow marine environment (marenssi 1995; marenssi et al. 1998a, b). the fossiliferous sediments belong to two groups, the lower marambio group of late cretaceous to paleocene age including the lopez de bertodano and sobral formations and the overlying seymour island group comprising the cross valley (middle – earliest late paleocene), la meseta (late paleocene – early middle eocene), and submeseta (middle eocene – early oligocene) formations (e.g., zinsmeister 1982; grande & chatterjee 1987; marenssi 2006; montes 2013). the la meseta and submeseta formations, which have yielded most vertebrate material up to now, are separated and bounded by a prominent erosional surface (marenssi 2006). the la meseta fm ranges from the thanetian to the lutetian, whereas the submeseta fm ranges from the late lutetian to the priabonian/rupelian. the la meseta fm is further subdivided into six allomembers, namely: valle de las focas (telm 1), acantillado i and ii (telms 2 and 3 in partem), campamento (telms 3 in partem and telm 4), and cucullaea i and ii (telms 5, and 6 brachycarcharias (lamniformes, odontaspididae) from the eocene of antarctica 285 in partem), respectively, whereas the submeseta fm includes three allomembers, submeseta i (telms 6 in partem and 7 in partem), submeseta ii (telm 7 in partem), and submeseta iii (upper telm 7) (montes et al. 2013). both stratigraphic schemes, allomembers and telms (= tertiary eocene la meseta; see sadler 1988), are widely used and we use both schemes to indicate where the material comes from to provide as much stratigraphic information as possible. the material that forms the focus of this study was recovered from three different telms. most of the material described and discussed herein (29 teeth, representing about 60% of the entire sample) was collected from telm 5 of the cucullaea i allomember (mostly at localities iaa [instituto antártico argentino] 1/90 and iaa 2/95) which is ypresian in age. from the underlying telm 4 of the cucullaea i allomember, (localities iaa 1/90, iaa 2/95 and nrm [naturhistoriska riksmuseet] 10), which also is ypresian in age, 16 teeth (32%) were collected. telm 2 produced the fewest material, comprising four teeth (8%), and is early ypresian in age. no teeth referable to brachycarcharias lerichei were recovered from telm 3 (ypresian), but this is probably due to sampling bias since very few samples were collected from this layer (m.r., and m.t., pers. obs.). on the contrary, the absence of b. lerichei teeth from telms 6 (lutetian) and 7 (priabonian) may represent a real paleoecological and stratigraphic signal since very few taxa adapted to cool waters have been recovered in these levels (kriwet et al. 2016). the cucullaea i allomember crops out all around the foothills of the meseta in the northern part of seymour island and has a maximum thickness of 90 m. it is composed of shelly channel fills, sand-mud alternations, and cross-bedded fine sands. the unit was deposited in estuarine to shallow marine estuary settings (marenssi et al. 1998a, b, 2002). the acantilados allomember crops out along the seacliffs and has a maximum thickness of 210 m. the remarkably faulted sediments are mainly composed of mudstones and very fine sandstones with prominent breccia horizons. the depositional setting is interpreted as a progradational/aggradational tide-dominated and wave-influenced delta front/delta plain which developed within a incised valley (marenssi et al. 1998a, b, 2002). material and methods the present study is based on 49 isolated teeth from the fossiliferous strata of the telms 2, 4 and 5 of the la meseta formation, seymour island, in antarctica, and collected during three summer campaigns in 2011, 2012 and 2013. all material is housed in the collections of the swedish natural history museum, stockholm, with nrm-pz collection numbers. all the specimens were extracted by hand-quarrying from the fig. 1 location and geological map of the seymour island, showing the ypresian horizons telms 2, 4 and 5 where teeth of brachycarcharias lerichei (casier, 1946) were found. marramà g., engelbrecht a., mörs t., reguero m.a. & kriwet j.286 matrix and some of them were mechanically prepared with needles to reveal fine details. the material was examined using a leica mz12 stereomicroscope and photographed with a reflex camera and/or a 3d digital microscope keyence vhx-1000d 3d. subsequently, all the teeth were measured to the nearest 0.01 mm by using the software package tpsdig 2.19 (rohlf 2005) and assigned to their respective positions using characters observed in brachycarcharias lerichei mostly following cappetta & nolf (2005), van den eeckhaut & de schutter (2009), cappetta (2012), and cappetta & case (2016). morphological tooth terminology follows cappetta (2012). morphometric terminology is adopted and modified from kriwet et al. (2015). systematic paleontology class chondrichthyes huxley, 1880 subclass elasmobranchii bonaparte, 1838 order lamniformes berg, 1958 family odontaspididae müller & henle, 1839 genus brachycarcharias cappetta & nolf, 2005 type species: lamna lerichei casier, 1946; from lower eocene, ypresian; forest-lez-bruxelles, belgium brachycarcharias lerichei (casier, 1946) figs 2, 3 see cappetta & nolf (2005) for a complete list of synonyms. 1946 lamna lerichei casier, p. 80, pl. 2, fig. 7a-b [after lamna vincenti winkler, 1874, pars, in leriche (1906)] 1952 odontaspis vincenti arambourg, p. 84, fig. 19 and pl. 13 1992a ?lamna cf. l. nasus long, p. 27, fig. 9c-d 2005 brachycarcharias lerichei cappetta & nolf, p. 241, pl.2 2009 brachycarcharias lerichei van den eeckhaut & de schutter, p. 5, pl. 7, figs. 1-11; pl. 8, figs. 1-5 2012 brachycarcharias lerichei cappetta, p. 193, fig. 182 2016 brachycarcharias lerichei cappetta & case, p. 49, pl. 3, figs. 8-22 2017b brachycarcharias lerichei marramà et al., p. 3, figs. 2, 3 referred material: 49 isolated teeth: telm 2, early ypresian, early eocene: south of nrm 9 (nrm-pz p15870), nrm 9 (nrm-pz p15883); nrm 7 (nrm-pz p15858) and natica-horizon, (nrm-pz p15838). telm 4, ypresian, early eocene: nrm 6 (nrm-pz p15846); nrm 1 (nrm-pz p15929, nrm-pz p15930, nrm-pz p15876, nrm-pz p15846, nrm-pz p15849, nrm-pz p15852), cucullaea i, (nrm-pz p15847, nrm-pz p15871), center of shark basin (nrm-pz p1886, nrm-pz p15774), below iaa 1/90 (nrm-pz p15773), jaw site (nrm-pz p15926), below iaa 1/90 (nrm-pz p15840) and iaa 1/11 (nrm-pz p15777). telm 5, ypresian, early eocene: nrm 10 (nrm-pz p15771, nrm-pz p15775, nrm-pz p15836, nrm-pz p15841, nrm-pz p15853, nrm-pz p15854), iaa 2/95 and iaa 1/95, ‘marsupial site’ (nrmpz p15772, nrm-pz p15837, nrm-pz p15839, nrm-pz p15806, nrm-pz p15878, nrm-pz p15789, nrm-pz p15842, nrm-pz p15778, nrm-pz p15798, nrm-pz p15796); nrm 11 (nrm-pz p15859, nrm-pz p15776, nrm-pz p 15885); iaa 1/90, ‘ungulate site’ (nrm-pz p15779, nrm-pz p15843, nrmpz p15845, nrm-pz p15872, nrm-pz p15887); natica-horizon (nrm-pz p15780, nrm-pz p15873, nrm-pz p15874); pass site (nrm-pz p15848); ‘monotreme site’ (nrm-pz p15882); dpv 6/84 (nrmpz p15857); south of marsupial site (nrm-pz p15888). locality and age: seymour island, antarctica; telms 2, 4 and 5 of the la meseta formation, ypresian, early eocene (see reguero et al. 2012). remarks. the genus brachycarcharias was introduced by cappetta & nolf (2005) based on the revision of the odontaspidid material from the ypresian of belgium previously referred to lamna vincenti winkler, 1876, or l. lerichei casier, 1946. the presence of l. vincenti had been reported from several other ypresian deposits of europe, north america, and africa (e.g., bassani 1897; woodward 1899; casier 1946; arambourg 1952; noubhani & cappetta 1997). in the revision of the ypresian material from belgium, casier (1946) assigned to l. lerichei the specimens previously assigned to l. vincenti. later, cappetta & nolf (2005) observed that teeth of l. lerichei are morphologically different from those of any other known odontaspidid or lamnid species and therefore erected the odontaspidid genus brachycarcharias to include lamna vincenti and l. lerichei. purdy & francis (2007) challenged the validity of brachycarcharias; however, according to cappetta (2012) evidence for synonymy with previously described genera or species was never provided. more recently, marramà & kriwet (2017) demonstrated through a quantitative morphometric analysis that teeth of brachycarcharias and lamna are morphologically different, excluding definitively the hypothesis of purdy & francis (2007). three other species within the genus brachycarcharias (b. atlasi, b. koerti, and b. mississippiensis) have been recognised based on some qualitative morphological differences. teeth from the thanetian to the ypresian of northern africa traditionally referred to odontaspis atlasi by arambourg (1952) were subsequently transferred to brachycarcharias by cappetta & nolf (2005). cappetta (2006) considered b. mississippiensis from the thanetian to the ypresian of north america (see also case 1994) to be a junior synonymy of b. lerichei; case et al. (2015) considers b. mississippiensis to be a valid species. finally, isolated teeth of lutetian to priabonian age of north africa, america, and asia traditionally assigned to otodus koerti (stromer, 1910) and to lamna or cretolamna twiggsensis (see e.g., case 1981) were included in brachycarcharias by underwood et al. (2011). the validity of b. twiggsensis was later questioned by adnet et al. (2011) and, more recently, cappetta & case (2016) transferred this species in tethylamna, a genus erected for material recovered from the lutetian of brachycarcharias (lamniformes, odontaspididae) from the eocene of antarctica 287 alabama. moreover, carcharias borodini and c. hynei from the thanetian to ypresian of mississippi (see case 1994) are considered to be junior synonyms of b. lerichei by cappetta (2006). long (1992a, fig. 9c-d) described and figured teeth as belonging to the porbeagle lamna cf. nasus from the early eocene strata of the la meseta formation of seymour island, although he regarded the entire succession as middle-late eocene in age. based on this occurrence, long (1992a) therefore supposed the presence of the earliest representative of the genus lamna in the eocene la meseta formation. it must be pointed out that this genus has been used traditionally as repository basket taxon for fossil teeth having similar morphologies (cappetta 2012). teeth of l. nasus are characterized by specimen telm bct bcw ch dcl ds lch lcw mcl pch pcw ra rw rh rt th position nrm-pz p15771 5 2.5 9.65 8.38 2.2 2.2 12.5 8.53 7.53 5.2 upper lateral nrm-pz p15772 5 2.6 11 9.71 10.6 8.14 2.8 1.8 11.9 8.56 6.91 121.7 11.7 3.4 5 13.1 upper antero-lateral nrm-pz p15773 4 2.6 10.4 9.41 9.94 13.7 2.3 2.2 11.8 8.56 6.31 122.7 11.7 3.6 3.9 13 upper lateral nrm-pz p15774 4 2.1 15.8 7.05 6.06 3.7 upper lateral nrm-pz p15775 5 2 8.62 4.19 1.9 1.3 9.86 7.57 5.11 3.5 4.1 12.1 upper antero-lateral nrm-pz p15776 5 0.5 2.1 1.6 6.31 3.3 lower lateral nrm-pz p15777 4 2.2 7.74 9 9.74 0.66 2.6 1.6 9.8 7.94 4.73 123.9 9.95 3 3.7 12 lower antero-lateral nrm-pz p15779 5 3 9.42 9.15 10.3 1.8 2.2 1.9 10.3 8.11 5.69 117.2 11.6 3.8 4.1 12.9 lower lateral nrm-pz p15780 5 2.4 8.44 3.2 4.88 17.6 1 1.8 5.83 2.71 5.05 112.8 10.8 5.1 4.3 8.31 upper lateralmost nrm-pz p15789 5 2.6 7.65 14 2.5 2.1 10.6 6.1 6.25 5.2 upper lateral nrm-pz p15796 5 2.9 11.5 11 11.6 13.4 2.5 2.1 13.4 9.99 7.2 133.6 15.4 4.4 4.7 15.4 upper lateral nrm-pz p15798 5 2.5 9.61 10 9.36 2.8 1.6 11.5 8.91 6.04 4.7 upper lateral nrm-pz p15806 5 1.8 8.93 7.42 8.38 6.64 1.6 1.6 8.99 6.66 5.85 130.7 2.9 2.9 10.4 upper lateral nrm-pz p15836 5 2.2 7.97 8.14 9.19 1.8 2 1.5 8.87 7.23 4.57 123.7 9.6 2.7 2.9 10.9 lower lateral nrm-pz p15837 5 2.8 11.4 8.85 3.2 2.1 13.8 9.26 6.78 5.1 5.4 16.4 intermediate nrm-pz p15838 2 upper lateral nrm-pz p15839 5 2.5 9.56 0.5 2.4 1.6 10.6 8.57 6.19 4 3.6 13.5 lower lateral nrm-pz p15840 4 2.1 10.2 10.6 16.2 2.4 1.8 9.29 6.7 3.6 4.3 13.8 upper lateral nrm-pz p15841 5 10.6 7.21 7.89 16.7 1.8 2.3 10.2 6.31 6.55 138 11.4 3.2 10.4 upper lateral nrm-pz p15842 5 2.9 10 10.8 0.2 2.6 1.5 5.34 4 anterior nrm-pz p15843 5 2 6.33 7.2 7.82 2 2 1.2 7.92 6.23 3.9 4 lower antero-lateral nrm-pz p15845 5 2.1 8.56 9.32 1.65 1.8 1.5 7.77 4.75 4.2 4.5 12.7 lower antero-lateral nrm-pz p15846 5 2.2 10.6 9.38 9.8 15.1 2.3 2 12.2 8.4 6.47 3.2 upper lateral nrm-pz p15847 4 2.7 9.76 10.7 0.3 2 1.5 8.51 6.16 2.6 12.4 lower antero-lateral nrm-pz p15848 5 2 11.5 10.2 10.6 16.7 2 2.2 13.3 9.39 6.96 132.3 14.2 3.3 3.7 13.5 upper lateral nrm-pz p15849 5 2.9 10.7 10.6 11.1 11.9 2.1 1.9 9.53 7.49 4.4 upper lateral nrm-pz p15852 5 10.1 14 17.2 2.2 2.3 9.32 9.18 upper lateral nrm-pz p15853 5 2.4 8.64 9.76 0.7 2.5 1.8 7.75 5.22 4 lower antero-lateral nrm-pz p15854 5 1.6 6.59 7.41 0.8 1.8 1.5 5.89 3.78 lower antero-lateral nrm-pz p15857 5 2.6 11.3 9.52 10 12.6 2.9 2.2 12.5 8.22 6.87 130.5 12.9 3.2 4.3 12.7 upper lateral nrm-pz p15858 2 nrm-pz p15859 5 3.1 8.98 9.58 10.5 1 2.5 1.7 10.6 8.47 5.23 3.4 13 lower antero-lateral nrm-pz p15870 2 2.6 10.6 11.5 0.5 2.2 1.6 11.5 9.41 5.94 2.8 3.7 13.5 lower antero-lateral nrm-pz p15871 4 2.5 9.57 8.36 0.3 2.3 1.9 7.96 4.3 4.1 13.9 lower antero-lateral nrm-pz p15872 5 3.3 12.3 12 12.2 18.5 2.8 2.1 15.4 10.2 8.1 5.4 5.8 17.4 upper lateral nrm-pz p15873 5 2.4 8.9 10.1 1.65 2.2 2.3 8 5.72 3.7 4 12.6 lower antero-lateral nrm-pz p15874 5 2.3 9.41 8.4 9.23 11.7 1.8 1.8 10.3 7.86 6.74 142.5 11.5 3.4 3.5 11.8 upper lateral nrm-pz p15876 4 3 10.2 9.63 10.4 5.05 2.3 2 11.4 8.44 6.34 130.3 2.6 12.2 upper antero-lateral nrm-pz p15878 5 3 12.9 13.6 14.9 11.8 8.08 5.2 upper lateral nrm-pz p15882 5 3.2 11.5 11.7 2.4 1.9 14.1 9.89 8.59 3.8 5.4 15.3 upper lateral nrm-pz p15883 2 2.9 10.3 13 2.3 2 13.2 9.23 6.98 5.8 upper lateral nrm-pz p15885 5 2.5 8.6 4.99 2.1 1.8 11 7.37 6.9 128.7 2.9 4.9 11.5 upper antero-lateral nrm-pz p15886 4 1.9 8.41 6.22 6.74 12.7 1.7 1.4 8.56 5.44 5.15 135.6 2.5 3.3 8.69 upper lateral nrm-pz p15887 5 2.5 9.25 13.2 2 1.7 11.6 8.18 7.46 147.4 2 3.6 11.2 upper lateral nrm-pz p15888 5 2.5 9.52 8.14 9.12 3.43 2.5 1.9 9.84 6.98 5.61 125.5 12.3 3.2 3.4 11.3 lower lateral nrm-pz p15926 4 2.6 8.88 9.97 1 1.9 1.9 7.76 5.64 3.5 3.9 12.4 lower antero-lateral nrm-pz p15928 5 nrm-pz p15929 4 2.5 7.95 8.23 8.9 3.96 1.8 1.3 9.46 7.44 5.3 3.7 3.7 11.9 lower lateral nrm-pz p15930 4 1.8 9.31 10 12.5 2.3 2.1 8.25 6.79 upper lateral tab. 1 tooth measurements (lengths in mm; angles in degree) and related position. abbreviations: bct , basal crown thickness; bcw, basal crown width; ch, crown height; dcl, distal crown edge length; ds, degree of slant; lch, height of lateral cusplets; lcw, width of lateral cusplets; mcl, mesial crown edge length; pch, height of principle cusp; pcw, width of principle cusp; ra , angle between root lobes; rh, root height; rw, root width; rh, root height; rt, root thickness; th, total height of tooth. marramà g., engelbrecht a., mörs t., reguero m.a. & kriwet j.288 small and not very high lateral cusplets that may be absent in anterior and lateral teeth (see cappetta 2012; marramà & kriwet 2017), whereas those described by long (1992a) appears in our opinion more consistent with the diagnosis of the lower antero-lateral or lateral teeth of brachycarcharias, although we do not exclude that they might belong to isurolamna. in this perspective, the presence of lamna in the la meseta formation and more generally in the eocene can be definitively excluded, thus supporting the hypothesis that the origin of the genus is probably oligocene or miocene (cappetta 2012), since the earliest fossil occurrences so far of true lamna nasus are from late miocene of the north sea (peters 2009; mollen 2010). based on the present study, the morphology of the isolated teeth from the ypresian of the la meseta formation here studied, as well as those reported by long (1992a) as lamna cf. nasus, are consistent with the diagnosis of teeth of brachycarcharias lerichei described and figured by cappetta & nolf (2005), van den eeckhaut & de schutter (2009), cappetta (2012), cappetta and case (2016), and marramà et al. (2017b). description. a detailed list of tooth measurements is given in table 1. the largest tooth is an upper lateral tooth of 17.41 mm total height (th), slightly smaller than the maximum size of 25 mm reported by cappetta (2012) for the species. teeth belonging to different jaw positions (upper and lower teeth, anteriors, antero-laterals and laterals) have different morphologies, therefore suggesting that the dentition is monognathic and dignathic heterodont and of tearing type. the single anterior tooth recognized in our sample (nrm-pz p15842; fig. 2a) shows a straight and triangular cusp, which is not very high and regularly decreasing in width from the base to the apex. the cutting edges reach the base of the crown, and fig. 2 brachycarcharias lerichei (casier, 1946) from the ypresian (early eocene) of la meseta formation, seymour island, antarctica. each tooth is shown (from left to right) in labial, profile and lingual view. anterior tooth: a) nrm-pz p15842. lower lateral teeth: b) nrm-pz p15843; c) nrm-pz p15854; d) nrm-pz p15777; e) nrm-pz p15859; f) nrm-pz p15836; g) nrm-pz p15839; h) nrm-pz p15929; i) nrmpz p15888; j) nrm-pz p15779. scale bars 2 mm. brachycarcharias (lamniformes, odontaspididae) from the eocene of antarctica 289 its profile is slightly curved. the lingual crown face of the cusp shows a strongly convex profile and short lingual folds are present. the labial crown face of the cusp overhangs the root by a little protruding bulge, and is characterised by the presence of very short and slender folds close to the crownroot junction. there was originally a single pair of high and upright cusplets, well separated from the main cusp, although only one is preserved in the specimen. the root is missing. the cusp of lower antero-lateral teeth is upright and almost symmetrical in labial and lingual views (e.g., fig. 2b–d). the base of the crown is larger than that of the anterior tooth. lower antero-lateral teeth bear one to two pairs of lateral cusplets: the proximal one is always large, triangular in shape, with apices often diverging from the main cusp. the distal lateral cusplets, when present, are smaller than the proximal ones. the holaulacorhizous root lobes have flat distal extremities. the nutritive axial furrow on the lingual protuberance is deep and strongly developed with a marked nutritive foramen. more lateral teeth of the lower jaw (e.g., fig. 2g–j) possess a lower cusp with respect to antero-lateral ones. a single probable upper intermediate tooth of about 10 mm th has been recognized in our sample (nrm-pz p15837; fig. 3a). the crown height is about 60% of th. the labial bulge of the crown is wide. the main cusp is distally bent. there are two lateral cusplets with the distal one being smaller than the proximal one. in upper antero-lateral teeth (e.g., fig. 3b) the main cusp is slightly distally inclined. the mesial cutting edge is oblique and almost straight, whereas the distal one is more vertical and slightly sigmoidal. a weak depression is recognizable at the base of the crown in labial view. short and faint folds are always present basally on lingual face. the proximal lateral cusplets are broad, and a smaller second pair of fig. 3 brachycarcharias lerichei (casier, 1946) from the ypresian (early eocene) of la meseta formation, seymour island, antarctica. each tooth is shown (from left to right) in labial, profile and lingual view. intermediate tooth: a) nrmpz p15837. upper lateral teeth: b) nrm-pz p15772; c) nrm-pz p15773; d) nrm-pz p15857; e) nrm-pz p15796; f) nrmpz p15848; g) nrmpz p15789; h) nrmpz p15806; i) nrm-pz p15841; j) nrm-pz p15780 (lateralmost tooth). scale bars 2 mm. marramà g., engelbrecht a., mörs t., reguero m.a. & kriwet j.290 distal cusplets can be recognised at least partially in nrm-pz p15772 (fig. 3b). the root is asymmetric with a mesial lobe that is longer than the distal one. upper teeth of more lateral position (fig. 3c–i) have a triangular cusp, which is strongly bent distally. the cusp appears slightly flattened labio-lingually. the labial face of the crown is smooth and almost flat with only a shallow median concavity at base. the lingual face is gently convex; short and weak lingual folds are present at the base of the crown. the cutting edges extend from the apex down to the base of the crown. the mesial cutting edge of the main cusp is oblique and slightly convex, whereas the distal one is concave. the labial bulge of the crown is wide and distinctly overhangs the root. there are often two pairs of lateral cusplets: the proximal lateral cusplets are broad and triangular, whereas the distal ones are significantly reduced. the cutting edges of the proximal cusplets are separated from those of the main cusp by a distinct notch. the root is low and broad, with well-separated lobes. the lingual furrow on the protuberance is weak or sometimes appears absent. in the single upper lateralmost tooth recognized (fig. 3j) the main cusp is strongly bent distally and its crown is reduced in size. the labial bulge of the crown is mesio-distally wide. the two pairs of lateral cusplets are low and the apices of the proximal ones appear rounded, although we cannot exclude that this is due to abrasion. the root is bulky and wider than the total height of the tooth. discussion comparisons. the analysis of the material from the la meseta formation on seymour island has revealed the presence of several characters that support the assignment of these teeth to the family odontaspididae, including a monognathic and dignathic heterodonty, a tearing type dentition, well-developed and separated root lobes, a marked nutritive furrow, and a tall and slender main cusp (compagno 1984, 1999; cappetta 2012). teeth of this ypresian sand tiger shark of antarctica can be distinguished from other eocene odontaspidids, and in particular from those occurring in the same horizons on seymour island (i.e., odontaspis, palaeohypotodus and striatolamia) by their unique combination of characters such as a triangular, not very high cusp, which decreases regularly in width distally. for the same reason they can also be easily separated from those of jaekelotodus (wide and bulky cusp), carcharias (very sharp and high cusp), and hypotodus (robust cusp) (see cappetta & nolf 2005; cappetta 2012). the presence of fine folds on the basal labial face of the main cusp distinguishes the teeth from those of araloselachus, glueckmanotodus, hypotodus, jaekelotodus, odontaspis, and orpodont which lack any labial or lingual crown ornamentation, whereas palaeohypotodus is characterized by a series of tubercles or strong ridge-like folds boarding the base of the labial face (leriche 1951; cappetta & nolf 2005). the cutting edges in b. lerichei extend the full height of the crown. this character additionally separates the teeth from the la meseta fm from those of araloselachus, carcharias, hypotodus, and odontaspis, in which the cutting edges do not reach the base of the crown (cappetta & nolf 2005; mannering & hiller 2008; cappetta 2012). although the cutting edges reach the base in jaekelotodus and sylvestrilamia, the teeth of the former are larger (up to 45 mm high) and have a wide and bulky main cusp that is almost the same height as the root, whereas sylvestrilamia possesses a main cusp with a more flexuous profile and always bears single pair of small lateral cusplets, a feature that also characterizes the teeth of hypotodus (cappetta & nolf 2005; cappetta 2012). the teeth from the la meseta fm can be easily separated from those of araloselachus, borealotodus, carcharias, glueckmanotodus, hypotodus, jaekelotodus, mennerotodus, sylvestrilamia, tethylamna, and turania by the presence of well-developed lateral cusplets (cappetta & nolf 2005; cappetta 2012; cappetta & case 2016). odontaspis, orpodon, and palaeohypotodus also possess tall lateral cusplets but, contrary to brachycarcharias, teeth of orpodon are much smaller (less than 12 mm), odontaspis possess a very high, sharp and non-sigmoidal main cusp that is often flanked by up to three pairs of cusplets in lateral teeth, whereas palaeohypotodus possesses short and strong folds on the labial bulge, and strong and irregular serrations at the base of the cutting edges (compagno 1984; cappetta 2012). finally, the stratigraphic age of the teeth can provide additional information for separating the teeth described here from araloselachus, orpodon, and tethylamna since these genera are unknown from ypresian deposits (cappetta & nolf 2005; cappetta 2012; cappetta & case 2016). howbrachycarcharias (lamniformes, odontaspididae) from the eocene of antarctica 291 ever, stratigraphic distribution might not be a valid reason for distinguishing taxa. currently, four valid species are recognized within the genus brachycarcharias: b. atlasi, b. koerti, b. lerichei, and b. mississippiensis (see also ‘remarks’ above). teeth from the la meseta fm can be easily separated from those of b. atlasi in the absence of strong lingual folds and a well-developed second pair of lateral cusplets (arambourg 1952) and from those of b. koerti for the presence of lower and broader lateral cusplets (see dartevelle & casier 1943; underwood et al. 2011). finally, although teeth of b. lerichei are morphologically very similar to those of b. mississippiensis, which cappetta (2006) considers to be synonymous with b. lerichei, we can exclude that our teeth pertain to this species mostly for biogeographic reasons, since b. mississippiensis was only restricted to the eastern coasts of north america according to our current knowledge (case 1994; case et al. 2015). the significance of b. lerichei in the eocene of antarctica. this report represents the southernmost occurrence of the extinct sand tiger shark brachycarcharias. the abundance of brachycarcharias lerichei teeth in telms 2, 4 and 5 of the la meseta fm on seymour island is in accordance with the increase in species richness of elasmobranchs and holocephalans detected from the earliest to the latest ypresian of the la meseta formation (kriwet et al. 2016). this ypresian occurrence in antarctica supports the hypothesis that this species was probably an opportunistic top predator having a wide range of habitats and feeding preferences (marramà et al. 2017b). remains of b. lerichei were in fact abundantly recovered from tropical shallowto deep-water deposits of the northern hemisphere (see e.g., arambourg 1952; ward & wiest 1990; case 1994; noubhani & cappetta 1997; smith et al. 1999; cappetta & nolf 2005; adnet & cappetta 2008; diedrich 2012; cicimurri & ebersole 2015; cappetta & case 2016). more recently, isolated teeth of this species have been also reported from the celebrated ypresian bolca lagerstätte of italy (marramà et al. 2017b, c). it has been suggested that the mean annual surface seawater paleotemperatures for this early eocene deposit located in tethys realm was very high (giusberti et al. 2014). in fact, the fossil occurrence of the specialized marine water strider halobates suggested that sea surface paleotemperatures in the bolca paleobiotope exceeded 20°c, representing the lower temperature tolerated by the extant species of this genus (andersen et al. 1994; cheng et al. 2012). this concurs to suggest a tropical environment with high mean annual surface paleotemperature for the bolca paleobiotopes (see also carnevale et al. 2014; marramà et al. 2016c). on the contrary, paleotemperatures detected for telms 2 to 5 of the la meseta fm of the antarctic peninsula were remarkably lower, around 10-11° c, as suggested by ivany et al. (2008), thus resulting in an association of chondrichthyans adapted to cooler waters (kriwet et al. 2016), although non-resident warm-water genera as carcharhinus and pristis have been also reported (kriwet 2005). the first unambiguous occurrence of b. lerichei from the southern hemisphere therefore supports the hypothesis of marramà et al. (2017b) that this species was an opportunistic cosmopolitan top predator having a wide range of habitat preferences, contrary to the hypothesis of maisch et al. (2014) who supposed a life style more similar to that of the living porbeagle shark, lamna nasus, a lamnid species today only restricted to temperate and cool waters and never occurring in tropical seas (compagno 1984; compagno et al. 2005), although more recent studies suggest that l. nasus might be more heat-tolerant than previously thought (campana et al. 2010). in this perspective, we also hypothesize that b. lerichei might have been an eurytherm species, tolerating wide ranges of temperatures, from tropical warm waters of the bolca lagerstätte to the cooler contexts of the la meseta formation. this remarkable adaptation probably enabled this lamniform shark to become one of the most successful odontaspidid taxa of the early paleogene, inhabiting a wide range of latitudes of the northern and southern hemisphere, which is also consistent with high vagility and a circum-global distribution as observed in living odontaspidids (musick et al. 2004). the danian marks the first occurrences of brachycarcharias, with at least three species (b. lerichei, b. atlasi, and b. mississippiensis) occurring in deposits of the northern hemisphere (see fig. 4). mannering & hiller (2008) reported an indeterminate species of brachycarcharias from the paleocene of the southern hemisphere (new zealand). however, the occurrence from the paleocene of new zealand appears, in our opinion, controversial. teeth figured by mannering & hiller (2008, fig. 11l–n) as lowmarramà g., engelbrecht a., mörs t., reguero m.a. & kriwet j.292 er laterals of brachycarcharias sp. possess only small, narrow and not very high lateral cusplets, a very sigmoidal main cusp in profile view, and strongly bent distally. conversely, lower lateral teeth of brachycarcharias always possess robust, high and upright cusplets and a main cusp that is only slightly sigmoidal and which is not (or only slightly) bent distally in lower teeth (cappetta & nolf 2005; van den eeckhaut & de schutter 2009; cappetta & case 2016). moreover, teeth of brachycarcharias sp. figured as upper anteriors by mannering & hiller (2008, fig. 11i– k) possess very small, low lateral cusplets, a low root with robust and very divergent lobes, a very robust and wide main cusp base. in comparison, upper anterior teeth of brachycarcharias show very large and upright lateral cusplets, a comparably higher root with more tapered lobes forming a smaller angle, and a crown decreasing regularly in width apically (see van den eeckhaut & de schutter 2009; marramà et al. 2017b). the exclusion of brachycarcharias from the paleocene of new zealand might be also supported by biogeographic reasons, since the high clustering of brachycarcharias occurrences in the paleocene of the northern hemisphere suggests that the centre of origin of this taxon would have taken place between north america and the tethys realm. if we exclude the doubtful brachycarcharias teeth from the paleocene of new zealand, the ypresian occurrence of b. lerichei in antarctica, beside representing the southernmost occurrence of the genus, also implies that this species reached the southern hemisphere in concomitance with its maximum geographic distribution (see also marramà et al. 2017b) via a dispersal corridor along the continental margins of the western atlantic ocean, as suggested for other elasmobranchs (long 1994). the widespread ypresian distribution of b. lerichei (well-before the maximum expansion of the genus with b. koerti in the lutetian of southwestern africa) was probably due to the concomitance of its opportunistic strategy, eurytherm adaptation, high vagility, and favourable global high temperatures in the context of the early eocene climatic optimum. brachycarcharias disappeared at the end of the eocene, with the last occurrences from priabonian deposits of georgia, usa (as ‘carcharias’ koerti in parmley et al. 2003), mexico (gonzález-barba 2003), morocco (gajić et al. 2014), japan (tanaka et al. 2006) and louisiana, u.s.a. (breard & stringer 1995). the absence of brachycarcharias in telms 6 (lutetian) and 7 (priabonian) of the la meseta formation on seymour island is consistent with the fig. 4 schematic simplified maps showing the paleobiogeographic distribution of brachycarcharias cappetta & nolf, 2005. localities: 1 mexico, 2 mississippi, 3 maryland, 4 france, 5 morocco, 6 new zealand, 7 tunisia, 8 south carolina, 9 virginia, 10 georgia, 11 italy, 12 england, 13 belgium, 14 austria, 15 algeria, 16 congo, 17 angola, 18 alabama, 19 egypt, 20 germany, 21 nigeria, 22 togo, 23 uzbekistan, 24 spain, 25 north carolina, 26 senegal, 27 texas, 28 louisiana, 29 japan. the yellow star marks the ypresian occurrence of b. lerichei in the la meseta fm of seymour island, antarctica. data from marramà et al. (2017b). maps modified from scotese (2002). brachycarcharias (lamniformes, odontaspididae) from the eocene of antarctica 293 reduction and disappearance of cartilaginous fishes during the latest phase of the eocene in this region that was related to the progressive cooling, reduction of shelf areas, and establishment of antarctic ice sheets (grande & eastman 1986; kriwet et al. 2016). in this perspective, the gradual disappearance and extinction of brachycarcharias, as well as other odontaspidids such as striatolamia and palaeohypotodus at the end of the eocene, might be related, at least in part, to a major wave of biotic turnovers that took place at the e–o boundary and that particularly involved marine top predators (prothero et al. 2003; marramà et al. 2017b). this turnover of taxa at the eocene-oligocene transition was the most severe extinction event in the cenozoic and was likely related to significant temperature declines and sea level falls (e.g., zachos et al. 2001; liu et al. 2009; harnik et al. 2012). it has also been suggested that eocene odontaspidids were partly replaced by other odontaspidids in the oligocene (e.g., araloselachus, new carcharias species) and predominantly by lamnids (e.g., carcharoides, isurus, lethenia) that might have been better adapted to cooler environments (marramà et al. 2017b). conclusions eocene shallow marine sediments of the la meseta fm on seymour island yielded the most diverse paleogene ichthyofauna from the southern hemisphere to date. this fauna provides important insights into diversity and distribution patterns of both cartilaginous and bony fishes during crucial time intervals in global climatic developments such as the early eocene climatic optimum (ca. 53-49 ma) resulting in extensive greenhouse conditions. the first report of the extinct sand tiger shark brachycarcharias from ypresian strata of the la meseta formation, seymour island, antarctica, helps to better understand the paleobiology and paleobiogeography of this paleogene taxon. teeth of brachycarcharias lerichei were recovered from telms 2 (early ypresian), 4 and 5 (late ypresian) and an increase in raw numbers from 4 teeth in the early ypresian to 45 teeth in the late ypresian is recognizable. the rather high abundance and increase of b. lerichei teeth in the la meseta fm agrees well with the general pattern of taxonomic diversity increase of chondrichthyans postulated for this assemblage. the wide habitat and geographic distribution of this species, ranging from deep-waters to shallow water environments and from tropical to cool-temperate climates, indicate that b. lerichei was a rather opportunistic, cosmopolitan, highly vagile, and eurytherm top predator in marine food-webs, which might have been the reasons for its success during the paleocene and eocene. at the end of the eocene, however, this species vanished, which correlates with decreasing global temperatures and the appearance of new, probably better adapted lamniform sharks. acknowledgements: the argentinian antarctic institute (iaadna), argentinian air force and swedish polar research secretariat (spfs) are acknowledged for logistic support for field-work on seymour island. the authors are grateful to martin de los reyes, museo de la plata, for picking the small fractions in the laboratory. thanks are also due to iris fuchs (naturhistorisches museum wien) for the photographs. we also thank todd d. cook (penn state behrend), alberto collareta (università di pisa), márton szabó (geological and geophysical institute of hungary), and an anonimous reviewer for their valuable comments that improved the quality of the manuscript. this work was supported by the austrian science fund (fwf) [m2368-b25 to g.m., and p26465-b25 to j.k.]; the graduation scholarship of the university of vienna to a.e.; the swedish research council grant [vr grant number 2009-4447 to t.m.]; the consejo nacional de investigaciones científicas y técnicas grant [conicet grant number pip 0462 to m.r.]; the argentinian national agency for promotion of science and technology grant [anpcyt grant number picto 0093/2010 to m.r.]. references adnet s. & cappetta h. 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(+'-&( 0�$)."&(� -%-#-.(&� #$.0�'%-& $.0 �&"%$(-0 (--(+ 4"! /�!����� ���!! �����!� 5 �� 2�!!��� 6�5� �� 78� �&����� � ����!�9 ��� lab. code archaeological layer cultural attribution sample conventional radiocarbon age (bp) calibrated radiocarbon age (bp) (2 sigma, 95%) dsh2816 3.1.l byzantine-late roman vegetal remain 1260±34 1320-1120 calbp dsh2814 3.2.c late roman charcoal 1332±26 1340-1220 calbp dsh2815 3.4.b seed 3244±42 3600-3360 calbp dsh1976 3.4.g human bone 3948±35 4500-4270 calbp beta-314642 4.2 charcoal 9450 ± 50 10860-10540 calbp early bronce-middle neolithic 3$'2 � � �'&"%9($)-& "4 (+�$%%".�.4-!." %-/-%&2 nisp mni % nisp mni % nisp mni % nisp mni % nisp mni % od oh w r wa taxa/levels erinaceus europaeus 0 0 0.0 0 0 0.0 1 1 5.0 7 3 2.4 0 0 0.0 0.25 0.75 crocidura cf. c. sicula 2 2 14.3 0 0 0.0 2 1 5.0 10 6 4.8 12 6 28.6 0.25 0.25 0.5 suncus etruscus 0 0 0.0 0 0 0.0 0 0 0.0 1 1 0.8 0 0 0.0 1 rhinolophus sp. 0 0 0.0 0 0 0.0 1 1 5.0 0 0 0.0 0 0 0.0 0.75 0.25 myotis gr. myotis-blythii 0 0 0.0 0 0 0.0 0 0 0.0 2 1 0.8 1 1 4.8 0.5 0.5 pipistrellus sp. 0 0 0.0 0 0 0.0 0 0 0.0 1 1 0.8 0 0 0.0 0.75 0.25 minopterus schreibersii 0 0 0.0 0 0 0.0 0 0 0.0 0 0 0.0 4 2 9.5 0.25 0.25 0.5 arvicola cf. a. amphibius 0 0 0.0 0 0 0.0 0 0 0.0 3 2 1.6 0 0 0.0 1 microtus (terricola ) savii 2 2 14.3 0 0 0.0 8 4 20.0 104 58 46.4 16 8 38.1 0.75 0.25 apodemus sylvaticus 5 4 28.6 0 0 0.0 9 5 25.0 57 29 23.2 8 4 19.0 1 rattus norvegicus 5 4 28.6 0 0 0.0 0 0 0.0 0 0 0.0 0 0 0.0 0.5 0.5 eliomys quercinus 1 1 7.1 0 0 0.0 0 0 0.0 2 1 0.8 0 0 0.0 0.5 0.5 glis glis 1 1 7.1 2 2 100 15 7 35.0 42 20 16.0 0 0 0.0 1 muscardinus avellanarius 0 0 0.0 0 0 0.0 1 1 5.0 5 3 2.4 0 0 0.0 1 total 16 14 100 2 2 100 37 20 100 234 125 100 41 21 100 2 1 7.25 1.75 2 4.23.1 3.2 3.3 3.4 3$'2 � � 3+.9#'-! 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(2017) calcareous nannoplankton response to the latest cenomanian oceanic anoxic event 2 perturbation. riv. it. paleontol. strat., 123(1): 159-176. rivista italiana di paleontologia e stratigrafia (research in paleontology and stratigraphy) vol. 123(1): 159-176. march 2017 abstract. morphometric analyses were performed on biscutum constans, zeugrhabdotus erectus, discorhabdus rotatorius and watznaueria barnesiae specimens from five sections spanning the cenomanian-turonian boundary interval including oceanic anoxic event (oae) 2 (~ 94 ma). the study provides evidence for size fluctuations and dwarfism of b. constans during oae 2, followed by a partial recovery at the end of the event: this taxon appears to be the most sensitive species, with similar and coeval size trends in all the analyzed sections. conversely, morphometry shows negligible or unsystematic coccolith variations in z. erectus, d. rotatorius and w. barnesiae. the comparison of oae 2 data with those available for the early aptian oae 1a and latest albian oae 1d, indicates that b. constans repeatedly underwent size reduction and temporary dwarfism, possibly implying that the same paleoenvironmental factors controlled calcification of b. constans during subsequent oaes although the amplitude of b. constans coccolith reduction is significantly larger for oae 1a than oae 2. paleoceanographic reconstructions suggest that ocean chemistry related to the amount of co 2 and toxic metal concentrations played a central role in b. constans coccolith secretion, while temperature and nutrient availability do not seem to have been crucial. contrary to oae 1a, z. erectus, d. rotatorius and w. barnesiae appear to be substantially unrelated to oae 2 paleoenvironmental stress, possibly because of different degrees of perturbation. received: november 15, 2016; accepted: january 15, 2017 key words: morphometry; coccoliths; oceanic anoxic event 2; ocean acidification. introduction the geological record of the cretaceous world documents profound changes in the oceanatmosphere system, including volcanic injection of large amounts of co 2 , super-greenhouse conditions, oceanic anoxia and eutrophication (e.g. schlanger & jenkyns 1976; erba 2004; hay 2009; jenkyns 2010; föllmi 2012; bottini et al. 2015; erba et al. 2015). intervals of prolonged global anoxia, known as oceanic anoxic events (oae)s (schlanger & jenkyns 1976) were characterized by burial of unusually large amounts of organic matter during profound perturbations of the ocean-atmosphere system, similarly to current (and future) global changes. the latest cenomanian oae 2 (~94 ma) is arguably regarded as the most extreme paleoenvironmental stress related to the warmest climate of the past 150 my, characterized by an accelerated hydrological cycle, enhanced production and burial of organic matter, very high concentrations of volcanically-produced co 2 and altered chemistry and structure of the oceans (see review provided by jenkyns 2010). in the time interval encompassing oae 2, calcareous nannoplankton were forced to face anomalous paleoceanographic and paleoclimatic conditions that induced extinctions and originations (e.g. bralower 1988; leckie et al. 2002; erba 2004). quantitative studies of nannofossil assemblages in various geological settings revealed differential abundance patterns of fertility-related species, in particular of biscutum constans and zeugrabdothus erectus. in general, an eutrophication episode was reconstructed at low latitudes (hardas & mutterlose 2007) while a shift to oligotrophic conditions was evidenced at mid-latitudes (gale et al. 2000; linnert et al. 2010, 2011). in the western interior seaway (wis) local changes controlled the trophic level across oae 2 (corbett & watkins 2013). in this work, we focus on size variations of four species, namely biscutum constans, discorhabdus rotatorius, watznaueria barnesiae and zeugrabdothus erectus in the cenomanian-turonian boundary interval (ctbi), and specifically across oae 2. these taxa were previously studied through the early aptian oae 1a (erba et al. 2010; lübke et al. 2015; lübke & mutterlose 2016), the latest albian oae 1d (borfaucher g., erba e., bottini c. & gambacorta g.160 nemann & mutterlose 2006) and the ctbi (linnert & mutterlose 2012) providing insights into coccolith size changes across oaes. in particular, erba et al. (2010) demonstrated that calcareous nannoplankton was extremely sensitive to ocean acidification associated to oae 1a at low latitude sites in the tethys and pacific oceans, allowing separation of most-, intermediate-, and least-tolerant taxa: dwarfism of b. constans, z. erectus and d. rotatorius coccoliths was documented during oae 1a but no significant changes in w. barnesiae dimension were detected, although more elliptical and malformed specimens were identified (erba et al. 2010). dwarfism of b. constans and z. erectus was documented also across oae 1a at higher latitudes (north sea and lower saxony basin, lübke & mutterlose 2016), but explained as the response to decreased light in surface waters under conditions of intense run off. in the same sections, w. barnesiae remained stable without significant size variations and interpreted as a robust species with respect to changes of sunlight penetration. during oae 1d, morphometry of b. constans specimens attested a decrease in mean length, while the average size of w. barnesiae coccoliths remained constant (bornemann & mutterlose 2006). linnert & mutterlose (2012) investigated nannofossil morphometry at two deep sea drilling project (dsdp) sites from goban spur in the cenomanian-turonian interval, although hiatuses across oae 2 hampered the reconstruction of detailed size variations in the ctbi. however, slightly smaller biscutum specimens were observed in the lower turonian compared to the upper cenomanian interval, while no significant changes in size were detected for genus watznaueria. sections from various oceanic basins were selected on the basis of their stratigraphic characterization, ensuring relatively high resolution and correlation at supra-regional scale. the main objectives of this study are: i) the assessment of size variations of b. constans, d. rotatorius, w. barnesiae and z. erectus across oae 2; ii) the evaluation of timing and type of reactions of phytoplankton calcification through progressively stressing conditions and their demise; iii) the identification of global, regional and local response to paleoenvironmental perturbations; iv) the evaluation of synchroneity or diachroneity of nannoplankton variations versus paleoenvironmental changes; v) implementation of paleoecological reconstructions of abiotic and biotic parameters influencing coccolith secretion across oaes. in particular, we want to test the following hypotheses: (1) did short-term fluctuations in coccolith morphometry follow transient changes in climate and ocean chemistry across oae 2?; (2) did b. constans, d. rotatorius, w. barnesiae and z. erectus respond to oae 2 perturbations similarly to other oaes?; (3) were single or concomitant paleoenvironmental factors influencing calcareous nannoplankton calcification?; (4) were changes in coccolith size abrupt or gradual, especially at the onset and demise of oae 2?; (5) were the paleolatitudes discriminating for coccolith size ? location/studied sections we studied five stratigraphic sections covering the ctbi (figs 1 and 2), including eastbourne (east sussex, england), clot chevalier (vocontian basin, france), novara di sicilia (northeastern sicily, italy), rock canyon (pueblo, colorado, usa) and cuba (northern-central kansas, usa) sections from the wis. these sections were chosen based on availability of integrated stratigraphy granting a good time control derived from c-isotopic stratigraphy and biostratigraphy. eastbourne is the thickest section covering the ctbi in the anglo-paris basin (gale 1995, 1996; gale et al. 1999; paul et al. 1999). the studied outcrop was sampled at gun gardens and consists of a 27 m-thick section comprising three lithological units: grey chalk (rhythmically bedded marly chalk), plenus marls (greenish grey marls and clay-rich chalk) and white chalk (coccolith and calcisphere-dominated nodular limestone). samples analyzed were collected during the ct-net project and nannofossil biostratigraphy combined with chemostratigraphy was published in tsikos et al. (2004): oae 2 is constrained by the last occurrences of corollithion kennedyi and axopodorhabdus albianus and the first occurrence of quadrum gartneri at oae 2 onset and end, respectively. these nannofossil events are consistent with more recent biostratigraphic data by linnert et al. (2011). the novara di sicilia section consists of a 19 m-thick stratigraphic interval measured through an olistolith within a melange of the argille varicolori (scopelliti et al. 2008). from a lithological oceanic anoxic event 2 161 point of view, the succession can be divided into three parts: a lower part with alternations of thick marly limestone with light and dark marlstone, an intermediate portion with thinner light-colored strata and an upper part of black shale alternating with dark marlstone and claystone. the nannofossil and planktonic foraminiferal biostratigraphy, integrated with c-isotopic stratigraphy (scopelliti et al. 2008) indicate that the novara di sicilia section contains only the lower part of the bonarelli level equivalent and the underlying interval. specifically, the outcrop ends shortly above δ13c org peak b indicating that only the onset and early part of oae 2 are represented. the clot chevalier section is located in the vocontian basin (southeastern france) that was a part of the hemipelagic intrashelf basin of the european tethyan passive margin (wilpshaar et al. 1997). the 35 m analyzed section consists of an alternation of dark grey marlstone and light grey limestones (falzoni et al. 2016). the local equivalent of the “bonarelli level” in the clot chevalier section is represented by the “thomel level”, which is characterized by laminated organic-rich sediments (from 3 to 31 meters from the base of the section) (falzoni et al. 2016). at the base of th1 unit, the presence of a borrowed sharp surface reveals the existence of a hiatus in the lower part of the section. furthermore, within units th1 and th2, a condensed stratigraphic level has been observed and interpreted as an interval of very low sedimentation rate or sedimentation interruption. nannofossil biostratigraphy was achieved by russo (2014). the rock canyon-pueblo and the cuba sections are both from the wis where the cenomanian/turonian boundary lies within the bridge creek limestones member of the greenhorn formation. the rock canyon section is located in southern-central colorado near pueblo, along the arkansas river; it was ratified as the global boundary stratotype section and point (gssp) for the base of the turonian stage (kennedy et al. 2005). here, the bridge creek limestones consist of alternating pelagic chalk and limestone and c org -rich beds (marlstone and calcareous shale), mainly bioturbated, with laminated and sublamintated units. the rock canyon section has been intensively investigated for litho-, bio-, chemostratigraphy and for paleoecological and paleoenvironmental reconstructions (leckie 1985; pratt 1985; pratt et al. 1993; bralower 1988; kennedy 2000; keller & pardo 2004; snow et al. 2005; sageman et al. 2006; corbett & watkins 2013). the analyzed interval goes from 0.3 m to 10.2 m: samples from the preoae 2 interval were not available. the cuba section in northern-central kansas, is about 600 km east of the rock canyon section. equivalent strata around cuba consist of more fine-grained calcareous to chalky shale within the jetmore chalk and pfeifer shale members of the greenhorn formation. these two members are dominated by marlstone and calcareous shale (bowman & bralower 2005). at the cuba section the stratigraphic control is based on integrated litho-, chemo-, biostratigraphy (hattin 1975, 1985; bowman & bralower 2005; eleson & bralower 2005; desmares et al. 2007; corbett & watkins 2013; corbett et al. 2014). the analyzed samples were previously investigated for chemoand biostratigraphy by bowman & bralower (2005) and eleson & bralower (2005). cc e nds western tethys wis proto north atlantic p c caribbean plateau 0 30° 30° 60° 60° fig. 1 paleo-location of the studied sections at ~ 94 ma (late cenomanian). e = eastbourne, cc = clot chevalier, nds = novara di sicilia, p = pueblo and c = cuba, wis = western interior seaway. the paleo-map is from du vivier et al. (2014). faucher g., erba e., bottini c. & gambacorta g.162 methods calcareous nannofossils were investigated in smear slides under light polarizing microscope, at 1250x magnification. smear slides were prepared using standard techniques (bown & young 1998) without centrifuging and/or ultrasonic cleaning in order to retain the original sediment composition. a small fraction of sediment was powdered in an agate mortar with bidistillate water; a few drops of the suspension were extracted with a pipette and spread over a slide, leaving to dry on a hot plate. the dried suspension was sealed on a glass slide with norland optical adhesive. the nannofossil preservation was carefully assessed in order to avoid diagenetically altered material. in fact, partial dissolution and/or overgrowth have the potential to artificially decrease or enlarge the coccolith size, respectively. this is particularly relevant for the delicate biscutum, zeugrhabdotus and discorhabdus coccoliths that are diagenetic-prone (e.g. thierstein 1980). accurate screening of preservation was achieved under light polarizing microscope to ascertain the degree of etching and/or overgrowth. moreover, individual b. constans, z. erectus, d. rotatorius and w. barnesiae coccoliths considered for morphometry were selected based on their complete and continuous outline and lack of crimping due to etching or overgrowth. we also tested the potential control and its degree of lithology on coccolith preservation, because marlstones normally contain the best preserved nannofossil assemblages, while coccoliths are often affected by overgrowth in limestones and by etching in carbonatepoor lithotypes (thierstein & roth 1991). morphometric analyses were performed on a total of 53 samples from eastbourne, 37 from clot chevalier, 40 from novara di sicilia, 21 from pueblo and 18 from cuba sections. for each sample 30 specimens of b. constans, z. erectus and d. rotatorius and 50 specimens of w. barnesiae were digitally photographed in random fields of view. samples containing less than 30 specimens of selected species were not included in the analyses. morphometry of w. barnesiae was conducted with a lower sampling resolution. images of coccolith specimens were taken using a camera mounted on a leitz laborlux light microscope. length and width (the diameter for d. rotatorius) of coccolith distal shield were measured on digital images using imagej64 software. a total of 4740 of b. constans, 4500 of z. erectus, 4770 of d. rotatorius and 7500 of w. barnesiae were measured. the error of measurements is of ± 0.08 µm. data were statistically elaborated, using r software, to obtain mean, median, maximum and minimum values, 25% and 75% percentile, 95% confidence intervals, standard deviations, pearson correlation coefficient (r) and the linear regression function. moreover, the analyses of variance (anova) and tukey post hoc tests were performed with r software in order to determine if size variations are statistically significant. results in all studied samples preservation is moderate and only very minor indications of etching and overgrowth were sporadically detected. our diagenetic estimates are in agreement with evaluation of dissolution and overgrowth previously published for nannofossil assemblages. in particular, at pueblo, bralower (1988) and corbett et al. (2014) documented moderate preservation as assessed in this study. similarly, linnert et al. (2011) and scopelliti et al. (2008) observed moderately preserved assemblages at eastbourne and novara di sicilia, respectively. good to excellent preservation was documented for nannofloras of the cuba section (eleson & bralower 2005). samples analyzed from the five sections randomly consist of various lithologies, all yielding very similar degree of preservation and there is no correlation between coccolith size and carbonate content. diagenetic modifications were certainly different for the studied sections were deposited at different paleo-water depths and in various geological settings, under diverse sedimentation regimes. however, similar patterns in nannofossil assemblages with coexistence of delicate and robust species points to negligible diagenetic influence. moreover, we underline that morphometric data were gathered only for complete specimens without signs of corrosion or overgrowth in the outline. the adopted stratigraphic framework is based on the δ13c curve integrated with nannofossil biostratigraphy (fig. 2). specifically, peaks a, b and c of δ13c curve (pratt et al. 1985; tsikos et al. 2004; jarvis et al. 2006; gambacorta et al. 2015) are taken as key-levels for dating and correlations. as discussed by jarvis et al. (2006, 2011), peak a represents the acme of the first shift towards δ13c positive values. it is followed by a trough and a subsequent increase culminating at δ13c peak b marking the onset of the so-called plateau. the end of the oae 2 c-isotopic anomaly correlates with δ13c peak c followed by the decrease back toward pre-excursion δ13c values (tsikos et al. 2004). in the next sub-chapters, the size fluctuations of b. constans, z. erectus, d. rotatorius and w. barnesiae coccoliths are presented for every section (fig. 3, supplementary tab. 1). since the correlation of length and width of b. constans, z. erectus and w. barnesiae coccoliths evidences a high pearson correlation coefficient (supplementary fig. s1, s2, s3), only the length is plotted and used for tracing size changes. eastbourne coccoliths of b. constans, z. erectus and d. rotatorius show similar size trends through the eastbourne section (fig. 3a). specifically, at the oae 2 onset, b. constans, z. erectus and d. rotatorius evidence a decrease in size becoming smaller than the mean values (blue line). subsequently, a slight increase oceanic anoxic event 2 163 in coccolith size is detected and higher values are reached across δ13c peak a. successively, coccoliths display a reduction in size reaching minimum values around δ13c peak b. in the uppermost part of the section, coccoliths evidence an increase in size becoming slightly larger than the mean size. w. barnesiae is rather constant through the event (see supplementary tab. 1), although a minor increase in size is observed in the interval between δ13c peaks c and d (fig. 3a). clot chevalier in the clot chevalier section, the interval preceding oae 2 is rather thin and only a few samples were available for the analyses. size trends are well expressed in b. constans and d. rotatorius (fig. 3b). from the base, b. constans and d. rotatorius show a relative increase in size compared to the mean length-diameter. b. constans and d. rotatorius maintain bigger sizes in the lowermost part of oae 2 and then sizes progressively decrease just after δ13c peak a, reaching relative minimum values around δ13c peak b. b. constans displays values close to average in the upper part of δ13c plateau up to peak c where, along with d. rotatorius, there’s a minor increase in size. size fluctuations are less evident in z. erectus and w. barnesiae that do not show significant trends through the event. however, z. erectus shows a minor increase in size just after δ13c peak c. novara di sicilia in the novara di sicilia section, only the preoae 2 interval and the first part of the event are represented (fig. 3c). in the pre-oae 2 interval size values of b. constans, z. erectus and d. rotatorius are erratic (fig. 3c) with successive increases and decreases in size trends. in the first part of oae 2, after a basal interval barren of calcareous nannofossils, b. constans shows coccoliths larger than the mean size and subsequently a trend towards smaller coccoliths, followed by a recovery in size around δ13c peak a. a final size decrease ends with a relative minimum around δ13c peak b. z. erectus and d. rotatorius coccoliths display erratic values through oae 2 and w. barnesiae specimens show values around average through the studied interval. cuba in the lowermost portion of this section, only a few samples were available to characterize the pre-oae 2 interval (fig. 3d). maximum coccolith c ub a c en om an ia n tu ro ni an n . j ud di i s . g ra ci le m . m os by en se r . c us hm an i w . a rc ha eo cr et ac ea h . h el ve tic a w . d ev on e as tb ou rn e c lo t c he va lie r p ue bl o n ov ar a di s ic ili a a. albianus h. chiastia e. octopetalus c. kennedyi n c 11 b* * n c 12 ** n c 13 a* * q. gartneri q. intermedium p la nk to ni c f or am in ife ra l z on es a m m on ite z on es s ta ge nannofossil zones -27-28 -24 -23-26 -25 δ13corg‰ 543 6 δ13ccarb‰ a o a e 2 b c dc c 11 u c 7 c c 10 uc6 u c 5 u c 4 321 fig. 2 stratigraphic ranges of the studied sections through the late cenomanian-early turonian time interval. nannofossil zonations discussed in gambacorta et al. (2015). 1 = sissingh (1977); 2 = burnett (1998), 3 = gambacorta et al. (2015). faucher g., erba e., bottini c. & gambacorta g.164 fig. 3 variations of b. constans, z. erectus and w. barnesiae length, and d. rotatorius diameter against δ13c curve during oae 2 at eastbourne, clot chevalier, novara di sicilia, pueblo rock canyon and cuba. δ13c and nannofossil zones at eastbourne after tsikos et al. (2004); clot chevalier δ13c curve after falzoni et al. (2016), nannofossil zones after russo (2014); δ13c and nannofossil zones at novara di sicilia after scopelliti et al. (2008); δ13c at pueblo rock canyon after snow et al. (2005), nannofossil zones after russo (2014); cuba δ13c after bowman and bralower (2005), nannofossil zones after eleson and bralower (2005) integrated following gambacorta et al. (2015). 62 δ13ccarb a b c d 25 20 15 10 5 0 m tu ro ni an c en om an ia n nc 12 ** n c 11 * n c 13 ** q . g . r . c . a . a . w .d . m . g c . g f . c . n .j. 0 10 15 m -28 -22 δ13corg ‰ b on ar el li le ve l b a c en om an ia n n c 11 * e . o . e . e . a . a . n c 12 ** tu ro ni an c en om an ia n n c 11 n c 12 a* q .g . r .a h . c . s. g. n . j . w .d v.b m . m os by en se m . n od os oi de s nc 12** b d c a 40 0 20 0 80 0 60 0 0 -2 00 10 00 δ13corg-28 -22 cm e as tb ou rn e c lo t c he va lie r n ov ar a di s ic ili a p ue bl o a b c e a b c d 10 0 20 30 2.5 4.5 δ13ccarb (%vpdb) tu ro ni an c en om an ia n n c 11 * n c 12 b* * n c 13 ** h . c q .g . a . a t h4 t h3 th2 n c 12 a* * m th1 c ub a d c c 10 a c c 10 b c c 11 u c 34b 5b -c u c 6a 6b u c 7 5a n c 1 3* * n c 1 2* * 16 m δ13ccarb δ13corg 8 9 10 11 12 13 14 15 tu ro ni an c en om an ia n -2 4 a b c -23-29 n c 1 1 o a e 2 diameter μm1.5 4.0 length μm b. constans z. erectus d. rotatorius w. barnesiae 6.0 4.5 6.06.0 4.5 4.5 4.5 6.0 6.0 6.0 2.0 4.51.5 1.5 1.5 1.5 1.5 2.0 2.0 2.0 2.0 l ith o lo g ic a l u n its 1.5 4.0 1.5 4.0 1.5 4.0 1.5 4.0 2.0 9.0 2.0 9.0 2.0 9.0 2.0 9.0 2.0 9.0 ' length μmlength μm ±1 standard deviation sample mean length total mean length pre-oae mean length 95% confidence interval o a e 2 o a e 2 o a e 2 o a e 2 barren interval oceanic anoxic event 2 165 values of b. constans are observed around δ13c peak a followed by a decrease in size reaching minimum values around δ13c peak b. in the upper part of the section, above δ13c peak c, an increase in coccoliths is observed with values similar to the mean. coccoliths of d. rotatorius display a well-defined size peak at the onset of oae 2 around δ13c peak a, whereas z. erectus dimensions are very similar to the mean size values with a gradual increase towards larger coccoliths recorded after δ13c peak c. w. barnesiae doesn’t show significant size changes, although an increase was observed in the lower part of oae 2. pueblo in the lowermost part of oae 2 at pueblo, an increase in size is observed in b. constans and d. rotatorius showing larger coccoliths around δ13c peak a (fig. 3e). both b. constans and d. rotatorius display a decrease in size, reaching minimum values around δ13c peak b and remaining small through the upper part of oae 2 and overlying interval. a return to larger coccoliths was recorded after δ13c peak d. z. erectus and w. barnesiae show minor changes in size; however, smaller coccoliths of z. erectus were observed at the end of the isotopic plateau and a slight decrease in size of w. barnesiae specimens correlates with δ13c peak b. coccolith morphometry among analyzed species, b. constans shows distinct and coeval size fluctuations through oae 2 in all studied sections, while d. rotatorius, z. erectus and w. barnesiae, record only minor and isolated size variations at different stratigraphic levels. mean length of b. constans (supplementary tab. 1) varies depending on latitude: coccoliths are bigger at highest latitude (i.e. eastbourne) and progressively decrease southwards, reaching minimum values at lowest latitude sections (i.e. novara di sicilia). within the studied sections, the main size features of b. constans can be summarized as follow: larger coccoliths characterize the pre-oae 2 interval; a minor decrease in size correlates with the onset of oae 2; an increase in size trend reaches a maximum around δ13c peak a; a decrease in size trends culminates with dwarf coccoliths at δ13c peak b; a relative recovery in size marks the post – oae 2 interval, although without reaching preevent sizes. the anova and tukey post hoc test were applied to coccolith morphometric data in order to evaluate if b. constans means are significantly different in the ten intervals identified within the δ13c curve (supplementary tab. 2, fig. s4): 1) pre-positive excursion; 2) initial rise; 3) peak a; 4) trough between peak a and peak b; 5) peak b; 6) plateau; 7) peak c; 8) decreasing values after peak c; 9) peak d; 10) further decrease after peak d. statistical analyses evidence that, in every section, b. constans is always significantly smaller (p < 0.05, supplementary tab. 2) in interval number 5 corresponding to δ13c peak b compared to interval number 1 (pre-oae 2) and interval number 3 (δ13c peak a). discussion calcareous nannoplankton calcification and oae 2 perturbation morphometric and statistical analyses indicate consistent and significant changes in coccolith size of b. constans in all studied sections. we exclude a diagenetic control since the investigated successions yield rather homogenous moderate preservation, belong to different settings and underwent diverse diagenetic paths. measured coccoliths are complete with no evidence of corrosion or overgrowth in the outline and the co-occurrence of incoherent size variations in different taxa is further evidence for disregarding diagenetic modifications. our morphometric data highlight latitude-related changes in b. constans size, being its coccoliths bigger at highest latitude (eastbourne section) and progressively smaller southwards, with minimum values at lowest latitude (novara di sicilia section). such findings are consistent with patterns observed in cenozoic (herrmann & thierstein 2012) and holocene (herrmann et al. 2012) assemblages indicating changes in coccolith size across latitudes. in order to understand the role of complex environmental stress of oae 2 on b. constans size variations, we first take under consideration the multiproxy data available for the wis and eastbourne sections (fig. 4). the record of pco 2 concentrations derived from stomata in fossil leaves faucher g., erba e., bottini c. & gambacorta g.166 of the wis area is limited to the first part of the c-isotopic anomaly (barclay et al. 2010) associated with a long-term increase in pco 2 interrupted by two relative lows. it was hypothesized that volcanic activity started to influence atmospheric co 2 levels before the oae 2 onset (fig. 4a; barclay et al. 2010): an increase by 20%, equivalent to a rise of 60-300 ppm, was attributed to a massive magmatic episode occurring some 500 kyrs in advance of the positive c-isotopic excursion that defines oae 2 (barclay et al. 2010). a distinct low in atmospheric co 2 of about 25-26% (barclay et al. 2010; sinninghe damstè et al. 2008) was documented at the stratigraphic level of δ13 c peak a, followed by a further increase up to the middle part of oae 2 (barclay et al. 2010). osmium isotopic signature moves towards less radiogenic values approximately 30-40 kyrs before oae 2 onset and was attributed to an input of unradiogenic os from a mantle source, thus suggestive of an intense submarine volcanism continuing through the first phase of oae 2 (du vivier et al. 2014). in the pueblo section, snow et al. (2005) documented a first relative increase and then a peak in trace metals at the onset of oae 2 and at δ13c peak b, respectively (fig. 4a). the presence of these metal abundance anomalies needs an explanation other than influx of terrigenous input and the most reliable hypothesis is the release of magmatic fluids in event plumes (snow et al. 2005). the metal-rich intervals are more abundant in the less volatile and more reactive elements (such as sc, co, mn and fe) and this is in good agreement with the position of the caribbean plateau, which was proximal to the wis area. cuba and pueblo sections are, as expected, rich in a wide range of near-field and far-field elements. in the eastbourne section, a relative increase in chromium (cr/al) was documented at the onset of oae 2 (fig. 4b) and a maximum precedes δ13c peak b (pearce et al. 2009). shape and stratigraphic levels of both cr enrichments at eastbourne and pueblo point to the caribbean hydrothermal events (pearce et al. 2009) associated with oae 2. a b a tu ro ni an c en om an ia n -2 00 δ13c c d ep th (c m a bo ve b as e) m 80 0 60 0 40 0 20 0 0 n c 11 n c 12 a* nc 12 b* * q . g ar tn er i r. a sp er h . c hi as tia s. g n .j. w .d v .b p.f m . m os by en se m . n od os oi de s -28 10 00 δ13corg pratt et al., 1993 cm 5.52.0 b. constans μm-22 δ44ca bralower al., 2003 unpublishised data. pratt et al., 1993 o a e 2 trace metals element/zr 0 9 laurus nobilis hypodaphnis zenkeri 95% ci 95% ci pco2 ppmv ppmv0 400 pco2 calibration pco2 400 0.0 1.0 (187os/188os)initial -1.6 -1.3 co/zr x10 cu/zr x 2 w/zr x100 800 b -5 -2 -5 -2 δ 18o plenus cold event thermal maximum t u ro n ia n c e n o m a n ia n q . g . r . c . a .a . w . d . m . g . c . g . n c 11 * n c 12 ** n c 13 **f . c . n . j . 2.0 5.5 b. constans length 0 0.45 δ44ca m 10 15 20 25 -24-27 δ 13corg(‰ vpdb) holywell event 28 29 δ 13c(‰vpdb) a b c d δ13ccarb 62 0 40 0 28 biscutum spp. % productivity index watnaueria spp. % 0 2.2 0.5 1.5 cr/al (mm/m) onset of caribbean volcanism o a e 2 fig. 4 synthesis of major geological events in the late cenomanian – early turonian interval a) wis δ13c curve (snow et al. 2005); pco 2 reconstruction (barclay et al. 2010); trace metals (snow et al. 2005); osmium-isotope signature (du vivier et al. 2014); calcium-isotope signature (du vivier et al. 2015); composite b. constans morphometric curve (this study): pre-oae 2 interval data from cuba section, oae 2 and post oae 2 interval from pueblo rock canyon; b) eastbourne section δ13c curve (tsikos et al. 2004) and δ13c curve (jarvis et al. 2011); δ18o (tsikos et al. 2004), cooler pulses during plenus cold event (jenkyns et al. 2016); relative abundance of watznaueria, biscutum and productivity index (linnert et al. 2011); cr/al (pearce et al. 2009); calcium-isotope signature (circle = δ44/42ca, blättler et al. 2013; diamonds = δ44/42ca du viver et al. 2015); b. constans morphometric curve (this study). oceanic anoxic event 2 167 at eastbourne, temperature fluctuations were reconstructed using the o-isotopic record (pearce et al. 2009; jarvis et al. 2011): a relative warming correlates with the onset of oae 2 and is interrupted by the plenus cold event (fig. 4b) associated with a temporary spread southwards of north boreal biota (including the index belemnite praectinocamax plenus) (pearce et al. 2009; jarvis et al. 2011; jenkyns et al. 2016). the middle-late part of oae 2 is then characterized by increased paleotemperature culminating with the “early turonian thermal maximum” just after δ13c peak d (jarvis et al. 2011). a shift towards lower δ44ca and δ7li values indicates accelerated weathering at the beginning of the oae 2 c-isotopic anomaly (blättler et al. 2011; pogge von stradmann et al. 2013), reaching a maximum around δ13c peak b, some 200-300 kyrs after the volcanism onset (pogge von stradmann et al. 2013). at eastbourne and in the wis areas, du vivier et al. (2015) documented a transient increase of δ44ca at the beginning of δ13c anomaly: co 2 induced ocean acidification during the caribbean lip eruption was proposed as the potential cause of such δ44ca positive shifts (du vivier et al. 2015). previous studies of nannofossil assemblages of the eastbourne (linnert et al. 2011) (fig. 4b) and wis (corbett & watkins 2013) sections showed partially contrasting data. yet, a general picture of suppressed abundances of the fertility-taxa b. constans and z. erectus was derived for mid-latitude shelf and epeiric settings through oae 2 (corbett & watkins 2013). however, in the cuba section, an increase of the higher-fertility nannofossil species was detected, suggesting local mesoto eutrophic conditions in the eastern margin of the wis (corbett & watkins 2013). paleoenvironmental perturbations in the ctbi reconstructed at eastbourne and in the wis are consistent with results gathered in other sections and used for global characterization of oae 2 (jenkyns 2010) in terms of paleoco 2 fluctuations, climate moving to global warming, enhanced primary productivity due to increased nutrient fluxes and upwelling as well as ocean acidification. our nannofossil morphometric data may provide insights into the comprehension of type and rate of reactions of phytoplankton calcification through progressively stressing conditions and their demise. we notice that short-term fluctuations in morphometry follow transient changes in climate and ocean chemistry. in particular, the reconstructed early increase in pco 2 shortly prior to and at the beginning of oae 2, associated with initial volcanism, correlates with the first decrease in size of b. constans coccoliths (fig. 4 and supplementary fig. s4). a short-lived return to relatively larger coccoliths falls around δ13c peak a, in the interval of the plenus cold event and reconstructed lower pco 2 . the subsequent decrease in b. constans size culminating in dwarfism across δ13c peak b (fig. 4 and supplementary fig. s4) correlates with metal enrichments and a significant temperature rise. only a partial return to pre-oae 2 sizes was observed in b. constans coccoliths after δ13c peak d, suggesting the persistence of stressing conditions in the earliest turonian characterized by high paleotemperatures known as the “early turonian thermal maximum” (jarvis et al. 2011). as for living coccolithophores (riebesell et al. 2000; beaufort et al. 2011; müller et al. 2011, 2012; hoffmann et al. 2012; sett et al. 2014), the main paleoenvironmental factors influencing calcareous nannoplankton calcification in the cretaceous were presumably nutrient availability, sea surface temperature, carbonate chemistry, ocean acidification, toxic trace metals, competition with other organism and salinity (erba 2004, 2006; mutterlose et al. 2015). linnert & mutterlose (2012) highlighted a decrease in size of biscutum specimens during oae 2 at oceanic sites from northeastern atlantic ocean. the associated decrease in abundance of biscutum suggested that the reduction in size occurred under oligotrophic conditions (linnert & mutterlose 2012). we found a similar relationship at eastbourne and pueblo (figs 3, 5), but in the cuba (corbett & watkins 2013) and novara di sicilia (scopelliti et al. 2008) sections, the analogous decrease in b. constans coccolith size and dwarfism around δ13c peak b, occurred under high nutrient availability (scopelliti et al. 2008; corbett & watkins 2013). therefore, the trophic level of surface water cannot be ascribed as the trigger of reduced calcification. a cool-water preference was implied for b. constans (lees et al. 2005) and may explain the increase in size of b. constans coccoliths around δ13c peak a, during the plenus cold event (jenkyns et al. 2016) that is a global scale sst decrease of about 4°c (voigt et al. 2004; pearce et al. 2009). conversely, the abundance and size decrease recorded faucher g., erba e., bottini c. & gambacorta g.168 in b. constans coccoliths at δ13c peak b are associated to a sst increase of ~ 6°c (voigt et al. 2004; pearce et al. 2009). however, it is dubious that b. constans calcification was directly controlled by ssts, since the full recovery in size started just after δ13c peak d under hot conditions of the “early turonian thermal maximum”. indeed, the preference of b. constans for cooler surface-waters was previously questioned (e.g., linnert & mutterlose 2012) and, indeed, we notice that b. constans was very abundant during oae 2 in the tropical atlantic (hardas & mutterlose 2007) characterized by ssts up to 35°c and in the earliest turonian under the highest ssts (forster et al. 2007) recorded in the boreal area (linnert et al. 2010, 2011). it is feasible that some acidification of seawaters postulated for the early phase of oae 2 (du viver et al. 2015), affected b. constans calcification. as a matter of fact, in the studied sections, b. constans coccolith size fluctuations mimic co 2 reconstructions (fig. 4): a decreasing size trend is observed in the interval with higher co 2 concentration and, conversely, bigger coccoliths occur around δ13c peak a, where a ~25% reduction in co 2 was estimated (barclay et al. 2010; jarvis et al. 2011). the subsequent reduction of b. constans coccolith dimensions and dwarfism correlates with a further rise in pco 2 continuing through and after the late phase of oae 2 (jarvis et al. 2011). as previously inferred for mesozoic times (erba 2006), in the ctbi there was arguably a causal link between levels of co 2 , marine carbonate chemistry and calcite secretion by nannoplankton: when pco 2 in the ocean-atmosphere system was minimum, production of larger coccoliths was favored by high caco 3 saturation state. vice versa, under high co 2 concentration, a drop in carbonate saturation state induced production of smaller (and less calcified) coccoliths. in living coccolithophorids, calcification is ruled by an optimum curve; low inorganic carbon content reduces photosynthesis that indirectly limits calcification due to a deficiency of organic molecules needed to construct coccoliths (krug et al. 2010). increased inorganic carbon substrate stimulates photosynthesis and calcification reaching maximum values; excess co 2 release determines low ph that decreases calcification rates (krug et al. 2010; bach et al. 2015). the threshold values, below or above, which calcification becomes metabolically unfavorable differ among species due to a speciesspecific sensitivity to ph conditions (krug et al. 2010; bach et al. 2015). we are conscious that co 2 injections during oae 2 occurred during a time interval much longer than modern anthropogenic releases, thus the reduction in carbonate saturation should have been buffered by shallowing of the calcite compensation depth (ccd) with only a minor effect on surface water chemistry. however, the construction of the caribbean plateau was sustained for 1-2 million years with several pulses of co 2 releases and not as a single volcanic eruption. possibly, as for oae 1a (méhay et al. 2009; erba et al. 2010, 2015) the sum of subsequent large and geologically brief co 2 emission was responsible for major alteration of the ocean-atmosphere system. submarine volcanism of the caribbean plateau lip introduced high concentrations of biolimiting metals during oae 2 (snow et al. 2005) that possibly resulted in ocean fertilization and/or poisoning by toxic metals (leckie et al. 2002; erba 2004) perhaps influencing coccolith calcification as observed on single modern coccolithophorid species intolerant to cadmium and copper (brand 1994). we speculate that b. constans was sensitive to metal toxicity as previously hypothesized by erba (2004). our morphometric results show that this taxon was most receptive, while z. erectus, d. rotatorius and w. barnesiae were more tolerant. the metal peaks detected at pueblo (snow et al. 2005), and novara di sicilia (duncan et al. 2013) are coeval with reduction and dwarfism of b. constans coccoliths, remaining small after the metal enrichment around δ13c peak b through the earliest turonian. thus, high metal abundances were possibly a concurring even if not necessarily the prime factor controlling b. constans calcification. summarizing, in the ctbi, nutrient availability in surface waters per se does not seem to be crucial for coccolith calcification because b. constans evidence coeval size reductions in the analyzed sections where different trophic levels were reconstructed. fluctuations in paleotemperatures might be partly responsible for b. constans coccolith sizes across oae 2, with reduced coccolith size under warmer conditions and vice versa, although this correlation does not hold in the aftermath of oae 2. morphometric data suggest that ocean chemistry related to the amount of carbon dioxide and/ oceanic anoxic event 2 169 or large quantities of toxic metals played a central role at species-specific level and particularly in b. constans coccolith size. however, excess co 2 in the ocean-atmosphere system was seemingly linked to volcanically-induced warming as well as metal fertilization via hydrothermal plumes during (pulses) of submarine plateau construction. comparison among cretaceous oaes records previous studies performed across other oaes highlighted size variations of selected nannofossil species. during the cretaceous, w. barnesiae was a cosmopolitan species, common in tropical and subtropical regions and dominant especially in oceanic gyres. it is described as a r-selected opportunistic species (hardas & mutterlose 2007) or used as a low productivity indicator (roth & krumbach 1986; erba et al. 1992; erba 1992, 2004). the data from the early aptian oae 1a (erba et al. 2010; lübke & mutterlose 2016), latest albian oae 1d (bornemann & mutterlose 2006) and oae 2 (linnert & mutterlose 2012; this study) are showing that w. barnesiae remains statistically steady in size. however, during oae 1a, several coccoliths of w. barnesiae were affected by some malformation evidenced by strong ellipticity and asymmetry in the interval of most extreme paleoenvironmental stress (erba et al. 2010). instead, in all studied sections, no signs of stronger ellipticity or asymmetry were found for the oae 2 interval. the very little variability measured in w. barnesiae specimens in both oae 1a (erba et al. 2010; lübke & mutterlose 2016) and oae 2 (linnert & mutterlose 2012; this study) possibly suggests that this taxon was most adaptable and only marginally affected by the paleoenvironmental perturbations characterizing cretaceous oaes. during both oae 1a and oae 2, b. constans appears to be the most sensitive species to the environmental perturbation. the morphometric analyses conducted on b. constans and z. erectus across oae 1a in the tethys and pacific oceans (erba et al. 2010) as well as in the north sea and lower saxony basin (lübke & mutterlose 2016) highlighted size reduction of their coccoliths during the early stage of oae 1a in the core of the negative carbon isotope excursion. contrarily to oae 2, b. constans sizes are smaller in average at higher latitudes (lübke et al. 2015; lübke & mutterlose 2016) compared to lower latitudes (erba et al. 2010). however, the drastic size decrease of b. constans is coeval at high median25% 75% min. max. pre oae post oae δ13 c peak b interval average length alstӓtte1 average length north jens-1 average length adda 2 cismon core dspd 463 novara di sicilia clot chevalier eastbourne cuba pueblo pre oae post oaeoae 1a 7 2 b . c on st an s le ng th fig. 5 box plots of b. constans length across oae 1a and oae 2. data for oae 1a are from the cismon core and dsdp site 463 (erba et al. 2010). the average length of b. constans across oae 1a are from adda-2, alstätte1 and north jens-1 cores (lübke & mutterlose 2016); data for oae 2 are from eastbourne, clot chevalier, novara di sicilia, pueblo and cuba sections. intervals pre-oae, oae 1a, δ13c peak b, post-oae, are based on δ13 c stratigraphy. faucher g., erba e., bottini c. & gambacorta g.170 and low latitudes and occurred in the core of the negative carbon-isotope interval (carbon isotope segment c3) (erba et al. 2010; lübke & mutterlose 2016) reaching comparable minimum b. constans coccolith size. the amplitude of size reductions during the earliest phase of oae 1a is significantly different at lower and higher latitudes, of the order of ~30% versus ~17%, respectively, relative to the interval preceding oae 1a. in fig. 5 the size measurements of b. constans across oae 1a (erba et al. 2010; lübke & mutterlose 2016) and oae 2 (this study) are compared. b. constans display very similar variations in both events, with a drastic decrease during oaes. a size decrease of b. constans coccoliths by 15 % on average, occurred during oae 2. the size reduction of b. constans coccoliths during oae 1d was ascribed to cooler surface-waters as indicated by nannofossil-based temperature (bornemann & mutterlose 2006). such temperature influence, however, contradict the conclusions of linnert & mutterlose (2012) who observed smaller biscutum coccoliths in the lower turonian interval characterized by very warm surface temperature. additionally, during oae 1a dwarfism occurred in the core of the negative carbon-isotope interval where the highest temperature is reconstructed. therefore, we do not see a correspondence between lower temperature and smaller size, but possibly the opposite correlation (fig. 4). the potential influence of nutrient availability on coccolith size, discussed for oae 1a (erba et al. 2010, 2011; lübke et al. 2015) and oae 1d (bornemann & mutterlose 2006) was documented by general smaller sizes under enhanced fertility. this interpretation is in contrast with the reconstruction by linnert & mutterlose (2012) who found larger biscutum coccoliths associated with higher nutrient concentrations in the ctbi. nutrient availability of surface waters was considered by corbett & watkins (2013) to explain changes in nannofossil assemblages across oae 2 in the wis. indeed, the cuba and pueblo sections were analyzed and paleoceanographic reconstructions suggest opposite conditions dominated by oligotrophy and eutrophy for the pueblo and cuba sections, respectively, during oae 2. our data (fig. 3d, e) document very similar sizes for b. constans coccoliths at these two locations and, thus, we infer that fertility was not the primary cause of coccolith size control. although oae 1a was also a time of ocean eutrophication (erba 1994, 2004; bottini et al. 2015), the time interval of dwarfism corresponds to the climax in the volcanic activity of the ontong java plateau as indicated by os-isotope (tejada et al. 2009; bottini et al. 2012), maximum in co 2 (méhay et al. 2009), trace metal concentrations (erba et al. 2015) and maximum warming (ando et al. 2008; bottini et al. 2015) but not to the highest fertility (bottini et al. 2015). high run-off due to intensified continental weathering was supposed to have introduced more clastic particles and fresh waters into the sea: muddy and restricted photic zone reduced the availability of light favoring the dominance of small specimens at a regional scale (lübke & mutterlose 2016). however, the coeval reduction in sizes at both higher and lower latitudes and at highly diverse oceanic settings during oae 1a and specifically in the carbon isotope segment c3 contradict such interpretation and unambiguously evidences a global paleoenvironmental drive for synchronous coccolith dwarfism of b. constans. the data collected by erba et al. (2010) and lübke & mutterlose (2016) indicate that minimum coccolith size corresponds to most extreme perturbations as recorded by δ13c negative spike in the early part of oae 1a. excess co 2 concentrations related to the volcanic activity of the ontong java plateau remains the most plausible explanation for coccolith dwarfism in b. constans, z. erectus and d. rotatorius, and some malformation in w. barnesiae (erba et al. 2010). such modifications in size and morphology were interpreted as the species-specific transient response to survive progressively increasing surface-water acidification. it is clear that a combination of paleoclimatic and paleoceanographic triggers concurred in stressing the oceanic ecosystem, but, as discussed by erba et al. (2011), ocean acidification under excess volcanic co 2 seems to be the requisite or determinant of species-specific size reduction and malformation during oae 1a, because coccolith dwarfism has not been documented during other cretaceous cases of high fertility independent from oaes. during oae 2, reconstruction of pco 2 from the stomatal index suggests an increase of about 600 ppm reaching values of about 900-1000 ppm (barclay et al. 2010). pogge von strandmann et al. (2013) calculated the amount of co 2 sequestered from the continental weathering (~ 4-8 x 104 gt oceanic anoxic event 2 171 co 2 ) in agreement with previous calculation of the amount of co 2 emitted in the first phase of construction of the caribbean plateau (7-12 x 104 gt co 2 ) (kuroda et al. 2007). it is therefore conceivable that size reduction of b. constans coccoliths (fig. 4), reflects a progressive acidification of surfacewaters induced by large emissions of volcanic co 2 . proxy data (e.g. erba & tremolada 2004; méhay et al. 2009) suggest that during oae 1a co 2 was emitted in subsequent individual volcanic pulses reaching a concentration of about 1000-2000 ppm. in both oaes, thus, the occurrence of b. constans dwarf coccoliths correlates with excess co 2 suggesting that concentrations above threshold values possibly become adverse for calcification of sensitive cretaceous coccolithophore species, in analogy to results of laboratory experiments (riebesell et al. 2000; krug et al. 2011; bach et al. 2012; bach et al. 2015). calcification must be a beneficial process for coccolithophore algae because this high energyconsuming process remained stable in coccolithophore evolution from its appearance in the triassic (young & henriksen 2003). however, during the most extreme intervals such as oaes, a decrease of the ph might have favored other groups other than coccolithophore algae (bach et al. 2015). during oae 2, in fact, in the interval characterized by the major perturbation (δ13c peak b) the decrease in b. constans is paralleled by a strong increase in calcisphere percentages (pearce et al. 2009). this can be interpreted as an inability of b. constans to properly calcify and adapt to a new carbonate chemistry conditions, facilitating the spread of other more competitive functional groups such as dinoflagellates. one potential factor that may have affected coccolith calcification in addition to co 2 -induced ocean acidification is the concentration of toxic metals. indeed, we notice that during both oae 1a (larson & erba 1999; erba et al. 2015) and oae 2 (snow et al. 2005; duncan et al. 2013) there were large injections of trace metals associated with hydrothermal plumes during the construction of submarine lips (neal et al. 2008). abnormal quantities of biolimiting but also toxic metals might have had positive and negative feedbacks in coccolith production and, particularly, on size of b. constans coccoliths. metal toxicity is known for living nannoplankton: cadmium can inhibit calcification and copper is not tolerated by some coccolithophore species (brand 1994). as speculated by erba (2004) perhaps, poisoning by toxic metals affected calcareous nannoplankton in the cretaceous, inducing some extinctions, abundance drop and species-specific coccolith dwarfism. conclusions morphometric analyses of selected nannofossil taxa across oae 2 in various geological settings revealed differential species-specific patterns. coccolith size variations of d. rotatorius, w. barnesiae and z. erectus result to be minimal and substantially unrelated to oae 2 paleoenvironmental stresses. conversely, b. constans coccolith morphometric analyses identified size variations coeval with specific phases of environmental pressure. size fluctuations across oae 2 are similar and synchronous in all the analyzed sections located at great distance, in different oceans and settings. the quantitative characterization of coccolith size and morphology in the intervals preceding, coeval and postdating oae 2 assessed the occurrence of a relationship between calcification of b. constans and paleoenvironmental perturbations. in fact, during oae 2, b. constans coccoliths show a general decrease in mean size followed by a partial recovery at the end of the event, but dimensions remained smaller than in the interval preceding the main perturbation. at higher resolution, coeval decreases and increases in coccolith size correspond to individual paleoenvironmental changes across oae 2. we underline that b. constans dwarfism is observed in complete specimens with no corrosion in their outline and, therefore, cannot be explained as an artifact of diagenesis. the comparison with morphometric data available for the early aptian and latest albian oaes confirms that b. constans was most affected by stressing conditions and repeatedly underwent size reduction with evidence of temporary dwarfism. this suggests that presumably the same paleoenvironmental factors (or the same combination of ecosystem changes) controlled calcification of b. constans coccoliths during oae 1a, oae 1d and oae 2. however, the amplitude of b. constans coccolith reduction is different for oae 1a and oae 2, being larger for the former event. z. erectus and d. rotatorius display a less-expressed decrease in size during oae 1a and insigfaucher g., erba e., bottini c. & gambacorta g.172 nificant or scattered changes across oae 2; dwarfism of these taxa was detected only in the interval of climax perturbation during the early aptian selli event. the very little variability measured in w. barnesiae size indicate that this taxon was most adaptable and only marginally affected by the paleoenvironmental stress characterizing cretaceous oaes: while pronounced ellipticity was taken as evidence of malformation of w. barnesiae coccoliths during oae 1a, negligible variations in ellipticity were observed across oae 2, perhaps because of an attenuated degrees of perturbation. our results reinforce and implement paleoecological reconstructions of abiotic and biotic parameters somehow influencing coccolith secretion across oae 2, oae 1d and oae 1a allowing the evaluation of the factor, or combination of factors, inducing b. constans coccolith dwarfism in subsequent episodes of paleoenvironmental perturbation. temperature and nutrient availability in surface waters do not seem to have been decisive for b. constans coccolith size, but ocean chemistry related to the amount of co 2 concentrations and carbonate saturation state, played a central role in coccolith production by b. constans with a repetitive reduction in size. species-specific coccolith dwarfism seems to be the response of nannoplankton to ocean acidification during both oae 1a and oae 2 to survive lower ph. massive emissions of volcanic co 2 during the caribbean plateau construction across oae 2 (neal et al. 2008) arguably induced ocean acidification that was detrimental to some calcareous nannoplankton species, with temporary production of dwarf b. constans coccoliths. hydrothermal plumes introduced biolimiting metals that fertilized the global ocean, but also pumped in toxic metals that might have disturbed the functioning of some intolerant species, thus affecting their biocalcification. comparison of our results with data available for oae 1a and oae 1d indicate that analogous causes have induced similar response at different times in the cretaceous. the oaes indicate that past oceanic perturbations were severe and that the species-specific production of dwarf coccoliths possibly represents adjustment of some individual taxa to survive hostile ecosystems. this study further suggests that b. constans was the mid-cretaceous taxon most sensitive to ocean chemistry, while other species (z. erectus, d. rotatorius and w. barnesiae) were affected only at highest degrees of perturbation. it seems that calcareous nannoplankton have been resilient to ph changes, excess toxic metals, global warming and nutrification episodes, but it is certainly possible that the rate of change was decisive for coccolithophore adaptation. the geological record indicates that, at wide spatial scale, calcareous nannoplankton could adjust to high pco 2 , but past changes occurred over tens of thousands of years, giving enough time to adapt. acknowledgements. we thank tim bralower and bob duncan for sharing samples from the wis sections, and hugh jenkyns for providing samples from the clot chevalier section. we thank the editor, isabella raffi, and two anonymous reviewers for their valuable comments and suggestions on the manuscript. the research was funded through miur-prin 2011 (ministero dell’istruzione, dell’università e della ricerca–progetti di ricerca di interesse nazionale) to e. erba, and through sir-2014 (ministero dell’istruzione, dell’università e della ricerca–scientific independence of young researchers) to c. bottini. g. faucher was supported by fondazione fratelli confalonieri. references ando a., kaiho k., kawahata h. & kakegawa t. 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(2003) biomineralization within vesicles: the calcite of coccoliths. reviews mineral. geochem., 54(1): 189-215, doi: 10.2113/0540189. zurita 57..66 ��� ������� � ��� ���� ��� ��� �� ��� � ����� � ������� ���������� ���������� ��� ������������ ��� �� ���� ��� �� �� ������ �� ��� �������� ������ ������� ������� ��� �� � �� � �� �������� ��������� ��� � � � ��� � ��������� ������� ��� ��� ��� �� ������������������ �����!�"���������� ������� � ��� ��������� �� ������ ������� � � ���� �� #��$#�"���� %�&���� "�'���� �"'������� ��� ���� ������ ������� � ������������ � ()�*"�"�+ ,"�& �!� #��$#�"��� (���� ���-���+ �, �!� %�&���� "�'���� .�/ 0��.� �"�1 2��#/�� ���� ,"�& �!� ���� &���"���� !� "�-��� ,�����' �, � .��� �"�1 /�"2�� /��-�&��� &����� �!� -�#��� �#*� ��� ��&* *���/� ����./ �&1 �"�2�� 0��.���'� �, ��/ &�"�!���'� ��� -�&��"�/�� .��! 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"���# ��. -���# "��.� <� �>?.�a9� �-. $ ���!-# riv. it. paleont. srat. v.96 n.4 pp. 501-504 febbraio 1991 presenza di sedimenti lacustrt intercalati in vulcaniti oligo-miocemche della sardegna mertdionale marco murru (1) 8ú alessandra salvadori (2) key-uords: lactstrine sediments, oligo-miocene, southern sardinia. aknact sediments mainly carbonatic, with fossil remains, interbedded in calc-alkaline formations related to early oligo-miocene volcanic cycles, have been detected in three areas of sourhern sardinia. it is thought that these sediments may have been deposited in small lake basins, during periods ofvolcanic inactivicv. nelle vulcaniti calco-alcaline della sardegna meridionale, di età compr esa fra 35-32 ma e 13-11 ma (savelli et al.,1979i montigny er al., 1981; beccaluva et al., 1982), sono stati rinvenuti in tre settori (fig. t) depositi continentali sedimentari, intercalati nelle vulcaniti, contenenti resti fossili. la loro presenza consente di formulare ipotesi sull'ambiente deposizionale di questi sedimenti. gli affioramenti, di cui viene data qui di seguito una breve descrizione, sono ubicati rispettivamente nel sulcis (monte arruda-f. 233iii ne e cortoghiana-f. 233 iv so) e nel bordo orientale del campidano (serrenti-f. 226iii no). monte arruda (perdaxius). alla base delle pendici vulcaniche del monte arruda, al di sotto della grotta di tuvu mannu, a quota 60 m si rinviene una lente tufitica granodecrescente di circa 30 cm di spessore, intercalata a brecce andesitiche di probabile origine esplosiva. in questa lente sono presenti ostracodi in cattivo stato di conservazione e pertanro non classificabili. questo episodio vulcano-sedimentario è probabilmente legato all'instaurarsi di piccole conche lacustri durante periodi di stasi dell'attività vulcanica. (1) dipartimentp di scienze della terra dell'università di cagliari, via trentino 51,09127 cagliari. (2) vir ausonia 73, 09126 cagliari lavoro eseguito con il contributo del m.p.l 60.6. 502 m. murru & a. salpadori. cortoghiana. in alcuni sondaggi profondi effetruad dalla società carbosulcis s.p.a. (1), ubicati nei dintorni.di cortoghiana, sono stati rinvenuti episodi sedimentari intercalati nelle ignimbriti riolitiche e quarzotrachitiche che, come è noto, in quest'area rappresentano 1'episodio vulcanico più recente. nel sondaggio n. 20 le vulcaniti si incontrano sino a -154.i7 m; rispettivamente alle quote comprese fra -744.q5 e -744.60 m e fra -145.25 e -145.70 m si rinvengono due intercalazioni carbonatiche. nel sondaggío n.21,le ignimbriti arrivano sino a -278.40 m ed i calcari, con sottili intercalazioni tufitiche, si rinvengono fra -247.4o e -244.10 m. infine nel sondaggio n.24 le vulcaniti sono state incontrate sino a -196.60 m e fra -185.20 e -186.40 m si rinviene un'intercalazione calcarea. questi episodi sedimentari sono rappresentati da calcari parzialmente ricristallizzati, in cui tuttavia sono riconoscibili, in sezione sottile, oogoni di characeae talora perfettamente conservati (sondaggio n. 21). serrenti. i calcari selciferi del distretto di serrenti sono noti da antico tempo; il primo che li segnalò fu il la marmora (1s57), che li attribuì "molro dubitativamente" al giurassico. successivamente vardabasso & grimaldi (1935) attribuirono questi affioramenti in parte al miocene ed in parte al giurassico. infine cavinato (tlsl) cartografò nel foglio geologíco 226 "mandas" l'affioramento di calcare selcifero di monte ollasru come calcare miocenico. i principali affioramenti, ognuno di dimensioni inferiori al km2, si rinvengono ai piedi delle pendici vulcaniche di monte candidu, monte mannu, monte ollastu e monte porceddu. i relativi sedimenti sono costiruiti da calcari selciferi grigio-chiari, con stratificazione ben marcata, e sono spesso tettonicamente dislocati. i rapporti con le brecce andesitiche sono particolarmente chiari negli affioramenti ubicati sulle pendici meridionali del monte candidu, ove è possibile osservare la loro posizione intravulcanica. i calcari selciferi sono generalmente sterili, ma talora si rinvengono livelli centimetrici con rari oogoni di carofite e frammenti di gasteropodi polmonati. fanno eccezione gli affioramenti di monte candidu e monte mannu, che sono invece più fossiliferì ed il contenuto paleontologico è costituito quasi esclusivamente da oogoni di characeae, subordinatamente da gasteropodi polmonati integri e da rari ostracodi a valve articolate. (1) gli aa esprimono il più vivo ringraziamento alla società carbosulcis per aver messo a disposizione il materiale in studio. kdi.menti lacwtri intercalati a oulcaniti 503 zo km =mmffiilrn fig. 1 schema geologico della sardegna meridionale ed ubicazione dei sedimenti lacusrri. 1) basamenro paleozoico-paleogenico; 2) vulcaniti calco-alcaline oligo-mioceniche; 3) depositi prevalenremenre marini dell'oligocene superiore-miocene; 4) basalti plio-pleistocenici; 5) depositi continentali plioquaternari; 6) perdaxius; 7) cortoghiana; 8) serrenti, conclusionl. nei tre settori della sardegna meridionale sopra descritti (fig. l) è stata accertata la presenza di depositi sedimentari fossiliferi intercalati nelle vulcaniti oligo-mioceniche. questi sedimenti sono cosdtuiti da tufiti (perdaxius), da calcari parzialmente ricristallizzati (cortoghiana) e da calcari selciferi (serrenti). il contenuto paleontologico è abban î 8 o* ^-o/ "o 504 m. murru &a, saloadori stanza simile in tutti gli episodi ed è rappresentato da oogoni di carofite e subordinatamente da ostracodi e gasteropodi polmonati. l'ambiente di deposizione può essere considerato di tipo lacustre, instauratosi in piccole conche formatesi durante periodi di stasi dell'attività vulcanica nelle fasi iniziali vulcaniche, in quanto i depositi sedimentari si rinvengono nella parte basale delle successioni vulcaniche. opere citate beccaluva l., brotzu p., macciotta g., morbidelli l., serri g. 8c traversa g. (1982) cainozoic tectono-magmatic evolution and inferred mantle sources in the sardo-tyrrhenian area. in boriani et al. (eds.). the lithosphere in italy. advances in earth sciences research. acc. naz. lincei., pp.229-248,2 fig., roma. cavinato a. (1959) foglio geologico n. 226-mandas. litografaartistica cartografica,firenze. la marmora f.a. de (1857) voyage en sardaigne. troisième partie. description géologique. 3ème ed. v.2, pp.xx+7o6,18 tav., torino. montigny r., edel j.b. 6r thuizat r. (1981) oligo-miocene rotation of sardinia. k-ar ages and paleomagnetic data of tertiary volcanics. earth planet. sc. ktt., v. 54, pp. 261-271,4 fig., amsterdam. savelli c., beccaluva l., deriu m., macciotta g. 6c maccioni l. (l979) k-ar geochronology and evolution of the tertiary calc-alkaline volcanism of sardinia (italy). journ. volcanol. geotbum. res., t. 5, pp. 257-269,5 fig., amsterdam. vardabasso s. ec grimaldi g. (1935) contributo allo studio dei giacimenti caoliniferi di serrentifurtei (cagliari).,4s. min, sarda, v. 40, n. 4, pp. l9-29,iglesias. dalla_vecchia 309..322 ��� ����� ����� �� ��������� � ����� ������ ���� �� � ����� �� ����� ����� � ���� � ��� � ��������� ����� � �� ����� ��������� �� �� � ���� ��� �� ��� ��������� ������ ������� �!��"#$� ����$%���&� �� ������� � � ��������� ������ �� ���#� ��#�&�#�%$� '� &((� !�$$� $��#� )&�$� ' &�#&%� � �� ���� � $#��&�#� ����� !��" ��# ����&��&�*+&�����&� ,���#� ��#�&�#�%$'� &((� !�$$� $��# ,)&�$�� � ��& ��$ ��# !��$� �#���� �! & ����$%���&� ������� �!��" !��" ��& �� � ���%.� �� $�&�#$ $��&��"�����#$ /��� �##�� �#!#��#� �� ��# %���#&� .#�%$ �� ������ &�� /��� ��# 0���/&�&� .#�%$ � � ���������� �� �$ ��$����� �� ��# %�%$%& ��"1��&���� �! !#&�%�#$� $%..#$���. ��# ��#$#��# �! & �#� %�* �#����#� ����$%���&� �&2�� �� ��# ����&���*���&��� �&�1��&�# ' &�* !��" ��&�#� �� ��# 3#���$ 1#�/##� ��# �!��&�&1�&� ������#�� &�� ��# ����� %���#&� &��"&$$ �%���. �&�# ��#�&�#�%$ ��"#$� ����$%* ���&�$ /#�# �����& � �#��#$���& ������� �!��"$� $%�������. ��# ��#* $#��# �! 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ro s p o ra fl o ri d a 8 1 d u p le x is p o ri te s g e n e ra li s 8 2 d u p le x is p o ri te s 8 3 c ic a tr ic o s is p o ri te s b re v il a e s u ra tu s 8 4 a e q u it ri ra d it e s s p in u lo s u s 8 5 a e q u it ri ra d it e s v e rr u c o s u s 8 6 c ic a tr ic o s is p o ri te s k e d v e s ii 8 7 c ic a tr ic o s is p o ri te s s p ru m o n ti i 8 8 im p a rd e c is p o ra a p iv e rr u c a ta s p . s p p . s p in a te s p o re s ( -l ik e ) ? s p . s p . c f. s p . g y m n o s p e rm s 8 9 a ra u c a ri a c it e s a u s tr a li s 9 0 e p h e d ri p it e s 9 1 b a lm e io p s is li m b a tu s 9 2 c a ll ia la s p o ri te s d a m p ie ri 9 3 c la s s o p o ll is to ro s u s 9 4 c la s s o p o ll is c la s s o id e s 9 5 c la s s o p o ll is 9 6 c y c a d o p it e s 9 7 e u c o m m ii d it e s tr o e d s s o n ii 9 8 s p h e ri p o ll e n it e s s p p . s p . s p p . s p . 9 2 9 4 9 6 9 8 6 8 7 0 7 2 7 4 7 6 7 8 8 0 8 2 8 4 8 6 8 8 b e rr ia s ia n -b a rr e m ia n a p ti 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(2018) sivalhippus ptychodus and sivalhippus platyodus (perissodactyla, mammalia) from the late miocene of china. riv. it. paleontol. strat., 124(1): 1-22. rivista italiana di paleontologia e stratigrafia (research in paleontology and stratigraphy) vol. 124(1): 1-22. march 2018 abstract. herein, the authors report on skulls, mandibles and postcranial specimens of two species of chinese sivalhippus, s. ptychodus and s. platyodus. we frame our description and analyses within the context of newly described characters of the cheek teeth of hippotherium from the pannonian c of the vienna basin, the oldest and most primitive old world hipparions. our report includes original skull, mandibular and limited postcranial material of sivalhippus ptychodus and skulls and dentitions of sivalhippus platyodus from the paleontological museum of uppsala (pmu, uppsala, sweden), the american museum of natural history (amnh, new york, usa) and the licent collection in tianjin natural history museum (tianjin, china). the skull, maxillary and mandibular material we attribute to sivalhippus ptychodus and sivalhippus platyodus exhibit some primitive features for old world hipparions and synapamorphies of the face and dentition that unite it with the sivalhippus clade. our analysis shows that s. ptychodus and s. platyodus differ significantly from the cormohipparion occidentale – hippotherium primigenium clade. species belonging to the sivalhippus clade are found in indopakistan (s. nagriensis, s. theobaldi, s. perimensis and s. anwari), libya and kenya (s. turkanensis) and uganda (s. macrodon). we hypothesize that the sivalhippus clade originated in south asia where it is earliest represented by sivalhipus nagriensis, ca. 10.4 ma and underwent range extension into africa and china circa 9-7 ma. received: august 11, 2017; accepted: october 18, 2017 keywords: sivalhippus ptychodus; sivalhippus platyodus; late miocene; evolution; biogeography. introduction sivalhippus is a derived lineage of eurasian and african hipparion horses (bernor & hussain 1985; wolf et al. 2013). this genus first appears in indopakistan and appeared later in the late miocene in china and africa. lydekker (1877a) originally erected the genus sivalhippus and the taxon has been recognized by a number of authors since then (lydekker 1877b, 1882, 1885; pilgrim 1910, 1913; matthew 1929; colbert 1935; forsten 1968; hussain 1971; skinner & macfadden 1977; macfadden & bakr 1979; macfadden & woodburne 1982; hussain & bernor 1984; bernor & hussain 1985; bernor et al. 2010; wolf et al. 2013; sun 2013; bernor & sun 2015). wolf et al. (2013) recognized that sivalhippus is a clade in indopakistan (s. nagriensis, s. theobaldi, s. perimensis and s. anwari), libya and kenya (s. turkanensis) and uganda (s. macrodon). however, two chinese taxa have been referred to the sivalhippus group and are in need of description within a contemporary morphological and evolutionary context: sivalhippus platyodus and sivalhippus ptychodus (following bernor et al. 1990; bernor & lipcomb 1991; wolf et al. 2013). although some authors such as bernor et al. (1990) suggested a possible affinity of these chinese taxa with sivalhippus (sensu bernor et al. 1990; bernor & lipcomb 1991; wolf et al. 2013), their formal recognition, characterization and evolutionary relationships have not occurred until this contribution. we identify four skulls and two fragmental mandibles of sivalhippus ptychodus housed in museum of evolution of uppsala university, uppsala, sweden (pmu). in addition, we have recently discovered a mandible from qingyang, china in the licent collection housed in tianjin natural history museum. the qingyang area was discovered by the mailto:rbernor@howard.edu sun b., zhang x., liu y. & bernor r.l.2 french jesuit priest, emile licent (figure 1 – locator map). as a missionary, licent was sent to china in 1914 to initiate natural science investigations (qiu et al. 2013). early on, licent worked in the qingyang area, eastern gansu province, and reported his discoveries in the french newspaper la politique de peking (august, 1920). in october 1921, licent selected some well-preserved and representative specimens, and sent them to the museum national d’histoire naturelle, paris. based on this material, teilhard de chardin (1922) submitted a report to the geological society of france under the title: “sur une faune de mammiferes pontiens provenant de la chine septentrionale”. nevertheless, licent deposited several qingyang specimens in the musée hoang-ho pai-ho de tientsin (now tianjin natural history museum, thp). these specimens include hyena material studied by qiu et al. (1979), giraffe material (lacking any systematic report) and hipparion specimens most of which have been studied by qiu et al. (1987). the mandible described in this manuscript has not previously been reported or described. its dental characters such as small size, short p2 and triangular metaconid and metastylid are very similar as s. ptychodus. we also recently have recognized specimens of sivalhippus ptychodus in the american museum of natural history, including a complete skull and associated jaw. it was identified as “hipparion” cf. platyodus by bernor et al. (1990). however, its facial morphology reveals its identity as sivalhippus ptychodus and is the best preserved specimen of this species including complete skull and mandible. material that we recognize as belonging to sivalhippus platyodus include one skull and associated mandible in uppsala and one skull in tianjin. our report leads us to provide the morphological distinction of s. ptychodus and s. platyodus and their geographic and chronologic ranges in china (fig. 1), and their evolutionary and biogeographic relationships with indo-pakistan and african species of sivalhippus. systematic conventions the nomen hipparion has been used in a variety of ways by different authors. we follow characterizations and definitions for hipparionine horses recently provided in bernor et al. (1996, 1997). the taxon hipparion has been applied in a variety of ways by different authors. we utilize the following definitions in this work: hipparionini (or, the common name hipparionines) a tribe of equidae with an isolated protocone on maxillary premolar and molar teeth and, as far as known, tridactyl feet. bernor et al. (1996, 2010, 2016) has recognized the following north american, eurasian and african lineages that they have held to generic rank, including: cormohipparion, neohipparion, nannippus, pseudhipparion, hippotherium, cremohipparion, hipparion, sivalhippus, eurygnathohippus (= a senior synonym of stylohipparion), proboscidipparion and plesiohipparion. these hipparionine lineages have recently been reviewed by qiu et al. fig. 1 distribution of s. ptychodus (black horse) and s. platyodus (red horse) in china. mixed color shows coexistence. sivalhippus (perissodactyla, mammalia) from the late miocene of china 3 (1987), bernor & white (2009), bernor et al. (2010, 2013, 2014, 2015), armour-chelu & bernor (2011), wolf et al. (2013) and bernor & sun (2015). qiu et al. (1987) and deng (2012) have held these taxonomic rankings to the subgenus rank, such as hipparion (proboscidipparion) pater. we follow bernor and co-authors recognizing these as multi-species lineages warranting genus-level ranking. metric procedures measurements are all given in millimeters and rounded to 0.1 mm. measurement numbers (m1, m2, m3, etc.) refer to those published by eisenmann et al. (1988) and bernor et al. (1997) for the skulls and postcrania and bernor et al. (1997) for maxillary and mandibular dentitions. those specimens that we refer to sivalhippus ptychodus and sivalhippus platyodus are listed in tab. 1 with accompanying information on their geographic location and chronologic age. tabs 2-6 list standard measurements for specimens (after bernor et al. 1997) referred to these two chinese species of sivalhippus: tab. 2, skulls; tab. 3, maxillary dentition; tab. 4, mandibles; tab. 5, mandibular dentitions; tab. 6, metapodial measurements. tab. 7 provides measurements on key features of sivalhippus skulls used in our following analyses. we cite these measurements in our description of skulls. character state distribution tab. 8 summarizes the characters state distribution of sivalhippus ptychodus and sivalhippus platyodus updated from bernor et al. (1990) including new character state attributes from bernor et al. (2017). the new additional characters of the maxillary and mandibular cheek teeth follow the definition in bernor et al. (2017), and are recorded as unordered states (a, b, c, . . . etc.). when a character state is followed by a slash (/), we record a state that as intermediate (i.e., a/ is intermediate between states a and b). when variable, we record 2 or more states without a / but use of an intervening comma (, i.e. a, b) in the species hypodigm. these tables have ordered specimens into morphological groups based on their shared states. morphological groups have been cross checked by size comparisons and the stage of cheek tooth wear was evaluated so that a given specimen was not excluded from a group because of ontogenetic variation. abbreviations we use the following abbreviations in this manuscript: table 1 accompanying information on specimens of chinese species of sivalhippus id taxon quarry sex bone side age (ma) pmum347 s. ptychodus lectotype wuxiang, shanxi 3 skull lt. 7-6.5 pmum350 s. ptychodus hequ, shanxi 3 skull lt. pmum593 s. ptychodus hequ, shanxi 3 skull lt. pmum3684 s. ptychodus yushe, shanxi male skull lt. 6.5 amnh143267 s. ptychodus shouyang, shanxi male skull with mandible lt. thp01839 s. ptychodus qingyang, gansu 3 mandible lt. 7 pmum353 s. ptychodus wuxiang, shanxi male mandible lt. 7-6.5 pmum356 s. ptychodus xinan, henan 3 mandible rt. pmum3691 s. platyodus lectotype wuxiang, shanxi male skull with mandible lt. 7-6.5 thp22708 s. platyodus yushe, shanxi female skull lt. 6.5 tab. 1 accompanying information on specimens of chinese species of sivalhippus. table 2 standard skull measurements for specimens of chinese species of sivalhippus (mm) id pmum 347 pmum 3684 pmum 350 pmum 593 amnh 143267 pmum 3691 thp 22708 taxon s. ptychodus s. platyodus m1 102.5 108.8 98.8 97.0 m2 91.2 106.8 99.6 111.9 101.3 111.7 86.8 m3 121.0 97.0 84.6 m4 m5 174.8 m6 379.9 m7 73.0 76.3 77.2 75.3 73.6 82.5 77.6 m8 62.2 62.8 67.1 60.4 66.5 64.7 m9 134.8 136.8 142.2 133.8 149.2 140.0 m10 70.7 61.0 56.5 65.8 51.6 m11 32.8 38.8 29.5 22.0 31.9 29.7 m12 35.7 38.1 31.2 31.0 29.1 31.8 37.3 m13 68.6 66.0 53.4 28.3 56.8 57.6 m14 42.0 34.0 26.8 33.5 35.3 m15 44.7 33.3 45.7 47.2 m16 78.0 77.5 m17 121.0 m18 148.0 117.0 63.0 142.0 m19 181.0 146.0 171.0 m20 m21 m22 62.2 m23 284.8 295.6 292.4 218.3 m24 163.2 m25 89.0 80.7 65.4 89.0 95.5 78.1 77.2 m26 105.0 74.0 93.6 80.7 m27 m28 56.1 54.6 58.1 59.0 55.5 m29 45.8 55.1 45.3 52.5 48.8 m30 0.0 131.6 113.5 m31 0.0 126.6 149.1 m32 50.2 47.0 41.6 44.7 43.5 43.6 m33 55.4 47.6 65.3 60.1 47.6 70.3 m34 60.3 60.7 65.2 55.2 64.1 m35 33.3 27.9 31.0 37.7 32.8 49.7 m36 38.8 29.5 24.3 40.8 28.8 17.5 m37 54.7 37.6 29.8 56.8 24.6 m38 59.1 46.4 57.8 57.4 43.6 tab. 2 standard skull measurements for specimens of chinese species of sivalhippus (mm). sun b., zhang x., liu y. & bernor r.l.4 ma: mega-annum in the geochronologic time scale. ages in m.y. are based on radioisotopic analyses or magnetostratigraphic analyses. measurement table abbreviations sex: m = male; f = female; 3 = unknown. sex can be defined by the size of a canine tooth, male being large, female being small. side: lt. = left; rt. = right. cranial abbreviations: iof = infraorbital foramen; pob = preorbital bar; pof = preorbital fossa. element abbreviations in tables: tx = maxillary tooth; tm = mandibular tooth; skull = skull; mand = mandible; mciii = metacarpal iii. m1-m38 refers to measurements as described by eisenmann et al. (1988) and bernor et al. (1997). c1-c52 refers to characters 1-52 of tab. 8. institutional abbreviations amnh american museum of natural history, new york gsi geological survey, calcutta, india. ivpp institute of vertebrate paleontology and paleoanthropology; v = vertebrate specimen pmu paleontological museum of uppsala. m = vertebrate specimen. currently named “the museum of evolution of uppsala university”. thp tientsin hoangho paiho, from old name of tianjin natural history museum terminology and measurements: all follow sisson (1953), eisenmann et al. (1988) and bernor et al. (1997). systematics order perissodactyla owen, 1848 suborder hippomorpha wood, 1937 superfamily equoidea hay, 1902 family equidae gray, 1821 subfamily equinae steinmann & doderlein, 1890 tribe hipparionini quinn, 1955 genus sivalhippus lydekker, 1877 sivalhippus ptychodus (sefve, 1927) hipparion ptychodus sefve, 1927. hipparion platyodus qiu et al., 1987 (in part). “hipparion” ptychodus bernor et al., 1990. “hipparion” cf. platyodus bernor et al., 1990. “sivalhippus” ptychodus bernor and lipscomb, 1991. lectotype (bernor et al. 1990): pmum347, an adult skull with right and left p2-m3, lacking the snout, anterior dentition and most of the cranium, housed in the museum of evolution of uppsala university (fig. 2). type locality: locality 73 (andersson locality number; fig. 1), doujiaogou, dongcun village, wuxiang county, shanxi province, china. locality of new material: in 1920, licent initiated field work in qingyang, gansu and collected abundant vertebrate fossils from two localities, xingjiagou and zhaojiacha (licent 1936). the mandible thp01839 described herein likely comes from one of these two localities. stratigraphy and age: 76.5 ma. geographic distribution (fig. 1): yushe county, shanxi province; shouyang county, shanxi province; wuxiang county, shanxi province; hequ county, shanxi province; xinan county, henan province; qingyang county, gansu province. revised diagnosis (synapomorphies with sivalhippus indicated in bold *): medium-size hipparion with cheek tooth row 134.8 142.2 mm. pob relatively long (44.7-50.2 mm in adult individuals); anteriormost limit of lacrimal placed well posterior to pof (c1g*); nasolacrimal fossa absent (c2c); orbital surface of lacrimal bone with reduced foramen (c3b); pof placed anterodorsally on face, anteroposteriorly oriented (c4k*), posterior pocketing slight (c5b), medially deep (c6a), medial wall morphology without internal pits (c7a), peripheral border moderately expressed (c8b), anterior rim faint (c9a); infraorbital foramen inferior to, or encroaching upon anteroventral border of the preorbital fossa (c10b); buccinator fossa distinctly limited (c11b) and not pocketed (c12a); caninus fossa absent (c13a); malar fossa absent (c14a); nasal notch approximately half the distance between canine and p2 (c15b) maxillary and mandibular dp1 absent (c16c). maxillary p2 anterostyle short (c17a*); curvature of maxillary cheek teeth moderate (c18b); maximum cheek tooth height ca. 60 mm (c19c/d); cheek teeth with very complexly plicated fossettes (c20a,b), pre and postfossette opposing borders separate (c21b), posterior wall of postfossette always distinct (c22b), pli caballins bifid or complex (c23a,b,c), hypoglyphs moderately to deeply incised (c24b,c), protocones short, lingually flattened and labially rounded (predominately c25e*, also g and d; c26a*), protocone always isolated from protoloph (c27b) with no protocone spur (c28c), protocone more lingually placed than hypocone in premolars (c29b) and molars (c30b). mandibular p2 with short and rounded anterostylid (c31b); mandibular incisors are not grooved (c32a) and straight (c33b); i3 lateral aspect is elongate, not labiolingually constricted (c34a); premolar metaconid may be angular on the distal surface (c35a,c*), molar metaconid angular on the distal surface (c36a,c*), premolar metastylid angular on the mesial surface (c37c*), premolar metastylid spur may be absent or present (c38a,b), molar metastylid angular on sivalhippus (perissodactyla, mammalia) from the late miocene of china 5 the mesial surface (c39c*), molars lacking metastylid spur (c40b), premolar ectoflexid may or may not separate metaconid and metastylid (c41a,b); molar ectoflexid separates metaconid and metastylid after early adult wear (c42 a,b); pli caballinid varies from being rudimentary or single to absent (c43b,c); protostylid varies from being placed on mesio-labial surface of the tooth to being a small pointed projection (c44b,f); protostylid orientation is vertically placed, lying lateral to protoconid band (c45c); ectostylids are absent (c46b); premolar linguaflexid varies from being shallow u-shaped to deep, broad u-shape (c47c,d*); molar linguaflexid varies from being deep, broad u-shape to very broad and deep u-shape (c48d,e*); preflexid morphology varies from having simple to complex margins (c49a,b,c); postflexid morphology varies from being simple to very complex (c50a,c); postflexid does not invade metaconid/metastylid junction by anteriormost portion bending sharply lingually (c51a); protoconid enamel band rounded (c52a). description pmum347 (fig. 2a, b) is the lectotype skull of sivalhippus ptychodus (bernor et al. 1990). the skull is medium size with p2-m3, 134.8 mm in length. the preorbital bar is very long, 50.2 mm with the anterior edge of the lacrimal placed less than ½ the distance from the anterior orbital rim to the posterior rim of the fossa; nasomaxillary fossa is absent; orbital surface of the lacrimal bone has a reduced foramen; pof is a small, rounded structure, pof ventral border distance to facial-maxillary crest, 38.8 mm; pof pocketing reduced with moderate to slight depth; fossa medial depth is deep, greater than 15 mm; preorbital fossa medial wall without internal pits; preorbital fossa peripheral border outline moderately delineated around the periphery; anterior rim is present; infraorbital foramen inferior to anteroventral border of pof; buccinator fossa not confluent with a caninus fossa; buccinator fossa not pocketed posteriorly; caninus fossa absent; malar fossa absent; nasal notch position cannot be determined. maxillary dp1 is absent; p2 anterostyle is short; curvature of the cheek teeth is moderate; maximum cheek tooth crown height is estimated to be approximately 60 mm; maxillary cheek tooth fossette ornamentation is complex with several deeply amplified plications; preand postfossette opposing borders are not linked; posterior wall of postfossette is always distinct; pli caballin morphology is double to complex; hypoglyph is moderately deeply incised; protocone is lingually flattened and labially rounded on all cheek teeth; no protocone is connected to the protoloph; there are no protoconal spurs; premolar protocone more lingually placed than hypocone; molar protocone more lingually placed than hypocone. pmum3684 (fig. 3 a, b) is an adult skull with a complete snout, lacking the posterior cranium. muzzle length is 102.5 mm. p2-m3 is similar in length to pmum347, 136.8 mm. pob length is slightly shorter than the lectotype, 47.0 mm. pof fig. 2 skull of s. ptychodus, pmum 347. a) left lateral view; b) occlusal view of left cheek tooth row. scale bar = 2 cm. sun b., zhang x., liu y. & bernor r.l.6 is shorter than in the lectotype,47.6 mm and has a dorsal-ventral height slightly less than the lectotype, 27.9 mm. the distance from the ventral border of the pof to facial maxillary crest is less, 29.5 mm. as with the lectotype, pmum3684 has a pof that is placed high and far anteriorly on the face. the maxillary cheek teeth are lacking dp1, have a short anterostyle, cheek teeth are moderately curved, have an estimated maximum crown height of about 60 mm, fossette ornamentation is very complex; opposing borders of preand postfossette are separate; posterior wall of postfossette is always distinct; pli caballin is double to complex (on p3 and p4), hypoglyph is only moderately deeply incised; protocone shows some degree of lingual flattening, especially on p4-m3; protocone is moderately flattened, but less so than in the lectotype; protocone is isolated from the protoloph on all cheek teeth; there are no protoconal spurs; premolar protocone is more lingually placed than the hypocone; molar protocone is more lingually placed than the hypocone. pmum350 is a juvenile skull fragment with p2-m2, lacking cranium, anterior nasals and snout (bernor et al. 1990, fig. 12d from locality 114). pob length is shorter than the lectotype, 41.6 mm; pof is longer than in the lectotype, 65.3 mm; and has a dorsal-ventral height slightly less than the lectotype, 31.0 mm. the distance from the ventral border of the pof to facial maxillary crest is much less, 24.3 mm. the cheek teeth reveal less plication complexity of the preand postfossettes, due to the very early stage of their wear. the pli caballins are single on p2 and m2, probably due to their early stage of wear, but bifid on p4-m1. hypoglyphs are deeply incised. protocones are lingually flattened on p4-m1, and elongate on the remaining cheek teeth, again probably due to the very early stage of wear (bernor et al. 1990). pmum593 (re: bernor et al. 1990, fig 12c) is an old adult individual with right p2-m3, left p3 (fragmentary)-m3, lacking posterior cranium, nasals and snout. length of p2-m3 is 142.2 mm; pob length is shorter than the lectotype, 44.7 mm; pof is longer than in the lectotype, 60.1 mm; and has a dorsal-ventral height greater than the lectotype, 37.7 mm. the distance from the ventral border of the pof to facial maxillary crest is greater than in the lectotype, 40.8 mm. the cheek teeth are heavily worn due to the advanced age of the individual. this has resulted in increased rounding of the protocones, except on m3, which has retained a lingually flattened and labially rounded morphology, and simpler enamel plications. amnh143267 (= amnh field number 91b979 of bernor et al. 1990, tab. 4; fig.4 a, b) is a well preserved skull with an associated mandible of an adult male individual. it is the most complete specimen of sivalhippus ptychodus. skull muzzle length is 108.8, significantly longer than m3684. p2-m3 length is very similar to the lectotype, pmum347, 133.8 mm. pob length is shorter than the lectotype, 43.5 mm. pof is shorter than in the lectotype, 47.6 mm and has a dorsal-ventral height fig. 3 skull of s. ptychodus, pmum 3684. a) right lateral view; b) occlusal view of left cheek tooth row. scale bar = 2 cm. sivalhippus (perissodactyla, mammalia) from the late miocene of china 7 lesser than the lectotype, 32.8 mm. the distance from the ventral border of the pof to facial maxillary crest is less, 28.8 mm, slightly less than m3684. the cranial features are identical to the lectotype. the maxillary tooth characters are similar to those seen in the lectotype, but the maxillary cheek tooth plications are less complex than those of the lectotype (fig. 4; tab. 3). sefve (1927: text fig. 22, pg. 42) figured a right mandibular p2-m1 (occlusal view) of sivalhippus ptychodus from locality 73, yushe basin. an unfigured specimen pmum353 is a left p2-m1 plus mesial portion of m2 that is likely the same individual as text fig. 22 (fig. 7a herein). characteristics of the specimen figured by sefve (1927: text fig. 22, pg. 42) dentition include: premolar metaconid is angular on the distal surface of p3 and p4 and metastylid is angular on the mesial surface; p2-p4 ectoflexids are deep and closely approach the metaconid-metastylid isthmus; pli caballinid is rudimentary on p2-p4. the smallto medium length of the cheek tooth row is a character congruent with the maxillary dentitions of the sivalhippus ptychodus hypodigm described above. sefve (1927) also referred a right mandibular cheek tooth series, pmum356 with p3m3 to s. ptychodus with a similar morphology but having ectoflexid being shorter in p4, perhaps due to advanced wear (fig. 7b). this is a later stage of wear dentition with rounded metaconids and angular metastylids, shallow premolar ectoflexids and very deep molar ectoflexids. amnh143267 (fig. 5; 7d) is the best preserved mandible specimen of sivalhippus ptychodus, with complete dentition including incisors and canines and the left cheek tooth row. mandibular incisors are not grooved; they are straight; i3 lateral aspect is elongate, not labiolingually constricted. the p2 anterostylid is short and rounded. premolar and molar metaconid are rounded; premolar and molar metastylid are angular on mesial surface; premolarmolar metastylid spurs are absent; premolar ectoflexid does not separate metaconid and metastylid whereas molar ectoflexid separates metaconid and metastylid; pli caballinid is rudimentary or single; protostylid is absent on occlusal surface, but may be on side of crown buried in cement; ectostylids are absent; premolar linguaflexid is shallow u-shaped; molar linguaflexid is deep, broad u-shape; preflexid and postflexid have simple margins; postflexid does not invade metaconid/metastylid junction by anteriormost portion bending sharply lingually; protoconid enamel band is rounded. thp 01839 (fig. 6; 7c; tab. 4) is a mandibular fragment with left p2 to m3 and right p2 to m2 of a young adult with m3 in an early stage of occlusion. while the mandible is fragmentary, this specimen includes both right and left cheek tooth rows and has been preserved as early adult wear stage. the length of lower cheek tooth row is 146.0 mm which is slightly exaggerated by the slight separation of the teeth’s interstitial spaces; the actual length would be somewhat less. as such, the size is close to the maxillary cheek tooth rows of the skulls described above. the p2 anterostylid is short and rounded; metaconid and metastylid are rounded; metastylid spur is present and strongly developed; ectoflexid is shallow and does not separate metaconid – metastylid; pli caballinid is strongly developed and single; protostylid and ectostylid are absent; linguaflexid is deep and u-shaped; preflexid and postflexid have fig. 4 skull of s. ptychodus, amnh 143267. a) left lateral view; b) occlusal view of left cheek tooth row. scale bar = 2 cm. sun b., zhang x., liu y. & bernor r.l.8 complex margins; postflexid does not invade metaconid/metastylid junction by the rounded anteriormost portion bending sharply lingually; protoconid enamel band is rounded. the p3 metaconid is angular on the distal surface; metastylid is nearly triangular; preand postflexid are less complex than p2; other features are similar to p2 except that metaconid spur is weakly developed. the p4 is similar to p3 except for more complex preand postflexid margins, and flat protoconid enamel band; it is also the only tooth in the mandibular row with the postflexid invading metaconid/metastylid junction by rounded anteriormost portion bending sharply lingually. the m1 metaconid is subtriangular on the distal surface; metastylid is triangular on the mesial surface; metastylid spur is absent; ectoflexid is deep and separates metaconid and metastylid; preflexid has moderately complex margins; postflexid has simple margins; other features are similar to the premolar series. the m2 is similar to the m1 except for having a very complex preflexid margin. the m3 is of an early wear adult having a very shallow ectoflexid, complex postflexid margin, protoconid enamel band flattened. thp 01839 has very strongly developed pli caballinids on p2-p4 while pmum353 and amnh143267 have rudimentary pli caballinids due to its more advanced age. pmum353 also has priid pmum347 pmum350 pmum593 taxon s. ptychodus lectotype s. ptychodus s. ptychodus tooth p2 p3 p4 m1 m2 m3 p2 p3 p4 m1 m2 p2 p3 p4 m1 m2 m3 m1 28.2 23.1 22.8 20.4 19.4 21.1 31.5 25 23.5 23 22.6 13.5 21.6 23 20.7 21.8 23.5 m2 m3 22.5 23.6 23 21.4 20.9 17.7 20.7 23.9 21.6 20.8 19.4 24.5 25.8 26.5 24.7 23.5 19.8 m4 m5 m6 m7 m8 m9 m10 6.2 6.3 6.6 7 6 6.8 7.9 7.9 7.3 7.3 8.9 9.1 8.7 8.9 9.5 m11 4.5 4.2 4.4 4.4 4.3 3.8 4.4 4.4 4.4 4.4 3.7 7.1 5.8 6 4.9 id pmum3684 amnh143267 taxon s. ptychodus s. ptychodus tooth i1 i2 i3 c p2 p3 p4 m1 m2 m3 i1 i2 i3 c p2 p3 p4 m1 m2 m3 m1 14.7 16.2 16.9 9.2 28.9 24.6 25.3 20.1 21.0 20.2 15.9 15.6 16.6 12.1 27 23 22.6 19.5 19.9 20.9 m2 m3 8.8 7 7.9 6.2 23.2 23.6 23 21.8 20.3 17.7 9.9 8.6 7.9 6.6 21.3 20.9 20.8 19.3 19.8 16.4 m4 m5 33 m6 2 3 2 1 2 m7 3 4 4 4 4 3 m8 3 2 2 3 3 2 m9 1 1 1 1 1 2 m10 7.5 7.1 6.8 6.4 7.2 6.4 5.9 5.9 6.3 6.2 6.2 6.2 m11 4.5 4.6 4.9 4.7 4.5 4.5 4 3.4 3.6 3.5 3.6 3.3 id pmum3691 thp22708 taxon s. platyodus lectotype s. platyodus tooth i1 i2 i3 c p2 p3 p4 m1 m2 m3 p2 p3 p4 m1 m2 m3 m1 15.4 16.2 15.6 12.9 33.1 26.1 25 20.9 21.4 23.4 32.2 24.2 22.4 22 21.5 19.1 m2 m3 9.2 7.1 6.5 7.3 25.1 26.6 26.3 23.5 22.7 19.7 20.1 21.6 21.1 20.6 19.5 15.5 m4 m5 m6 4 2 3 4 5 3 m7 7 7 7 9 8 5 m8 2 3 4 7 6 2 m9 1 1 1 3 2 m10 7 6.1 6.9 6.4 7.1 8.2 7.6 7.3 7.2 7.2 7 8.2 m11 5 4.7 4.1 4.1 4.1 3.6 3.8 3.4 4 3.7 3.3 3.1 tab. 3 standard maxillary cheek tooth measurements for specimens of chinese species of sivalhippus (mm). sivalhippus (perissodactyla, mammalia) from the late miocene of china 9 mitive, deeply projecting ectoflexids on the premolars whereas thp 01839 has shallow ectoflexids on p2-p4 likely due to its young age. the hypodigm of skulls, maxillary and mandibular dentitions of s. ptychodus are very similar to each other in their size. remarks qiu et al. (1979) suggested that the geologic age of the xingjiagou and zhaojiacha faunas originated from the same stratigraphic horizon (fig. 1). deng (2006) correlated the qingyang fauna, gansu with european mn 12, ca.7 ma, based on mammal biochronological correlations. the lectotype of sivalhippus ptychodus, pmum347, was collected from the wuxiang basin, shanxi. another skull pmum3684 originated from the yushe basin, shanxi. opdyke et al. (2013) correlated the base of the mahui fm., lowest formation of the yushe basin is ca. 6.5 ma based on magnetostratigraphic data. tedford et al. (2013) considered that the basal sediments of the wuxiang basin to be correlative with the mahui fm. because of similarities in their mammalian fauna, except for some subtle differences, such as the presence of eostyloceros blainvillei (well known from older baode fm. faunas). tedford et al. (2013) suggested that the age of the base of wuxiang basin is probably a little earlier than that of yushe basin. therefore the age of pmum347 and pmum3684 is circa 7-6.5 ma. sivalhippus ptychodus has very long preorbital bar, strong preorbital fossa placed dorsally high and far anteriorly (with long pob), shallow nasal notch, complex upper cheek teeth fossettes, lingually flattened and labially rounded protocones, deep hypoglyphs, double or complex pli caballins, triangular metaconid and metastylid and broad u-shaped linguaflexid. these features are strongly similar to sivalhippus perimensis from the siwaliks (bernor & hussain 1985; bernor et al. 1990 and wolf et al. 2013). sivalhippus platyodus is similar to s. ptychodus differing in its larger pof dorsoventral dimension. we discuss this matter further below. fig. 5 mandible of s. ptychodus, amnh 143267. a) left lateral view; b) occlusal view. scale bar = 2 cm. id pmum353 amnh 143268 thp 01839 pmum3691 taxon s. ptychodus s. ptychodus s. ptychodus s. platyodus m1 m2 94.7 105.7 93.7 m3 69.3 69.1 77.5 m4 63.2 69 m5 132.5 146 m6 m7 47.4 36.0 46 m8 m9 m10 89.9 m11 66.1 58.8 m12 42.9 51.9 40.3 41.8 m13 63.6 71.7 79.5 m14 31.5 29.8 tab. 4 standard mandible measurements for specimens of chinese species of sivalhippus (mm). sun b., zhang x., liu y. & bernor r.l.10 sivalhippus platyodus (sefve, 1927) hipparion platyodus sefve, 1927. hipparion platyodus qiu et al., 1987 (in part). “hipparion” cf. platyodus bernor et al., 1990. “sivalhippus” platyodus bernor and lipscomb, 1991. hipparion platyodus deng et al., 2011. lectotype (bernor et al. 1990): pmum3691, an adult skull with right and left c-m3, housed in the paleontological, museum, university of uppsala, sweden (fig. 8). type locality: locality 70 (andersson locality number; fig. 1), wuxiang county, shanxi province, china stratigraphy and age: 76.5 ma. geographic distribution (fig. 1): yushe county, shanxi province; wuxiang county, shanxi province. revised diagnosis (synapomorphies with sivalhippus indicated in bold *): medium-size hipparion with cheek tooth row 140.0-149.0 mm. pob relatively long (43.6 mm in the lectotype pmum3691); anteriormost limit of lacrimal placed well posterior to pof (c1c, primitive as in s. nagriensis); nasomaxillary fossa absent (c2c); orbital surface of lacrimal bone with reduced foramen (c3b); pof subtriangular shape and anteroventrally oriented (c4d), posterior pocketing slight (c5b), moderately deep (c6b), medial wall morphology without internal pits (c7a), peripheral border moderately expressed (c8b), anterior rim faint (c9a); infraorbital foramen inferior to, or encroaching upon anteroventral border of the preorbital fossa (c10b); buccinator fossa distinctly limited (c11b) and not pocketed (c12a); caninus fossa absent (c13a); malar fossa absent (c14a); nasal notch at or near the anterior border of p2 (c15c) maxillary and mandibular dp1 absent (c16c). maxillary p2 anterostyle elongated (c17b); curvature of maxillary cheek teeth moderate (c18b); maximum cheek tooth height nearly 60 mm (c19c); cheek teeth with complexly plicated fossettes (c20a), preand postfossette opposing borders separate (c21b), posterior wall of postfossette always distinct (c22b), pli caballins mostly bifid or complex (c23a,c), hypoglyphs deeply incised (c24b), protocones short, lingually flattened and labially rounded (predominately c25e*, also g and d; c26a*), protocone always isolated from protoloph (c27b) with no protocone spur (c28c), protocone more lingually placed than hypocone in premolars (c29b) and molars (c30b). mandibular p2 with modestly elongate anterostylid in the lectotype pmum3691 (c31a); mandibular incisors are not grooved (c32a) and straight (c33b); i3 lateral aspect is elongate, not fig. 6 mandible of s. ptychodus, thp 01839. a) right lateral view; b) occlusal view. scale bar = 2 cm. sivalhippus (perissodactyla, mammalia) from the late miocene of china 11 labiolingually constricted (c34a); premolar metaconid is rounded on the distal surface (c35a), molar metaconid angular on the distal surface (c36c), premolar metastylid angular on the mesial surface (c37c*), premolar metastylid spur is absent (c38b), molar metastylid angular on the mesial surface (c39c*), molars lacking metastylid spur (c40b), premolar ectoflexid does not separate metaconid and metastylid (c41a); molar ectoflexid separates metaconid and metastylid (c42b); pli caballinid varies from being rudimentrary or single (c43b); protostylid present on occlusal surface as an enclosed enamel ring (c44a); protostylid orientation unknown (c45); ectostylids are absent (c46b); premolar linguaflexid shallow ushape (c47c*); molar linguaflexid is a very deep broad u-shape(c48e*); preflexids have complex margins (c49b); postflexids have complex margins (c50b); postflexid does not invade metaconid/ metastylid junction by anteriormost portion bending sharply lingually (c51a); protoconid enamel band is rounded (c52a). fig. 7 occlusal view of cheek tooth row of s. ptychodus. a) pmum353, left side; b) pmum353, right side; c) thp 01839, left side; d) amnh 143267, left side. scale bar = 2 cm. fig. 8 skull of s. platyodus, pmum 3691. a) right lateral view; b) occlusal view of left cheek tooth row. scale bar = 2 cm. sun b., zhang x., liu y. & bernor r.l.12 description pmum3691 (fig. 8) is the lectotype skull of sivalhippus platyodus (bernor et al. 1990). the skull is of an adult male, medium size with snout being 98.8 mm in length. the p2-m3 length is 149.2 mm. the nasomaxillary fossa is absent; orbital surface of the lacrimal bone has a reduced foramen; pob is 43.6 mm in length. pof is 70.3 mm in length with a dorso-ventral height of 49.7 mm, subtriangular shaped and anteroventrally oriented; pof ventral border id amnh 143269 pmum3691 taxon s. ptychodus s. platyodus lectotype tooth i1 i2 i3 c p2 p3 p4 m1 m2 m3 i1 i2 i3 c p2 p3 p4 m1 m2 m3 m1 13.5 14.5 13.5 10.2 24.8 21.2 21.1 20.5 19.8 22.2 12.5 14.1 14.1 11.0 28.7 24.5 23.8 22 22.1 24.6 m2 m3 9.9 8.8 7.6 7.6 9.9 12.3 11.7 11.5 11.3 9.4 9.6 9.6 8.3 8.3 11.9 15.3 14.7 13.8 12.6 10.7 m4 7.8 7.5 11.3 5.7 5.5 5.6 8.4 6.3 6.1 6.2 6.3 6.1 m5 11.3 11.6 10.2 8.4 8.4 7.1 12.6 12.8 12 8.4 8.7 8.4 m6 12.9 13.9 13.5 11.7 11 9.7 13.2 16.8 17.3 12.7 12.1 12 m7 m8 9.9 11 11 9.9 9.5 7.5 10.1 13.3 13.1 11.1 11.2 10.1 m9 11.5 11.4 10.8 9.1 8 6.7 11.8 13.4 12.2 10.2 9.8 8.5 id thp 01839 pmum353 pmum356 taxon s. ptychodus s. ptychodus s. ptychodus tooth p2 p3 p4 m1 m2 m3 i1 i2 i3 c p2 p3 p4 m1 p3 p4 m1 m2 m3 m1 26.0 23.7 23.1 22.8 23.9 22.5 10.3 12.2 12.3 9.4 26.1 22.4 21.7 19.0 23.4 23.8 21.6 22.1 m2 m3 9.6 12.2 10.8 12.5 11.4 9.2 9.6 9.7 7.9 7.4 9.9 13.2 12.1 11.6 13.4 14.3 12.4 12.9 11 m4 8.1 8.3 7.1 7.7 7 6.8 6 7.8 7.9 5.8 7.5 8.1 5.8 6.6 6.4 m5 11.6 11.8 10.8 9.5 9.2 6.6 12 11.5 11.3 6.4 11 11.3 7.1 7.9 9.5 m6 11.8 13.6 12 10.1 9.4 8.6 12 13.8 13.3 11.4 15.9 16 13.9 14.4 13.6 m7 m8 9.4 10.9 10.2 9.5 9.1 7.9 9.5 11.7 12.1 10.5 13.7 13.4 12.3 11 10 m9 10.5 11.2 9.5 9.3 9.2 7.1 11.3 11.4 10.8 10.1 13 12.3 10.6 10.1 9 tab. 5 standard mandibular teeth for specimens of chinese species of sivalhippus (mm). fig. 9 occlusal view of mandible of s. platyodus, pmum3691. scale bar = 2 cm. id nhmw a4229 fam 71800 amnh 19761 pmum 3691 thp 22708 pmum 347 pmum 3684 pmum 350 pmum 593 amnh 143267 taxon h. primigenium c. occidentale s. perimensis s.platyodus s.ptychodus country austria usa indopakistan china china china china china china china age(ma) 10.5 10.5 8.0 7.0 7.0 7.0 7.0 7.0 7.0 7.0 m1 136.6 110.7 116.8 98.8 97.0 102.5 108.8 m9 160.0 140.3 153.0 149.2 140.0 134.8 136.8 142.2 133.8 m32 47.2 39.7 56.2 43.6 50.2 47.0 41.6 44.7 43.5 m33 76.8 71.3 49.2 70.3 64.9 55.4 47.6 65.3 60.1 47.6 m35 56.2 39.3 29.1 49.7 44.9 33.3 27.9 31.0 37.7 32.8 m36 32.7 32.0 35.0 17.5 29.7 38.8 29.5 24.3 40.8 28.8 tab. 6 measurements of key characters of skull for cormohipparion occidentale (fam: 71800), sivalhippus perimensis (amnh19761) and chinese species of sivalhippus (mm), hippotherium primigenium (vienna basin pannonian c-d, nhmwa4229) standard. sivalhippus (perissodactyla, mammalia) from the late miocene of china 13 distance to facial-maxillary crest is 17.5 mm; pof pocketing reduced with moderate to slight depth; fossa medial depth is deep, greater than 15 mm; preorbital fossa medial wall without internal pits; preorbital fossa peripheral border outline is moderately well delineated around the periphery; anterior rim is present; infraorbital foramen inferior to anteroventral border of pof; buccinator fossa not confluent with a caninus fossa; buccinator fossa is not pocketed posteriorly; caninus fossa absent; malar fossa absent; nasal notch position cannot be determined. maxillary dp1 is absent; p2 anterostyle is moderately elongated; curvature of the cheek teeth is moderate; maximum cheek tooth crown height is estimated to be approximately 40-60 mm; maxillary cheek tooth fossette ornamentation is complex with several deeply amplified plications; preand postfossette opposing borders are not linked; posterior wall of postfossette is always distinct; pli caballin morphology varies from double/complex to single (on m2 and m3); hypoglyph is deeply incised; protocone is lingually flattened and labially rounded; no protocone is connected to the protoloph; there are no protoconal spurs; premolar protocone more lingually placed than hypocone; molar protocone more lingually placed than hypocone. pmum3691 (fig. 9) mandibular incisors are not grooved; they are straight; i3 lateral aspect is elongate, not labiolingually constricted. the p2 anterostylid is elongated. premolar and molar metaconids are rounded; premolar and molar metastylid are angular on mesial surface except for the right m3; premolar-molar metastylid spurs are absent; premolar ectoflexid closely approaches but does not separate metaconid and metastylid whereas molar ectoflexid separates metaconid and metastylid; pli caballinid is rudimentary or lacking; protostylid is present on occlusal surface often as an enclosed enamel ring; ectostylids are absent; premolar linguaflexid is shallow u-shape; molar linguaflexid is a deep u-shape; preflexid and postflexids have complex margins; postflexid does not invade metaconid/metastylid junction by anteriormost portion bending sharply lingually; protoconid enamel band is rounded. thp22708 (fig. 10) is a medium-sized adult female skull with a complete snout, lacking the posterior cranium. compared to the lectotype of sivalhippus platyodus, muzzle length virtually the same, 97.0 mm; p2-m3 is shorter, 140.0 mm.; pof is shorter, 64.9 mm and has a dorsal-ventral height shorter than the lectotype, 44.9 mm; distance from the ventral border of the pof to facial maxillary crest is greater, 29.7 mm. as with the lectotype, fig. 10 skull of s. platyodus, thp 22708. a) left oblique view; b) occlusal view of right cheek tooth row. scale bar = 2 cm. sun b., zhang x., liu y. & bernor r.l.14 pmum3684 has a pof that is placed high and far anteriorly on the face. the other cranial and dental features are as in the lectotype. ivppv8247 is a left mciii found from changyin locality, yushe basin (fig. 11). it is very short and robust (tab. 7), resembles those of typical sivalhippus (wolf et al. 2013). articular facets with mcii and mciv are large, which show that it has very robust medial and lateral metacarpals. remarks sivalhippus platyodus is morphologically similar to sivalhippus ptychodus (bernor et al. 1990; wolf et al. 2013). these two species also overlap in their geographic distribution (fig. 1). the long preorbital bar, large and subtriangular preorbital fossa, shallow nasal notch, complex upper cheek tooth fossettes, protocones that are lingually flattened and labially rounded, metaconid-metastylid with angular facing border and broad u-shaped linguaflexid of this species support its referral to the genus sivalhippus (fig. 5). zhegallo (1978) regarded s. platyodus as a subspecies of s. theobaldi. wolf et al. (2013) demonstrated that sivalhippus theobaldi is much larger with very robust metapodials but referred s. platyodus to the genus sivalhippus because of a number of synapomorphies: shallow nasal notch, strong subtriangular preorbital fossa, long preorbital bar, complex folds of upper cheek teeth, lingually flattened and labially rounded protocones and triangular double knots of lower cheek teeth. according to known s. platyodus material described by qiu et al. (1987) and bernor et al. (1990), this species has many similarities as siwaliks hipparion horses listed above. the highly robust metapodial is another diagnostic character of s. platyodus. qiu et al. (1987) reported a very short and robust mc iii from yushe basin (fig.11). as listed by wolf et al. (2013), the metapodial specimens of sivalhippus perimense, s. theobaldi and s. anwari are all robust. qiu et al. (1987) regarded sivalhippus ptychodus as a synonym of s. platyodus. later bernor et al. (1990) reiterated the validity of s. ptychodus, stating that this species is unique compared to other chinese hipparionines in its placement of the facial fossa far anteriorly and high dorsally above the plane of the facial-maxillary crest. with our observations herein, the differences between s. ptychodus and s. platyodus are observed on the cranium. the lectotype sivalhippus platyodus pmum3691 (bernor et al. 1990 fig. 4a&b) has a shorter pob, longer and deeper pof positioned closer to the facial-maxillary crest. the pof is larger and not positioned high on the face. the maxillary cheek teeth of s. platyodus have persistently labially rounded and lingually flattened protocones as in siwalik sivalhippus perimensis (wolf et al. 2013) and like s. ptychodus. the lectotype mandible of sivalhippus platyodus (pmum3691; bernor et al. 1990, fig 4c) has rounded premolar and molar metaconids while having angular premolar and molar metastylids; whereas most specimens of s. ptychodus have more angular metaconid and metastylid. however, the mandible amnh143267 has rounded metaconids both on premolar and molar (fig. 7d). this makes it difficult to distinguish s. ptychodus and s. platyodus fig. 11 left metacarpal iii of s. platyodus, ivpp v 8247. a) cranial view; b) caudal view; c) proximal view; d) distal view. scale bar = 2 cm. tab. 7 measurements of left mciii of sivalhippus platyodus, v8247 (mm). m1 m2 m3 m4 m5 m6 m7 171.3 165.4 27.0 19.4 36.3 24.2 31.9 m8 m9 m10 m11 m12 m13 m14 9.8 5.6 37.8 34.9 24.0 20.4 23.7 sivalhippus (perissodactyla, mammalia) from the late miocene of china 15 based on mandibular cheek teeth alone. the mandibular cheek teeth of sivalhippus ptychodus also have p2s with shorter anterostylids that are seen on all of our specimens: it is a stable feature. whereas the lectotype of s. platyodus, m3691 has elongated anteorstylid on p2 it is shorter on specimens of s. ptychodus. the specimens of sivalhippus perimensis, amnh19466 and amnh19761 crania are larger than both s. ptychodus and s. platyodus (wolf et al. 2013). chinese species of plesiohipparion and proboscidipparion and african eurygnathohippus evolve even more extreme angular metaconids and metastylids on the cheek teeth (bernor & sun 2015). discussion sivalhippus ptychodus shares with sivalhippus perimensis a pof that is placed dorso-ventrally high and far anterior on the face. other lineages of hipparionine horses have long preorbital bars and large, dorsoventrally extensive pof such as is found in s. platyodus: cormohipparion spp. and hippotherium spp (bernor et al. 1997, 2011; bernor & white 2009). most species of cormohipparion group are distributed in north america. cormohipparion was the lineage that populated eurasia and africa and is known from sinap, turkey, (bernor et al. 2003), bou hanifia, algeria (bernor & white 2009) and ethiopia (bernor et al. 2010; 2017). hippotherium was a successful clade that ranged from central europe eastward to the balkans and turkey including the following taxa: hippotherium sp. (vienna basin, pannonian c; bernor et al. 2017), hippotherium primigenium (central europe, bernor et al. 1988; 1997), hippotherium brachypus (greece and iran, koufos 1987; bernor et al. 2016), hippotherium intrans (hungary, kretzoi 1983; bernor et al. 2004); hippotherium microdon (hungary, kormos 1914), hippotherium kammerschmitti (germany, kaiser et al. 2003) and hippotherium malpassi (italy, bernor et al. 2011). chinese hipparion species similar to hippotherium include “hippotherium” weihoense and “hippotherium” chiai (qiu et al. 1987). bernor and white (2009) analyzed key measurements for distinguishing clades of old world hipparions: m1 = snout length; m9 = cheek tooth row length; m32 = pob length; m33 = pof length; m35 = pof dorsoventral height; m36 = distance from ventral border of pof to facial-maxillary crest. table 6 provides the key measurements that we use to compare north american cormohipparion occidentale (fam 71800), siwalik sivalhippus perimensis (amnh19761) and china sivalhippus ptychodus and sivalhippus platyodus specimens discussed herein. we use the hippotherium primigenium from inzersdorf (nhmwa4229) as our standard and plot the log10 fig. 12 ratio diagrams of skull measurements of sivalhippus and other hipparion horses. log10 standard is inzersdorf hipparion, hippotherium primigenium (zero line). a) critical measurements on fam71800 cormohipparion occidentale, sivalhippus perimensis and chinese sivalhippus, log10 ratios. b) critical skull measurements on fam71800 cormohipparion occidentale, sivalhippus perimensis and sivalhippus ptychodus. c) critical skull measurements in cormohipparion occidentale, sivalhippus perimense and sivalhippus platyodus. measurement numbers: m9, upper cheek teeth length; m32, distance between the orbit and the preorbital fossa; m33, length of the preorbital fossa; m35, height of the preorbital fossa; m36, distance between the preorbital fossa and the facial crest. sun b., zhang x., liu y. & bernor r.l.16 ratios for the comparative samples (fig. 12 a-c). fig. 12 plots skulls of chinese specimens of sivalhippus ptychodus and sivalhippus platyodus compared to north american c. occidentale (fam: 71800 following bernor et al. 2003; bernor & white 2009), siwalik s. perimensis (amnh19761) with pannonian d-e h. primigenium from inzersdorf (nhmwa4229) being the log10 standard. fig. 12a plots all chinese sivalhippus together with north american c. occidentale and siwalik s. perimensis. fig. 12b plots c. occidentale and s. perimensis compared to chinese sivalhippus ptychodus only and fig. 12c plots c. occidentale and s. perimensis compared to sivalhippus platyodus only. a number of clear morphological distinctions can be made between the taxa under consideration (tab. 6 and fig. 12a-c). the h. primigenium standard from inzersdorf, austria is the largest taxon with the longest cheek tooth row (m9), s. perimensis has a slightly shorter tooth row and all chinese sivalhippus have shorter cheek tooth rows than both. siwalik s. perimensis is the most derived taxon in the length of the pob (m32 = 56.2mm) and restriction of the pof high on the face (m35 = 29.1, m36 = 35.0 mm). figures 12b and 12c exhibit clear distinctions between sivalhippus ptychodus (fig. 12b) and s. platyodus (fig. 12c): s. ptychodus has all measurements for pof height (m35) falling below north american c. occidentale as is found for siwalik s. perimensis; s. platyodus has all measurements for pof height falling above that for c. occidentale. lectotype of sivalhippus ptychodus (pmum347) is of only moderate size (m9 = 134.8 mm) but has a very long pob (m32 = 50.2 mm) comparing closely with s. perimensis, albeit being smaller. bernor et al. (1988; 1996) and woodburne (2007) argued that cormohipparion was the ancestral group from which old world hipparion evolved. cormohipparion occidentale was believed to be the ancestral type for old world hipparionine horses (bernor et al. 2003). cormohipparion sinapensis (sinap, tab. 8 summary character state distributions of sivalhippus ptychodus and sivalhippus platyodus skulls, maxillary and mandibular dentitions. taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 pmum347 g c b k b a a b a b b a a a c a b pmum3684 g c b k b a a b a b b a a a b c a b pmum350 g c b k b a a b a b b a a a pmum593 g c b k b a a b a b b a a a pmum353 pmum356 thp01839 amnh143267 g c b k b a a b a b b a a a c c a b pmum3691 c c b d b b a b a b b a a a c c b b thp22708 c b d b b a b a b b a a a c c b b taxa 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 pmum347 c/d a b b a,c c e a b c b b pmum3684 c/d a b b a,c c e a b c b b pmum350 b a,b b d,e pmum593 b g,e pmum353 b c pmum356 c c thp01839 b c c amnh143267 a b b a,b b e a b c b b b a b a a a pmum3691 c a b b a,c b e a b c b b a a b a a a thp22708 c a b b a,c b e a b c b b taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 pmum347 g c b k b a a b a b b a a a c a b pmum3684 g c b k b a a b a b b a a a b c a b pmum350 g c b k b a a b a b b a a a pmum593 g c b k b a a b a b b a a a pmum353 pmum356 thp01839 amnh143267 g c b k b a a b a b b a a a c c a b pmum3691 c c b d b b a b a b b a a a c c b b thp22708 c b d b b a b a b b a a a c c b b taxa 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 pmum347 c/d a b b a,c c e a b c b b pmum3684 c/d a b b a,c c e a b c b b pmum350 b a,b b d,e pmum593 b g,e pmum353 b c pmum356 c c thp01839 b c c amnh143267 a b b a,b b e a b c b b b a b a a a pmum3691 c a b b a,c b e a b c b b a a b a a a thp22708 c a b b a,c b e a b c b b taxa 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 pmum347 pmum3684 pmum350 pmum593 pmum353 c b c b b c b b d a a a a pmum356 c b c b a b c f c b d e a a a a thp01839 c a c b a a b b b c d b,c c a a amnh143267 c b c b a b b b b c d a a a a pmum3691 c b c b a b b a b c e b b a a thp22708 sivalhippus (perissodactyla, mammalia) from the late miocene of china 17 turkey 10.8 ma) and hippotherium primigenium (central europe, ca. 11.4-11.0 ma.) are the two oldest and most primitive old world hipparion lineages; c. occidentale occurred in indopakistan, turkey and africa while h. primigenium evolved originally in central europe (bernor et al. 2017). in these lineages the pof is characterized as being large, dorsoventrally and anteroposteriorly extensive, medially deep and posteriorly pocketed with accompanying long pob. species of these primitive lineages also have a pof positioned closer to the facial-maxillary crest. morphometric analysis by bernor et al. (2003) and phylogenetic analysis of woodburne (2007) showed that north american early late miocene c. matthewi and c. occidentale were sister taxa and formed the most derived clade within the american cormohipparion. woodburne (2009) considered that another cormohipparion, cormohipparion sp. from the punchbowl formation, california, was the likely source of the old world “hipparion” datum and radiation. this was supported by bernor et al.’s (2017) analysis of vienna basin pannonian c hipparions. the siwaliks, indopakistan, underwent a regional evolutionary radiation of the genus sivalhippus (wolf et al. 2013). the oldest occurring member of the sivalhippus clade is sivalhippus nagriensis which occurred 10.5-9.3 ma. the very large and very robust species sivalhippus theobaldi ranged in age from 9.3 to 7.8 ma. the large and robust species, sivalhippus perimensis ranged in age from 8.5-7.3 ma. the last locally occurring potwar plateau species, also large, s. anwari ranged from 7.4-7.2 ma. in china, sivalhippus cf. theobaldi is reported from yunan, ca. 8.2-7.2 ma (sun 2013). sivalhippus ptychodus occurred in qingyang, gansu ca. 7 ma, and in yushe and wuxiang basins, shanxi in 7-6.5 ma. sivalhippus platyodus also occurred in the yushe and wuxiang basins, shanxi in 7-6.5 ma and the wulanhua area circa 7 ma. sivalhippus cf. theobaldi occured in myanmar ca. 8 ma and yuanmou and lufeng, yunnan, china between 8 and 7 ma (qi et al. 2006; sun 2013). flynn and qi (1982) argued the age of lufeng fauna was 8 ma. sivalhippus also occured in africa: in uganda s. macrodon is circa 9 ma, and is likely related to siwalik s. theobaldi; s. turkanensis occurs at lothagam, ca. 6.5 ma (bernor & harris 2003) and plausibly sahabi, circa 7 ma (bernor et al. 2010) and is related to sivalhippus perimensis (fig. 13).the phylogenetic analysis of wolf et al. (2013) hypothesized that s. platyodus was the sister taxon of siwalik s. theobaldi. however, we believe that eventhough the type specimen of siwalik s. nagriensis is fragmentary, it is morphologically closer to s. platyodus than either s. theobaldi or s. ptychodus. both s. nagriensis and s. platyodus have dorsoventrally extensive, subtriangular pof placed close to the facial-maxillary crest. we believe that at least these two species share a very close relationship. on the other hand, s. ptychodus is most similar to siwalik s. perimensis. although s. ptychodus and s. platyodus have overlapping geographic and choronologic distributions in china, they likely represent two separate dispersal events of hipparion horse from indopakistan (fig. 13). conclusions sivalhippus ptychodus has derived features including a long pob, strongly developed pof placed dorsally high and far anteriorly shallow nasal notch, complexly plicated upper cheek tooth fossettes, lingually flattened and labially rounded protocones, variably occurring pli caballinids, rounded to triangular metaconids, triangular metastylid and broad fig. 13 geological distribution of sivalhippus lineages in indopakistan, china and africa showing their phylogenetic relation and dispersal events. sivalhippus nagriensis lineage (s. nagriensis and s. platyodus) in black; sivalhippus theobaldi lineage in red (s. theobaldi , s. macrodon, and s. cf. theobaldi); sivalhippus perimensis lineage (s. perimensis, s. turkanensis, s. anwari and s. ptychodus) in blue. sun b., zhang x., liu y. & bernor r.l.18 u-shaped linguaflexid on the mandibular molars typical for the genus sivalhippus. sivalhippus ptychodus differs from s. platyodus in having a pof placed high on the face with a shorter pof dorsoventral height. sivalhippus ptychodus is a valid species and not a synonym of s. platyodus. the facial morphology of s. ptychodus most resembles that of siwalik s. perimensis and to a limited extent, s. turkanensis, whereas that of s. platyodus is more similar to sivalhippus nagriensis. sivalhippus spp. likely extended its range from south asia into china between 9 and 7 ma. acknowledgements: we thank prof. deng tao for his comments. we thank dr. jan-ove r. ebbestad and dr. benjamin kear of museum of evolution of uppsala university, dr. meng jin and ms judy galkin of american museum of natural history for providing the facilities and opportunity for studying the chinese hipparionine material in their care. we thank gao wei for his photographs and su dan for preparation of the specimen. we thank sun danhui for her support on photography and measurement work of metacarpal specimen. we thank editor prof. lorenzo rook, reviewer dr. wang shiqi and two anonymous reviewers for their review and important comments for our manuscript. this work was supported by national natural science foundation of china (41430102 and 41402003), the strategic priority cultivating research program, cas (xdpb05), and the key research program of frontier sciences, cas (qyzdy-ssw-dqc022). sun also acknowledges funding from the china scholarship council for 1 year research in the bernor lab at howard university. bernor wishes to acknowledge research funding by nsf ear grants 8806645, 0125009, 1113175, 1558586 and bcs0321893 (to f.c. howell and t.d. white) which provided data and background investigations critical to this manuscript. references armour-chelu m. & bernor, r. l. 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(1978) gippariony tsentral’noj azii. tr sovmest sov-mong paleont eksped, 7: 1-152. appendix hipparionine character states (bernor et al. 1990; 2017) 1) relationship of lacrimal to the preorbital fossa: a = lacrimal large, rectangularly shaped, invades medial wall and posterior aspect of preorbital fossa; b = lacrimal reduced in size, slightly invades or touches posterior border of preorbital fossa; c = preorbital bar (pob) long with the anterior edge of the lacrimal placed more than half the distance from the anterior orbital rim to the posterior rim of the fossa; d = pob reduced slightly in length but with the anterior edge of the lacrimal placed still more than 1/2 the distance from the anterior orbital rim to the posterior rim of the fossa; e = pob vestigial, but lacrimal as in d; f = pob absent; g = pob very long with anterior edge of lacrimal placed less than 1/2 the distance from the anterior orbital rim to the posterior rim of the fossa. 2) nasolacrimal fossa: a = pof large, ovoid shape and separated by a distinct medially placed, dorsoventrally oriented ridge, dividing pof into equal anterior (nasomaxillary) and posterior (nasolacrimal) fossae; b = nasomaxillary fossa sharply reduced compared to nasolacrimal fossa; c = nasomaxillary fossa absent (lost), leaving only nasolacrimal portion (when a pof is present). 3) orbital surface of lacrimal bone: a = with foramen; b = reduced foramen. 4) preorbital fossa morphology: a = large, ovoid shape, anteroposteriorly oriented; b = pof truncated anteriorly; c = pof further truncated, dorsoventrally restricted at anterior limit; (note csk a-c are pre-old world hipparionine characters); d = subtriangular shaped and anteroventrally oriented; e = subtriangularly shaped and anteroposteriorly oriented; f = egg-shaped and anteroposteriorly oriented; g = c-shaped and anteroposteriorly oriented; h = vestigial but with a c-shaped or egg-shaped outline; i = vestigial without c-shape outline, or absent; j = elongate, anteroposteriorly oriented; k = small, rounded structure; l = posterior rim straight, with non-oriented medial depression. 5) fossa posterior pocketing: a = deeply pocketed, greater than 15 mm in deepest place; b = pocketing reduced, moderate to slight depth, less than 15 mm; c = not pocketed but with a posterior rim; d = absent, no rim but a remnant depression; e = absent. 6) fossa medial depth; a = deep, greater than 15 mm. in deepest place; b = moderate depth, 10-15 mm in deepest place; c = shallow depth, less than 10 mm in deepest place; d = absent. 7) preorbital fossa medial wall morphology: a = without internal pits; b = with internal pits. 8) fossa peripheral border outline: a = strong, strongly delineated around entire periphery; b = moderately delineated around periphery; c = weakly defined around periphery; d = absent with a remnant depression; e = absent, no remnant depression. 9) anterior rim morphology: a = present; b = absent. 10) placement of infraorbital foramen: a = placed distinctly ventral to approximately 1/2 the distance between the preorbital fossa’s anteriormost and posteriormost extent (preold world hipparionine condition); b = inferior to, or encroaching upon anteroventral border of the preorbital fossa. 11) confluence of buccinator and canine fossae: a = present (preold world hipparionine condition); b = absent, buccinator fossa is distinctly delimited. 12) buccinator fossa: a = not pocketed posteriorly; b = pocketed posteriorly. 13) caninus (= intermediate) fossa: a = absent; b = present. 14) malar fossa: a = absent; b = present. 15) nasal notch position: a = at posterior border of canine or slightly posterior to canine border; b = approximately half the distance between canine and p2; c = at or near the anterior border of p2; d = above p2; e = above p3; f = above p4; g = above ml; h = posterior to ml. 16) presence of dp1: a = persistent and functional; b = reduced and non-functional; c = absent. 17) p2 anterostyle: a = short; b = elongate 18) curvature of maxillary cheek teeth: a = very curved; b = moderately curved; c = straight. 19) maximum cheek tooth crown height: a = < 30 mm; b = 30 40 mm; c = 40 60 mm; d = 60 75 mm; e = 75+ maximum crown height. 20) maxillary cheek tooth fossette ornamentation: a = complex, with several deeply amplified plications; b = moderately complex with fewer, more shortly amplified, thinly banded plications; c = simple complexity with few, shortly amplified plications; d = generally no plis; e = very complex. 21) confluence (linkage) of preand postfosstte opposing sivalhippus (perissodactyla, mammalia) from the late miocene of china 21 borders: a = linked; b = separate. 22) posterior wall of postfossette: a = may not be distinct; b = always distinct. 23) pli caballin morphology: a = double; b = single or occasionally poorly defined double; c = complex; d = plis not well formed. 24) hypoglyph: a = hypocone frequently encircled by hypoglyph; b = deeply incised, infrequently encircled hypocone; c = moderately deeply incised; d = shallowly incised. 25) protocone shape: a = round q-shape; b = oval q-shape; c = oval; d = elongate-oval; e = lingually flattened-labially rounded; f = compressed or ovate; g = rounded; h = triangular; i = triangular-elongate; j = lenticular; k = triangular with rounded corners. 26) protocone flattened: a = yes; b = no 27) isolation of protocone: a = connected to protoloph; b = isolated from protoloph. 28) protoconal spur: a = elongate, strongly present; b = reduced; c = absent. 29) premolar protocone/hypocone alignment (preold world hipparionine condition): a = anteroposteriorly aligned; b = protocone more lingually placed. 30) molar protocone/hypocone alignment (preold world hipparionine condition): a = anteroposteriorly aligned; b = protocone more lingually placed. 31) p2 anterostylid: a = elongate; b = short and rounded. 32) mandibular incisor morphology: a = not grooved; b = grooved. 33) mandibular incisor curvature: a = curved; b = straight. 34) i3 lateral aspect: a = elongate, not labiolingually constricted; b = very elongate, labiolingually constricted distally; c = atrophied. 35) premolar metaconid: a = rounded; b = elongated; c = angular on distal surface; d = irregular shaped; e = square shaped; f = pointed. 36) molar metaconid: a = rounded; b = elongated; c = angular on distal surface; d = irregular shaped; e = square shaped; f = pointed. 37) premolar metastylid: a = rounded; b = elongate; c = angular on mesial surface; d = irregular shaped; e = square shaped; f = pointed. 38) premolar metastylid spur: a = present; b = absent 39) molar metastylid: a = rounded; b = elongate; c = angular on mesial surface; d = irregular shaped; e = square shaped; f = pointed. 40) molar metastylid spur: a = present; b = absent 41) premolar ectoflexid: a = does not separate metaconid and metastylid; b = separates metaconid and metastylid. 42) molar ectoflexid: a = does not separate metaconid and metastylid; b = separates metaconid and metastylid; c = converges with preflexid and postflexid to abut against metaconid and metastylid. 43) pli caballinid: a = complex; b = rudimentary or single; c = absent. 44) protostylid: a = present on occlusal surface often as an enclosed enamel ring; b = absent on occlusal surface, but may be on side of crown buried in cement; c = strong, columnar; d = a loop; e = a small, poorly developed loop; f = a small, pointed projection continuous with the buccal cingulum. 45) protostylid orientation: a = courses obliquely to anterior surface of tooth; b = less oblique coursing, placed on mesio-labial of tooth; c = vertically placed, lies flush with protoconid enamel band; d = vertically placed, lying lateral to protoconid band; e = open loop extending posterolabially. 46) ectostylids: a = present; b = absent. 47) premolar linguaflexid: a = shallow; b = deeper, v-shaped; c = shallow u-shaped; d = deep, broad u-shape; e = very broad and deep. 48) molar linguaflexid: a = shallow; b = v-shaped; c = shallow u-shaped; d = deep, broad u-shape; e = very broad and deep u-shape. 49) preflexid morphology: a = simple margins; b = complex margins; c = very complex. 50) postflexid morphology: a = simple margins; b = complex margins; c = very complex. 51) postflexid invades metaconid/metastylid junction by anteriormost portion bending sharply lingually: a = no; b = yes. 52) protoconid enamel band morphology: a = rounded; b = flattened. _goback _goback _goback _goback _goback _goback _goback _goback _goback _goback seko 49..60 ��������������� ��� ������ ���� �� � � ����� �� ������������ ������ �� �������� � ������ ���� ��� ������ ������� �� ������ ��� ��� ������������ �� ��� ���� �� ���� ����� ���� � � ����� ��� �������� � �� ������� � ������ ��� � �� ��� ������ � ��������� � ��� �� ���� ���� ��������� �� ������� ��� � � ��� � ����������� ��������� ��������� �� � �� ����� ��������� ����� � �� ���� ��� �� ��� ��� !"# � �$"�%&$� �'"!$($� � �& �)' ( *"�$� #$$&� �$(�� + � � �$�)%�� �$#'�$�� ($ (� � +$",'"-$"-� &)%� � +$"$� !"+"-%. ���� ���. �/$+0$ �&$!'$� ���',1�$# �" �)$ -$(1� �!����� � � ) 0$ ,$$( �$!"�#$# 2�"3 �)$ $&'($�'�'!4($�'�'! 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�% �" ���������" ��� ���������" +"��!� ,�� �� � �% �����$ ������ ��� � ����������-: ���)�!� �� �� >�'# ;# 7 8 9 10 11 12 13 2,4 2,5 2,6 2,7 2,8 2,9 3,0 protoconch whorls pr ot oc on ch d ia m et er /d ia m et er o f th e fi rs t h al f w ho rl >�'# 0 ( � !������"�� ) �+ � ���������" ���� � �c���� � � �% �" %���� "�!% +"��! ����� ��� ���������" +"��!� ,�� �� � �% ����������� ��� � ����������-: ���)�!� �� �� >�'# ;# %�� ���� &' ( ����� )'* virguloides depressofuniculata epigloafuniculata fredianii fulgurata obliquicallosa plicatula propinqua pseudoepiglottina raropunctata strictiumbilicata sulcogradata undata vittata dillwyni koeneni adansoni astensis filosa prietoi rizzae tectula n atica virguloides 0,3 -0,2 0,3 0,3 0,3 -1,6 0,3 -0,7 0,3 0,3 0,3 -0,3 -0,7 -1,8 -0,5 -1,3 -1,3 -1,2 -1,1 -1,0 -1,4 c ochlis depressofuniculata -0,4 0,0 0,0 0,0 -1,9 0,0 -0,9 0,0 0,0 0,0 -0,5 -1,0 -2,0 -0,8 -1,5 -1,6 -1,5 -1,4 -1,3 -1,7 epigloafuniculata 0,4 0,4 0,4 -1,5 0,4 -0,5 0,4 0,4 0,4 -0,1 -0,6 -1,6 -0,3 -1,1 -1,1 -1,1 -0,9 -0,8 -1,2 fredianii 0,0 0,0 -1,9 0,0 -0,9 0,0 0,0 0,0 -0,5 -1,0 -2,0 -0,8 -1,5 -1,6 -1,5 -1,4 -1,3 -1,7 fulgurata 0,0 -1,9 0,0 -0,9 0,0 0,0 0,0 -0,5 -1,0 -2,0 -0,8 -1,5 -1,6 -1,5 -1,4 -1,3 -1,7 obliquicallosa -1,9 0,0 -0,9 0,0 0,0 0,0 -0,5 -1,0 -2,0 -0,8 -1,5 -1,6 -1,5 -1,4 -1,3 -1,7 plicatula 1,9 1,0 1,9 1,9 1,9 1,4 1,0 -0,1 1,1 0,4 0,3 0,3 0,5 0,6 0,2 propinqua -0,9 0,0 0,0 0,0 -0,5 -1,0 -2,0 -0,8 -1,5 -1,6 -1,5 -1,4 -1,3 -1,7 pseudoepiglottina 0,9 0,9 0,9 0,4 0,0 -1,1 0,2 -0,6 -0,6 -0,6 -0,4 -0,3 -0,7 raropunctata 0,0 0,0 -0,5 -1,0 -2,0 -0,8 -1,5 -1,6 -1,5 -1,4 -1,3 -1,7 strictium bilicata 0,0 -0,5 -1,0 -2,0 -0,8 -1,5 -1,6 -1,5 -1,4 -1,3 -1,7 sulcogradata -0,5 -1,0 -2,0 -0,8 -1,5 -1,6 -1,5 -1,4 -1,3 -1,7 undata -0,5 -1,5 -0,3 -1,0 -1,1 -1,0 -0,9 -0,8 -1,2 vittata -1,0 0,2 -0,5 -0,6 -0,6 -0,4 -0,3 -0,7 t anea dillw yni 1,3 0,5 0,4 0,5 0,6 0,8 0,3 koeneni -0,8 -0,8 -0,8 -0,6 -0,5 0,9 t ectonatica/c ryptonatica adansoni -0,1 0,0 0,1 0,3 -0,2 astensis 0,0 0,2 0,3 -0,1 filosa 0,2 0,3 -0,1 prietoi 0,1 -0,3 rizzae -0,4 tectula � �) # ( 1 ����� �" � + �� ' � �%% � � � �� � & � ) � � % � �� �� �� � �" + " � �!� + ��" �� �� �� � � ����: ��' � �%���� � $ �!& � �� ) � !� %�� � # � !! �" � ������ ��� � %� & � � �� � ��& � �� �" � !�� � � ,� � ���!� � � ) ) � 5 5 = : � � � � � � �� ��� !��� � : � ���� � � �� � � � � �� � �� "��� �� �� � �, , � � �!�� �!� � � %� � �� � � ����� � # ������� �" ��� %������� �������� �" �� ��� � ��� ���� �� *����' **' ��� � �"����� ��������� *3 virguloides depressofuniculata epigloafuniculata fredianii fulgurata obliquicallosa plicatula propinqua pseudoepiglottina raropunctata strictiumbilicata sulcogradata undata vittata dillwyni koeneni adansoni astensis filosa prietoi rizzae tectula n at ic a vi rg ul oi de s 17 -1 1 14 23 1, 3 -4 3 9 -2 10 -6 -2 0 -2 2 24 -2 5 -4 1 -4 4 -4 5 -3 2 -2 5 -3 8 -4 5 c oc hl is de pr es so fu ni cu la ta -2 6 -3 7 -1 6 -5 2 -8 -1 8 -7 -2 2 -3 4 -3 5 8 -3 8 -5 1 -5 3 -5 4 -4 4 -3 8 -4 9 -5 4 ep ig lo af un ic ul at a 24 31 13 -3 5 20 10 20 9 -1 0 -1 2 33 -1 6 -3 4 -3 7 -3 8 -2 4 -1 6 -3 0 -3 8 fr ed ia ni i 10 -1 3 -5 0 -5 -1 5 -5 -1 6 -3 1 -3 3 11 -3 6 -5 0 -5 2 -5 2 -4 2 -3 6 -4 7 -5 2 fu lg ur at a -2 2 -5 5 -1 5 -2 4 -1 4 -2 7 -3 8 -4 0 1 -4 2 -5 5 -5 6 -5 7 -4 8 -4 2 -5 2 -5 7 ob liq ui ca llo sa -4 3 8 -3 9 -7 -2 1 -2 3 23 -2 6 -4 2 -4 4 -4 5 -3 3 -2 6 -3 9 -4 5 pl ic at ul a 48 41 48 37 28 26 56 23 2 -2 -4 15 23 -7 -4 pr op in qu a -1 1 1 -1 5 -2 8 -2 9 16 -3 3 -4 7 -4 9 -5 0 -3 9 -3 3 -4 4 -5 0 ps eu do ep ig lo tti na 12 -4 -1 9 -2 1 25 -2 4 -4 0 -4 3 -4 4 -3 1 -2 4 -3 7 -4 4 ra ro pu nc ta ta -1 5 -2 8 -3 0 15 -3 3 -4 7 -4 9 -5 0 -3 9 -3 3 -4 4 -5 0 st ri ct iu m bi lic at a -1 5 -1 7 28 -2 3 -4 0 -4 2 -4 3 -3 0 -2 3 -3 7 -4 3 su lc og ra da ta -3 39 -6 -2 6 -2 9 -3 0 -1 5 -6 -2 3 -3 0 un da ta 41 -4 -2 5 -2 8 -3 0 -1 3 -4 -2 1 -3 0 vi tta ta -4 3 -5 5 -5 7 -5 9 -4 9 -4 3 -5 3 -5 9 t an ea di llw yn i -2 1 -2 5 -2 6 -1 0 0 -1 7 -2 6 ko en en i -4 -5 13 21 5 -5 t ec to na ti ca /c ry pt on at ic a ad an so ni -2 17 25 9 -2 as te ns is 18 26 10 18 fil os a 10 -9 -1 8 pr ie to i -1 7 -2 6 ri zz ae -1 0 te ct ul a � �) # ( 1 �� �� � �" � + �� ' � � � � � � �% % � � � �� � �� � � � � % �" !� �$ �! �" ! ! + �� " �� �� � � � � �� �� : �� ' � �% �� �� � $ �! & � �� ) � !� %� � � # � !! �" � �� �� �� �� � � %� & � � �� � �� & � �� �" � !� � � � ,� � �� �! � � � ) ) � 5 5 = : � � � � � � �� � �� !� �� � : � �� �� � � �� � � � � �� � �� "� �� �� �� � � , , � � �! �� � !� � � %� � �� � � �� �� � � # %�� ���� &' ( ����� )'*? virguloides depressofuniculata epigloafuniculata fredianii fulgurata obliquicallosa plicatula propinqua pseudoepiglottina raropunctata strictiumbilicata sulcogradata undata vittata dillwyni koeneni adansoni astensis filosa prietoi rizzae tectula n atica virguloides 7 38 26 47 -6 71 -21 67 -6 -19 -27 -1 74 79 47 71 68 71 65 59 71 c ochlis depressofuniculata 34 22 43 -13 68 -27 64 -13 -25 -32 8 72 78 44 69 67 69 63 56 69 epigloafuniculata -16 14 -42 52 -51 43 -42 -50 -55 -38 57 67 14 52 48 52 43 33 52 fredianii 28 -31 60 -42 56 -31 -40 -47 -26 64 72 28 60 56 60 52 44 60 fulgurata -50 44 -58 37 -50 -57 -61 -48 51 61 0 44 39 44 33 22 44 obliquicallosa 72 -16 69 -1 -14 -23 5 76 81 50 72 69 72 67 61 72 plicatula -77 -12 -72 -76 -79 -71 12 30 -44 0 -9 0 -17 -29 0 propinqua 74 16 2 -8 20 80 84 58 77 74 77 72 67 77 pseudoepiglottina -69 -73 -76 -67 22 36 -39 9 0 9 -8 -21 9 raropunctata -14 -22 5 76 81 50 72 69 72 67 61 72 strictium bilicata -10 19 79 83 57 76 74 76 71 67 76 sulcogradata 26 81 85 62 79 77 79 74 70 79 undata 74 79 47 71 68 71 65 59 71 vittata 22 -50 -10 -18 -10 -25 -36 -10 t anea dillw yni -61 -30 -36 -30 -42 -50 -30 koeneni 44 39 44 33 22 44 t ectonatica/c ryptonatica adansoni -9 0 -17 -29 0 astensis 9 -8 -21 9 filosa -17 -29 0 prietoi -14 17 rizzae 29 tectula � �) # 3 ( 1 ����� �" � + �� ' � �� � � � �%% � � � �� � ��� � � � % �" %���� " �!% + " � �! 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" �'"� ,�� �� � �% ����������� ��� � ����������-: ���)�!� �� �� >�'# 5# 0 2 4 6 8 10 12 0 5 10 15 20 25 30 35 40 45 50 shell height (mm) sp ir e he ig ht (m m ) >�'# 3 ( � !������"�� ) �+ � ���� " �'"� ��� �" !! " �'"� ,���� %��� � ����!� ��)( )� 55= ��� �� � �� ��� �-: �� � �6&�� �� ����+ ������$ ��: ��!�� �����'! �� ������ ��: � � �� � ��������� � �$ � ������ '��&� ����+ ����$ ���� ��!&� �: �� � ����! �� ����������� ��� � ������$ ���� ������� ��������' 0 5 10 15 20 25 0 5 10 15 20 25 aperture height (mm) ap er tu re w id th (m m ) >�'# ? ( � !������"��� ) �+ � �� ��&� +���" ��� �� ��&� " �'"� ,�� �� � �% ������� ��� ������ �"�� ��&��-: ��!�� �����'! �� ���!���" ��� ���� ��� " ������� ���)�� �: �� � �6&�� �� #������# " � -� �� ��'��%����� ��%% � �� ��&!� ) � � �� � ) ( �+ � ���!���" ��� ����� " ������� ��� #������ �� + !! �� ) �+ � �" � ��� �" ��" � ������� ���� � �!� +��" )� � ����!� ��))� , 55=-# >��� >�'# = �"��&'" / ,� '� ����� !�� � ����� � �� �� � �$ �!������� ��� ��� �"�� ����������� �� �� � ��� "����� � ��" � ��%% � %��� �� ����" �� ��� �" � ��%% � %��� �" ���� ������ � � )� �" �)�$ ��� � �&�"���# �" �� %%��� ��� �% ���� !�( ���� ��!�&!�� � %�� �" �"� ������ � � � !������ � ( �&!� � �� ) "�'" �� �!! �� �� � ,� e 5#*/(5#00-# �" ��� � !�� �!�� ,���!������� �% �" ��� � !�� �� �" �" !! ����$��� � ��� �� ! �� '� ��!� ��������' �� �� �� �# �" 0=b ���%�� �� ��� �$�!� ,��)# =!��' !� �$ �!�� �� ����" � ��� �� ��� �!!�+ ��� � !��)! � ( �������� ) �+ � �" �!��� � ������� ���� ,� ����!� ��))� 55=: �� � �� ��&��-# �" ���� ��! ��!!&� �% �" ��� ���� ��� ��� �� ) "���!� &��)! �� ������'&��"��' �"����� �# �� ����� �� ��� ) ��� � �"�� �" ���� ��! ��!!&� �% �������� "�� �" ��� ���� !�) ���!! ��� �&)( ��&�� � �� ���������� ,��)# ;-# .�������� ��� ���� �# �" &�)�!��&� +���" ���$ � �� ) ��'��%�����!� ���� !�� � +��" �" �����&� ���( � � � �% �" �" !! �� �!! �� �� � ,� e 5#;/(5#00-# d" � �" &�)�!��&� +���" �� � '� �� � �'����� �����&� ���( � � �� �� ��� ��� �"�� ���!���" ��� ����� " ������� ��� #������ ��� ) ������'&��" � %��� �� ����" �# �� %���� �" '���" ,>�'# *�"�+� �"�� �" !�� � %�� ���!���" ��$ � ���� ��� #������ "�$ ����!�� �!�� � )&� ��'��%�����!� ��%% � �� ! $������ +" � �� �" !�� %�� " �������� ���( ��� � �� �" ��" � �+�� "�� ��'��%�����!� ��%% � �� �!�� # �"&�� %�� � '�$ � ���� � � �% �" �" !!� �" &�)�!��&� �% #������ �� �!+��� '� �� � �"�� �"�� �% ���!���" ��� ���� ���� ��������!�� �!�� �"�� �"�� �% " �������# 1�� �$ �� �" ���!��&� �% �" &�)�!���! �� ���' ���� �� � �&���' '��+�" ��� %��� � �� �" !��� � ����� �"�� �� �" %��� � �+�# 4"�&!� �" '���" �% >�'# * ) �&� ������ � �� �"�� �&)!��" � )� � ����!� ��))� , 55=� >�'# �%�� �"�� ��� � !������ �� +�!! ��� �� �"�� �" �!�� � �% �" �� � �� ���� �� ��'��%�����!� �"�!!�+ � ���c�� ! �� �� � �"�� �"�� �% �" �� �� � � �!� +��" )� �" ��� � �&�"���# �� �&������ �" &�)�!��&�� ������ � �� �" ��9 �% �" )��� +"��!� �� ��������!� ���!! � �� ���!���" ��� ����� " ������� ��� #������ �"�� �� �" ��" � �!��� � �� �� � ������� �" ��� %������� �������� �" �� ��� � ��� ���� �� *����' **' ��� � �"����� ��������� */ standard 95% confidence deviation interval natica virguloides 141° 8 125°-157° cochlis depressofuniculata 128° 7 114°-142° epigloafuniculata 126° 7 112°-140° fredianii 121° 10 101°-141° fulgurata 128° 9 110°-146° propinqua 125° 7 111°-139° raropunctata 125° 9 107°-143° obliquicallosa 134° 10 114°-154° strictiumbilicata 115° 5 105°-125° sulcogradata 130° 7 116°-144° undata 129° 6 117°-141° plicatula 122° 8 106°-138° pseudoepiglottina 116° 8 100°-132° vittata 117° 8 101°-133° tanea koeneni 120° 5 110°-130° tectonatica astensis 110° 18 74°-146° prietoi 113° 6 101°-125° tectula 113° 7 99°-127° mean ��)# ? ( 4��� ��'! �% �� �� � ��$�!$ � �� �" ����������! ���!���� ,� ����!� ��))� 55=: �� � �� ��� �-# 0 1 2 3 4 5 6 7 8 9 0 2 4 6 8 10 12 aperture height (mm) ap er tu re w id th (m m ) >�'# = ( � !������"��� ) �+ � �� �( �&� +���" ��� �� ��&� " �'"� ,�� �� � �% ���������$ �� ��� � ����������-: ���( )�!� �� �� >�'# 5# �% �������� �&�" �� �� ���!���" ��� ����# �" �!�� �% &�)�!��&� +���" �'����� �����&� ���� � � %�� �����$ ������ �� �� � ��� "����� ,��� �!!&����� � " � �"�+� �"�� �" ����� �� �% �������� ��� � ����� �� ��� ��'��%�����!� ��%% � �� %��� �� ����" � �� �� �"�� �% "����� ��� ���� ��# �" ���)�� � � '� ����� �% �" %��� � �+� ���� ��� �"�� �!�� ���)�� � �% "����� ��� ���� �� ,>�'# 0-� "�$��' ���" � ����!�� �!�� �� ��%% � ��'��%�����!� �� ! $�( ����: %�� � '�$ � ��9 �% �" )��� +"��!� �" �" !!� �% �������� ��� � ����� "�$ �" &�)�!��&� ��������!� ���!! � �"�� �"�� �% "����� ��� ���� ��# �" ���� !����� ) �+ � +���" �% �" &�)�!���! ��!!&� ��� +���" �% �" &�)�!��&� � �&!� � �� ) ��'��%�( ����!� "�'" �� ���!���" ��� ���� ,� e 5#**-� ���!! ��'��%�( ���� )&� !�+ � �� " ������� ,� e 5#3 ��� #������ ,� e 5#; ���� �� �" !��� � �+� ���� �" +���" �% �" &�)�( !���! �� ���' �� �&�" ��� $����)! �"�� �" ��� �'�" �% �" %&���! # �� � '���� �" %�&� �� �� � �� �" �������$ ����c� ���������� '��&�� �" ���� !����� �� $ � ��� ��'��%����� ,� e 5#0/( -# �" �!�� �% +���" �% �" &�)�( !���! ��!!&� �'����� +���" �% �" &�)�!��&� ,>�'# 5�"�+� �"�� #������ ��� ) � ���!� ������'&��" � %��� �" ��" � �+� �� �� � )� �" ��'��%�����!� '� �� � ! $�( ����: %�� � '�$ � &�)�!���! ���!��&� � �" &�)�!���! ��!( !&� �% #������ �� ��'��%�����!� +�� �# �� � '���� ���!���$ " ��� ���� ��� " �������� �" � �� ���$ � '� ����� !�� � �"�)�� ��'��%�����!� ��%% � �� �!�� �: �" &�)�!���! ��!!&� �% �" %���� �� �� � +�� �� ���2 �!� %��� � �"�� �"�� �% �" � ���� �� �" &�)�!��&� �!��' �# 4"�&!� �" ����� �� �% >�'# 5 ) �&� ������ � �� �"�� �� � �� � )� � �( ���!� ��))� , 55=� � ��(%�'# 3 �%�� �" ��� ���� �% �"����� ��� �� +�!! ) � � �"�� #������ �� ��� ��'��%�( %�� ���� &' ( ����� )'** 0 2 4 6 8 10 12 0 2 4 6 8 10 12 14 16 18 shell heigth (mm) ap er tu re h ei gh t ( m m ) >�'# ; ( � !������"�� ) �+ � �� �( �&� " �'"� ��� �" !! " �'"� ,�� �� � �% ����������� ��� � ����������-: ���)�!� �� �� >�'# 5# 0 1 2 3 4 5 6 7 8 9 0 2 4 6 8 10 12 14 16 18 maximum diameter (mm) ap er tu re w id th (m m ) >�'# / ( � !������"�� ) �+ � �� �( �&� +���" ��� �����&� ���� � � ,�� �� � �% �����$ ������ ��� � ����������-: ���)�!� �� �� >�'# 5# ����!� ��%% � �� %��� ������� �� ���� ��������� ,4����� *0 -� �" !�� %�� ���!���" ��� ���� )�����!!� ���( %���� �� �"�� �% ������� ������ �� ,������ *3 -� ��� 3" ������� ��%% �� %��� �!! �" ��" � ���� )� ��� ���2( �!� ! �� �� � �!�� # �" ������ � �� � ����' �" �" !!� �% ��������� "������ � ����� ��� ���� �� %��� !��'�� ����( � �� �"�+��' � �!�� ����!����� ) �+ � �������� ��� � ����� �� + !! �� ) �+ � "����� ��� ���� �� ,>�'# -# �" %��� � �� �� � ���� � ��� �� "�$ �" &�)�!���! ��!!&� �!�'"�!� ���!! � �"�� �"�� �% "����� ��� ���� ��� �"�� ������� �" ��� %������� �������� �" �� ��� � ��� ���� �� *����' **' ��� � �"����� ��������� *0 standard 95% confidence deviation interval natica virguloides 22° 2 18°-26° cochlis depressofuniculata 30° 6 18°-42° epigloafuniculata 24° 5 14°-34° fredianii 22° 5 12°-32° fulgurata 25° 5 15°-35° propinqua 30° 6 18°-42° raropunctata 27° 6 15°-39° obliquicallosa 25° 6 13°-37° strictiumbilicata 26° 4 18°-34° sulcogradata 26° 5 16°-36° undata 25° 7 11°-39° plicatula 28° 7 14°-42° pseudoepiglottina 33° 6 21°-45° vittata 30° 7 16°-44° tanea koeneni 18° 4 10°-26° tectonatica astensis 25° 5 15°-35° prietoi 22° 3 16°-28° tectula 24° 7 10°-38° mean ��)# = ( 4!�� �% ��� � !�� �% �� �� � ��$�!$ � �� �" ����������! ���!���� ,� ����!� ��))� 55=: �� � �� ��� �-# natica virguloides very thick, overlapping the basal fasciole; anterior lobe well developed, tongue-shaped, obscuring the adapical part of the umbilicus. cochlis depressofuniculata moderately thick to thick, subquadrangular, ending some distance from the basal fasciole; anterior lobe indistinct. epigloafuniculata thick, subquadrangular, reaching the basal fasciole; anterior lobe absent. fredianii moderately thin, slightly narrowing abapically, ending near the basal fasciole but not touching it; anterior lobe indistinct. fulgurata thick, short and broad, ending some distance from the basal fasciole; anterior lobe indistinct. obliquicallosa thick, slightly narrowing abapically, ending some distance from the basal fasciole; anterior lobe indistinct. plicatula quadrangular, rather wide and thin, nearly reaching the basal fasciole; anterior lobe indistinct. propinqua rather thick, subquadrangular, never reaching the basal fasciole; anterior lobe indistinct. pseudoepiglottina moderately thick, narrowing abapically, ending close to but not in touch with the basal fasciole; anterior lobe indistinct. raropunctata moderately thick, slightly narrowing abapically, ending near the basal fasciole but not touching it; anterior lobe very small to indistinct. strictiumbilicata thick, narrowing abapically, ending close to but not reaching the basal fasciole; anterior lobe indistinct. sulcogradata rather thin and short, ending some distance from the basal fasciole; anterior lobe indistinct. undata rather thick, subquadrangular, ending some distance from the basal fasciole; anterior lobe indistinct. vittata thin to moderately thick, ending some distance from the basal fasciole; anterior lobe small, pointed. tanea koeneni quadrangular, rather wide and thick, ending some distance from the basal fasciole; anterior lobe absent. tectonatica astensis quadrangular, rather wide and thick, ending at the level of the basal fasciole; anterior lobe small, subrounded, or indistinct. prietoi subrectangular, moderately wide and rather thin, ending at the level of the basal fasciole; anterior lobe absent. tectula subquadrate, rather thick; anterior lobe absent. ��)# ; ( > ��&� � �% ���� ��! ��!!&� �% �" ������� ���� %�&�� �� ���&� �� �" �!��� � ,� ����!� ��))� 55=: �� � �� ��� �-# 0 5 10 15 20 25 5 10 15 20 25 30 maximum diameter (mm) um bi lic us w id th (m m ) >�'# * ( � !������"��� ) �+ � &�)�!��&� +���" ��� �����&� ���� � � ,�� �� � �% ������� ��� ������ �"�� ��&��-: �� � ����! �� ���!���" ��� ����: �� � �����'! �� " �������: �� � �6&�� �� #������# >�'# 0 ( � !������"��� ) �+ � &�( )�!��&� +���" ��� �����&� ���� � � ,�� �� � �% �����$ ������ ��� � ����������-: �� � ����! �� "����� ��� ���$ � �� ��������� ����� �� ����� �������� ��� � ����� ���( )�� �# 0 1 2 3 4 5 6 7 0 2 4 6 8 10 12 14 16 18 maximum diameter (mm) um bi lic us w id th (m m ) 0 1 2 3 4 5 6 7 0 2 4 6 8 10 umbilicus width (mm) w id th o f t he u m bi lic al c al lu s (m m ) >�'# 5 ( � !������"�� ) �+ � �" +���" �% �" &�)�!���! ��!!&� ��� �" &�)�!��&� +���" ,�� �� � �% ������� ��� ������ �"�� ��&��-: ���)�!� �� �� >�'# *# %�� ���� &' ( ����� )'05 0 1 2 3 4 5 6 7 0 1 2 3 4 5 6 7 umbilicus width (mm) w id th o f t he u m bi lic al c al lu s (m m ) >�'# ( � !������"�� ) �+ � �" +���" �% �" &�)�!���! 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%�����! �% �" ������� ���� %�&�� �� ���&� �� �" �!��� � ,� ����!� ��))� 55=: �� � �� ��� �-# !��'��&����! %&���+�: �"�� �% ���� �� )� �" $ �� %�� �� %���� ��� � '���$ # ��������� ������� �� ��� ���������� �� ��� �������� ��������� �"�� �"��� � � ����� �!! �" �2 ! ��! �"����� �� � �!� +��" �� �" �� $��&� �� �� ��� � �� ����� �&� +" �" � ��� +" � �" � �� ��'��%����� �� �� �� � � ��'( ������� +��" � % � �� �� �" �'"� � ���� �% �" ����( ���� �)���� � %��� �" ��$ ���'�� � �!��� � � ������ ,� ����!� ��))� 55=: �� � �� ��&��-# >�� ���$ ��( �� �% �" � �� �� + � ��!! �"�� �" ���� ��$�!$ � �� �"�� ���!���� �� � ������ �� ! ������ 4����� *05� ������� ��� ����" ��� ���� ,4����� *0 -� �# ���!���" ��� ���� ,4����� *05-� �# " ������� ��# �#� �# " �! ��� ,1 � ( '"���� �� � ��"��!�� *;?-� �# ������ �� ,������ *3 -� �# � ����+ � ,� ��"��!�� *;?-� �# ��� �����!������� ,4��( ��� *05-� �# � �� ������ � �� ������ ,4����� * /-� �# � �� ������ ����+ �������� � ����!� ��))�� 55=� �# �� ���� ��������� ,4����� *0 -� �# � ���! ����� � ����!� ��))�� 55=� �# ����� ,4����� * /-� �# ������� ,�� !��� /0 -� ����� �������� #������ ,4����� *0 -� %�� ���� &' ( ����� )'0 background color pattern natica virguloides not observable 14. reddish brown dots and 2 spiral rows of large, approximated, irregular spots cochlis depressofuniculata light brown reddish pattern which may consist of: 1. uneven, irregularly arranged spots 2. spirally elongated oval spots 3. triangular spots and/or chevron marks which may fuse to form collabral stripes epigloafuniculata light brown reddish pattern of: 15. uneven, subquadrangular spots, irregularly arranged into spiral rows fredianii light brown reddish pattern of: 10. moderately large, even spots irregularly arranged into collabral rows fulgurata pale yellowish brown reddish pattern which may consist of: 4. usually crowded, small and even dots often arranged in collabral rows 5. axial zigzag lines or stripes 6. uneven, variously sized dots and spots arranged in irregular collabral rows 7. spots replaced by chevron markings during growth plicatula light gray reddish brown pattern of 4 spiral rows of squarish or chevron spots propinqua light brown reddish pattern of: 8. dense, undulating, collabral lines or stripes; spots may replace lines in later growth stages (rare occurrence) pseudoepiglottina uniform pale brown to pinkish brown without any color pattern pale reddish brown red pattern which may consist of: 4. small and even dots roughly arranged in collabral rows 9. crowded, uneven and irregularly arranged small dots which may fuse adapically to form subsutural chevron markings 1. uneven, variously sized and irregularly arranged spots raropunctata brown to light brown reddish brown pattern which may consist of: 4. usually crowded, small and even dots often arranged in collabral rows 10. large, even spots irregularly arranged in collabral rows 1. uneven, irregularly arranged, large spots 11. spirally elongated spots 3. spots and chevron marks occasionally fused to form collabral stripes 12. irregular spiral broken lines obliquicallosa pale brown 4. usually crowded, small and even dots often arranged in collabral rows 1. uneven, irregularly arranged spots 11. spirally elongated spots 12. irregular spiral broken lines strictiumbilicata pale brown reddish pattern of: 1. uneven, irregularly arranged spots sulcogradata light brown reddish brown pattern which may consist of: 4. usually crowded, small and even dots often arranged in collabral rows 10. large, even spots irregularly arranged in collabral rows 11. spirally elongated spots undata pale brown reddish pattern which may consist of: 8. gently undulating collabral lines occasionally interrupted or partially replaced by variously shaped spots 13. axially elongated, oval spots arranged in collabral rows vittata pinkish white 16. reddish brown irregular reticulated pattern of interconnected polygons and 2-3 spiral rows of brown spots; a subsutural dark band is also present tanea koeneni uniform pale brown apparently without any color pattern tectonatica astensis pale brown; protoconch light grey 17. dark brown subsutural band prietoi mottled pale brown 18. alternating brown and whitish spots forming 3 weak spiral rows tectula pale brown 19. reddish brown, undulating collabral lines associated with a dark brown subsutural band ��)# * ( ���2'��&�� ��!�� ��� ��!�� ���� ��� �% �" ������� ���� %�&�� �� ���&� �� �" �!��� � ,� ����!� ��))� 55=: �� � �� ��� �-# ����������� �������� ,4����� *05-� �# � ����� ,@���!'�� */3��� �# ���� �� ,4����� *05-# # % ��������# >��� ��)# 5 �� +�!! 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m ar gi na l a re a ho ri zo nt al ra th er w id e, m od er at el y de ep , s om et im es ob so le sc en t, m ay b ea r lo ng itu di an l r ow s of gr an ul es us ua lly n ar ro w er a nd le ss e xc av at ed , so m et im es a tte nu at ed or o bs ol et e sh ar p to ro un dto pp ed us ua lly w el l d ev el op ed , th in to th ic k, w ith s ha rp to fl at to p, s el do m ob so le te ob liq ui ca llo sa op er cu lu m in di st in gu is ha bl e fr om th at o f r ar op un ct at a fr ed ia ni i ra th er s ho rt 2 gr oo ve s, 2 ri dg es ; m ar gi na l a re a sl op in g in w ar d w id e an d sh al lo w a fa in t d ep re ss io n as ym m et ri ca l i n cr os s se ct io n sh ar p an d m od er at el y pr om in en t po or ly d ev el op ed pr op in qu a br oa d 2 gr oo ve s, 2 ri dg es ; m ar gi na l a re a be ar in g ou te r gr oo ve a nd r id ge s lo pi ng ou tw ar d m od er at el y ex ca va te d, an gu la r i n cr os s se ct io n na rr ow er a nd s ha llo w er , at te nu at ed to o bs ol et e in la rg er s pe ci m en s sh ar p sh ar pl y or ro un dl y ed ge d, re cl in at e to w ar d th e in ne r g ro ov e ������� �" ��� %������� �������� �" �� ��� � ��� ���� �� *����' **' ��� � �"����� ��������� 0= de pr es so fu ni cu la ta el on ga te 2 gr oo ve s, 2 ri dg es ; 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number of whorls diameter diameter of the initial half whorl natica 1. virguloides 7, 9, 15, 16, 17, 18 5, 7, 12, 13, 14, 15, 16, 17, 18 3, 4, 5, 7, 8, 9, 12, 14, 15, 16, 17, 18 cochlis 2. depressofuniculata 7, 9, 13, 14, 15, 16, 17, 18 3, 7, 11, 12, 13, 15, 16, 17, 18 3, 4, 5, 7, 8, 9, 11, 12, 14, 15, 16, 17, 18 3. epigloafuniculata 7, 9, 14, 16, 17, 18 2, 4, 5, 7, 8, 10, 14, 15, 16, 18 1, 2, 6, 7, 8, 9, 10, 11, 12, 13, 14, 16, 17, 18 4. fredianii 7, 9, 13, 14, 15, 16, 17, 18 3, 7, 12, 13, 15, 16, 17, 18 1, 2, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18 5. f ulgurata 7, 9, 13, 14, 15, 16, 17, 18 1, 3, 6, 7, 9, 11, 12, 13, 15, 16, 17, 18 1, 2, 4, 6, 7, 8, 9, 10, 11, 12, 13, 14, 16, 17, 18 6. obliquicallosa 7, 9, 13, 14, 15, 16, 17, 18 5, 7, 12, 13, 14, 15, 16, 17, 18 3, 4, 5, 7, 9, 12, 14, 15, 16, 17, 18 7. plicatula 1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 15, 17 1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 17 1, 2, 3, 4, 5, 6, 8, 10, 11, 12, 13, 15 8. propinqua 7, 9, 13, 14, 15, 16, 17, 18 3, 7, 12, 13, 15, 16, 17, 18 1, 2, 3, 4, 5, 7, 9, 13, 14, 15, 16, 17, 18 9. pseudoepiglottina 1, 2, 3, 4, 5, 6, 7, 8, 10, 11, 12, 16, 18 5, 7, 13, 14, 15, 16, 17, 18 1, 2, 3, 4, 5, 6, 8, 10, 11, 12, 13, 14, 15 10. raropunctata 7, 9, 13, 14, 15, 16, 17, 18 3, 7, 12, 13, 15, 16, 17, 18 3, 4, 5, 7, 9, 12, 14, 15, 16, 17, 18 11. strictiumbilicata 7, 9, 13, 14, 15, 16, 17, 18 2, 5, 7, 14, 15, 16, 17, 18 2, 3, 4, 5, 7, 9, 14, 15, 16, 17, 18 12. sulcogradata 7, 9, 13, 14, 15, 16, 17, 18 1, 2, 4, 5, 6, 7, 8, 10, 14, 15, 16, 18 1, 2, 3, 4, 5, 6, 7, 9, 10, 13, 14, 15, 16, 17, 18 13. undata 2, 4, 5, 6, 7, 8, 10, 11, 12, 14, 16, 17, 18 1, 2, 4, 5, 6, 7, 8, 9, 10, 14, 15, 16, 18 3, 4, 5, 7, 8, 9, 12, 14, 15, 16, 17, 18 14. vittata 1, 2, 3, 4, 5, 6, 7, 8, 10, 11, 12, 13, 16, 18 1, 3, 6, 7, 9, 11, 12, 13, 15, 16, 17, 18 1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 13, 15, 17 tanea 15. koeneni 1, 2, 4, 5, 6, 7, 8, 10, 11, 12, 16, 17, 18 1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 17 1, 2, 4, 6, 7, 8, 9, 10, 11, 12, 13, 14, 16, 17, 18 tectonatica 16. astensis 1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 15 1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 17 1, 2, 3, 4, 5, 6, 8, 10, 11, 12, 13, 15 17. prietoi 1, 2, 3, 4, 5, 6, 7, 8, 10, 11, 12, 13, 15 1, 2, 4, 5, 6, 7, 8, 9, 10, 11, 14, 15, 16, 18 1, 2, 3, 4, 5, 6, 8, 10, 11, 12, 13, 14, 15 18. tectula 1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 15 1, 2, 3, 4, 5, 6, 8, 9, 10, 11, 12, 13, 14, 17 1, 2, 3, 4, 5, 6, 8, 10, 11, 12, 13, 15 ��)# 5 ( 4&�����9 � ��%% � �� � �� ���������" � ��&� � ���# >�'&� � � % � �� �� �� � ,! %� �% %���� ��!&����� � ��� �"�� �� �� � +"��" �� ������'&��" � %��� �"�� �� �" ! %� �% ��" ��+� �� �" )���� �% �" ������ � � ���������" � ��&� � ��� ,��!&��� �"��&'" ?-# �!! �" ��&�� � �!��� � �������� ,� ����!� ��))� 55=: �� � �� ��� ��� !��� �# >�$ �� �� �� �# # ��� ����" ��� ����� � ����+ �� � �$ � ������� �� ���� ��������� ��� � ���! ����� !��2 �������( ��$ �" !! �"����� �� ��� �� � 6&�� �� ��&!�� �� ��( � �� �� ��� � �� ) ���%�� ��!� �� ���%� �# �� ��� �� �� ) ��� �� ��)# +" � �" �� ��&!&� �% #��$ ���� �� ��� ������� � �� ���'������ ���� �� �� ����2��'!� ����!�� �� �"�� �% �������� ,� �)�$ -: "�+ $ �� �" �� ��&!&� ������'&��" � #������ %��� �!! �" ��" � �!��� � ������� �� �� � !��� � �� ��)# # ������� �������������� �"�� �"��� � ���� �� ����&�� �" ' � ��� ����'�( � �� �% �" �� �� � ��$ � � �� �"�� ��� � ��� %�!!�+� �" ��� �������" ����� � )� � ����!� ��))� , 55=-� �# # �� ������ �� �" ' � ��� �!!������� �% �" �� �� � ��� )� �� $��&� �&�"���� �" ��� ��� ������ �% �" ������ � � ' � �� ���$�� � �� ��a�� ����&��� �&)!��" � �� %��� ��� 3�" ��� (�� �� � �% �" ��$�!$ � ' � �� �� ��� � �� � %�� �" �"����� �� �% �" � !��� �# �� �" %�!!�+��'� ���! � % � �� +�!! ) ��� �� ��&�" � d�� � , 003+"�� �� �" %��� �% � � $����� �% � � �� ����" ��� ��!����� )��"��! ��� �)����! 1 ��'���������� �"���&'"!� �� �� � �" ��������� �� ��� ��a�� � $������ ���� ����' )��" ��� �� ��� � ��9��� �������� ,g�!( )&�� 0/;: 1�����"�$��" 0//: 1�a��� 0*0-� ��� �� g�)�� , 00 +"� � $� + � �" ' �&� ! $ ! ��� � �% �" �������� # �" ��� ���� ��&�� � " � �� �"�)�� 6&�� ��%% � �� ��&!��&��! % ��&� � �% �" �&� � �&�%�� �% �" �� ��&!&� ���� �� �"�� )����� ��� ) ������)&� � ���� �"� !���# ��� ���!&� � ���!���" ��� ���� ��� " ������� �"�� "�$ �" �� ��&!&� ��&!��&� � +��" ���'���! 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� �������"#�� ��$����% �� # �� �� �� ����� �� & �� � ����������� ��� � ���' ()-. !� �,� # ��� ���� ��� ���# ,1��� �4#5 0#5 #53*c53-# ����� � ���! � �/ �!��%# �&����� k ��"��� ����# ,���$�� ��!! ������-: ������� ������� ?* ����# ,1��1 0?//-� ? ����# ,1��1 0?/*(0?* -� 3 ����# ,�� 0=;?-� ����# ,1�� =/=-� = ����# ,���$�� ��!! �����-# � � ����# % ��������� ���!!� � �� �� � �&�)���%���� �% #=( #/= �!�'"�!� ���$ �� �����" +"��!�� ��� ���!!# -����� '!�)�� � ��� ��� !� � �� �� �� ��!��� "���!� �� ���' = �� �� " �'"�# -�� �� !�+(������!� ���" � � �� �� �� +"��!� ��� ��� !� ���$ �# � �� %�� � ����� �� ���� �� �# /��� ��� �� ��%!�� �� ��� +"�� � �� �� �� ��� ��� !� ���( �&� � ��� ����� � ��+��� �" �� ��&� � +��" �������� �&)�&�&��! �" !%# �� � �� �(�"�� �� " �'"� �$ ��'��' ��� � �" +���"# %� ����� ���� �� �"��2� �&)6&�����'&!��� � ��"��' �" )���! %��( ���! : ��� ���� !�) �)� ��# .������ �� ���" � ���!!# � ������ � )����� $ �� !�+ ���� � ����� � %��� �" )���! %�����! )� � +�� ��� �"�!!�+ � �� �����# .�������� ���� �� ���2 �!� � �� �� � ��� �"��2� !���� � �� �" ����! �% �" ��!&� !!�� !��� +��" �!�'"�!� ���&�&� �&�!�� �)!�6& !� �� �� � �������!!� �� � �' ���� �" ���� ��! ��!!&�# /���� "�������� ��� ��� !� +�� � ���2 �!� � �� �� �� � %�� � �)����!!� )� �" �&�� � � $������ �% '��+�" !�� �# "���� +��" � �� ��� %�� � ' ��!� ������!�� '��+�" !�� �� �!�'"�!� �����' � �� �&)�&�&��! �" !% ��� )���! %�����! : � %���� �����! ��������� �� ��� � �� �" )��� +"��!# ���� � )��2'��&�� !�'"�()��+� +��" � ����" ���� �� �% &� $ �� �&)6&�����'&!�� ������ ��� '&!��!� �����' � ���� �����! ��+�# 0�� � � �� ������� �" ��� %������� �������� �" �� ��� � ��� ���� �� *����' **' ��� � �"����� ��������� 00 %�� ���� &' ( ����� )' 55 species reference umbilical callus operculum remarks natica (cryptonatica) clausa broderip & sowerby, 1829 marincovich, 1977 cryptonatica clausa (broderip & sowerby, 1829) majima, 1989 cryptonatica affinis (gmelin, 1791) = natica clausa broderip & sowerby, 1829 bouchet & waren, 1993 cryptonatica clausa (broderip & sowerby, 1829) saito in okutani, 2000 large, semicircular, completely filling the umbilicus or separated from the umbilical wall by a narrow groove smooth type-species of cryptonatica natica (tectonatica) bougei sowerby, 1908 cernohorsky, 1972 large, semicircular, completely filling the umbilicus smooth tectonatica filosa (philippi, 1845) bouchet & waren, 1993 large, semicircular, separated from the umbilical wall by a narrow groove, slightly wider abapically smooth cryptonatica ichishiana (shibata, 1970) majima, 1989 large, semicircular, completely filling the umbilicus smooth natica (tectonatica) janthostoma deshayes, 1839 marincovich, 1977 cryptonatica janthostoma (deshayes, 1839) majima, 1989 cryptonatica janthostoma (deshayes, 1839) saito in okutani, 2000 large, semicircular, separated from the umbilical wall by a groove of variable breadth, wider adapically smooth or with weak marginal striations natica (cryptonatica) pusilla say, 1822 abbott, 1954 tectonatica pusilla say, 1822 warmke & abbott, 1961 large, semicircular, separated from the umbilical wall by a narrow groove smooth cryptonatica ranzii (kuroda, 1961) saito in okutani, 2000 large, semicircular, completely filling the umbilicus smooth natica (tectonatica) robillardi sowerby, 1893 cernohorsky, 1972 large, semicircular, completely filling the umbilicus smooth natica (tectonatica) tecta anton, 1839 kilburn, 1976 large, semicircular, completely filling the umbilicus smooth cryptonatica wakkanaiensis habe & ito, 1984 saito in okutani, 2000 large, semicircular, completely filling the umbilicus smooth cryptonatica zenryumaruae habe & ito, 1976 saito in okutani, 2000 large, semicircular, completely filling the umbilicus smooth tectonatica tectula (sacco, 1891) pavia, 1980; our specimens large, semicircular, separated from the umbilical wall by a narrow to moderate groove in the mid-abapical part 1 fine, shallow groove bounding the smooth marginal area type-species of tectonatica tectonatica astensis (sacco, 1891) pavia, 1980; our specimens rather large, subtriangular, separated from the umbilical wall by a moderate to wide groove in the mid-abapical part 1 fine, shallow groove bounding the smooth marginal area cryptonatica figurata (sowerby, 1914) saito in okutani, 2000 large, semicircular, separated from the umbilical wall by a narrow groove 1 fine, shallow groove bounding the smooth marginal area cryptonatica hirasei (pilsbry, 1905) saito in okutani, 2000 large, semicircular, completely filling the umbilicus 1 fine, shallow groove bounding the smooth marginal area cryptonatica operculata (jeffreys, 1885) bouchet & waren, 1993 large, semicircular, separated from the umbilical wall by a narrow groove 1 fine, shallow groove bounding the smooth marginal area � ���" � �"��: � � ����! ��!!&� ��������� ��� ��� !� !��'�� � ���'& (�"�� �: � ��� � ���'�� � ��!� �����'"�� +��" )!&�� '���&! �: � ��� � �&�%�� %!��� �&�! &� ��� �����&���': � �&� � �&�%�� �!�'"�!� �����$ � +��" ���'���! %&���+� ��� ��)�: �&� � %&���+ ��� ��)� �� � ��������!� ! $�� �� "���9����! �" !%: � �&� � '���$ ���" � �����+ ��� ��� ��� !� � �� ) ����' '���&! � �"�� � �� �� %��� �)!�6& ��+�: � ��� � '���$ $ �� +�� � ���" � � �� +��" �!�'"�!� �����$ )�����: � �&� � ���' �"�� ��� ������ ��� �"���( �' �: � ��� � ���' �"��2� �!�'"�!� ) �� ��+��� �" ��� � %&���+� %!��( ���� �� �� +�� �� �� "�!% �" ��� � '���$ # �!���#!��# 0��1� �@d �� @ � 4@ �@ �d 5# 0(5# 3 5#/;(5#*? *#33( /#33 /#;5( ;#5? 5#;5(3#3 /# ;( ?#?; 3#=;(/#0 5# 5#*5 #*3 #* #0; 5#*; =#/? �d d�� d�� d�� �4 4� # /(3#*3 5#;5( # 5 5# /( # 3 5#5;(5#0? ?f(3?f f( ?5f #== #?5 5#;= 5#=5 ?f ;f ��� �2## �" �� � �� ����� +�� ���'���!!� ���( ��� � ,4���� *05�� � $��� �� �% ������ ������ ������ ������2� * # �� �" )���� �% �" ���������$ �� ��&!�� % ��&� �� �� ��� ��� �"�� �� �� ����% ��!� &�� !�� � �� ������ ������� ��� �� �� ��$���)! �� �� �� ���!���" ��$ � ���� �� � �������� �� �� � ) !��'��' �� �" ' �&� ��$ ����� ��.���'� /0*# 4���� , *0 �"���!� � ����) � ��� ��� � ������ ���!������� $��# �3" ��� ���� �"�� &���( )�'&�&�!� ��� ��� �� "�$ �&)�� ��%�� ���2 %��� ��� ���( '���! �&)!������� ,��l� 000� ���# ?=#;-# d "�$ �( ���� � 4����8� ��� ���! �� 1��� ��� + � ��� �)! �� %��� �&� ��� ��'��%����� ��%% � �� �� � �� �� �� ������� ���!���" ��� ����# ��������'!�� + ���!&� �3" ��� ���� �� �" �������� �% �" �� � �� �� �� �# �� �� �% ��� �"�� �" ��� ���!���" ��� ���� ) ��� �������� "�$��' ) � $�!��!� ������ � �� � �� ��!� �# ����' �" �!��� � ������� �� �� � �% ���!� � ( ����) � �� %�� ,� ����!� ��))� 55=: �"�� ��� �-� ��$ ����� ��� ����" ��� ���� ,4���� *0 �� �" ��!� �� +��" +"��" ������� ���!���" ��� ���� ��� ) ������ � ������� �" ��� %������� �������� �" �� ��� � ��� ���� �� *����' **' ��� � �"����� ��������� 5 tectonatica prietoi (hidalgo, 1873) our specimens moderate to small, separated from the umbilical wall by a wide groove 1 fine, shallow groove bounding the marginal area; marginal area with 2-3 fine grooves tectonatica rizzae (philippi, 1844) bouchet & waren, 1993 moderate, subtriangular, separated from the umbilical wall by a rather wide groove in the midabapical part 1 fine, shallow groove bounding the smooth marginal area cryptonatica adamsiana (dunker, 1859) majima, 1989 small to large, nearly semicircular, separated from the umbilical wall by a moderate to broad groove with 2 well developed marginal grooves neither cryptonatica nor tectonatica; according to the fugure of the operculum, probably tanea cryptonatica bathybii (friele, 1879) bouchet & waren, 1993 poorly developed undescribed umbilical characters not consistent with neither cryptonatica nor tectonatica natica (tectonatica) janthomostoides (kuroda & habe, 1949) marincovich, 1977 cryptonatica andoi (nomura, 1935) = tectonatica janthomostoides kuroda & habe, 1949 majima, 1989; saito in okutani, 2000 large, semicircular, usually separated from the umbilical wall by a groove of variable breadth with 2 well developed marginal grooves neither cryptonatica nor tectonatica; according to the fugure of the operculum, probably tanea natica (cryptonatica) oregonensis (conrad, 1865) marincovich, 1977 large, semicircular, separated from the umbilical wall by a narrow groove with 1 low marginal ridge probably neither cryptonatica nor tectonatica natica (tectonatica) simplex sowerby, 1897 kilburn, 1976 large, semicircular, usually separated from the umbilical wall by a narrow groove at least abapically, completely filling the umbilicus in some shells with numerous, fine marginal threads figure of the operculum needed ��)# ( ��)�!���! ��!!&� ,�"�� � ��'���&� ��� �� ��&!�� % ��&� � �% �� �� � ����'� � �� � ���������� �� ����������� )� � ! �� � �&�"���# ��!� + !! � ���)! �!!&��������� ���c�� �� ��� � ���������� + � ������ � �# species reference umbilical callus operculum remarks �� ����&�� �% � ' � ��! �" !! ����!�����# ������� ���!���$ " ��� ���� ��%% �� %��� ������� ��� ����" ��� ���� �� "�$��'� ���������" +��" ��'��%�����!� '� �� � ���( � � � , ;b ��%% � �� ��� ���!! � ���� � � �% �" %���� "�!% +"��! ,3?b ��%% � �� -: ���!! � � ! �����" ��9 : 3���� ��! ��!!&� ��� +"�� ��� !��'�� : ?���!! � &�)�!��&�: =&�)�!���! ��!!&� � �'��' ���� �" ���� ��! ��!!&� +��"�&� ��� ����" �� ) �+ �: ;�� ��&!&� ��( ��!��!� ���&��&� �� )&� +��" � � � �&� � '���$ � �&�" +�� � ��� � '���$ � ��� ���2 �!� �"��2 � ��� � ���' # �" �� ��&!&� ������ �� �" ���� � ! $��� �"����� � ��� ������'&��" � ������� ���!���" ��� ���� %��� �" ��" � ������� �� �� �# ��� �!3� ��!� ���"������# �' ���!���" ��� ���� ��� ��� �� ���&� %�� �" %���� ��� �� �" ��������� �% �� �����# �!��� � � ����� �� %��� l���! 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"#� #��) k �& ��)"� "� ��, �55c1, ����) � " �$�##" ����"� � " �:(��#��� !'&&"!!��� ! �2! �"�� +����� #"�"��+"# ��#)��'�#! !'))"!���) � �� � �! ��"� ��b"�� �"+�"!"��"# � " #"+�&"���" �( � " -�!��, �� � " &������� �� =�!�#� ��# ���'�# ����!&��� � " ��#)��'�#! ��" -"��"� #"�"�� !�"�� �� � �����&����" ����� ���( ��'� �& �'� ������ �� ������ )*�# �"��� + ���� , "&c7� 1p. cf. enigma� 2 schizosphaerella sp.� 8 f. multicolumnatus�7 f. multicolumnatus� 3 schizosphaerella sp.� 9 c. mexicana minor� 4 m. quadratus� 5 m. quadratus� 6 m. quadratus� = ��� � �1 2# &(, ���( � ��?* �1 ��'�1���'�� �""� !+, �7a* .1 ��'�1���'�� �""� !+, �5�* c1 *# 3 �� �� � �ac* 71 *# 3 �� �� � �7a* ?1 *# 3 �� �� � /��# � (��)%"�� �( ������� � �� ��2"� ��) �1 �7�* a1 �# "����" ��� � /(��)%"��1 �5�* 61 �# "����" ��� � /(��)%"��1 �67* 51 !# �4����� ��� ����, �&��" -�� �"+�"!"��! 7 �%, !���""��� !#$#� ���� " �# % ���& ���'� �#c7� s'adde section lithology thickness environmental interpretation lithostratigraphy (previous authors) lithostratigraphy (this study) age dorgali fm. dolomitic fine to medium quartzarenite, fe-stained ooids & coated grain dolomitic grainstone in metre thick, often amalgamated beds. 25m coastal highenergy shoal, locally lagoon and beach dorgali dolostone dorgali fm. late bathonian condensed succession (s'adde lithozone 1) dolomitic mudstone and peloidal wacke-packstone with dolomitic marly dolostone, rare silty sandstone interbeds. hg are both present at the base and at the top of the unit. 10m middle-outer ramp with low sedimentation rate dorgali dolostone s'adde lms. late bathonian\ early callovian s'adde lithozone 2 mudstone to crinoid-micropeloidal wacke-packstone, rare fine grained peloidal, oo-bioclastic grainstone intercalations. thickening up trend (15-40 cm to 30-70 cm at the top). 87m mainly outer ramp/ intraplatformal basin s'adde lms. s'adde lms. upper callovian to (early) kimmeridgian s'adde lithozone 3 mudstone, wackestone and peloidal bioclastic packstone with chert nodules locally ammonites. 30 to 50 cm beds, l cm-thick marly interbeds at the top. 40m outer ramp/ intraplatformal basin s'adde lms. s'adde lms. (late) kimmeridgian to earliest tithonian mt. bardia fm. peloidal-crinoid packstone to oobioclastic grainstone in m-thick beds, often amalgamated locally low angle clinostratified. 65m (top missing) mainly middle ramp/slope mt. bardia fm. mt. bardia fm. early tithonian siniscola section lithology thickness environmental interpretation lithostratigraphy (previous authors) lithostratigraphy (this study) age uppermost dorgali fm. bio-intraclastic, peloidal packstone with open marine benthic\planktonic microfacies. 1m (base missing) open subtidal, drowning of the dorgali platform dorgali dolostone dorgali fm. late bathonian ? condensed succession (s'adde lithozone 1) fe-hardgrounds, bio-lithoclastic packstone, with qz extraclasts and feoxides, phosphates. 1m carbonate swell, condensed sedimentation dorgali dolostone s'adde lms. early callovian ? s'adde lowermost lithozone 2 bio-intraclastic, peloidal packstone, wackestone with open marine benthic\planktonic microfacies. neptunian dykes. 4\5m carbonate swell, low sedimentation rate s'adde limestone s'adde lms. late callovian ? posada section lithology thickness environmental interpretation lithostratigraphy (previous authors) lithostratigraphy (this study) age uppermost dorgali fm. oolitic grainstone, fe-stained at the top. 5m oolitic shoals dorgali dolostone dorgali fm. bathonian condensed succession (s'adde lithozone 1) fe-hardgrounds, bio-lithoclastic bioturbated packstones\rudstone with qz extraclasts and fe-oxides. about 13 m carbonate swell, condensed sedimentation dorgali dolostone s'adde lms. late bathonian callovian s'adde lithozone 2 nodular bio-intraclastic, peloidal packstone, wackestone with open marine benthic\planktonic microfacies, local pelagic oncoid and ammonoids. 25m (top missing) middle-outer ramp, low sedimentation rate s'adde lms. s'adde lms. oxfordiankimmeridgian ��-, � � ���� "!�! �( �$�##" %!, ��� �(�&�"! �!!�&������!, �+"# /!"" ��"�� "� ��, �5?? 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�"��� + ���� , "&c7a sample j4 element app. concentr. fit index element % sigma % atomic % point 15 na 0,5525 73,5000 1,1258 0,1270 1,9878 na2o 1,5175 mg 0,4615 2,8056 0,9248 0,0886 1,5441 mgo 1,5334 al 0,9937 0,2162 1,6439 0,0844 2,4731 al2o3 3,1059 si 1,8536 0,3077 2,4611 0,0807 3,5571 sio2 5,2650 p 1,2716 0,4762 1,1784 0,0772 1,5444 p2o5 2,7002 cl 0,0791 0,1957 0,0857 0,0523 0,0981 0,0000 k 0,2494 0,2308 0,2335 0,0547 0,2424 k2o 0,2813 ca 3,0254 0,3214 2,9774 0,0851 3,0155 cao 4,1659 ti 0,9838 0,2344 1,0305 0,0812 0,8733 tio2 1,7188 fe 39,4871 2,2125 43,0326 0,4457 31,2790 feo 55,3606 o 21,0406 0,3115 53,3852 point 16 na 0,3364 52,2059 0,6937 0,1108 1,2542 na2o 0,9351 mg 0,3143 1,2778 0,6323 0,0770 1,0809 mgo 1,0483 al 0,6231 0,1081 1,0296 0,0727 1,5860 al2o3 1,9453 si 1,5462 0,2051 2,0336 0,0735 3,0095 sio2 4,3504 p 1,4813 1,7857 1,3547 0,0747 1,8179 p2o5 3,1041 cl 0,0621 0,5870 0,0667 0,0496 0,0783 0,0000 k 0,1205 0,2115 0,1118 0,0538 0,1189 k2o 0,1347 ca 4,5457 0,4286 4,4494 0,0997 4,6142 cao 6,2255 ti 0,9323 0,3906 0,9816 0,0805 0,8518 tio2 1,6373 fe 39,8192 3,3125 43,3726 0,4457 32,2802 feo 55,7980 o 20,5194 0,3034 53,3081 point 17 na 0,4131 77,2353 0,8685 0,1289 1,5604 na2o 1,1707 mg 0,3485 2,3889 0,7146 0,0889 1,2141 mgo 1,1848 al 0,8807 0,2703 1,4802 0,0843 2,2661 al2o3 2,7968 si 1,9747 0,5641 2,6478 0,0825 3,8942 sio2 5,6644 p 0,5655 1,1667 0,5285 0,0707 0,7049 p2o5 1,2111 cl 0,0312 0,4565 0,0337 0,0530 0,0393 0,0000 k 0,1824 0,4038 0,1702 0,0547 0,1798 k2o 0,2050 ca 1,3533 0,5179 1,3206 0,0716 1,3610 cao 1,8477 ti 0,8985 0,8906 0,9184 0,0797 0,7920 tio2 1,5320 fe 43,8635 1,2125 47,3269 0,4645 35,0048 feo 60,8850 o 20,5218 0,3166 52,9836 point 18 na 0,4557 73,2353 0,9219 0,1267 1,6194 na2o 1,2426 mg 0,4085 2,0278 0,8112 0,0889 1,3476 mgo 1,3451 al 1,1864 0,1892 1,9455 0,0878 2,9120 al2o3 3,6759 si 2,5965 0,1026 3,4399 0,0886 4,9465 sio2 7,3590 p 1,0543 0,7381 0,9883 0,0775 1,2886 p2o5 2,2645 cl 0,0543 0,3261 0,0592 0,0511 0,0675 0,0000 k 0,4925 0,3269 0,4640 0,0592 0,4792 k2o 0,5589 ca 2,0835 0,4821 2,0627 0,0787 2,0785 cao 2,8861 ti 0,9178 0,2031 0,9606 0,0778 0,8100 tio2 1,6024 fe 38,5722 1,0125 42,0883 0,4415 30,4366 feo 54,1458 o 21,3977 0,3121 54,0143 point 19 na 0,4648 70,0000 0,9545 0,1203 1,6892 na2o 1,2866 mg 0,3881 2,3611 0,7808 0,0821 1,3068 mgo 1,2947 al 0,8658 0,0541 1,4337 0,0796 2,1620 al2o3 2,7088 si 1,5876 0,0513 2,1026 0,0776 3,0460 sio2 4,4981 p 1,5421 1,5238 1,4193 0,0781 1,8644 p2o5 3,2521 cl 0,0150 0,1957 0,0162 0,0516 0,0186 0,0000 k (0,0564) 0,0962 (0,0526) 0,0540 (0,0547) k2o (0,0633) ca 3,5545 0,1071 3,4821 0,0910 3,5350 cao 4,8721 ti 1,1192 0,4844 1,1715 0,0811 0,9951 tio2 1,9541 fe 40,2116 1,4875 43,7792 0,4495 31,8960 feo 56,3211 o 21,0530 0,3099 53,5415 point 20 na 0,3412 68,8824 0,7034 0,1157 1,2422 na2o 0,9482 mg 0,4079 1,7222 0,8208 0,0841 1,3707 mgo 1,3610 al 1,1970 0,1351 1,9848 0,0871 2,9863 al2o3 3,7501 si 3,1918 0,0769 4,2683 0,0910 6,1697 sio2 9,1311 p 0,2392 0,1190 0,2280 0,0659 0,2988 p2o5 0,5223 cl 0,0117 0,1522 0,0128 0,0508 0,0147 0,0000 k 0,5410 0,6154 0,5107 0,0612 0,5302 k2o 0,6152 ca 0,4162 0,2321 0,4112 0,0599 0,4165 cao 0,5753 ti 0,4827 0,2656 0,4959 0,0714 0,4203 tio2 0,8272 fe 41,5528 0,6625 45,0318 0,4531 32,7354 feo 57,9324 o 21,2080 0,3091 53,8152 point 21 na 0,2772 56,5588 0,5969 0,1130 1,1444 na2o 0,8046 mg 0,2576 1,8889 0,5374 0,0785 0,9744 mgo 0,8911 al 0,6228 0,1351 1,0593 0,0749 1,7304 al2o3 2,0014 si 1,6914 0,1795 2,2774 0,0754 3,5741 sio2 4,8721 p 0,2556 0,5476 0,2386 0,0622 0,3395 p2o5 0,5467 cl 0,0279 0,3696 0,0300 0,0508 0,0373 0,0000 k 0,1668 0,4808 0,1548 0,0546 0,1745 k2o 0,1865 ca 0,7875 0,5000 0,7630 0,0647 0,8391 cao 1,0676 ti 0,6417 1,0938 0,6465 0,0749 0,5949 tio2 1,0784 fe 45,1436 1,1500 48,3599 0,4663 38,1680 feo 62,2140 o 19,0285 0,3054 52,4234 point 22 na 0,2435 49,7647 0,5284 0,0975 1,0113 na2o 0,7123 mg 0,2373 1,0833 0,4981 0,0697 0,9013 mgo 0,8258 al 0,5714 0,0541 0,9760 0,0693 1,5916 al2o3 1,8441 si 1,6537 0,0513 2,2313 0,0725 3,4955 sio2 4,7734 p 0,1088 0,2857 0,1016 0,0596 0,1443 p2o5 0,2328 cl 0,0644 0,1739 0,0691 0,0487 0,0857 0,0000 k 0,2490 0,3654 0,2308 0,0545 0,2597 k2o 0,2780 ca 0,5223 0,4286 0,5052 0,0610 0,5546 cao 0,7069 ti 0,4493 0,2813 0,4501 0,0697 0,4134 tio2 0,7508 fe 46,8670 0,9000 50,0640 0,4740 39,4426 feo 64,4063 o 18,9449 0,3018 52,1000 compound % �������� � ��-�"! �"+��� ������� !"%�;'���������" �����!"! &��� ��"# �'� �� &��-�� &���"# � �� !"&����! /!�%+�" �c� ��# �71 (��% �0�� �( ����!&��� !"&���� /��), 50��1, �����!"! ��" -""� +"�(��%"# �� �-���� � " "�"%"���� &�%+�!����� �( !"�"&�"# +����! �( � " &�����) !'���'�#��) ;'���3 ":���&��!�!, !���""��� !#$#� ���� " �# % ���& ���'� �#c76 s 'a d d e v a ll e y s e c ti o n s tu d ie d s a m p le s ( 1 ) p r e s e r v a t io n t o t a l a b u n d a n c e p . c f. e n ig m a s . p u n c tu la ta w . b a rn e s ia e w . m a n iv it ia e w . m a n iv it ia e l a r g e w . c o m m u n is w . fo s s a c in c ta w . b ri ta n n ic a l . h a 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schmidt, 1965 family uncinulidae glossinulus schmidt, 1942 subfamily uncinulinae family glossinotoechiidae havlí ek, 1992 obturamentella amsden, 1958 not described ? obturamentella family obturamentellidae savage, 1996 plethorhyncha hall & clarke, 1893 subfamily uncinulinae family uncinulidae sphaerirhynchia cooper & muir-wood, 1951 not described sphaerirhynchia subfamily sphaerirhynchiinae savage, 1996 * ? cassidirostrum mclaren, 1961 subfamily beckmaniinae savage, 1996 * estonirhynchia schmidt, 1954 subfamily sphaerirhynchiinae * ? pectorhyncha mclearn, 1918 family obturamentellidae tadschikia nikiforova, 1937 subfamily sphaerirhynchiinae * gerrhynx baranov, 1991 lapradella baranov, 1989 mongolorhynx erlanger, 1992 voskopitoechia havlí ek, 1992 cerveratoechia garcía-alcalde, 1998 ** assigned to different family or subfamily * subfamily of the family hebetoechiidae ** added in savage (2007) lebanzuella garcía-alcalde, 1999 ** ���� ( � �! ����#������� ������ �� ��� ������ �� ��� ������ 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[2400 2400] /pagesize [595.000 842.000] >> setpagedevice balini_kry 47..84 late carnian-early norian ammonoids from the gssp candidate section pizzo mondello (sicani mountains, sicily) marco balini 1, leopold krystyn 2, marco levera1 & angelo tripodo 3 this paper is dedicated to the memory of paola de capoa, the first paleontologist since the time of g.g.gemmellaro to understand the great paleontologic potential of the ‘‘calcari con selce’’ of western sicily. received: january 9, 2012; accepted: february 27, 2012 key words: upper triassic, carnian/norian boundary, ammonoids, biostratigraphy, sicily, sicanian domain, scillato formation. abstract. a small collection of ammonoids from the upper triassic scillato formation at pizzo mondello (agrigento, sicily) is studied. these specimens were collected within the framework of a project intended to provide integrated high-resolution bio-chronostratigraphic support for the upper carnian-norian magnetostratigraphic scale defined at this site, which is located in an historical area from which g.g. gemmellaro collected and monographed upper triassic ammonoids at the beginning of the 20th century. the specimens, which were collected utilizing the bed-by-bed sampling method, represent the first collection of upper triassic ammonoids described from western sicily since gemmellaro’s time. several levels of the pizzo mondello section yielded ammonoids, but very few beds have provided more than one specimen. this scarcity of specimens has resulted in a complex taxonomic analysis because gemmellaro, who described 166 new species, did not explain the variability of many of his taxa. sixteen taxa belonging to eleven genera are described. they include placites sp. ind., discotropites plinii (mojsisovics), anatropites sp. ind., microtropites cf. paronai, metathisbites cf. affinis, hyattites cf. praefloridus, projuvavites boehmi (gemmellaro), projuvavites inflatus (gemmellaro), gonionotites cf. italicus, gonionotites aff. recuperoi, pregriesbachites n. gen., p. bukowskii (gemmellaro), dimorphites noricus n. sp., dimorphites selectus mojsisovics, dimorphites sp. and discophyllites insignis. among the new taxa, dimorphites noricus n. sp. formalizes the nomen nudum ‘‘dimorphites n. sp. 1’’, which has been quoted in the literature for several years as the index ammonoid for the lowest subzone of the norian stage. the small collection documents the discotropites plinii and gonionotites italicus subzones of the uppermost carnian spinosus zone as well as the dimorphites noricus and d. selectus subzones of the jandianus zone, the first zone of the lower norian. this chronostratigraphic classification not only confidently ties the pizzo mondello succession to the tethyan chronostratigraphic scale, but it is also crucial for the calibration of the halobia and conodont bioevents identified in the section. also discussed is the chronostratigraphic ammonoid-based correlation of the pizzo mondello section with feuerkogel (austria), jomsom (nepal), west union canyon (nevada, usa), black bear ridge (british columbia, canada) and the yana okhotskaya river (siberia, russia). these are the most complete ammonoid-bearing sections in the world that span the carnian/norian boundary, and they exhibit a discrete distribution of ammonoid-bearing beds across the boundary. ammonoids will not provide the primary marker event for the definition of the gssp of the norian, but they are crucial for the selection of the most significant events based on other groups. riassunto. in questo lavoro viene studiata una piccola collezione di ammonoidi proveniente dalla formazione scillato (triassico superiore) di pizzo mondello (agrigento, sicilia). gli esemplari sono stati raccolti nel quadro di un progetto finalizzato alla taratura bio-cronostratigrafica integrata e ad alta risoluzione della scala magnetostratigrafica del carnico superiore e norico riconosciuta in questa località. pizzo mondello è una località fossilifera nuova, ubicata in un’area storica della sicilia da cui g.g. gemmellaro raccolse gli ammonoidi del triassico superiore che descrisse in un’importante monografia all’inizio del xx secolo. gli esemplari di pizzo mondello sono stati raccolti strato-per-strato e rappresentano la prima raccolta di ammonoidi del triassico superiore della sicilia occidentale descritta in oltre un secolo dalla monografia di gemmellaro. gli esemplari provengono da numerosi livelli della sezione, all’interno dei quali però spesso gli ammonoidi sono molto rari. la classificazione dei nuovi esemplari è risultata complessa in quanto gemmellaro, che nella sua monografia descrisse 166 specie nuove, non fornı̀ indicazioni chiare sulla variabilità dei suoi taxa. vengono descritti 16 taxa, appartenenti a 11 generi: placites sp. ind., discotropites plinii (mojsisovics), anatropites sp. ind., microtropites cf. paronai, metathisbites cf. affinis, hyattites cf. praefloridus, projuvavites boehmi (gemmellaro), projuvavites inflatus (gemmellaro), gonionotites cf. italicus, gonionotites aff. recuperoi, pregriesbachites rivista italiana di paleontologia e stratigrafia volume 118 no. 1 3 pls. pp. 47-84 march 2012 1 dipartimento di scienze della terra ‘‘ardito desio’’, via mangiagalli 34, 20133 milano, italy. e-mail: marco.balini@unimi.it 2 department of palaeontology, vienna university, althanstraße 14, 1090 wien, austria. 3 dipartimento di scienze della terra e del mare, università di palermo, via archirafi 22, 90123 palermo, italy. n. gen., p. bukowskii (gemmellaro), dimorphites noricus n. sp., dimorphites selectus mojsisovics, dimorphites sp. e discophyllites insignis. dimorphites noricus n. sp. è il più importante tra i nuovi taxa e rappresenta la descrizione di un nomen nudum riportato nella letteratura recente come ‘‘dimorphites n. sp. 1’’. questa specie è indice della prima sottozona del norico. la collezione studiata permette di riconoscere le sottozone a discotropites plinii e a gonionotites italicus della zona a spinosus del carnico sommitale, e le sottozone a dimorphites noricus e d. selectus della zona a jandianus, la prima zona del norico inferiore. questa classificazione cronostratigrafica data in modo chiaro la successione di pizzo mondello con la scala cronostratigrafica della tetide, ed è fondamentale per la taratura dei bioeventi ad halobia ed a conodonti individuati nella stessa sezione. vengono discusse le correlazioni cronostratigrafiche ad ammonoidi con le migliori sezioni stratigrafiche del mondo, per quanto riguarda il limite carnico/norico. queste sono feuerkogel (austria), jomsom (nepal), west union canyon (nevada, usa), black bear ridge (british columbia, canada) e del fiume yana okhotskaya (siberia, russia). tutte queste sezioni sono caratterizzate da una distribuzione discontinua di ammonoidi, soprattutto in corrispondenza del limite carnico/norico. di conseguenza, molto probabilmente gli ammonoidi non forniranno l’evento primario per la definizione del gssp del norico, ma sono fondamentali nel processo di ricerca e valutazione dei bioeventi riconosciuti in altri gruppi sistematici. introduction the ‘‘cherty limestone’’ (‘‘calcari con selce’’ or halobia limestone auct., scillato formation: schmidt di friedberg et al. 1960) of western sicily are been well known since the second half of the 19th century for its rich macrofossil record of upper triassic pelagic bivalves (halobiids) and ammonoids, first described by g.g. gemmellaro (1882 and 1904). following gemmellaro’s pioneering works, the investigation of halobiids continued and a few authors (montanari & renda 1976; de wever et al. 1979; cafiero & de capoa bonardi 1982; de capoa bonardi 1984) provided the first picture of the stratigraphic distribution of the various species within the ‘‘cherty limestone’’ as well as the refinement of their taxonomy (see especially cafiero & de capoa bonardi 1982 and de capoa bonardi 1984). curiously, not even one specialist has chosen to follow in the ‘‘footprints’’ of gemmellaro and continue the study of triassic ammonoids from western sicily, and since gemmellaro’s 1904 monograph, the literature regarding this fossil group is restricted to just the report of a single specimen of hoplotropites from the monte cammarata section by cafiero & de capoa bonardi (1982: fig. 2 and pl. 6, fig 4-5). such a lack of literature is surprising because on the one hand, the collection from western sicily described by gemmellaro consisted of several hundreds of specimens from several localities scattered over six different areas (gemmellaro 1904; tripodo 2011). on the other hand, a few new but very small collections have been made during the 20th century from localities in sicily that did not provide near as much material as found during gemmellaro’s time (e.g., mufara formation [ladinian-carnian] from palermo area: zia 1956; ‘‘cherty limestone’’ from monte judica, eastern sicily: lentini 1974). in this paper we describe and illustrate ammonoids collected during a four-year long research program, whose purpose was to provide a high-resolution integrated bio-chronostratigraphy for the upper triassic magnetostratigraphic scale and stable isotope curve by muttoni et al. (2001 and 2004) in the pizzo mondello section (sicani mountains, agrigento). in just a very few years, this section has become known worldwide for its unusually thick and uniform late carnian to rhaetian pelagic record as well as for the good preservation of its primary magnetization. such a rare combination of features lead hounslow & muttoni (2010) to propose this section as a standard reference for the calibration of the upper triassic gpts (geomagnetic polarity time scale). this extensive bio-chronostratigraphic investigation, based on a very thorough bed-by-bed sampling program for ammonoids, halobiids, conodonts and radiolarians was carried out by specialists from five italian universities in cooperation with foreign specialists within the framework of the activities of the carnian-norian boundary task group of the subcommission on triassic stratigraphy. due to the importance of the pizzo mondello section for the definition of the gssp (global stratotype section and point) of the norian stage, the investigation focused on the lower part of the scillato formation, which is the formal name for the ‘‘cherty limestone’’ auct., straddling the carnian/ norian boundary. we herein provide taxonomic descriptions for the ammonoids collected during the project, which complement the taxonomic descriptions for the halobiids (levera 2012) and the conodonts (mazza et al. 2012). lithostratigraphy and stratigraphic section as shown in fig. 1, the sicani mountains of western sicily are characterized by a typical permian to cenozoic pelagic succession (mascle 1979) of the sicanian domain, which is divided into several tectonic units (sicanian structural units: catalano et al. 1995) that are overthrust onto a thick allochthonous complex of neogene mudstones and evaporites of the gela nappe (bellanca et al. 1993; bellanca et al. 1995; guaiumi et al. 2007 and references therein). the pizzo mondello section, located in the lowermost sicanian thrust sheet (pizzo mondello tectonic unit) that overthursts upper tortonian-messinian clays, consists of three triassic units, the mufara formation, the scillato formation and the portella gebbia limestone (di stefano 1990; gullo 1996; di stefano & gullo balini m., krystyn l., levera m. & tripodo a.48 1997; guaiumi et al. 2007). the scillato formation (schmidt di friedberg et al. 1960) is the formal name accepted and codified by the ispra (italian geological survey) for the unit known as ‘‘cherty limestone’’ or ‘‘calcari con selce’’ or halobia limestone in sicily. relatively new, the pizzo mondello locality was first described in the mid 1990s (bellanca et al. 1995 and gullo 1996), but it is located within gemmellaro’s ‘‘regione ex-feudo votano’’, one of the source areas for a large part of his collection of halobia and ammonoids. within this wide area (fig. 1) the two best known localities for the scillato formation are monte triona and monte cammarata, both of which were studied in the 1970s and early 1980s (montanari & renda 1976; mascle 1979; de wever et al. 1979; cafiero & de capoa bonardi 1982; de capoa bonardi 1984). monte triona is located about 14 km northwest of pizzo mondello, and monte cammarata is about 19 km to the east. outcrops at pizzo mondello are of much better quality than at monte triona and monte cammarata, and the locality is much better suited for a high resolution bed-by-bed sampling study. this observation applies especially to the lower part of the 430 m thick scillato formation, which straddles the carnian/norian boundary. this part is easy accessible at pizzo mondello because it is exposed in an abandoned quarry that was active in the 1970s (‘‘la cava’’ on some topographic maps). this 143 m thick interval (fig 2), equivalent to interval ii of muttoni et al. (2001, 2004), has been extensively sampled for ammonoids along two segments a and b, which in part overlap (fig. 3). the distance between the two segments is about 200 m, and correlation is achieved simply by following the beds along strike. bed thickness does not change between the segments and the thicker beds serve as excellent marker levels. lithologically, the succession consists mostly of evenly bedded to nodular halobia-bearing cherty calcilutites. this particular sequence was recently described by guaiumi et al. (2007), nicora et al. (2007), balini et al. (2008) and balini et al. (2010). sampling during the past 10 year period, the pizzo mondello section has been sampled several times by specialists from a wide spectrum of fields including magnetostratigraphy, sedimentology, microand macropaleontology. during this time, sampling has become more and more thorough and several types of labels and numbering schemes have been used (for details see balini et al. 2010). beginning in 2006, the basis for all samplings has been an extremely detailed stratigraphic log measured by preto and guaiumi in the lower 143 m of the section underlying the slump breccia (interval iii of muttoni et carnian-norian ammonoids from pizzo mondello 49 fig. 1 geological and structural map of the monti sicani (sicani mountains) showing the distribution of the sicanian triassic to oligocene pelagic successions, as well as the position of pizzo mondello with respect to monte triona and monte cammarata (from cacciatore et al. 2010). balini m., krystyn l., levera m. & tripodo a.50 fig. 2 stratigraphic log of the pizzo mondello section. a) the complete development of the section, from the base of the scillato formation to the portella gebbia limestone, with magnetostratigraphy (muttoni et al. 2001, 2004) and position of the conodont samples studied by nicora et al. (2007), mazza et al. (2010), mazza et al. (2011) and mazza et al. (2012). the log is from balini et al. (2010) and is partly modified from muttoni et al. (2004). b) detail of the lower part of the scillato formation (143 m) sampled for ammonoids. al. 2001, 2004). all previous samples taken by muttoni et al., (2001 and 2004) that could be correlated with numbers painted on the beds, were also plotted on the log. when possible, ammonoids were labelled with the bed number from which they were collected, provided the bed had been previously sampled. ammonoids from beds with no label have been numbered as pmamxx. the sequence of these samples is merely chronologic and has no relation to the stratigraphic succession. the ammonoid record six field trips to pizzo mondello, each averaging about one week duration, were carried out between spring 2007 and fall 2010. from the beginning, ammonoids were found throughout the succession, but their occurrence was found to be quite rare (fig. 4). because of this scarcity, it was necessary to follow the beds along strike for tens of meters searching for ammonoid crosssections on the outcrop. the resulting collection was combined with ammonoid specimens found during the thorough bed-by-bed search for halobia that even included beds without visible macrofossils on the surface. a total number of 166 beds were sampled utilizing this approach (see levera 2012). overall, the final collection consists of 130 ammonoids collected from a stratigraphic interval of about 100 m. on rare occasions, two or more specimens were collected from the same bed (fig. 4b), and even though the resulting collection is representative of the ammonoid record of the section, it does not allow for a population analysis. most of the ammonoids are about 1 or 2 cm in diameter (fig. 4a) and the frequency of larger specimens decreases with increasing size. the largest specimen (fig. 4d) is a gonionotites of about 10 cm in diameter. quite often, specimens are preserved with a strongly recrystallized test, but some are also preserved as internal molds (fig. 4c). the latter are usually very easy to extract from the rock matrix. ammonoid taxonomy: the complex heritage of g.g. gemmellaro as mentioned in the introduction, our knowledge of triassic ammonoids from western sicily is based entirely on the monograph by gaetano giorgio gemmellaro (1832-1904) that was published just after his death by his collaborators g. di stefano, l. schopen and a. carapezza (gemmellaro 1904: p. iii). this monograph is based on a large collection of ammonoids from the carnian-norian ammonoids from pizzo mondello 51 fig. 3 general view of the lower part of the scillato formation at ‘‘la cava’’, showing the position of the two segments of the composite section. the beds of the upper part of the segment a can be followed along strike to the lower part of the segment b (see also fig. 2). following areas: palermo (‘‘regione giacalone’’, ‘‘cave di billiemi’’), trapani (‘‘castellammare del golfo’’), bisacquino (‘‘regione madonna del balzo’’= monte triona), s. stefano quisquina s.l. (‘‘regione ex-feudo votano’’) and castronuovo (‘‘regione modanesi’’), of which the major part is preserved in the museo geologico g.g. gemmellaro of the university of palermo. gemmellaro did not always provide the total number of specimens attributed to each of his taxa, but the agreement between the number of specimens mentioned by him and the specimens stored in the museo is generally quite good. the registered collection includes 780 specimens. gemmellaro’s monograph is devoted entirely to taxonomy, with the description of 166 new species (and 84 taxa in open nomenclature), attributed to 49 genera and subgenera. the genera siculites, palicites, mojsisovicsites and gonionotites were also erected in this monograph. however, even with this impressive and well illustrated taxonomy (30 high quality plates), two main weak points are immediately obvious. the first is the lack of detailed geographic and geological information regarding the successions from which the fossils were collected and the number and position of the fossil-bearing levels. the second major problem is the lack of a biostratigraphic analysis of the faunas. this second point is somewhat surprising because by the end of the 19th century the knowledge of ammonoid biochronostratigrahic scales was well developed (i.e., mojsisovics et al. 1895), and the common practice followed by all authors of that time was to include in every ammonoid monograph a chapter on faunal subdivision and correlation. this absence of critical information is difficult to understand; it is possible that gemmellaro did not collect the major portion of the material by himself, or maybe he was unable to complete his monograph. no matter the reason, this information was not reported, and this may well be one of the reasons for the relatively low scientific impact of gemmellaro’s monograph. balini m., krystyn l., levera m. & tripodo a.52 fig. 4 field pictures of ammonoids. a) cross section of a small specimen from level pmam12. b) rare occurrence of two specimens from the same level, fnp317b. c) rarely the specimens are exposed in 3d and easy to be extracted: projuvavites inflatus mpum 10974, level pmam49. d) sometimes the specimens identified on the basis of the cross section are also easy to be collected, here gonionotites cf. italicus mpum 10975 from level pmam7 (pl. 2, fig. 1a-b). new genera described by gemmellaro have generally been accepted (e.g., arkell et al. 1957; tozer 1981a), but very few workers have tried to improve his taxonomy at the species level. spath (1951) described the new genus euisculites with isculites bittneri gemmellaro as type species and tozer (1994) included styrites disciformis gemmellaro in his new genus discostyrites. some authors have not even mentioned gemmellaro’s monograph diener’s (1906: 6) criticism was in some respects ‘‘extreme’’, when he wrote that gemmellaro’s monograph could not be used because ‘‘the types are represented in a manner which defies every attempt to realize their true shape’’. surprisingly, for taxa with no stratigraphic position, a few of gemmellaro’s taxa have recently been recognized as having stratigraphic significance. gonionotites italicus gemmellaro was proposed by krystyn in 1980 as an index ammonoid for the latest subzone of the carnian stage. since then, the gonionotites italicus subzone of the spinosus zone has became part of the upper triassic tethyan scale (krystyn et al. 2002; balini et al. 2010). the genus mojsisovicsites gemmellaro, which is not found elsewhere in the tethys realm, was recognized in north america by silberling (1959), who attributed stikinoceras kerri mclearn, 1930 to gemmellaro’s genus as the index taxon for the first zone of the norian stage. tozer (1994) later rejected this attribution, but mojsisovicsites and stikinoceras are still considered to be very closely related forms. the lack of new discoveries and taxonomic revisions, especially at the species level, implies that the state of the taxonomy of ammonoids from the scillato formation/‘‘cherty limestone’’ remains the same as it was in 1904. this suggests that species are defined with a typologic approach, sometimes based on single specimens or even juveniles, or based on specimens grouped together on the basis of assumptions and not on ontogeny and population variability analyses. last but not least, there is a lack of information about the stratigraphic position of the specimens. as so ably summarized by tozer (1971), such a wide range of problems were rather normal for monographs published in the second half of the 19th century, but in the case of gemmellaro’s work the problem is much more severe because of the much greater uncertainty about the stratigraphic position of his taxa. the interval from which his taxa were collected is not restricted to just one or two substages, but rather it encompasses much of the upper carnian through the entire norian stage. taxonomic results the classification of the present collection from pizzo mondello and its comparison with the type specimens of the gemmellaro collection, recently re-described by tripodo (2011), provides the opportunity to test gemmellaro’s taxonomy with all its flaws as well as to evaluate the significance of his collection. a few points must be stressed: a) twelve genera have been identified in the new collection: placites, discotropites, anatropites, microtropites, metathisbites, hyattites, projuvavites, gonionotites, dimorphites, discophyllites, rhacophyllites and pinacoceras. in addition, we describe the new genus pregriesbachites in the systematic paleontology. small ammonoids are also referred to the families sandlingitidae and arcestidae. all genera identified herein are documented in the list of taxa described by gemmellaro. the genus discotropites hyatt & smith, 1905 includes almost all species described by gemmellaro as eutomoceras. another tropitidae well represented in the gemmellaro collection is hoplotropites spath, 1929, which is the new name for margarites mojsisovics, 1889. this genus, typical of the discotropites plinii subzone, has not yet been found at pizzo mondello. a group of juvavitidae that is well documented in the gemmellaro collection (37 species) is anatomites mojsisovics, 1893. however, several species originally attributed to this genus would be better included in projuvavites tozer, 1971. b) the taxonomic study of ammonoids from pizzo mondello provided an opportunity to describe two new taxa already outlined by krystyn (1980, 1982) during an investigation of classic localities in the salzkammergut area (northern alps) and a new site in the nepalese himalaya. dimorphites noricus n sp. is a new species quoted in several previous papers as dimorphites n. sp. 1, and the new genus pregriesbachites is erected for the ‘‘projuvavites n. gen. with nodes on the body chamber’’ mentioned by krystyn (1982). both new taxa are also documented in the scillato formation at pizzo mondello. c) classification of our specimens to a species rank has often revealed the limits of gemmellaro’s typologic approach to taxonomy. ammonoids are rare in the sedimentary successions, and thus far it has proven very difficult to collect more than one specimen per bed. comparison of these newly collected single specimens to gemmellaro’s species, which are often illustrated by only one or two types, is usually quite frustrating. information regarding species variability is not available, and often the new specimens do not correspond well with one particular type specimen, but rather, they more commonly are similar to two or more type specimens of different species. for instance, this is the case for the single microtropites specimen from level pmam52a that in some respects resembles m. paronai (gemmellaro), while for others it is more similar to m. brancoi (gemmellaro). similarly, the best preserved gocarnian-norian ammonoids from pizzo mondello 53 nionotites from level pmam7 does not correspond well with any of the 11 species of gonionotites described by gemmellaro. d) for a modern taxonomic revision of many of gemmellaro’s groups, it would be absolutely necessary to require a definition of their variability. however, very few beds of the scillato formation would provide enough specimens for even a simplified population analysis. a group that may potentially benefit from this approach is gonionotites, which is relatively common in the succession. however, the main difficulty with this group is the large size of its type specimens, which contrasts with the extreme scarcity of these larger specimens and the high frequency occurrence of smallor mediumsized specimens in the succession. several of the eleven species attributed to this genus by gemmellaro seem to be distinguished only by the adult body chamber. e) overall, gemmellaro’s studied collection is clearly representative of the ammonoid record of the scillato formation. a few of his genera have not been found in the field (e.g., jovites) but it is worth noting that a number of very rare, small forms were actually collected by him or by his collaborators. dimorphites, for instance, belongs to this group. thus far it has been found only in five beds at pizzo mondello, and the specimens are of rather small size. this scarcity, notwithstanding that dimorphites is documented by several specimens in gemmellaro’s collection. in the next chapter, we provide descriptions for the most significant and representative taxa, but a few specimens are left in open nomenclature. this approach strictly follows the recommendations of salvador (1994: 91) and remane et al. (1996) concerning the publication and illustration of the fossil record documented in gssp sections. in addition to this formal reasoning, the description of the ammonoid record is useful for calibration of the halobia and conodont bioevents (levera 2012; mazza et al. 2012). repository of specimens. specimens described in this contribution are housed in three institutions in italy and austria as follows: mpum: museo di paleontologia, dipartimento di scienze della terra ‘‘ardito desio’’, università degli studi di milano, via mangiagalli 34, 20133 milano, italy. mgup: museo geologico g.g. gemmellaro, università degli studi di palermo, corso tukory 131, palermo, italy. gba: sammlungen der geologischen bundesanstalt, neulinggasse 38, a-1030 wien, österreich. numbering of the specimens. every specimen mentioned in the text is identified by both registration number (e.g., mpum....) and collection number. the latter, in parenthesis, indicates the bed of collection and the individual number of the specimen (e.g., fnp144-2: 2nd specimen from bed fnp144). dimensions. d= diameter; h= max. whorl height in d; h= min. whorl height in d; u= umbilical width in d; w= whorl width in h; sgr= (h-h)/h x 100. all measurements are in mm. sgr is in percent. measurements in italics represent partly damaged specimens. occurrence. in the systematic paleontology and chronostratigraphy sections, the nomenclature for chronozones follows hedberg (1976). this nomenclature is more synthetic than the one required by salvador (1994): jandianus zone, instead of guembelites jandianus chronozone. for subzones, biostratigraphic nomenclature is used because there is no chronostratigraphic counterpart for such a short subunit. systematic paleontology family-group taxonomy follows that of tozer (1981a and 1994). order ceratitida hyatt, 1884 superfamily pinacocerataceae mojsisovics, 1879 family gymnitidae waagen, 1895 subfamily gymnitinae waagen, 1895 genus placites mojsisovics, 1896 type species: pinacoceras platyphyllum mojsisovics, 1873 placites sp. ind. pl. 1, fig. 1; fig. 5 material: one specimen mpum 10966 (na49.1-1). description. small specimen (d= 15.3mm) preserved as a three dimensional internal mold. specimen is a juvenile, last 70º of coiling belongs to the body chamber. involute platycone with nearly closed umbilicus, compressed elliptical whorl section and very narrow venter. suture line (fig. 5) is typical of placites. discussion. species of placites may reach a diameter of 100 mm and are distinguished (cf. mojsisovics 1873) mainly on the basis of their whorl section and suture line at medium to large size, i.e., adult stage of growth. the specimen from pizzo mondello (level na49.1) is clearly a juvenile and therefore is left in open nomenclature. occurrence. as with most of gymnitidae, placites is a long-ranging form; its range spans the entire norian to rhaetian stages. the specimen described here was collected from a bed yielding halobia mediterranea (see levera 2012), a species usually referred to the paulckei zone (krystyn et al. 2002; levera 2012). balini m., krystyn l., levera m. & tripodo a.54 fig. 5 suture line of placites sp. ind., specimen mpum 10966 (na 49.1.-1). bar scale 2 mm. superfamily tropitaceae mojsisovics, 1875 family tropitidae mojsisovics, 1875 genus discotropites hyatt & smith, 1905 type species: ammonites sandlingensis hauer, 1850 discotropites plinii (mojsisovics, 1893) pl. 1, fig. 2a-b 1893 eutomoceras plinii mojsisovics, p. 289, pl. 130, fig. 4, 5, 6. 1982 discotropites plinii – krystyn, p. 34, pl. 7, fig. 3-4. [cum syn.] 2000 discotropites plinii de zanche et al., pl. 1, fig. 1a-b. stratum typicum: hällstatter kalk, at feuerkogel (salzkammergut, austria). lectotype: original of mojsisovics 1893, pl. 130, fig. 5, gba1893/01/264, selected by krystyn (1982: 34). material: one specimen mpum 10967 (pmam17-1). description. slightly elliptical, extremely crushed internal mold of large-sized specimen with diameter of about 90 mm. about 130º of the body chamber are preserved. despite the extreme crushing, the keel is clearly visible. ribbing consists of slightly prorsiradiate, gently sinuous-falcoid ribs. organization of ribs is visible only on the last quarter of the preserved body chamber. ribs are organized in bundles, each consisting of three primary ribs (pl. 1, fig. 2b). usually one intercalatory/bifurcate rib appears a few mm from the umbilical margin. additional branching at a little less than ¼ of whorl height, and very rarely (once in the last half whorl) intercalatory ribs appears at 1/3 of whorl height. near the ventral shoulder, flat ribs are 3 to 4 times wider than the narrow inter-rib intervals. suture lines are strongly serrated, but not well preserved. remarks. this species was revised by krystyn (1982), who discussed the synonymy and its relationship with discotropites theron (dittmar, 1866), the most similar species. in d. theron the primary ribs start from node-like periumbilical swellings (mojsisovics 1893: 288; krystyn 1982: 34), that are not developed in d. plinii. the difference between d. plinii and d. theron is even greater, if we follow the interpretation of d. theron by tozer (1994), who extended its diagnosis to forms with several spiral rows of nodes. occurrence. the distribution of discotropites plinii is widespread in the tethys realm, from the northern alps to japan, through sicily, turkey, himalaya and timor. its stratigraphic position is so well constrained that this species has been selected by krystyn (1980, 1982) as the index ammonoid for the first subzone of the uppermost carnian spinosus zone (see also krystyn et al. 2002). genus anatropites mojsisovics, 1893 type species: tropites (anatropites) spinosus mojsisovics, 1893 anatropites sp. ind. pl. 1, fig. 3, 4a-b material: two specimens, mpum 10968 (fnp112-1) and mpum 10969 (pmam22bis). remarks. the two specimens attributed to anatropites were collected from two different levels and most probably are not conspecific. following the stratigraphic succession, the first specimen (mpum 10968 [fnp112-1]: pl. 1, fig. 4a-b) is preserved in three dimensions for one quarter of a whorl. the original diameter is about 20 mm, and the whorl cross section is typically semielliptical with lateral sides converging towards the rounded venter and the prominent keel. ornamentation is also typical of the genus, consisting of prorsiradiate concave primary ribs strongly elevated/subspiny at the periumbilical border and fading towards the shoulder. the second specimen, from level pmam22bis (pl. 1, fig. 3), is preserved only as a whorl cross-section that is unextractable from the rock matix. despite its very incomplete preservation, the specimen fits perfectly with anatropites because of its peculiar whorl section with lateral sides converging to a rounded venter, its maximum width at the umbilical margin, its elevated keel without side furrows and its weak but distinct small projections at the umbilical margin, suggesting the umbilical spines typical of anatropites. both specimens are left in open nomenclature, the first because of its rather small size and the second because details of its ornamentation are not visible. occurrence. the genus anatropites occurs in the tethys realm and north america. in both areas it is restricted to the last zone of the carnian stage, namely the spinosus zone in the tethys (krystyn 1973, 1980; krystyn & gallet 2002; krystyn et al. 2002) and the macrolobatus zone of central nevada (silberling 1959) and british columbia (tozer 1994). genus microtropites mojsisovics, 1893 type species: ammonites galeolus hauer, 1860 preliminary remarks. the genus microtropites mojsisovics, 1893 includes small-sized (dwarf) tropitidae that exhibit a change in coiling during growth, a weak to very weak ornamentation and an ammonitic suture line (e.g., mojsisovics 1893: 188; arkell et al. 1957: l169). species attributed to this genus show some differences in their whorl section, which varies from carnian-norian ammonoids from pizzo mondello 55 slightly compressed with a nearly semi-elliptical outline (e.g., the type m. galeolus [hauer, 1860], m. lepsiusi mojsisovics, 1893 and m. tubercularis mojsisovics, 1893) to slightly depressed with a broadly rounded venter (e.g., microtropites sp. ind. in krystyn 1982). taking into consideration the rarity of specimens from this group and the lack of pertinent literature plus the fact that their distribution is restricted to the latest carnian (krystyn 1980, 1982), we for now consider these differences in whorl section to be part of the variability of the genus. microtropites cf. paronai (gemmellaro, 1904) pl. 1, fig. 5a-c remarks on the type series. gemmellaro described the new species tropites paronai based on only three specimens (gemmellaro 1904: 96), but in the collection stored at museo gemmellaro, two groups of specimens are labelled as ‘‘tropites paronai’’. the first group (mgup 002.96.1-4) consists of four specimens and includes the two figured types (mgup 002.96.1= pl. 7, fig. 27-28; mgup 002.96.3= pl. 7, fig. 25-26). the second group includes six specimens that are referred to a different catalogue number (mgup 002.95.1-6) than the first group. we consider the types of the species to be only the specimens of the first group (mgup 002.96.1-4). material: one specimen mpum 10970 (pmam52a-1). description. specimen preserved with strongly recrystallized test, but internal mold is occasionally visible because test was lost in a few places during preparation. body chamber consists of 170º of the last whorl. coiling is involute, but from approximately 4-5 mm of h it becomes evolute due to umbilical egression. whorl cross-section is depressed with wide, rounded venter giving specimen a near cadicone shape. ventral keel without significant bordering furrows. specimen appears to be nearly smooth except for strong forward-projected, concave, extremely weak and fading ribs that are visible under certain light conditions on last 90º of preserved body chamber. suture line not visible. dimensions (mm) discussion. the genus microtropites was recognized in sicily by gemmellaro (1904), who classified a few specimens from his collection as m. lepsiusi mojsisovics, 1893. however, certain other taxa included by gemmallaro in tropites may well be better referred to microtropites; these are tropites paronai gemmellaro, t. brancoi gemmallaro, t. carapezzai gemmellaro and tropites n.f. indet. ex aff. t. brancoi. according to the original descriptions and plates, these taxa are very similar with only slight differences in the degree of whorl compression, type of ribbing and whether or not umbilical nodes and weak furrows bordering the keel are present. an examination of gemmellaro’s type material reveals that he emphasized these differences in his descriptions and also that some details of the drawings were slightly exaggerated. a revision of the whole group is beyond the scope of this paper and would require much new material to test the population variability of these very small, poorly known tropitiidae. the pizzo mondello specimen is closer to microtropites paronai (gemmellaro, 1904) than to m. brancoi (gemmellaro, 1904), which is characterized by ribs (visible on gemmellaro’s plate 25, fig. 1-4) and by a slightly more compressed whorl section. its similarity with m. paronai includes the depressed whorl section (gemmellaro’s plate 7 fig. 27-28) and faded ribs. the specific attribution of the pizzo mondello specimen is challenged by certain differences from the syntype shown in pl. 7, fig. 25-26 (mgup 002.95.3), which shows a narrowing of the whorl section that apparently is related to umbilical egression. our specimen is slightly smaller than this particular syntype and it exhibits umbilical egression, but its venter does not become narrower. the same syntype has weak umbilical nodes (weaker than shown in gemmellaro’s figure) that are neither visible on the other figured syntype (pl. 7, fig. 27-28; mgup 002.96.1), nor on the pizzo mondello specimen. our specimen does not have furrows along side the keel, but this feature is often visible on gemmellaro’s types. we prefer not to over-emphasize this difference since it may simply be related to different types of preservation. the test of our specimen is nearly complete, whereas gemmellaro’s types are preserved as internal molds. occurrence. the genus microtropites is known only from the tethys realm, where it is regarded as a marker for the gonionotites italicus subzone, upper part of the spinosus zone, uppermost carnian (krystyn, 1980, 1982). the stratigraphic positions of the taxa described by gemmellaro (1904) from sicily are unknown, but the new pizzo mondello specimen from level pmam52a is herein used to mark the lower boundary of the gonionotites italicus subzone (fig. 6). family thisbitidae spath, 1951 genus metathisbites tozer, 1994 type species: buchites hilaris var. dawsoni mclearn, 1940 balini m., krystyn l., levera m. & tripodo a.56 carnian-norian ammonoids from pizzo mondello 57 fig. 6 distribution of the upper carnian-lower norian ammonoids in the studied interval of pizzo mondello section. the position of all the ammonoid samples is shown, but the occurrence of small sized specimens not classified in this work is differentiated with a short dash. for the sake of completeness the position of the conodont samples of muttoni et al. (2001 and 2004) and of the halobiids described by levera (2012) is also shown. metathisbites cf. affinis (gemmellaro, 1904) pl. 1, fig. 6a-b material: one specimen mpum 10971 (fnp145-1). description. rather small specimen (diameter about 18.5 mm) that consists of an internal mold of a body chamber and preserved only on one side with a small part of the venter exposed. rate of coiling very low with h~u. flank slightly convex suggesting a compressed whorl section with well developed shoulder. no keel visible on exposed part of venter. ornamentation consists of gentle, wavy-like, prorsiradiate, slightly sinuous primary ribs. only one bifurcate rib occurs in about one half whorl. all ribs end on ventral shoulder with small but distinct rounded node (see pl. 1, fig. 6b). suture line not visible. discussion. the specimen is very close to thisbites mojsisovics, 1893 and metathisbites tozer, 1994. metathisbites is characterized (tozer 1994: 232) by an ovoid whorl section, a weak keel that sometimes is lacking, mostly simple ribs and nodes that are weaker than in thisbites. the available specimen does not show an ovoid section, but its cross-section seems to be more subrectangular/subquadrate. apart from this feature, the other characters suggest attribution to metathisbites. ribbing is rather simple and most of the ribs are primary. the ventrolateral nodes are very weak and for what is visible on the small part of the ventral side, there is no prominent keel. thisbites includes species with more dense ribbing, a much higher frequency of secondary ribs and usually a distinct and strong keel. as regard the species, the specimen is referred by cf. to ceratites (buchites) affinis gemmellaro on the basis of ribbing and nodes. this species is characterized by weak ventrolateral nodes and a very weak keel, both features not consistent with buchites mojsisovics, 1893. the attribution by cf. is due to the incomplete preservation of the ventral side of the specimen from pizzo mondello. occurrence. according to tozer (1994) metathisbites is documented in both the tethys realm and north america. its range is calibrated only in north america where it is restricted to the lower norian kerri zone, more precisely to the stikinoceras kerri subzone ii. the specimen from pizzo mondello, in open nomenclature, has been found together with dimorphites selectus, index of the 2nd subzone of the lower norian jandianus zone. this finding is consistent with the stratigraphic position of the genus in north america. family juvavitidae tozer, 1971 subfamily juvavitinae tozer, 1971 genus hyattites mojsisovics, 1902 type species: pinacoceras praefloridum mojsisovics, 1873 preliminary remarks. hyattites mojsisovics, 1902, one of the poorest known ammonoid genera from the upper triassic of the tethys, was established by mojsisovics (1902) to accommodate three species with quite different stratigraphic positions that were previously referred to beyrichites waagen, 1895 by philippi (1901:110). pinacoceras praefloridum mojsisovics, 1873, meekoceras maturum mojsisovics, 1882 and meekoceras emmerichi mojsisovics, 1882 are of relatively medium size and they exhibit involute coiling and a lack of ornamentation, but mojsisovics (1902: 306) emphasized the suture line of hyattites with its phylloid habitus and broad median hump in the external lobe (litt. ‘‘medianhöckers im externlobus’’) that was said to suggest a family connection with sturia mojsisovics, 1882. in his description of the genus, mojsisovics selected pinacoceras praefloridum mojsisovics, 1873 (of carnian age) to be representative of the development of the suture line of hyattites, and this is probably the reason this taxon has long been considered to be the type species of the genus (diener 1915: 154; spath 1951: 154; arkell et al. 1957: l182). krystyn 1982 (p. 43) closely examined the type specimen of the species and recognized that the suture line drawing in mojsisovics 1893 was incorrect due to excessive grinding of the test. in reality the external suture is actually juvavitid and hyattites is closely related to gonionotites from which it differs by the early fading of its sculpture. spath (1951) pointed out the relatively large age differences of the three species grouped together by mojsisovics and therefore, reduced the composition of hyattites. he confirmed the attribution of anisian aged m. maturum to beyrichites, as already suggested by philippi (1901), and he selected ladinian aged m. emmerichi as type species for the new genus parasturia. thus, spath left only the type species in hyattites, and until now it has remained the composition of the genus, because the only two taxa referred to this genus, hyattites salomoni gemmellaro, 1904 and hyattites (?) nepalensis jeannet, 1959 apparently are not conspecific with h. praefloridum. gemmellaro described the new species hyattites salomoni on the basis of two specimens from ‘‘feudo modanesi,’’ western sicily. however, his figured suture line (1904: pl. 27, fig. 29) contains simpler indentations than h. praefloridus. hyattites (?) nepalensis jeannet, 1959 from the carnian of nepal is referred with question to hyattites, but apparently it is not conspecific with h. praefloridus because of its ribbed ornamentabalini m., krystyn l., levera m. & tripodo a.58 tion. gonionotites rarus mclearn, 1940, as figured in tozer (1994, pl. 119), lacks ornamentation and could well be included in the genus hyattites. occurrence and age. neither the occurrence nor the age of hyattites is well defined in literature. arkell et al. (1957), probably following spath’s lead (1951), reported hyattites from the carnian of sicily and the northern alps, whereas tozer (1981a and b) provided an early norian age and a much wider paleogeographic distribution, from the tethys to the east pacific. evidence in support of tozer’s conclusions is not clear because he does not cite the genus from north america, although it may very well occur there (e.g., gonionotites rarus). the tethys occurrence may be supported by the distribution of hyattites praefloridus (mojsisovics) and h. salomoni (gemmellaro), but there have been no reports of hyattites from north america, neither before tozer 1981a nor after (e.g., tozer 1994). h. praefloridus is known from ‘‘schichten mit lobites ellipticus’’ of röthelstein (mojsisovics 1873: 58), a fossil site containing a condensed fauna ranging in age from carnian to early norian, according to krystyn (1980). hyattites cf. praefloridus (mojsisovics, 1873) pl. 1, fig. 7a-b; fig. 7 material: one specimen, mpum 10972 (fnp333). description. specimen is preserved as an internal mold, but retains small areas of recrystallized test. last 240º of whorl belong to the body chamber. involute conch with very narrow umbilicus and compressed whorl section, with nearly flat flanks and rounded venter. surface of specimen is smooth on both the body chamber and portion of the last whorl that is phragmocone. suture lines are strongly serrated, specimen appears to be an adult. suture is ammonitic with a gonionotitic pattern of indentitions (fig. 7). external lobe has a wide internal saddle. discussion. one peculiar feature of the pizzo mondello specimen is the lack of ornamentation on the inner part of the last whorl at relatively small values of h. at this size, nearly all juvavitidae have ribs and/or constrictions. the attribution of the specimen to hyattites relies mainly on these features and on the gonionotiticlike suture line that is consistent with the general pattern of the type specimen’s suture line, re-described by krystyn (1982: 42). however, the suture line of the type specimen of h. praefloridus is more indented, but this magnitude of difference is not fully justified by the slightly larger size of the type specimen (d about 50 mm) with respect to specimen mpum 10972 (d about 39 mm). because of this difference the specimen from pizzo mondello is attributed to this species, with cf. hyattites salomoni gemmellaro from ‘‘feudo modanesi’’ differs from hyattites cf. praefloridus by its suture line with seven saddles on the lateral sides, very simple indentitions, and ribs developed on the external part of the lateral side. occurrence. hyattites cf. praefloridus is the oldest ammonoid collected to date from the scillato formation at pizzo mondello. it was collected from the upper carnian part of the section, about 9 m below the single occurrence of discotropites plinii. genus projuvavites tozer, 1971 type species: juvavites (anatomites) brockensis smith, 1927 projuvavites boehmi (gemmellaro, 1904) pl. 1, fig. 8a-b; fig. 8 v 1904 juvavites (anatomites) böhmi gemmellaro, p. 207, pl. 27, fig. 9. partim carnian-norian ammonoids from pizzo mondello 59 fig. 7 suture line of hyattites cf. praefloridus, specimen mpum 10972 (fnp333). bar scale 5 mm. ?v 1904 juvavites (anatomites) böhmi gemmellaro, p. 207, pl. 27, fig. 10. stratum typicum and locus typicus: light gray limestone with cherty nodules, ‘‘feudo modanesi’’ near castronuovo (gemmellaro 1904:208). type series: gemmellaro based his description of the species on two specimens, both of which are figured. their repository is the museo gemmellaro under registration number mgup 002.183.1 and.2. specimen mgup 002.183.1 (gemmellaro 1904: pl. 27, fig. 9) is here designated as the lectotype. material: one specimen mpum 10973 (pmam27-1). description. specimen consists of slightly more than half whorl, representing last part of phragmocone and beginning of body chamber (about 100º). test on phragmocone lost during extraction. juvavitid with involute coiling and slightly compressed whorl section. whorl section of inner whorls is semioval, but then apparently becomes almost subtriangular on body chamber. however, since body chamber is slightly crushed, this feature may be partly secondary. two concave, projected constrictions are visible, one on phragmocone and the second located just after last visible suture line, at about 80º from first. ornamentation consists of slightly prorsiradiate, concave ribs with projuvavites-like organization on the phragmocone (see krystyn 1982). at the shoulder, about 13 ribs are visible between the two constrictions. ribbing on body chamber is much weaker, consisting of widely-spaced primary ribs and intercalatory ribs visible only at the shoulder. three intercalatory ribs are usually present between two primary ribs. suture line ammonitic (fig. 8) with projuvaviteslike pattern of intentations. discussion. relatively few species of the large group attributed by gemmellaro to the subgenus juvavites (anatomites) have a large size and relatively thick whorl section. within this small group, the species most similar to the pizzo mondello specimen is j. (a.) boehmi gemmellaro, which exhibits a fading of the ribs on the body chamber, especially on the inner part of the lateral side. this feature is well visible on gemmellaro’s syntype figured in pl. 27, fig. 9 (mgup 002-182.1), which is designated as lectotype. the second specimen, now a paralectotype (pl. 27, fig. 10, mgup 002-183.2), is a juvenile that does not show the typical features of j.(a.) boehmi. an examination of the lectotype of j.(a.) boehmi reveals that gemmellaro’s figure of the whorl section is not correct. the specimen is more compressed than shown in the illustration in pl. 27, fig. 9, and in this feature the lectotype is more similar to the new specimen from pizzo mondello. the only difference between mpum 10973 and the lectotype is the width of the venter, which seems to be wider on the lectotype of p. boehmi. information regarding population variability of p. boehmi is not available, but it is well known from tens of papers on triassic ammonoids that population variability is rather wide for several groups, and thus, the shape and width of the venter may also be affected. given that the difference in the width of the venter is rather small, and that the relatively narrow venter of mpum 10973 may be due to a slight crushing of the body chamber, we have decided to include the new specimen in p. boehmi (gemmellaro). occurrence. no literature is available for projuvavites boehmi (gemmellaro). this species is apparently restricted to sicily, where its stratigraphic position is unknown. the new specimen from pizzo mondello comes from level pmam27, just slightly above pmam17, which yielded discotropites plinii. p. boehmi could probably be referred to this subzone. projuvavites inflatus (gemmellaro, 1904) pl. 1, fig. 9a-d v 1904 juvavites (anatomites) inflatus gemmellaro, p. 211, pl. 5, fig. 19-20. ?v 1904 juvavites (anatomites) gelonis gemmellaro, p. 213, pl. 17, fig. 3-4, pl. 21, fig. 10. stratum typicum: light gray limestone with chert nodules from ‘‘feudo modanesi’’ near castronuovo (palermo) (gemmellaro 1904:212). type series: the description of this species is based on two specimens, both registered in the museo gemmellaro under number mgup 002.185.1-2. the only figured specimen (mgup 002.185.1) is here designated as the lectotype. material: one specimen mpum 10974 (pmam49-1). description. the specimen consists of an internal mold of phragmocone and at least 270º of body chamber. juvavitid with very involute coiling (u= 9% of d) and slightly depressed whorl section (h/w from 0.86 to 0.94) with semielliptical outline that becomes subtriangular on last whorl. rounded venter is crossed by constrictions, but not by ribs, at least on phragmocone. balini m., krystyn l., levera m. & tripodo a.60 fig. 8 suture line of projuvavites boehmi (gemmellaro, 1904), specimen mpum 10973 (pmam 27-1). bar scale 5 mm. constrictions spaced about 80º-90º apart. ribs nearly straight, but bend forward at ventrolateral shoulder and then usually fade without crossing venter. organization of ribs between constrictions follows the typical pattern of projuvavites without trifurcation of ribs and about four primary ribs between constrictions. ribbing also developed on beginning of body chamber. suture line ammonitic, not well enough preserved for drawing. dimensions discussion. the pizzo mondello specimen is attributed to juvavites (anatomites) inflatus gemmellaro, 1904 in part because of its involute coiling and slightly depressed whorl section, but especially for the change in its whorl section, which becomes nearly subtriangular on the outer whorl and also for the development of ribs on the body chamber. the constrictions are not wellrepresented on gemmellaro’s drawings, but they are clearly mentioned in the text (p. 211) and also quite visible on the lectotype. juvavites (anatomites) gelonis gemmellaro, 1904 is cogeneric with projuvavites inflatus, and probably could be synonymized with this species. the size of the largest types (gemmellaro 1904, pl. 17, fig. 3-4) is nearly equivalent to the inner whorl of the type specimen of p. inflatus, and their coiling and whorl sections are also very similar. ribbing is slightly different, but this may not be of great significance at the species level. gemmellaro’s drawing (1904, pl. 17, fig 5) of p. gelonis appears to show a few trifurcations, but this feature is not very obvious on the original. among the large group of ‘‘anatomites’’ described by gemmellaro, projuvavites inflatus is also similar to j. (a.) archimedis gemmellaro, 1904, j. (a.) mariani gemmellaro, 1904 and j. (a.) boehmi gemmellaro, 1904, but all three species can be attributed to projuvavites tozer, 1971. juvavites (a.) archimedis (gemmellaro 1904: pl. 9, fig. 1-2) differs by the reduction of ribbing on its body chamber and its nearly semielliptical whorl section. juvavites (a.) mariani has a totally different body chamber (gemmellaro 1904: pl. 16, fig. 1-2). its venter becomes wider (this feature is more developed on the syntype than shown on the plate), and its ribs are much reduced and wavy-like, but they persist in strength and cross the venter. juvavites (a.) boehmi (gemmellaro 1904: pl. 27, fig. 9-10) also exhibits a reduction of ribbing on the body chamber, especially on the inner part of the lateral side. occurrence. no information is available regarding the age and stratigraphic position of juvavites (anatomites) inflatus as described by gemmellaro from western sicily. in this respect, the discovery of the single specimen from pizzo mondello provides at least a partial solution to its age. the new specimen was found in a level just slightly above the base of the gonionotites italicus subzone (spinosus zone, tuvalian 3, upper carnian) that is marked by the occurrence of microtropites in level pmam52a. genus gonionotites gemmellaro, 1904 type species: gonionotites italicus gemmellaro, 1904 preliminary remarks. gonionotites gemmellaro, 1904 has been recognized in several tethyan localities and north america, but despite a relative abundance of literature, the taxonomy of this genus at the species level is still complex. gemmellaro described eleven new species of gonionotites and two indeterminate taxa from the ‘‘cherty limestone’’ of sicily. since then, the genus has been recognized in the tethys himalaya (diener 1906 and 1908; wang & he 1976; krystyn 1982), northern alps (diener 1921; krystyn 1973 and 1980), iran (besse et al. 1998) and turkey (krystyn & gallet 2002), but it is worth noting that most of the species identified from these areas are different than those from sicily. the first report of gonionotites from north america was by smith (1927). afterwards a number of species, most of which are different from the tethyan faunas, were described from canada by mclearn (1940, 1947 and 1960) and tozer (1962). then, in 1994 tozer provided a comprehensive revision of the canadian gonionotites species. no recent literature is available from the united states apart from the illustration of two specimens by kristan-tollmann & tollmann (1983). tethyan gonionotites are not as well known as the north american faunas. krystyn (1982) described g. haugi gemmellaro, 1904, g. gemmellaroi diener, 1906, and g. noricus diener, 1921 from nepal. he also revised g. tingriensis wang & he, 1976 and transferred this species to malayites welter, 1914, based on its spiral strigations (krystyn 1982). however, no updates are available for most of the eleven species erected by gemmellaro, including the index species g. italicus. an examination of gemmellaro’s species and their comparison with other species from the tethys and north america suggests the following brief comments. a) as already mentioned for other ammonoid groups of gemmellaro’s collection, several of his species appear to be morphologically very close. surprisingly, a syntype of a given species is often more similar to a syntype of another species than to the other members of the same type-series. this is the case for gonionotites carnian-norian ammonoids from pizzo mondello 61 mojsisovicsi in gemmellaro’s pl. 21, fig. 1-3, which is more similar to the compressed inner whorls of the syntype of g. italicus in pl. 21 (fig. 4-6) than to the syntype of g. mojsisovicsi figured in pl. 13, fig. 1-2. b) many of gemmellaro’s species exhibit compressed to very compressed inner whorls with a subtriangular whorl section followed by a widening venter and a thickening of the whorl width on the body chamber. this modification of the whorl section on the mature body chamber is much less common in other tethyan and north american species. c) few sicilian species of gonionotites exhibits a thickening of the ribs on the widest part of the flank before the shoulder, which resembles radially elongated and forward concave nodes. these bulges are shown in g. recuperoi gemmellaro (pl. 7, fig. 43-44), g. diblasii gemmellaro (pl. 5, fig. 1-2) and possibly g. dubius gemmellaro (pl. 6, fig. 9-10), but they also occur in a few very compressed species that gemmellaro attributed to the subgenus anatomites, such as juvavites (anatomites) pulcher gemmellaro (pl. 5, fig. 3-4) and j. (a.) formosus gemmellaro (pl. 20, fig. 6-7). a clarification of these problems would require a population analysis of many bed-by-bed collected specimens, but thus far the scarcity of specimens from pizzo mondello does not allow such a revision. pending a thorough revision of the group, we can only compare our new specimens with gemmellaro’s taxonomy. gonionotites cf. italicus gemmellaro, 1904 pl. 2, fig. 1a-c v 2007 gonionotites maurolicoi nicora et al., pl. 2, fig. 6a-b. remark on the type series: gemmellaro (1904: 159) clearly stated that the new species was based on five well preserved specimens plus several fragments. seven specimens are housed in the museo gemmellaro collections, under registration number mgup 002. 147.1-4 and mgup 002.148.1-3. material: two specimens, mpum 10975 (pmam7-1) and mpum 10976 (pmam36-1). description. the two specimens attributed to gonionotites cf. italicus are of rather large size. one is slightly less than half one whorl (about 170º) of body chamber (estimated diameter of about 100 mm) with inner whorls visible in cross section, while the second is an incomplete fragment of less than 45º of body chamber. both specimens have a compressed, slender subtriangular whorl section with very narrowly rounded venter and a smooth surface. the most complete specimen (pl. 2, fig. 1) exhibits a gradual thickening of the whorl section that is not accompanied by a widening of the venter. at the end of the preserved part of the body chamber (pl. 2, fig. 1b-c), the venter is still relatively narrow, and the maximum width of the whorl section is located just slightly above onehalf of the whorl height. dimensions discussion. as emphasized in the preliminary remarks, the difference between some of gemmellaro’s species is rather obscure. we do not doubt that g. italicus, the first species described by gemmellaro, is a valid species, and accordingly, we place slightly more emphasis on this taxon than to the other species. specimen mpum 10976 is too incomplete for a fully accurate identification, while the attribution of the figured specimen (mpum 10975; pl. 2, fig. 1) requires additional discussion. comparison of this specimen with the gonionotites of gemmellaro’s collection leads us to conclude that only part of the body chamber of the new specimen is preserved, with about 70-90º missing. a second observation resulting from the examination of gonionotites of gemmellaro’s collection is that the thicknening of the whorl and widening of the venter usually take place on the last quarter of the whorl of the mature body chamber. apparently, this particular part of the whorl is not preserved in specimen mpum 10975. this specimen definitely exhibits a thickening of the whorl, but apparently its growth was not yet sufficient for the venter to become wider. instead its venter is still rather narrow at the end of the preserved part of the body chamber (pl. 2, fig. 1c). in this aspect, balini m., krystyn l., levera m. & tripodo a.62 plate 1 upper carnian and lower norian ammonoids from pizzo mondello. 1) placites sp. ind., specimen mpum 10966 (na49.1-1). 2) discotropites plinii (mojsisovics, 1893), specimen mpum 10967 (pmam171): a, lateral view, b enlargement showing branching of the ribs at the umbilical margin. 3) anatropites sp. ind., specimen mpum 10969 (pmam22bis). 4) anatropites sp. ind., specimen mpum 10968 (fnp112-1): a, lateral view, b) ventral view. 5) microtropites cf. paronai, specimen mpum 10970 (pmam52a-1): a, lateravl view; b, ventral view; c, apertural view. 6) metathisbites cf. affinis, specimen mpum 10971 (fnp145-1), a, lateral view; b, detail of the weak ventrolateral nodes. 7) hyattites cf. praefloridus, specimen mpum 10972 (fnp333), a) lateral view; b, view of the cross section. 8) projuvavites boehmi (gemmellaro), specimen mpum 10973 (pmam22-1), a, lateral view; b, ventral view. 9) projuvavites inflatus (gemmellaro), specimen mpum 10974 (pmam49-1), a, lateral view; b, lateral view after removal of the last part of body chamber; c, apertural view; d, ventral view; dots mark the constrictions. all specimens whitened with ammonium chloride except fig. 3. bar scale is 1 cm, that in lower right corner is for all the figures except for 2b and 6b. asterisk marks the last septum. carnian-norian ammonoids from pizzo mondello 63 the specimen seems to differ from g. italicus, but on the other hand, none of gemmellaro’s other type specimens appear to be close to specimen mpum 10975. the type specimen of g. discus (see 1904: pl. 13, fig. 3-6) has a body chamber that exhibits a flattening of the venter, a trend that is opposite with respect to the venter of mpum 10975. the type specimen of g. maurolicoi also has a narrow venter, but it is characterized by much thicker whorls, whose maximum width is very close to the umbilical margin (e.g., 1904: pl. 14, fig. 5); therefore, the previous attribution of specimen mpum 10975 to this species (nicora et al. 2007) is here emended. one of the syntypes of g. mojsisovicsi shows some similarity with the new specimen, with respect for its narrow venter, but unfortunately this feature is accompanied by a maximum whorl width very close to the umbilical margin, instead of at about 50 % of the whorl height. this syntype (mgup 002.153.2) was not figured by gemmellaro, and thus it cannot be considered to be especially representative of g. mojsisovicsi. according to gemmellaro (1904), the characteristic syntypes of this species include the large, much thicker specimen figured in pl. 13, fig. 1-2 with a more rounded venter than the present specimen, and the syntype in pl. 21, fig. 1-3, with a broad venter and maximum whorl width located at about ½ of whorl height. occurrence. both specimens were collected from levels pmam36 and pmam7. the first level is located about 3 m below the occurrence of microtropites cf. paronai, which presently marks the lower boundary of the gonionotites italicus subzone. the second specimen was collected from the middle part of the g. italicus subzone. gonionoites aff. recuperoi gemmellaro, 1904 pl. 2, fig. 2a-b; fig. 9 material: one specimen mpum 10977 (pmam43-1). description. specimen, which is preserved as an internal mold partly covered by recystallized test, consists of a phragmocone with about 270º of body chamber. whorl section is compressed and slender for first 160º of body chamber, but then venter rapidly widens and maximum width of whorl section moves from the umbilical margin to 1/3 of whorl height. a weak constriction is visible at beginning of the body chamber. ribbing follows pattern typical of gonionotites. on visible part of the phragmocone, ribbing consists of extremely weak prorsiradiate, primary ribs on the flank, and three to four secondary ribs are visible very close to the shoulder. on body chamber, ribs are barely visible and noticeable only on the outer half of the whorl. these ribs are widely spaced (about 4-5 ribs in 90º) and develop node-like elongated thickenings very close to the shoulder, which tend to follow the course of the ribs, first radially, but then bending forward and fading on the venter. suture line is ammonitic (fig. 9), with well-developed gonionotites-like indentations. three sutures are visible and show a serration. dimensions discussion. the specimen, attributed to gonionotites, agrees well with gemmellaro’s interpretation of the genus. an analysis of the taxonomic significance of the thicknening of the ribs on the outer part of the flank may be important, but would require a much larger collection of specimens. at the species level, the specimen from level pmam43 is referred by aff. to g. recuperoi gemmellaro, 1904 because this particular species exhibits the balini m., krystyn l., levera m. & tripodo a.64 fig. 9 suture line of gonionotites aff. recuperoi, specimen mpum 10977 (pmam 431). bar scale 5 mm. thickening whorl section and widening of the venter at a size that is comparable to the size of the pizzo mondello specimen. the new specimen from pizzo mondello, however, shows the thickening of ribs on the outer part of the flank at a larger size with respect to the only figured syntype of g. recuperoi, after the weak ventral ribs have disappeared. g. diblasii gemmellaro, 1904 reveals a ventral thickening of the whorl section that occurs at much larger size, while the ‘‘nodes’’ of g. dubius gemmellaro, 1904 are only barely visible on the figured syntype. occurrence. gonionotites recuperoi gemmellaro has not been found since gemmellaro’s (1904) time and it is unknown outside of western sicily, where its age might span from late carnian to early norian. the herein described specimen was collected from a level slightly above the younger of the two anatropites sp. ind. collected thus far from the pizzo mondello section. the co-occurrence of halobia radiata (cf. levera 2012) suggests a late carnian age. pregriesbachites gen. n. type species: juvavites (anatomites) bukowskii gemmellaro, 1904 etymology: from the suffix ‘‘pre’’ and griesbachites. the compound name emphasizes the evolutionary link with the genus griesbachites mojsisovics, 1896. diagnosis: small to middle-sized, relatively slim juvavitids with highly differing sculpture from juvenile to adult stage. phragmocone bears prorsiradiate, paulostome constrictions and multi-branching, externally alternating or interrupted ribs. body chamber is characterized by fading ribs and development of numerous pointed ventrolateral nodes. suture line projuvavitid with low and medium indented saddles similar to griesbachites. composition of the genus: juvavites (anatomites) bukowskii gemellaro, 1904 (type), juvavites timaei gemmellaro, 1904, griesbachites auctoris tozer, 1994, g. pinensis tozer, 1994, g. laevis tozer, 1994, g. humi (mclearn, 1937) and g. selwyni (mclearn, 1940). remarks. the new genus pregriesbachites formalizes a taxonomic entity that was outlined as ‘‘nov. gen. (= projuvavites mit wohnkammerknoten)’’ (=projuvavites with nodes on body chamber) by krystyn (1982: 46, 59-60). the previous interpretation of this taxon as a descendent of projuvavites and forerunner of griesbachites is here confirmed. comparisons. pregriesbachites has inner whorls similar to projuvavites tozer, 1971, but is distinguished from that genus by the presence of ventrolateral nodes on its body chamber. the shape and size of the nodes permits separation from the slightly younger but timely overlapping genus griesbachites mojsisovics, 1896. these nodes are rounded and pointed in pregriesbachites n. gen., whereas in griesbachites, they are larger and clavate. guembelites mojsisovics, 1896 lacks constrictions and bears nodes in a more external position on the phragmocone as well as on the body chamber. the inner whorls of pregriesbachites may show similiarity with those of anatomites. however, the type species, a. rotundus mojsisovics, 1893, brings to mind the morphology of the inner whorls of griesbachites and thereby, is suspected of representing the phragmocone stage of the latter genus (krystyn 1982: 46). occurrence. the new genus is documented in the tethys realm and north america. in canada the species herein included in pregriesbachites n. gen. document a latest carnian, late tuvalian 3 macrolobatus zone to early norian kerri zone range. an equivalent stratigraphic distribution (g. italicus subzone of late tuvalian 3 to jandianus zone of lacian 1 is known for the tethys (krystyn 1982: 58). at feuerkogel pregriesbachites n. gen. was collected from level iv of f4 section (fig. 10). the pizzo mondello occurrence (see description of species) most likely is from the gonionotites italicus subzone, spinosus zone. pregriesbachites bukowskii (gemmellaro, 1904) pl. 2, fig. 3a-b, 4a-b v 1904 juvavites (anatomites) bukowskii gemmellaro, p. 224, pl. 11, fig. 7-8. ?v 1904 juvavites (anatomites) bukowskii gemmellaro, p. 224, partim. stratum typicum and locus typicus: whitish limestone with cherty nodules, ‘‘feudo modanesi’’ near castronuovo (gemmellaro 1904: 225). type series: gemmellaro (1904: 225) cited seven specimens, but eight specimens are deposited in the museo geologico gemmellaro, under registration number mgup 002.193.1-8. gemmellaro figured only one of the type specimens (pl. 11, fig. 7-8). this specimen, which is much better preserved than the other four large sized types, is here designated as lectotype. material: one specimen mpum 10986 (pmam42-1). description. specimen is preserved as an internal mold that retains small patches of recrystallized test. body chamber appears to occupy about 260º of whorl from last visible suture line. very involute coiling platicone (u/d~0.06). whorl section, with maximum width very close to umbilical margin, exhibits a gradual change in the degree of compression combined with a very slight modification in the outline. whorl section at end of phragmocone is semielliptical, with convex lateral sides and h/w ratio = 1.50. just slightly past beginning of body chamber, the h/w ratio increases to 1.59, flanks become more flatter and shoulders become much more distinct. towards the end of the body chamber, whorl section becomes more inflated and h/w decreases to 1.48. this variation is even more evident when viewing the specimen because the appearance of two more distinctive shoulders on the body chamber makes the venter appear to be even wider. carnian-norian ammonoids from pizzo mondello 65 two constrictions are visible, one about 35º before the end of the phragmocone and the other on the body chamber about 100º after the first. ornamentation consists of slightly concave primary and secondary ribs with a general projuvavites-like organization between the constrictions. ribs are developed between the umbilical margin and the shoulder on the first part of the last whorl before the first constriction. first branching of ribs occurs very close to the umbilical margin, while the second subdivision is located near the middle of the flank. between the two constrictions, there are four widely-spaced weak primary ribs, each followed by four to five secondary ribs that are developed only ventrolaterally. ventral ribs also continue on the body chamber and all ribs cross the venter. ventrolateral rounded nodes, which appear on the body chamber at whorl height of about 28 mm are located on flank, very close to the ventral shoulder. suture line ammonitic. dimensions discussion. the type series consists of eight specimens (mgup002.193.1-8) of different size. four of them are very small, and their size does not exceed the innermost part of the last whorl of the four medium to large sized individuals (mgup002.193.1-4). since we have no evidence that the four small-sized specimens truly represent the juvenile growth stage of the larger individuals, these specimens are separated in the synonymy. the single specimen figured by gemmellaro, designated as lectotype, is much better preserved and fits quite well with the original figure. compared to the new specimen from pizzo mondello, the lectotype is balini m., krystyn l., levera m. & tripodo a.66 fig. 10 ammonoid and halobia record at feuerkogel sections f4 and f5 (determination by lk). preliminary data for these sections were previously reported by krystyn (1973 and 1980). plate 2 upper carnian ammonoids from pizzo mondello. 1) gonionotites cf. italicus, specimen mpum 10975 (pmam7-1), a, lateral view; b, ventral view; c, apertural view. 2) gonionotites aff. recuperoi, specimen mpum 10977 (pmam43-1), a, lateral view; b, ventral view. 3) pregriesbachites bukowskii (gemmellaro), specimen mpum 10986 (pmam42-1), a lateral view; b, ventral view. 4) pregriesbachites bukowskii (gemmellaro), lectotype mgup 002.193.1, a lateral view; b, ventral view. all specimens whitened with ammonium chloride except fig. 4. bar scale is 1 cm for all specimens. asterisk marks the last septum. carnian-norian ammonoids from pizzo mondello 67 slightly more compressed, but exhibits the same variation in the whorl section, especially on the body chamber, with nearly flat flanks meeting the shoulder on each side of the rounded venter. the ribbing pattern is also the same, but the nodes are already obvious at the beginning of the body chamber. occurrence. this is the first report of this species since gemmellaro’s 100 plus year old description. therefore, the stratigraphic position of pregriesbachites bukowskii (gemmellaro) must rely on the present specimen from pizzo mondello. this specimen is from level pmam42, which also yields halobia radiata and rare h. lenticularis (levera 2012). since it was found slightly below the second level with anatropites (pmam22bis), p. bukowskii is thus referred to the upper carnian gonionotites italicus subzone, spinosus zone. genus dimorphites mojsisovics, 1893 type species: juvavites (dimorphites) selectus mojsisovics, 1893 dimorphites noricus n. sp. pl. 3, fig. 1-5 v 1980 dimorphites n. sp. 1 krystyn, p. 73. v 2002 dimorphites n. sp. 1 krystyn et al., p. 344, fig. 1. derivatio nominis: from noricus, -a, -um. the name emphasizes the stratigraphic position of the species as a marker for the basal part of the norian stage. stratum typicum and locus typicus: hallstatt limestone, feuerkogel (northern alps, austria), section f5, bed iii (krystyn 1980, fig. 12; fig. 10). material: type series composed of four specimens. holotype gba 2011/055/0001; paratypes gba 2011/055/0002-0004. one additional specimen from pizzo mondello is attributed to the new species mpum 10978 (na42.1). diagnosis: relatively compressed dimorphites with subdiscoidal inner whorls and semi-circular venter on the mature body chamber. ornamentation consists of wide, flat ribs separated by narrow, furrowlike inter-spaces. description. since all available specimens are preserved with the test, the description is based on features visible on outer surface of shell. specimen from pizzo mondello compares well with paratype gba 2011/055/ 0002 that is most similar. coiling very involute (u/d decreases with growth from 7.85 to 5.4) with medium sgr. whorl section compressed (h/w increases during the growth from 1.47 to 2.29) with maximum width very close to the umbilical margin. flanks slightly convex, slowly converging to narrow venter. general shape of shell is subdiscoidal until first half of adult body chamber, at which point venter begins to widen. ornamentation consists of rather wide, flat ribs separated by narrow, furrow-like inter-spaces. at all stages of growth, ribs begin slightly above umbilical margin, which is rounded and smooth. innermost whorls differ from medium to large sized individuals by wavy-like ribbing. at whorl height less than 7-7.5 mm (specimens gba 2011/055/0002 and 0003; mpum 10978: pl. 3, fig. 1,2 and 5), ribs are slightly round rather than flat and about 1.5 times wider than the inter-rib spaces. at whorl height greater than 7.5 mm, ribsgradually become flat and wider than the inter-space furrows. ribs are sinuous (falcoid sensu arkell et al. 1957), with a rather peculiar course. flat primary ribs begin just above the umbilical margin and gradually become wider. bifurcation of primary ribs occurs at the beginning of a narrow furrow in the middle of rib, which is often very weak, but after a few millimeters gradually reaches depth and width of furrows delimitating primary rib. in such case, it is more common for the subdivisions to occur at the middle of the flank, the branching point of the secondary rib can be located only with some approximation. subdivision of ribs mainly occurs at two positions on the flank. the first is between the starting point of the primary rib and about 10% of whorl height. the second position is about 40 to 60% of whorl height, but may change from one rib to the next. occasionally, additional branching may also occur on the external part of the flank, at about 70-75% of whorl height. all ribs are projected forward slightly at the ventral shoulder, but then fade higher on the shoulder in an alternating manner such that an undulating effect is imparted to the course of the relatively narrow subtabulate venter. this feature is typical of the genus dimorphites. for medium to largesized specimens the frequency of ribs per half whorl is rather constant at about 33-35 ribs at the ventral shoulder. suture line not exposed on the type specimens, except for one single ammonitic 1st lateral saddle visible on one side of paratype gba 2011/055/0002. dimensions remarks. the description of dimorphites noricus n. sp. formalizes a species previously mentioned in the literature (krystyn 1980) as dimorphites n. sp. 1, and designated as an index ammonoid for the first norian ammonoid subzone (cf. krystyn et al. 2002). discussion. with respect to its coiling and pattern of rib subdivision, dimorphites noricus n. sp. is morbalini m., krystyn l., levera m. & tripodo a.68 phologically so close to the stratigraphically younger d. selectus mojsisovics, 1893 that the two species may represent an evolutionary lineage. despite d. selectus being slightly more compressed than d. noricus, the main difference between the two species is their ribs, which at all growth stages are much wider and flatter in d. noricus n. sp. than in d. selectus. accordingly, the frequency of ribs per half whorl is distinctly lower in d. noricus than in d. selectus. the rib frequency on the type specimens of d. noricus n. sp. compared to that of the ype specimens of d. selectus is as follows: a) small-sized specimens: 24/25 ribs (gba 2011/ 055/0002: pl. 3, fig. 1) versus 35 (mojsisovics 1893, pl. 127, fig. 2a, d= 14mm), b) small medium-sized specimens: 32 ribs (gba 2011/055/0003: pl. 3, fig. 2) versus 53 (mojsisovics 1893, pl. 127, fig. 5a, d= 27mm), c) medium to large-sized specimens: 33-35 ribs (gba 2011/055/0001 and 0004: pl. 3, fig. 3 and 4) versus 41-54 (mojsisovics 1893, pl. 127, fig. 3a [lectotype]: 41 at d= 30mm; fig. 4a: 46 at d= 41mm, 54 at d= 50mm). occurrence. the species has been recognized at feuerkogel (hallstatt, northern alps), section f5, level iii, and at pizzo mondello (sicani mountains, western sicily) level na42. with regard to its chronostratigraphic position at feuerkogel, this species was selected as the index ammonoid for the first subzone of the jandianus zone (lacian 1: krystyn 1980, 1982), which is the first norian ammonoid zone of the tethyan chronostratigraphic scale. the occurrence at pizzo mondello is also of earliest norian age. dimorphites selectus mojsisovics, 1893 pl. 3, fig. 6-7 1893 juvavites (dimorphites) selectus mojsisovics, p. 145, pl. 127, fig. 1-9. v 1904 juvavites (dimorphites) mariae gemmellaro, p. 243, pl. 17, fig. 19-20. 1921 dimorphites selectus – diener, p. 487. 1925 dimorphites selectus diener, p. 65, pl. 16, fig. 9. ? 1940 juvavites (dimorphites?) pardonetiensis mclearn, p. 48, pl. 1, fig. 13. ? 1960 dimorphites pardonetiensis – mclearn, p. 92, pl. 16, fig. 1-2. v 1982 dimorphites selectus – krystyn, p. 45, pl. 11, fig. 6. ? 1994 dimorphites pardonetiensis mclearn, tozer, p. 240, pl. 114, fig. 1-3, text-fig. 91c. v 2007 dimorphites sp. ind. – nicora et al., pl. 2, fig. 7. stratum typicum: hällstatter kalk, at feuerkogel (salzkammergut, austria). lectotype: original of mojsisovics 1893, pl. 127, fig. 3, gba1893/01/131, designated by diener 1925:65). material: ten specimens. three are relatively complete: mpum 10979 (fnp145-19), mpum 10980 (fnp145-20) and mpum 10981 (det-1). seven are fragmentary: mpum 10982 (fnp144-2); mpum 10983 (fnp144b-1); mpum 10984 (fnp145-2, -7, -8, -14, -21). description. the three best preserved specimens are of medium size (~30u), an elliptical whorl section and a suture line with triphyllic symmetrical saddles clearly different with respect to the asymmetrical d. patenslike saddles figured by mojsisovics (1896: pl. 19, fig. 6). an additional difference in the suture line of diecarnian-norian ammonoids from pizzo mondello 71 ner’s specimen is its very deep external lobe, which differs notably from the external lobe of d. patens and d. insignis. however, this particular portion of the suture line is not preserved on the type specimen figured by mojsisovics (1896); hence a comparison is impossible. we cannot exclude that it may be possibile to synonymize d. insignis gemmellaro, 1904 with d. ebneri mojsisovics, 1896 but, pending a revision of the latter, we prefer to keep d. insignis gemmellaro, 1904 separated. occurrence. the species occurs in sicily (gemmellaro 1904), but it has also been collected from the tuvalian 3 at feuerkogel in the northern alps (krystyn 1973: 120; fig. 10), in the spinosus zone. both new specimens from pizzo mondello were collected from levels pmam52 and fnp166. level pmam52 most likely belongs to the gonionotites italicus subzone, spinosus zone, tuvalian 3 upper carnian because of the occurrence of microtropites in the immediately overlying level, while fnp166 is located within the range of halobia mediterranea, a species usually referred to the paulckei zone (lacian 2) (see ammonoid bio-chronostratigraphy and levera 2012). ammonoid chronostratigraphy bio-chronostratigraphic subdivision of the pizzo mondello section ammonoids collected from the pizzo mondello section include short ranging age-diagnostic forms as well as long ranging taxa. among the first group the most interesting taxa, in stratigraphical order, are discotropites plinii; anatropites, microtropites, dimorphites noricus n. sp. and dimorphites selectus. gonionotites would also be of some interest, but only on the species level. this genus is reported in the literature from the last zone of the carnian to the first zone of the norian (krystyn 1982; tozer 1994); this long range is in part confirmed by the distribution at pizzo mondello, where it has been identified in levels fnp80a, pmam36, pmam54, pmam7, pmam42, pmam43, pmam35, pm28a and pm 28d (fig. 6). these short-ranging taxa document all four subzones of the uppermost carnian and lowermost norian and firmly tie the carnian/norian boundary succession under study to the tethyan chronostratigraphic scale. ammonoid rarity does not allow for continuous zonation of the section, but the discrete intervals with ammonoid-controlled chronostratigraphic standardization are presented below (from bottom to top; fig. 6), together with pertinent comments on the intermediate intervals without age diagnostic ammonoids. – the lower 25 m of the section has thus far yielded only very small ammonoids, including one sandlingitidae, one arcestidae and hyattites cf. praefloridus. the stratigraphic significance of hyattites has not yet been defined as the genus is rare and the literature quite inconclusive (see systematic paleontology). those levels yielding sandlingitidae and h. cf. praefloridus are referred to the upper carnian, and the single level with hyattites may well represent tuvalian 3 due to its close relationship with gonionotites. discotropites plinii subzone, spinosus zone (tuvalian 3, i) the index ammonoid d. plinii was identified in level pmam17 about 25 m above the base of the section. projuvavites boehmi was collected from level pmam27, just slightly above the level with d. plinii. however, the stratigrahic position of p. boehmi is not defined with respect to a particular bio-chronostratigraphic scale, so only level pmam17 is definitely referred to the discotropites plinii subzone. gonionotites italicus subzone, spinosus zone (tuvalian 3, ii) this subzone is characterized by microtropites and anatropites. microtropites occurs only in the g. italicus subzone (krystyn 1982) while anatropites first occurs in the underlying discotropites spinosus subzone (krystyn 1982). the anatropites specimens from pizzo mondello were found in beds overlying level pmam52a, which yields microtropites aff. paronai; hence, these levels are useful for tracing the upper boundary of the g. italicus subzone. based on our available collection, the boundaries of the spinosus zone are drawn at balini m., krystyn l., levera m. & tripodo a.72 plate 3 upper carnian and lower norian ammonoids from pizzo mondello and feuerkogel (northern alps, austria). 1) dimorphites noricus n. sp., paratype gba 2011/055/0002, lateral view. 2) dimorphites noricus n. sp., paratype gba 2011/055/0003, lateral view. 3) dimorphites noricus n. sp., holotype gba 2011/055/0001, a, lateral view; b, apertural view. 4) dimorphites noricus n. sp., paratype gba 2011/055/ 0004, a, lateral view; b, apertural view. 5) dimorphites noricus n. sp., specimen mpum 10978 (na42.1), lateral view. 6) dimorphites selectus mojsisovics, specimen mpum 10981 (det-1), lateral view. 7) dimorphites selectus mojsisovics, specimen mpum 10979 (fnp145-19), lateral view. 8) dimorphites sp., specimen mpum 10985 (pm36.1), lateral view. 9) discophyllites insignis gemmellaro, specimen mpum 10987 (pmam52-1), lateral view. all specimens whitened with ammonium chloride except fig. 5. bar scale is always 1 cm, that in the lower right corner is for all the figures except for 1, 5 and 8. asterisk marks the last septum. carnian-norian ammonoids from pizzo mondello 73 the base of level pmam52a and at the top of level pmam22bis. the fauna includes frequent gonionotites sp., as well as gonionotites cf. italicus, discophyllites insignis (gemmellaro), projuvavites inflatus (gemmellaro), pinacoceras sp. and pregriesbachites n. gen. bukowskii (gemmellaro). – the interval between level pmam22bis and na42.1 yielded very few ammonoids, especially from the lower and very uppermost parts. gonionotites cf. recuperoi was collected less than 1 m above the last occurrence of anatropites and a small tropiceltites specimen was collected one-half meter below the level yielding dimorphites noricus n. sp. no stratigraphic information is available for g. recuperoi gemmellaro and the bio-chronostratigraphic significance of tropiceltites is not fully clear. in north america tozer (1994) reported the genus from the macrolobatus to kerri zones (upper carnian-lower norian). very little information is available from the tethyan successions apart from the report by krystyn (1973) concerning the lacian 1 jandianus zone at feuerkogel and a new identification from the top of spinosus zone at f4 (fig. 10). because of the lack of reports in the literature, we do not consider tropiceltites to be a marker genus. dimorphites noricus n. sp. subzone, jandianus zone (lacian 1 i) the index species occurs only in level na42, at about 99.5 m above the base of the section. dimorphites selectus subzone, jandianus zone (lacian 1 ii) dimorphites selectus, index species of the subzone, was recorded from levels fnp144, fnp144b and fnp145, which range from about 99.5 to about 102 m above the base of the section, in direct stratigraphic succession with respect to the dimorphites noricus n. sp. subzone. this subzone also includes metathisbites cf. affinis gemmellaro. – level na43, located about 2 m above the last occurrence of dimorphites selectus yielded one cross section that could possibly be referred to tropiceltites. this occurrence may indicate a lacian age since the genus has not been recorded from younger beds. paulckei zone (lacian 2) dimorphites sp. ind., discophyllites insignis (gemmellaro) and placites sp. ind. occur in levels fnp166 and na49.1. these taxa are not bio-chronostratigraphically significant, but they occurr with halobia mediterranea (see levera 2012), which is considered as a good marker for the paulckei zone (krystyn 1980; krystyn et al. 2002). chronostratigraphic correlations of the pizzo mondello section based on ammonoids ammonoid-bearing successions documenting the latest carnian through the earliest norian are very rare in the world and only a few have been studied utilizing the bed-by-bed sampling method. in the tethys realm, pizzo mondello can be compared and correlated with a small number of localities in the salzkammergut area, above all with feuerkogel, and with jomsom in the tethys himalaya. in north america the most important localities are west union canyon (shoshone range, central nevada) and black bear ridge on williston lake (british columbia, canada). there are a few localities in northeastern russia that are of interest for comparison with the boreal realm, especially in the yana okhotskaya river basin. with respect to such a global approach to correlation, it is obvious that the faunal composition of the different sites will be strongly influenced by provincialism (see tozer 1981b, 1994; dagys 1988; zakharov 1997; konstantinov 2008), but minor changes are also the result of paleoecologic control. tethys realm tethyan successions have provided huge collections of upper carnian to lower norian ammonoids, especially from the hallstatt red limestone facies of the northern alps and timor, which were first documented in the literature in the mid 19th century. upper triassic successions from the himalayas have also contributed significantly to advance the knowledge of ammonoid taxonomy, notwithstanding the fact that very few sites have been subjected to the bed-by-bed sampling approach. the best site in the northern alps is feuerkogel, in the salzkammergut area (krystyn 1973 and 1980; fig. 10), and in the himalayas it is jomsom in the kali gandaki (nepal; krystyn 1982). a limited quantity of ammonoid data are available from sections encompassing the carnian/norian boundary in antalya, turkey (kavaalani: gallet et al. 2000; erenkolu mezarlik 2: krystyn & gallet 2002). other localities have recently provided stratigraphically controlled collections, but these are only upper carnian (eastern southern alps: de zanche et al. 2000; gianolla et al. 2003) or lower norian (alborz, iran: seyed-emami et al. 2009). interesting upper carnian faunas have also been described in the 1970s from okinawa, japan (ishibashi 1970, 1973 and 1975). correlation of the pizzo mondello ammonoid record with the record documented at feuerkogel f5 and jomsom is illustrated in fig. 12. at both pizzo mondello and jomsom, the successions are expanded, whereas at feuerkogel the hallstatt limestone is condensed. this notwithstanding, the discotropites plinii, gonionotites italicus, dimorphites noricus n. sp. and dimorphites selectus subzones are easily correlated by the balini m., krystyn l., levera m. & tripodo a.74 carnian-norian ammonoids from pizzo mondello 75 f ig . 1 2 a m m o n o id re co rd o f p iz z o m o n d el lo se ct io n co m p ar ed to th e re co rd o f f eu er k o g el f 5 se ct io n (h al ls ta tt l im es to n e, n o rt h er n a lp s, a u st ri a; k ry st y n 1 9 8 0 ) an d jo m so m (t h in ig ao n f o rm at io n , n ep al ; k ry st y n 1 9 8 2 ). jo m so m lo g is b as ed o n th e ra n g e ch ar t o f k ry st y n ’s fi g . 4 , b u t th e sc al e is ad d ed h er e. f eu er k o g el f 5 se ct io n is d es cr ib ed in k ry st y n (1 9 8 0 ) an d it h as b ee n in cl u d ed in th is co rr el at io n s ch ar t b ec au se it is th e ty p e lo ca li ty o f d im o rp h it es n o ri cu s n . sp . f eu er k o g el f 4 re co rd is m o re co m p le te an d sh o w n in f ig . 1 0 . t h e m o st si g n if ic an t ag ed ia g n o st ic ta x a ar e em p h as iz ed in b o ld . s ee te x t fo r d is cu ss io n . l ab el s t 1 , t 2 an d t 3 re fe r to co n o d o n t fa u n al tu rn o v er s d es cr ib ed b y m az z a et al . (2 0 1 0 ). occurrence of nominal species, or by the occurrence of genera (e.g., microtropites), whose range is limited to a single subzone. when documenting the wide paleogeographic distribution and correlation of certain ammonoid groups such as discotropites, anatropites, microtropites and dimorphites over large distances, it is not surprising that some other groups show some differences. the pizzo mondello fauna lacks certain genera, such as griesbachites and guembelites, that are well documented not only at feuerkogel and jomsom, but also in other tethyan successions of the himalayas (bordet et al. 1971), tibet (wang & he 1976), timor (welter 1914) and alborz (seyed-emami et al. 2009). even though faunal distribution is quite wide from a paleogeographic point of view, the frequency of biostratigraphically complete successions is rather low. not even one of the three sections illustrated in fig. 12 includes a direct superposition of lowermost norian ammonoids on uppermost carnian faunas. at pizzo mondello, the uppermost carnian g. italicus and the lowermost norian d. noricus n. sp. subzones are documented, but the actual carnian/norian boundary falls within an interval of about 18 meters without age diagnostic ammonoids. at feuerkogel f5, level iii is the stratum typicum of dimorphites noricus n. sp., but g. italicus occurs only in the underlying level iv(fig. 10 and 12), thereby demonstrating the high rate of sedimentary condensation. at jomsom the unfossiliferous interval between the latest carnian level 63 (g. italicus subzone) and earliest norian level 78 (jandianus zone, dimorphites selectus subzone) is more than 55 m thick and unfortunately lacks documentation of the d. noricus n. sp. subzone. west union canyon (central nevada, u.s.a.) the most important ammonoid locality in north america for the carnian/norian boundary is west union canyon in the shoshone range (central nevada). this is the type locality of both the schucherti and macrolobatus zones, the two latest carnian ammonoid zones of the north american triassic standard scale (silberling 1959; silberling & tozer 1968). at this locality there is also a good record of the earliest norian kerri zone, which was actually defined at brown hill (peace river valley, british columbia; tozer 1965, 1967). ammonoid faunas from west union canyon were described by silberling (1959), who also provided a good description of the sedimentary succession (luning formation). silberling collected ammonoids and carefully documented the faunal changes from bed to bed (at least for the macrolobatus zone), but unfortunately he did not include in his paper the log with the bed-by-bed distribution of the faunas. this log provided the basis for a preliminary sampling of the site carried out in 2010 (balini et al. 2011), which resulted in the collection of the same faunas reported by silberling in 1959. the ammonoid record of the luning formation is not continuous and fossil rich intervals are separated by shaly intervals without macrofossils, at least on the basis of surface sampling. ammonoid faunas of the schucherti and macrolobatus zones are dominated by tropitidae (tropites and anatropites), but they lack juvavitinae. at pizzo mondello section, the faunal composition of this particular part of the carnian is reversed, with juvavitinae (projuvavites, gonionotites, pregriesbachites) dominant over tropitidae (hoplotropites [from literature], discotropites, anatropites, microtropites). ammonoids of the kerri zone include of guembelites and stikinoceras, but dimorphites apparently is not present. as previously emphasized in the discussion regarding correlation with tethyan localities, guembelites has not been reported from the scillato formation, and gonionotites has not been found at west union canyon, neither by silberling (1959) nor by balini et al. (2011). thus, direct correlation between west union canyon and pizzo mondello is possible for the upper carnian, but not for the lower norian, at least on the basis of the available ammonoid data. black bear ridge (british columbia, canada) the peace river valley (british columbia, canada) is an important area for middle to upper triassic ammonoids. several localities were found and first described by mclearn from the 1940s through the early 1960s, and then tozer spent the next 40 years or so expanding the ammonoid collections, improving their taxonomy and revising the biostratigraphy of the area. although construction of the wac bennet dam (1968) and the subsequent flooding of the valley (1970s) destroyed several of mclearn and tozer’s paleontologic sites, the new shorelines of this vast lake have provided the opportunity to collect from new and perfectly washed triassic exposures, often of tens of kilometers in lenght. black bear ridge, on the northern shore of the lake, is the best locality for study of the carnian/ norian boundary because it is the only site in the williston lake area, where the succession encompassing the boundary is not affected by a lithologic change (zonneveld et al. 2010; cf. mcroberts 2011: fig. 3). at the wellknown pardonet hill site on the southern lake shore, just in front of black bear ridge, the carnian-norian boundary nearly coincides with the lithologic boundary between the baldonel formation and the pardonet formation (zonneveld & orchard 2002). a similar setting is documented at brown hill, on the northern shore of the lake, east of black bear ridge (zonneveld & orchard 2002). balini m., krystyn l., levera m. & tripodo a.76 carnian-norian ammonoids from pizzo mondello 77 f ig . 1 3 a m m o n o id re co rd o f th e c ar n ia n /n o ri an b o u n d ar y in te rv al at p iz z o m o n d el lo co m p ar ed to th e re co rd at b la ck b ea r r id g e (w il li st o n l ak e, b ri ti sh c o lu m b ia , c an ad a) . b ed d in g o f th e tw o se ct io n s is to sc al e. b la ck b ea r r id g e se ct io n is fr o m m cr o b er ts (2 0 1 1 : fi g . 9 ). t h e b b r le v el n u m b er s re fe r to in te rv al s o f b ed s. e v er y in te rv al is fu rt h er su b d iv id ed in to b ed s, ea ch o f w h ic h is id en ti fi ed b y a le tt er . t h e fi el d p ic tu re (r ig h t) sh o w s th e b ed s o f th e c ar n ia n /n o ri an b o u n d ar y in te rv al . m cr o b er ts & k ry st y n (2 0 1 1 ) p ro p o se d to d ef in e th e b as e o f th e n o ri an at th e b as e o f le v el 1 8 f , w h er e th e f o o f th e su g g es te d p ri m ar y m ar k er h a lo b ia a u st ri a ca is re co rd ed . a m m o n o id d et er m in at io n s fr o m b b r se ct io n b y l k . in b o th se ct io n s, am m o n o id s o cc u r in d is cr et e le v el s. a t p iz z o m o n d el lo , an in te rv al o f ab o u t 1 8 m et er s w it h o u t ag ed ia g n o st ic am m o n o id s ex is ts b et w ee n th e la st o cc u rr en ce o f la te c ar n ia n a n a tr o p it es an d th e fi rs t o cc u rr en ce o f n o ri an d im o rp h it es n o ri cu s n . sp . a t b la ck b ea r r id g e an in te rv al o f ab o u t 4 .5 m et er s ex is ts b et w ee n th e le v el y ie ld in g d ef in it e la te c ar n ia n a n a tr o p it es cf . m a cl ea rn i an d th e le v el p ro v id in g d ef in it e ea rl y n o ri an g u em b el it es cl a v a tu s. n o g o o d ag ed ia g n o st ic ta x a ar e d o cu m en te d w it h in th is in te rv al ; se e te x t fo r d is cu ss io n o f th e si g n if ic an ce o f p te ro si re n it es . ammonoids from black bear ridge (gsc collection, vancouver and a new, small collection 2010, herein classified by lk) are rather meager, documented by only few specimens. however, their stratigraphic position permits an accurate correlation (fig. 13). two ammonoids from levels 16a and 22b tie the succession to the north american chronostratigraphic scale and at the same time, allow for correlation with the tethyan scale. from the lower level (16a) a single specimen of anatropites cf. maclearni documents the uppermost carnian macrolobatus zone. level 22b is referred to the discostyrites ireneanus subzone, kerri zone (correlative with the lower jandianus zone of the tethys) based on the occurrence of guembelites clavatus, a species which, according to tozer, is restricted to this subzone. the interval between these two levels is not barren of ammonoids, but thus far, it has only yielded gonionotites and tropiceltites, both of which range through the carnian/norian boundary and pterosirenites, the only boreal element in the black bear ridge ammonoid fauna (fig. 13). this genus, introduced by tozer in 1980 from the kerri zone of northeastern british columbia, has been recognized since 1981 as a typical component of the norian boreal assemblage (tozer 1981b; dagys 1988; see below). at black bear ridge pterosirenites has been identified both below and above (18d and 18g) the fo of h. austriaca (mcroberts 2011; mcroberts & krystyn 2011; level 18f: fig. 13). however, the chronostratigraphic position of pterosirenites is not yet fully defined. in the boreal realm (see below) ammonoid faunal composition is very different with respect to that of british columbia, and the c/n boundary is based on an ammonoid faunal change that is different with respect to the faunal changes recorded in north america and the tethys realm. if the problem is approached from the opposite side of the panthalassa ocean, pterosirenites occurs in four localities in northeastern british columbia, including the williston lake shorelines (eastern british columbia) and mount mclearn (toad river area, northern british columbia), but even here its range is not well defined. according to its documented record, pterosirenites is not abundant and at the few lower norian sites (e.g. brown hill, type locality of the kerri zone) from which it has been reported. these occurrences are somewhat scattered (fig. 14), but more importantly, there is no record of uppermost carnian ammonoids underlying pterosirenites at these four localities. apparently the ammonoid record at these four sites has been influenced by the facies change that occurred at the boundary of the baldonel-pardonet formations. boreal realm ammonoid correlation of the tethyan successions with those of the boreal realm are very difficult because of their significantly different faunal composition, but it is possible to directly link the boreal successions to northeastern british columbia by the common occurrence of pterosirenites. the carnian/norian successions of northeastern russia are dominated by sirenitinae, with the minor contribution of a few additional genera from different families (dagys 1988; bychkov 1995; zakharov 1997; konstantinov 2008). carnian faunas are composed of yakutosirenites, neosirenites, ‘‘striatosirenites’’, sirenites, proarcestes, arctophyllites, arctoarpadites and obruchevites, while norian ammonoids include ‘‘striatosirenites’’, pterosirenites, norosirenites, wangoceras, yanotrachyceras, arctophyllites, arcestes and pinacoceras. based on this very general summary, the carnian/norian boundary is marked (konstantinov 2008) by the disappearance of sirenites, neosirenites, yakutosirenites and proarcestes, and by the first occurrence of pterosirenites, arcestes and pinacoceras. the best successions are located in the primorye region (summary in zakharov 1997) and the yana okhotskaya river area (bychkov 1995). while the former is characterized by a few tethyan elements (zakharov et al. 1996; zakharov 1997), boreal faunas are best developed in the latter, which also has the highest sedimentation rate. the uppermost carnian of the yana okhotskaya river, well described by bychkov (1995), is represented by the 290 m thick yakutensis zone (index ammonoid sirenites yakutensis kiparisova) and the lowermost norian is documented by the 300 m thick verchojanicum zone. pinacoceras verchojanicum archipov is the index taxon of the verchojanicum zone, but the most common taxa of this zone are pterosirenites and norosirenites. pterosirenites, which predates the fo of norosirenites, in particular is documented in the trans-baikal, amur, khabarovsk areas (see zakharov 1997) as well as in yana okhotskaya river basin (bychkov 1995), and is therefore of great potential as guide fossil for the base of the norian in the boreal realm. however, the fo of its range, at least on the basis of ammonoids, has not yet been calibrated. pterosirenites is part of the evolutionary lineage within the subfamily sirenitinae, but a stratigraphic interval with no ammonoid record exists (e.g., bychkov 1995) between the last occurrence of carnian sirenitinae and the fo of pterosirenites. in addition to the fo of pterosirenites, konstantinov (2008) emphasized that the base of the norian in the boreal realm is also characterized by the fo of arcestes and of pinacoceras. these two genera are common in the tethys realm and they also occur in north america, but they are long ranging and their fo in the boreal realm is clearly younger than in the tethys and north america. balini m., krystyn l., levera m. & tripodo a.78 data herein summarized suggest the need for an intergrated approach to the correlation of the various localities of the boreal realm because ammonoids alone will not facilitate a final solution. conodont, halobiid and integrated ammonoid data may provide this solution, and the key area is probably southern primorye, where the ammonoid record shows a tethyan influence, according to literature (zakharov et al. 1996; zakharov 1997). unfortunately, the occurrence of pterosirenites in the southern primorye successions, first mentioned by zakharov (1997), has been refuted by markevitch & zakharov (2008). they have modified the generic attribution of the two species previously recognized in the area, pterosirenites kiparisovae (zharnikova) and p. evolutus zakharov & zharnikova, to the genus norosirenites tozer. conclusions: towards the definition of a world standard even now, one centrury after gemmellaro’s monograph on upper triassic ammonoids from western sicily, the ‘‘cherty limestone’’ (scillato formation) still provides a huge amount of paleontologic information, including not only ammonoids and halobiids first discovered by gemmellaro, but also conodonts, radiolarians and nannofossils. the results of the extensive investigations conducted during the last four years (preto et al. 2012; levera 2012; mazza et al. 2012) demonstrate the significance of the pelagic successions of western sicily as a world reference for the understanding of the late carnian to norian transition, in terms of both time and changes in marine biofacies. new ammonoid data improve the understanding of the faunal changes across the carnian/norian boundary, and are significant in perspective for the definition of the gssp of the base of the norian stage. recognition of the discotropites plinii, gonionotites italicus, dimorphites noricus n. sp. and dimorphites selectus subzones links without question certain parts of the pizzo mondello section to the upper carnian-lower norian standard scale and facilitates the chronostratigraphic calibration of conodont and halobia bioevents. conodont bioevents several conodont bioevents have been identified at pizzo mondello during the last few years (mazza et al. 2010; mazza et al. 2011; mazza et al. 2012). mazza et al. (2010) identified three main faunal turnovers (fig. 12) and selected two possible conodont biovents for the definition of the gssp of the norian stage as follows: turnover 1, at level fnp 88a (64.76 m above the base), where epigondolella replaces carniepigondolella; turnover 2, between am23 and na35 (80 m above the base), with a decreasing of epigondolella and increasing of metapolygnathus; and turnover 3, between samples na41 and pm30a, characterized by the disappearance of metapolygnathus and replacement with advanced specimens of epigondolella. mazza et al. (2010: 123) identified the fad of epigondolella quadrata at level fnp88a as the most suitable bioevent for the definition of the gssp of the norian stage, while the fad of metapolygnathus communisti at level na35 was considered an alternative solution. taxonomic studies of the conodont populations has lead mazza et al. 2012 to reconsider the position and significance of the most significant bioevents. the revision of e. quadrata and the separation of the primitive specimens of this group into the new species e. miettoi has resulted in the placement of the fad of e. quadrata from fnp88a up to fnp112. furthermore, the study of conodont populations around the fo of the bivalve halobia austriaca, recently proposed as marker event for the gssp (mcroberts & krystyn 2011), has lead mazza et al. to identify three new possible conodont bioevents as follows: the fo of metapolygnathus parvus at sample am24; the fo of metapolygnathus echinatus at sample na36; and the fad of carnepigondolella gulloae at sample fnp134. new ammonoid data presented in this paper allow the dating of these bioevents. the fad of e. quadrata, in both of its possible locations, and the conodont turnover t1 definitely occur together with carnian ammonoids of the spinosus zone, gonionotites italicus subzone. the fads of m. communisti, m. parvus and m. echinatus, as well as the conodont turnover t2 are located within the interval between the last record of the spinosus zone and the first record of the dimorphites noricus n. sp. subzone of the jandianus zone, but they are very close to the uppermost levels yielding ammonoids of the spinosus zone; hence, they are probably latest carnian in age, as also suggested by halobiids (see levera 2012). the fad of c. gulloae and conodont turnover t3 occur very close to the fo of halobia austriaca in level fnp135a, and are likely earliest norian in age. halobia bioevents at the present stage of the discussion within the carnian/norian boundary task group of the subcommission on triassic stratigraphy, the most interesting bioevent for the definition of the base of the norian stage appears to be the first occurrence of the bivalve halobia austriaca, as discussed during the palermo workshop (september 2010) and proposed by mcroberts & krystyn (2011). at pizzo mondello this bioevent is located within the 18 m thick interval with a very poor ammonoid record between the spinosus and the jandianus zones (fig. 12). more precisely the fo of carnian-norian ammonoids from pizzo mondello 79 h. austriaca is located 4 m below the level na42, that marks the top of the dimorphites noricus n. sp. subzone, the 1st subzone of the norian stage. for the purpose of a pure speculation, we assume that level fnp135a can not be far from the carnian/norian ammonoid boundary. the (weak) elements in support of this speculation are derived from the comparison of the range of certain species that suggests a reduction of the sedimentation rate from the tuvalian 3 to the lacian 1. the spinosus zone (tuvalian 3) is at least 55 m thick (fig. 6), whereas the interval between the fo of dimorphites selectus and the fo of halobia mediterranea, which is equivalent to the d. selectus subzone (lacian 1 ii, sensu krystyn et al. 2002), is only 6-7 m thick. in this respect, the position of fnp135a at four meters below the boundary between the d. noricus n. sp. and d. selectus subzones most likely corresponds to an earliest norian age. future steps at this point in time, comparison of the pizzo mondello ammonoid record to that from black bear ridge does not solve the problem of the definition of the carnian/norian boundary. both sections are relatively expanded and ammonoids are rather scarce; thus, it seems clear that the gssp of the norian stage cannot be defined on an ammonoid bioevent alone. however, due to their very high power of resolution, ammonoids are the unrivaled tool for calibration of bioevents of fossil groups with a lower power of resolution, such as conodonts and in part halobiids. the two gssp candidate sections can be accurately correlated with ammonoids (fig. 13), but not the interval around the fo of halobia austriaca. at pizzo mondello, ammonoids have not been collected from level fnp135a, which records the fo of halobia austriaca. thus, it is necessary to conduct a new ammonoid sampling program for the interval na41-na42 (i.e., from the fad of carniepigondolella gulloae to the fo of dimorphites noricus n. sp.). with regard to ammonoid data at black bear ridge, the fo of h. austriaca at level 18f, as suggested by mcroberts & krystyn (2011), falls within the range of the pterosirenites norian element according to tozer (1994), whose fo is recorded in bed 18b, slightly below 18f (fig. 13). however, it is conceptually wrong to use data from scattered localities, such as isolated outcrops sampled by mclearn and tozer in northern and eastern british columbia (fig. 14), to calibrate the cm-by-cm sampled section black bear ridge. the correct approach is to balini m., krystyn l., levera m. & tripodo a.80 fig. 14 stratigraphic distribution of pterosirenites tozer, 1980 from eastern and northern british columbia, based upon data from tozer (1994). all sites yielding pterosirenites are included, but the genus has not been found in a few other lower norian sites (e.g., brown hill, type locality of the kerri zone). note (*): position of gsc locality 98880 was described by tozer (1994:347) as ‘‘probably stratigraphically between gsc loc. 64628 (macrolobatus zone) and gsc loc. 64607 (kerri subzone ii)’’. only site gsc 98880 was used for this figure because the exact position of the three sites was not fully clear to tozer, who sampled gsc 64628 and gsc 64607 in 1964 (tozer 1967, 1994), and gsc 98880 in 1982. pterosirenites was not found in the samples taken in 1964. resample the relevant portion of the black bear ridge section such that the first occurrence of h. austriaca can be better constrained. acknowledgements. ammonoids described in this paper were collected during several field excursions carried out at pizzo mondello from 2007 to 2010 (mb and ml, with contribution of lk and at). we would like to warmly thank a. nicora and m. mazza (both unimi), m. rigo and c. guaiumi (both unipd) who were with us in the field and occasionally helped with the sampling. m. balini and m. levera are deeply indebted to c. d’arpa (unipa), curator of the collections of the museo geologico ‘‘g.g. gemmellaro’’, for her very kind assistance during several visits to examine the collections, and for stimulating discussions regarding the history of the gemmellaro collections. special thanks also extended to p. di stefano (unipa), who was always convinced of the great potential of the sicanian facies and warmly encouraged us to look for new localities. he was right! last but not least we would like to express our gratitude to g. muttoni (unimi), who was the first stratigrapher of milano university who invested weeks in sampling of pizzo mondello section. our work is complemental to his accurate investigations. our knowledge of carnian/norian ammonoids was notably improved by a visit to the collections housed at the geological survey of canada facilities in vancouver, which occurred in 2010 with the crucial help and support of m. orchard (gsc, vancouver). we are also very grateful to j.p. zonneveld (university of edmonton, alberta) who provided us the opportunity to visit the most important sites in the williston lake area in may 2010 within the framework of the activities of the carnian/norian boundary task group of the subcommission on the triassic stratigraphy. john-paul did a great job organizing the filed trip and also covered most of the cost of the logistics. we had an enjoyable time together on the lake. technical support by g. chiodi (unimi). special thanks to reviewers s. lucas (albuquerque, usa), paolo mietto (padova, italy) and y. zakharov (vladivostok, russia) for the many fruitful suggestions that improved the manuscript. a special warm thank to j. jenks (salt lake city, usa) who very carefully reviewed the final version of the manuscript. he spent quite a lot of time on it and his suggestions improved not only the style, but also some rather complex parts of systematic paleontology and chronostratigraphy chapters. this work is a contribution to prin 2008 project ‘‘stratigrafia integrata del triassico superiore: gssp e sezioni ausiliarie in italia’’ (p.i. m. balini). financial support has been provided by grants to m. balini (milano research unit). l. krystyn has been generously supported by the austrian national committee for igcp within the project 467 (triassic time and trans-panthalassan correlations). carnian-norian ammonoids from pizzo mondello 81 r e f e r e n c e s arkell w. j., kummel b. & wright c. w. 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"��% �/ � � !% ,'�, ! ���!!'�#����� � � ������ �� � ����� ������� �������� �)� � � .�! ! �2�*�, %� %)� .�!% �#, #��%)0.�!% �/ ��+"�&�!!� ?�'(! � �@� %)��� +�'# !��%'�#! .��� ��((�, �#, !�+"��, ?�'(�@� ����.'#( %)� ����#!%� �%'�# �/ � �'%)�!%��%'(��")'� ! ���!!'�# .)'�) ����)�! � %)'�$0 #�!! �/ �44 + ?�'(�@�� � "��%'���* '#%��/'#(��'#( �'%)�!%��%'(��")'� +�+&��! .��� ,'!%'#( '!)�, ?�'(�@� ���� ! ��!������"������ #�#$� .�! �&0 !��2�, %� #��%)0.�!% �/ ��!���'"��#�� ��+"�!�, �/ ��0 %��#�%� /��,!"�%)'� !�#,!%�#�! ?&�,! /��+ �4 �+ %� + %)'�$@ �#, *����.'!) !'�%* +���!� .'%) 2��* ���� ��,,'!) !'�%'%�!�# ! ��!���������%��#� !��� #�#$� '! .',��* �1"�!�, '# %)� ! ��� #,'#(! �/ ��!���'"��#� 2'���(� �#, %� %)� #��%) �#, #��%)0.�!% �/ �"'#�%��)'! +�+&�� ��#!'!%! (�#�����* �/ ����!�0(��'#�, /��,!"�%)'� +�!!'2� %������� �& '� � (& %��� ��� )& %�������� *& + ,������ &�a8 author area age biostratigraphy sedimentary setting crostella & vezzani (1964) san bartolomeo in galdo (bn) elvetian (serravallian) tortonian not reported data foredeep basin iacobacci et al. (1967) san bartolomeo in galdo (bn) late miocene (f ) (not specified assemblages) foredeep basin di nocera et al. (1988) celenza valfortore langhian serravallian (f) g . siakenis zone of iaccarino (1985) piggy-back basin patacca et al. (1990, 1992) san bartolomeo in galdo (bn) ? latest tortonian early messinian not reported data piggy-back basin pescatore et al. (2000) san bartolomeo in galdo (bn) late tortonian messinian according to previous literature data wedge-top tardorogenic basin di nocera et al. (2006, 2011) irpinia daunia middle late tortonian messinian (n) cn9a-cn9b zones of okada & bukry (1980) wedge-top synorogenic basin patacca & scandone (2007) san bartolomeo in galdo (bn) ? latest tortonian ? early messinian not reported data thrust-sheet top basin pescatore et al. (2008) sannio mounts not older than late serravallian (n) (not specified assemblages) wedge-top synorogenic basin pieri et al. 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and reader 5.0 and later.) /jpn /deu /ptb /dan /nld /esp /suo /ita /nor /sve /kor /chs /cht >> >> setdistillerparams << /hwresolution [2400 2400] /pagesize [595.000 842.000] >> setpagedevice comparing the body mass variations in endemic insular species of the genus prolagus (ochotonidae, lagomorpha) in the pleistocene of sardinia (italy) blanca moncunill-solé1, caterinella tuveri2, marisa arca2 & chiara angelone1* 1institut català de paleontologia miquel crusafont, edifici z icta−icp, carrer de les columnes s/n, campus de la universitat autònoma de barcelona, 08193 cerdanyola del vallès, barcelona, spain. e-mail: blanca.moncunill@icp.cat, chiara.angelone@icp.cat. *corresponding author. 2soprintendenza archeologia della sardegna, via g. asproni 33, 08100 nuoro, italy. e-mail: caterinella.tuveri@beniculturali.it, marisa.arca@ beniculturali.it. to cite this article: moncunill-solé b., tuveri c., arca m. & angelone c. (2016) comparing the body mass variations in endemic insular species of the genus prolagus (ochotonidae, lagomorpha) in the pleistocene of sardinia (italy). riv. it. paleont. strat. 122(1): 25-36 rivista italiana di paleontologia e stratigrafia (research in paleontology and stratigraphy) vol. 122(1): 25-36. march 2016 abstract. prolagus figaro and p. sardus are part of an endemic insular anagenetic lineage that populated sardinia since the earliest late pliocene to holocene. bm of some populations of these two species was calculated using regression models. the best bm proxies for prolagus are: femur length, zeugopod measurements and distal humerus diameter. the anagenetic lineage shows a bm increase of ca 20% from the populations of p. figaro (398-436 g) to p. sardus (504-525 g). the trend shown by the size of lower third premolar, even if not directly comparable with bm, is opposite (ca -30% at the transition p. figaro-p. sardus). compared to p. cf. calpensis, a continental species of similar age, bm of p. figaro is ca +25%. the comparison with the insular endemic p. apricenicus evidenced differences in bm range and timespan required to attain it, due to the different size and palaeogeographical situation of the islands. insular endemic prolagus follow the small mammal pattern of island rule. mein’s (1983) biphasic model seems applicable to the evolution of p. figaro. a tachytelic phase followed by a bradytelic one seems to characterize also the appearance of p. sardus, at least for dental traits, a process probably triggered by important variations of abiotic and biotic traits of the environment, as indicated by the turnover that marks the onset of the dragonara subcomplex. the prediction of life history traits and other biological attributes of sardinian prolagus using bm should be considered with caution due to the complexity of ecological selective regimes of sardinia. received: september 27, 2015; accepted: january 12, 2016 keywords: body mass, regression models, postcranial bones, island rule, mediterranean islands, prolagus figaro, p. sardus. introduction body size is a fundamental trait in the biology and ecology of species as it shows tight correlation with several physiological, behavioral, morphological, ecological and life history attributes (peters 1983; calder 1984). the best proxy for quantifying the bs of individuals is their body mass or weight (gingerich et al. 1982). thus, predicting the bm of fossil species is of critical significance for knowing their biology as well as for understanding and quantifying their adaptations to habitats (palombo 2009a). for most of mammalian taxa, the allometric relationships among bm and bone/dental measurements of extant relative species allow the development of regression models to estimate the average weight of extinct ones (damuth & macfadden 1990; see palombo 2009a: tab. 1 for a synopsis). in spite of the abundance, diversity and ubiquity of fossil lagomorphs (leporids and ochotonids), models for estimating the bms of species belonging to this order were developed very recently. quintana cardona (2005) and quintana et al. (2011) provided the first models for leporids. subsequently, expanding the database of quintana cardona (2005) and adding measurements of extant ochotonids, moncunill-solé et al. (2015) developed general and specific equations for estimating the bm of lagomorphs based on a multiproxy approach (teeth, cranial and postcranial measurements). bm estimation models for lagomorphs are going to enhance data about palaeocommunity structures and their palaeoenvironmental interpretations. in view of the potential of this field, we decided to study the bm of the insular endemic ochotonids of the pleistocene of sardinia (italy): prolagus figaro and prolagus sardus. prolagus figaro is known from the latest pliocene/earliest pleistocene to the late early pleistocene of sardinia (capo figari/ moncunill-solé b., tuveri c., arca m. & angelone c. 26 orosei 1 subcomplex of the nesogoral fcorosei 2 subcomplex of the microtus (tyrrhenicola) fc; palombo 2009b). prolagus sardus is reported since the middle pleistocene (dragonara subcomplex of the microtus (tyrrhenicola) fc; palombo 2009b) until historical epoch in sardinia and also in corsica (vigne & valladas 1996; wilkens 2004). we aim to: 1) evaluate the bm trend of prolagus in an insular habitat from an evolutionary point of view, as the two species of prolagus from sardinia are part of an anagenetic evolutionary lineage (angelone et al. 2015). 2) assess the response of fossil ochotonid species to insular regimes (island rule) (see palombo 2009a and references therein for an update of the debate about this subject) as there are not extant relatives living on islands. 3) provide data to increase the scarce biological knowledge of sardinian prolagus. abbreviations bm: body mass; bs: body size; cmd1: capo mannu d1; ic: interval of confidence; fc: faunal complex; fl: femur length; ftdd: distal femoral transversal diameter; ftdp: proximal femoral transversal diameter; hapdd: distal humeral anteroposterior diameter; fig. 1 bm predictions (y axis, in g) for prolagus figaro and p. sardus calculated on the basis of different postcranial measurements (x axis). bm average (black line), confidence interval (dotted lines) and number of individuals are shown. a) prolagus figaro, fissure infilling x3; b) p. figaro, fissure infilling ivm; c) p. figaro, fissure infilling x4; d) prolagus sardus, fissure infilling xir and e) p. sardus, fissure infilling vib. femora tibiae humeri n n n (coding) (coding) (coding) prolagus figaro x3 10 6 14 (ssn/x3/fe/1−10) (ssn/x3/ti/1−6) (ssn/x3/hu/1−14) ivm 5 − − (ssn/ivm/fe/1−5) x4 13 11 − (ssn/x4/fe/1−13) (ssn/x4/ti/1−11) prolagus sardus xir 74 42 60 (ssn/xir/fe/1−74) (ssn/xir/ti/1−42) (ssn/xir/hu/1−60) vi6 20 9 49 (ssn/vi6/fe/1−20) (ssn/vi6/ti/1−9) (ssn/vi6/hu/1−49) species fissure filling tab. 1 fossil material used for performing the study. prolagus (ochotonidae, lagomorpha) from the pleistocene of sardinia (italy) 27 hapdp: proximal humeral anteroposterior diameter; hl: humerus length; htdd: distal humeral transversal diameter; lp3: length of the third premolar; n: sample size; ssn: soprintendenza dei beni archeologici per le provincie di sassari e nuoro, sede di nuoro; tapdp: proximal tibia anteroposterior diameter; tl: tibia length; ttdd: distal tibia transversal diameter; ttdp: proximal tibia transversal diameter; : arithmetic mean;  w : weighted mean. material the samples of p. figaro and p. sardus come from the monte tuttavista karstic complex (e sardinia; abbazzi et al. 2004) (tab. 1). remains of p. figaro come from infillings x3, ivm and x4, pertaining to the capo figari/orosei 1 subcomplex of the nesogoral fc-orosei 2 subcomplex of the microtus (tyrrhenicola) fc (latest pliocene/earliest pleistocene to the late early pleistocene; palombo 2009b). in this context, notice that palombo (2009b) gave a different relative temporal arrangement of the aforementioned infillings (ivm-x4x3). remains of p. sardus have been sampled from infillings xir and vi6, included in the dragonara subcomplex of the microtus (tyrrhenicola) fc (middle and late pleistocene; palombo 2009b). infillings xir and vi6 were accumulated in a quite short time and their palaeontological contents are taxonomically homogeneous (see angelone et al. 2008 for discussion). preliminary analysis of prolagus remains and literature data based on other taxa (angelone et al. 2009; palombo 2009b and references therein) pointed out that x3, ivm and x4 tab. 2 bm predictions (in g) for the populations of prolagus figaro and p. sardus analyzed in this paper. last two rows, highlighted in gray, show the arithmetic mean of bm calculated for each site, and their weighted average. also are taxonomically homogeneous infillings. the fossil material is curated at the ssn. methods the skeletal maturation indicates the complete cessation of longitudinal growth and, consequently, the moment when animals achieve the final bs which is maintained until their death (peters 1983). thus, bm estimations of prolagus species were only carried out on individuals that have already attained skeletal maturity (fused epiphyses). specimens with unfused or broken epiphyses were not considered. ochotona, the extant relative of prolagus, shows a minimal sexual dimorphism (smith 1988; nowak 1999). for this reason, we did not assume bm differences between sexes in our fossil sample. for undertaking the bm estimations, we followed the methodology described and illustrated by moncunill-solé et al. (2015: fig. 1). the following measurements were taken on postcranial remains: 1) fl, ftdd and ftdp on femora; 2) hl, hapdp, hapdd and htdd on humeri; and 3) tl, tapdp, ttdp and ttdd on tibiae. moncunill-solé et al. (2015) observed that burrower species of ochotona and leporids show significant differences in the allometric models of the humerus, but not in other skeletal elements (femur and tibia). although ochotona is the closest relative of prolagus, we preferred to use a general regression model for humeri (i.e. models that include data of leporids and ochotona) because the locomotion of the species of prolagus is not well known. the equations are shown x̄ x̄ x̄ x̄ x̄ (ic) (ic) (ic) (ic) (ic) 537.57 1 444.74 1 − − 463.09 52 452.12 19 (448.98-477.20) (426.38−477.85) 441.19 6 484.96 4 474.51 10 624.91 55 566.28 18 (337.88-544.51) (432.23-537.69) (423.18-525.84) (605.96-643.86) (538.47-594.08) 406.21 3 289.40 2 306.13 3 440.50 58 424.84 19 (207.12-605.30) (178.91-399.90) (251.76-360.51) (421.79-459.21) (398.98-450.70) 364.47 3 − − − − 633.99 47 416.71 44 (360.73-368.22) (605.66-662.33) (403.42-430.00) 665.15 5 − − − − 633.99 37 709.82 35 (593.75-736.56) (605.66-662.33) (675.19-744.15) 345.03 11 − − − − 425.75 54 441.69 44 (311.26-378.80) (409.64-441.87) (421.16-462.25) 470.57 11 − − − − 519.21 53 528.79 44 (393.39-547.74) (493.6-544.77) (501.34-556.24) 342.28 2 − − − − 422.18 40 445.57 9 (296.52-388.03) (407.09-437.27) (415.82-475.32) 298.77 5 − − 545.27 6 530.28 35 578.26 8 (233.38-364.16) (454.86-635.69) (509.28-551.47) (530.81-625.71) 170.50 4 − − 400.62 6 536.58 32 586.70 8 (155.93-185.06) (312.36-488.68) (513.18-559.98) (558.05-615.35) 334.94 2 − − 452.19 5 540.94 38 624.73 9 (241.77-428.12) (353.44-550.93) (516.22-565.66) (551.11-698.36) 397.88 406.37 435.74 503.86 525.05 (320.68-475.08) (289.50-523.23) (357.59-513.90) (456.89-550.83) (468.12-581.97) weighted average 402.34 423.34 453.326 520.83 512.40 ttdp logbm =0.219++2.577logttdp ttdd logbm =0.461+2.584logttdd arithmetic mean htdd logbm =1.053+1.513loghtdd tl logbm =−1.271+2.254logtl tapdp logbm =0.599+2.265logtapdp hl logbm =−1.221+2.418loghl hapdp logbm =0.916+1.769loghapdp hapdd logbm =1.354+1.769loghapdd n fl logbm =−1.11+2.229logfl ftdp logbm =0.498+2.217logftdp n n n ftdd logbm =0.318+2.481logftdd measurement equation n bm prolagus figaro bm prolagus sardus x3 ivm x4 xir vi6 moncunill-solé b., tuveri c., arca m. & angelone c. 28 in tab. 2. once the regression models were applied, we eliminated outliers due to their potential for skewing the distributions. we followed the criterion of tukey (1977): outliers (y) were considered when y < (q1−1.5iqr) or y > (q3+1.5iqr) (where q1 is the 25th percentile, q3 is the 75th percentile, and iqr the interquartile range (q3−q1)) (quinn & keough 2002). for each specific measurement, it was calculated an arithmetic mean () and a confidence interval (ic) [±((σ/√n)z α /2)]. based on the bm of each measurement, we performed an arithmetic average () and a weighted average ( w ) [(x 1 w 1 +x 2 w 2 +…+x n w n )/(w 1 +w 2 +…+w n )]. in order to compare the different populations of prolagus and analyze the bm variation, we performed anova analyses and post hoc tests (tukey hsd) (α = 0.05) using the ibm spss statistics 19 software. results the results of bm estimations (means, ic, n) are shown in tab. 2 and are represented in fig. 1. for p. figaro, we estimate a weight of 397.88 g (320.68-475.08) in fissure filling x3, of 406.37 g (289.50-523.23) in ivm and of 435.74 g (357.59513.90) in x4. for p. sardus the results are greater, 503.86 g (456.89-550.83) in fissure filling xir and 525.05 g (468.12-581.97) in vi6. we do not observe significant differences between  and  w (their difference is ca 10-20 g) and the latter falls perfectly in the ic of the former (tab. 2). statistically, there are only significant differences (p < 0.05) between the oldest population of p. figaro (x3) and the youngest of p. sardus (vi6). when the bm estimations of each measurement are assessed, a similar pattern could be observed comparing the populations with the largest n (vi6 and xir) (fig. 1). the variables ftdp and hapdp estimate a bm far above the arithmetic mean (between 100-200 g greater), especially in vi6 population. the other variables fall next or inside the ic of the arithmetic mean (specially fl, tapdp, ttdp, ttdd and htdd). analyzing the results of the other populations (x3, ivm and x4), we observe more heterogeneous patterns. this may be consequence of: 1) few measurements taken in postcranial bones (3 in ivm and 5 in x4) and 2) small n (ranging from 1 to 11 individuals in x3). however, in this latter population (x3), it is already evident a large value of bm when hapdp measurement is used, but not in ftdp. discussion bm of sardinian prolagus: trends and best estimators. based on dental morphology, a relative temporal arrangement of the studied fissure has been attempted. preliminary results suggested the relative chronological arrangement x3-ivm-x4 (from older to younger) of populations of p. figaro (angelone et al. 2009). in the case of populations of p. sardus, infilling xir is older than vi6 on the basis of a morphological cline (angelone et al. 2008). in view of this and the bm results, the three selected populations of p. figaro show a total weight increase of ca 10% from the oldest fissure filling (x3) to the youngest (x4) (see tab. 2, fig. 1 and 2a). the bm of the oldest population of p. fig. 2 bm range a) and lp3 range b) showing max, average and min values of prolagus aff. figaro (cross, cmd1), p. figaro (diamonds, x3, ivm, x4) and p. sardus (circles, iv5, iv20, xir, vi6), with detail of average values and trends of bm (a’) and lp3 (b’). prolagus (ochotonidae, lagomorpha) from the pleistocene of sardinia (italy) 29 sardus here analyzed (xir) is ca 15% greater than the youngest of p. figaro (x4). the average bms of the two populations of p. sardus selected for this study show a very slight difference (average bm of vi6 is about 4% larger). finally, the total increase among the oldest (x3) to the youngest populations (vi6) of sardinian prolagus is of 32% (statistically significant, p < 0.05). thus, we can affirm that sardinian prolagus increased its bm (average) throughout the pleistocene. the best bm estimator for an extinct species not only depends on the accuracy of the model (statistical values), but also on a subjective judgment of the results of predictions (reynolds 2002). according to the fissure infilling with highest sample (xir) (fig. 1d), hindlimb bones seem to be the better bm estimators for prolagus species (as shown also in moncunill-solé et al. 2015), particularly fl, tapdp, ttdp and ttdd. however, htdd also gives adjusted estimations. all these measurements predicted bm that fall inside the ic of the arithmetic mean and, consequently, we can consider them good proxies for the estimation of bm in the genus prolagus. however, n must be taken into due account. for example, the bm predicted by fl (n=1) in x3 population is far above the arithmetic mean. it is recommendable to work with the largest sample possible in order to better represent the biological variability of the species and, thus, obtain more realistic values. the measurements regarded as the best bm estimators are surprising for two facts. firstly, zeugopods (tibiae), which are involved in the locomotion and lifestyle of the animal, normally predict worse the bm of mammals (scott 1990). secondly, the lengths of long bones are also considered as less accurate than diameters or perimeters (scott 1990). however, in the case of lagomorphs, the models that use length or zeugopodial measurements are as reliable (coefficient of determination or average absolute per cent prediction) as those that use other postcranial elements, in contrast to other mammalian orders (see also moncunill-solé et al. 2015). taking into consideration quantitative results, hapdp measurement overestimates in all populations the bm in prolagus and could not be considered a reliable proxy. ftdp does not show a clear pattern, being far above in the case of xir population (those with the largest n), but not in others (x3, ivm, x4, vi6). the bm overestimation observed when hapdp is used for prediction is indicative that this measurement does not only represent the bm of the species but also other biological attributes, such as locomotion or phylogeny. samuels & valkenburgh (2008) described some skeletal specializations of rodents depending on their locomotion style. for example, a broad and robust distal humerus is indicative of fossorial or semifossorial habits. we encourage the scientific community to perform new studies that analyze the locomotion, biomechanics and skeletal proportions of prolagus species in comparison with its extant relatives (ochotona spp.). this will increase the biological knowledge of prolagus and might help us to discard those measures that are correlated with their locomotion or phylogeny for predicting bm. bm and teeth size: the case of sardinian prolagus. it is interesting to compare the trends of bm vs lp3 in the p. figaro − p. sardus lineage. we take into consideration p3 because it is the most reliable tooth position for specific identification in lagomorphs. as shown in fig. 2b, average lp3 increases (ca 13%) when the oldest population of p. figaro (x3) is compared to p. aff. figaro from cmd1 site, the “founder” of the sardinian lineage. lp3 of p. figaro shows a maximum oscillation of 6% in the considered populations. a drastic drop of lp3 (almost 30%) is recorded between p. figaro (x4) and the oldest studied population of p. sardus (iv5). after iv5, lp3 values of p. sardus increase slightly through time (total increase of ca 11% in the studied populations) following an asymptotic pattern (see also angelone et al. 2008). when we analyzed the bm variation, the first thing that we observe is that its record is more incomplete than for teeth: bm estimations are not available for p. aff. figaro and older populations of p. sardus (fig. 2a). moreover, we have to take into due account that bm values have been obtained after complex data treatment, whereas lp3 are raw data. nevertheless, it is evident that bm and lp3 of sardinian prolagus follow quite different trends. the differences are not so evident among populations of p. figaro: average bm increases of ca 10%, whereas average lp3 fluctuates of ca 6%. evident discrepancies can be noticed with the appearance of p. sardus. average lp3 drops of ca 30% between youngest p. figaro (x4) and oldest p. sardus (iv5). then, lp3 average increases through time in p. sardus attesting to a value of 2.06 mm (vi6) which is ca 15% smaller than in x4. lacking moncunill-solé b., tuveri c., arca m. & angelone c. 30 data relative to older infillings (iv5 and iv20), we can only state that younger ones (xir and vi6) show a higher bm average (ca 15-20%) than p. figaro (x4). hypothesizing a dramatic bm decrease between p. figaro and p. sardus followed by an explosive increase to exceed p. figaro bm values is not realistic. the most parsimonious hypothesis is that bm followed a general increase trend through the transition p. figaro-p. sardus and throughout the evolution of p. sardus, countertrending lp3 drastic drop observed at the transition p. figaro − p. sardus. the fact that p3 dimensional trend shows evident discrepancies with bm pattern inferred through postcranial elements casts doubts about the usage of p3 as a proxy for bm estimation in insular endemic prolagus. compared to continental species of prolagus, insular endemic species show a noticeable enlargement of the size of p3 vs the size of molariform elements of the lower tooth row (see angelone 2005: fig. 6 for a qualitative comparison) probably due to a reassessment in jaw mechanics. at any rate, the reliability of teeth as bm proxies has been questioned also in studies that took into consideration a continental species of prolagus as well as a wider selection of fossil lagomorphs case studies (moncunill-solé et al. 2015). they prefer models based on postcranial bones, as directly related to weight bearing. timing and patterns of bm variations in sardinian prolagus. mein (1983) illustrated a biphasic pattern of evolution on islands consisting in a first tachytelic step in which the immigrant species undergoes sudden morpho-dimensional changes corresponding to its entrance to insular selective regimes and a second step in which the taxon undergoes a relatively long bradytelic phase. millien (2006) further corroborated and “quantified” mein’s rule. according to some authors (sondaar 1977; alcover et al. 1981; lister 1989, 1996), the tachytelic stage is a change in the “evolutionary direction” (sensu sondaar 1977; e.g. bs shift or low gear locomotion) whereas the bradytelic one is a further continuation of the existing “direction of the change” (ib., e.g. harvesting saving by increase of hypsodonty, changes in dentognathic feeding apparatus, or developing traits for searching fallback resources). evans et al. (2012) estimated a minimum of 4000 years for small mammals to become giants (ca 16000 generations). if we apply mein’s model to sardinian prolagus lineage, the first phase should have taken place during or short after the early/late pliocene boundary (age of the cmd1 fossil assemblage). indeed, p. aff. figaro from cmd1 shows very slight morphological modifications due to endemism, evidence of its very recent arrival from mainland (angelone et al. 2015). the lp3 of p. aff. figaro is comparable to the values of populations of continental italy from mn16 (absence record for mn15; angelone et al. 2015) and is between 7-13% smaller than p. figaro. there is no record of the possible changes of bm occurred in the 1 ma that separate cmd1 and the oldest populations of p. figaro from monte tuttavista. we have not enough data to clearly recognize the tachytelic stage of mein’s model in p. aff. figaro-p. figaro and to verify/quantify the dimensional changes and the time span needed to produce them. the populations of p. figaro here analyzed should cover a time span of ca 0.3-0.4 ma (inferred from palombo 2009b: fig 2). in this time span, slight weight fluctuations have been observed, which may correspond to the bradytelic phase of mein’s model. the appearance of p. sardus (closely related to p. figaro and not an immigrant from mainland; angelone et al. 2015) occurred during the transition from the orosei 2 subcomplex to the dragonara subcomplex (ca 0.8-0.7 ma; inferred from palombo 2009b: fig 2). this transition is characterized by the highest species turnover recorded in the quaternary of sardinia (palombo 2009b). leaving aside the reason of this dramatic change (see next section), it seems to have triggered a new biphasic evolutionary phenomenon which follows mein’s model too. in general the phyletic lineages of sardinian small mammals underwent outright (geologically speaking), an abrupt and noticeable increase in dental size (abbazzi et al. 2004). contrarily prolagus, as stated above, experienced a drastic lp3 decrease. the absence of data from iv5 and iv20 fissure fillings does not allow us to quantify changes in bm. teeth size and morphology in early populations of p. sardus underwent an evolution comparable to mein’s model first phase. the slight, asymptotic growth of p. sardus lp3 and postcranial measurements (angelone et al. 2008), that in our data covers the interval between ca 0.6-0.4 ma (inferred from palombo 2009b: fig. 2), parallelizes mein’s model second phase. indeed, mein’s rule focuses on the first stages of colonization of the island, making reference to the prolagus (ochotonidae, lagomorpha) from the pleistocene of sardinia (italy) 31 biological adaptation of the species to the new selective regimes. however later on, changes can also take place consequence of the variation in the environment. abiotic changes (climatic, topographic, among others) and variations of biotic traits (e.g. levels of predation, intraand interspecific competition), both have a significant role to drive evolution (alcover et al. 1981; brockhurst et al. 2014). our data seem to indicate that mein’s model can be applied several times to a taxon during its evolution of an island, if significant ecological changes occur (e.g. climatic changes or variation in levels of selective regimes). the study of other mammalian lineages of the pleistocene of sardinia or other islands may provide more case studies to support this hypothesis. driving factors in the evolution of sardinian prolagus: some hypotheses. the sea level low stand at the early/late pliocene transition allowed the migration of p. sorbinii from italian mainland towards sardinia (angelone & kotsakis 2001; angelone et al. 2015). insular selective regimes triggered the morpho-dimensional changes in the immigrant that led to p. figaro. prolagus figaro survived until the end of the orosei 2 subcomplex. the appearance of prolagus sardus marks the onset of the dragonara subcomplex, characterized by a complete turnover in the small mammals’ component of sardinian fauna (except for talpa): the leporid and the glirid tyrrhenoglis did not survive the transition; the insectivore nesiotites and the rodents tyrrhenicola and rhagamys, descendants of taxa already present in the orosei 2 subcomplex, underwent evident modifications of teeth morphology and a noticeable increase of teeth size probably coupled with a bs increase (abbazzi et al. 2004); the ochotonid prolagus underwent a decrease of lp3 but an increase of bm. the ancestors of the small mammal genera which survived into the dragonara subcomplex were present and already showed endemic traits since the orosei 2 subcomplex (i.e. tyrrhenicola) or at least since the capo figari/orosei 1 subcomplex. the competition among small mammal species as driving factor of the turnover between the orosei 2/dragonara subcomplexes can be ruled out and the extinction of glirids and of the leporid is not likely to have triggered a competition to occupy its niche in taxa with such a wide range of ecological requirements, and the arrival of tyrrhenicola already occurred earlier. the arrival of the canid cynotherium (once regarded as a specialized prolagus hunter, and recently considered a small-prey hunter, possibly also birds, without evident specialization in prolagus hunting; malatesta 1970; lyras et al. 2006, 2010) occurred at the onset of the orosei 2 subcomplex without triggering any sudden, evident change in small mammals, least of all in prolagus, which increases its bm throughout pleistocene. this fact apparently contradicts van der geer et al. (2013) who noticed that the bs increase in insular small mammals that occurs following colonization or first appearance, ceases or is reversed after the arrival of mammalian predators or competitors. probably the impact of a new predator was not so catastrophic because, contrarily to other islands, several carnivores were already present in sardinia prior to cynotherium (i.e. chasmaporthetes, mustela and pannonictis; the latter also coexisted with cynotherium for a while). in fact, due to its large area, sardinia had selective regimes more similar to mainland than other mediterranean islands. it could support the presence of terrestrial predators and had not a strong resource limitation as small islands (heaney 1978, 1984). thus, we can not affirm that the arrival of cynotherium sardoum increased the extrinsic mortality of sardinian pikas. the most important cause of the turnover at the onset of the dragonara subcomplex, and thus the trigger of the transition p. figaro-p. sardus is likely to be climate change, in particular those related to the mid-pleistocene transition. even after middle pleistocene, the evolution of p. sardus seem related to climate changes and to consequent specific modifications of the environment. preliminary data by boldrini & palombo (2010) suggested a correlation between limb length and temperature in p. sardus. effects of climate on bs have been highlighted in insular endemic fossil vertebrates of the mediterranean by van der geer et al. (2013), according to whom bs fluctuates over time linked to climatic oscillation. also millien & damuth (2004) noticed the influence of geographical climatic gradients and climatic change through time on fossil endemic insular species. regarding the extinction of sardinian prolagus, it probably occurred less than 2000 years ago, in the roman period, between the arrival of rattus rattus and the present time (vigne 1982). authors moncunill-solé b., tuveri c., arca m. & angelone c. 32 do not agree on the importance of men’s influence (directly by predation and indirectly by introduction of alien predators, competitors, parasites, infectious diseases, modification of the landscape by agricultural activities, among others) to the extinction of prolagus. insular endemic lagomorphs and the island rule. radical morpho-dimensional adaptations are observed in insular endemic organisms. in mammals, apart from modifications in dental, cranial and limbs morphology and relative proportions, it is observed a bs trend coined as island rule (foster 1964; van valen 1973): in general small-sized mammals considerably increase their size, whereas large-sized mammals show an opposite trend. this ecogeographic rule is also observed in insular endemic fossil mammals. in the neogene-quaternary of mediterranean islands and palaeoislands, insular gigantism and dwarfism have been the subject of several studies and debates (from the pioneer general studies, e.g. vaufrey 1929; thaler 1973; sondaar 1977; azzaroli 1982; to the most recent reviews, e.g. van der geer et al. 2010; lomolino et al. 2013 and references therein). lagomorphs are usually considered as small mammals together with rodents and insectivores. although they have a larger size than the average of the small mammals, this order is far from reaching the size of the great majority of large mammals (e.g. elephants, rhinos, etc.). their medium or intermediate bm undertakes a key position in ecosystems (valverde 1964) and compromises their response (adaptation) to island environments (island rule). actually in extant endemic insular leporids the bm trend reported in literature is variable, but mostly directed towards a reduction of the size (foster 1963, 1964; lawlor 1982; palacios & fernández 1992; tomida & otsuka 1993). in the case of ochotonids, no extant species are present on islands and their trend is unknown. when we deal with insular endemic fossil lagomorphs, it is not easy to determine their actual bm and its relative variation compared to the continental ancestor. this is consequence of two facts: 1) mainly most remains consist in teeth, whose size, at least in lagomorphs, does not directly reflect bm (see above and moncunill-solé et al. 2015); and 2) the supposed ancestor is often unknown or wrongly identified (e.g. p. sardus from p. michauxi and gymnesicolagus gelaberti from p. crusafonti in lomolino et al. 2013 and van der geer et al. 2013). a reliable bs estimation is available for an insular endemic fossil leporid, nuralagus rex (pliocene of menorca, spain), which bm has been calculated in 8 kg (moncunillsolé et al. 2015, who reconsidered the bm estimation of 12 kg by quintana et al. 2011). even if not quantifiable, the size increase with respect to its supposed continental ancestor, the relatively smallsized genus alilepus, should have been quite remarkable. for prolagus species it is not possible to quantify exactly the relative bm increase of insular endemic vs their mainland ancestors. in the case of sardinian prolagus, this is due to the lack of studies of postcranial remains of p. sorbinii, whereas in the case of apulian prolagus, their continental ancestor is not known yet (angelone 2007; angelone & čermák 2015; angelone et al. 2015). nevertheless, we can have a gauge of the bm difference between endemic insular prolagus and continental species taking into consideration the only available bm datum of a continental prolagus, i.e. the bm of p. cf. calpensis from the late pliocene of spain, estimated in ca 320 g (based on average of femurs and tibiae; moncunill-solé et al. 2015). thus, p. figaro from monte tuttavista x3 had a noticeably larger bm (ca 25%) than an almost coeval western european mainland prolagus. one among the oldest known populations of p. apricenicus (cava fina f1; bm = ca 282 g; moncunill-solé et al. in press) had a bm ca 13% smaller than p. cf. calpensis, not because it decreased its size in an insular domain, but probably because the continental ancestor of apulian prolagus was a pre-messinian, medium-sized species (see angelone 2007; angelone & čermák 2015). later populations of p. apricenicus, weighing ca 601 g (moncunill-solé et al. 2015), almost doubled the bm of p. cf. calpensis. the known average bm range of both sardinian species of prolagus (397.88-525.05 g; a bm of 800 g for mesolithic p. sardus inferred by sondaar & van der geer (2000) on a qualitative basis has to be verified) is smaller than the populations of p. apricenicus, whose bm range is ca 280-600 g (moncunillsolé et al. 2015, in press). the other apulian species, p. imperialis, is traditionally considered gigantic, because it has the largest p3. however, our analyses and results make clear that dental remains do not directly reflect actual bm, and sometimes even countertrend postcranial-based results. pending a study prolagus (ochotonidae, lagomorpha) from the pleistocene of sardinia (italy) 33 of postcranial remains of p. imperialis, we refrain to make inferences about its bm. millien (2011) argued that in smaller islands the evolutionary rate is higher. this possibly explains the explosive bm increase of p. apricenicus, confined to a very limited, fragmented area, in contrast to the p. figaro-p. sardus trend, which lived in a larger island. in general, the scanty available quantitative data indicate that both fossil leporids and ochotonids follow the small mammal island rule pattern. they underwent a bm increase which extent is highly variable, though. this trend is not in line with the variable response observed in extant insular endemic leporids (see above). bm and life history of insular endemic prolagus. in the last twenty years, several researches have been focused on the life history of insular fossil species, principally addressed to dwarfism (bromage et al. 2002; raia et al. 2003; raia & meiri 2006; köhler 2010; kubo et al. 2011; marín-moratalla et al. 2011; jordana et al. 2012, 2013; van der geer et al. 2014) although, newly, investigations regarding gigantism have been performed (moncunill-solé et al. in press; orlandi-oliveras et al. in press). bm scales with several traits of the life history of species such as life span, fecundity, age at maturity, among others (blueweiss et al. 1978; peters 1983; calder 1984). for this reason, at first sight, we could think in predicting some of these life history traits for p. figaro and p. sardus using the bm results of our analysis. however, moncunill-solé et al. (in press) analyzed the histology and bm of one of the apulian insular endemic species of prolagus (p. apricenicus) and suggested that it had a slower life history than expected from its bs. histologically, the longevity is estimated of at least 7 years for p. apricenicus from f1 fissure filling contrasting with the 4.5 years expected from its bs (around 300 g). the selective regimes of insular habitats (low levels of extrinsic mortality and resource limitation) are the most probable triggers of this shift (palkovacs 2003). this is also observed in extant ochotonids (ochotona spp.) that dwell in rocky habitats which are subjected to a low average yearly mortality. they show a slower life history (later age at maturity and longer longevity) than the species of ochotona that live in meadows, although both groups do not have steep differences in bm (smith 1988). taking in consideration of the aforementioned, extant and extinct relative species that dwell in habitats with low levels of extrinsic mortality show a slower life history than expected from their bs. in the case of p. figaro and p. sardus the levels of extrinsic mortality may not be as low as in the case of p. apricenicus from gargano consequence of the presence of predators. however, the ecological selective regimes of sardinia would not be like the mainland’s one. for this reason, the prediction of life history traits and other biological attributes using the estimated bm should be considered with caution. probably, we would underestimate the values of these traits. the absence of histological data of extant and extinct ochotonids encourages the studies focused on this field in order to improve the biological knowledge of insular and mainland lagomorphs. conclusions bms were estimated for p. figaro from x3 (397.88 g), ivm (406.37 g) and x4 (435.74 g); and for p. sardus from xir (503.86) and vi6 (525.05 g). these results allow us to state a significant increase of bm of the species prolagus from sardinia throughout the pleistocene. the best measurements for determining the bm of prolagus are fl, tapdp, ttdp, ttdd and htdd. in contrast, hapdp and ftdp appear to be unreliable proxies. the bm increase opposes to the pattern of the lp3, which shows a drastic drop at the transition between p. figaro and p. sardus. this is due to the fact that teeth are not weight-bearing elements, and thus, their use as bm proxies is not recommended. however, when the teeth dimensions are taken into account, the biphasic mein’s model (tachytelic and bradytelic stages) may be observed twice. this cannot be confirmed with bm estimations due to the absence of postcranial elements in some key sites as cmd1 (p. aff. figaro) and iv5 (oldest known population of p. sardus). the entrance of cynotherium and the presence of other species of carnivores during the pleistocene seem to not have influence on the pattern of adaption to insular ecological regimes of prolagus. currently, the absence of ochotonids on islands does not allow us to know the adaptions of this group to insular ecological regimes (island rule). in moncunill-solé b., tuveri c., arca m. & angelone c. 34 the fossil record, the two species of prolagus studied in our research and one of the two endemic insular apulian species (p. apricenicus) suggest a gigantism pattern for ochotonids. however, this latter species shows a more explosive increase of bm perhaps as a result that it dwelled in a smaller, fragmented area. it is observed in extant and extinct species that the environments with a lower extrinsic mortality can promote a lower life history (e.g. greater longevity than that expected for its bm). thus, the estimations of life history traits taking into account the bm results of our research should be considered with caution. acknowledgements. we are grateful to the firms that perform quarrying activities at monte tuttavista for their kind collaboration; to m. asole, p. catte, a. fancello, g. mercuriu, g. puligheddu, a. useli for the careful work of preparation of the analyzed fossils; to m.a. fadda and the superintendents f. lo schiavo and f. nicosia of the soprintendenza archeologica della sardegna who allowed the study of the material analyzed in this paper; to t. kotsakis, m.r. palombo and to the reviewers s. čermák, x. jordana, and j. quintana, and to the editor l. rook for their useful remarks. this research was supported by the spanish ministry of education, culture and sport (ap2010−2393, b.m-s.). references abbazzi l., angelone c., arca m., barisone g., bedetti c., delfino m., kotsakis t., marcolini f., palombo m.r., pavia m., piras p., rook l., torre d., tuveri n., valli a. & wilkens b. 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()� )("1�) '%+'�("�+ 6e 2/ 7+8� % $#%")(!" "# #"1�) "��"1 % 91 $1 "1�* ()� ,#)� #) ��!! !")( 21"/ �1� ,#�()! 4)#, �()(% $( ()� !#,�91(" �()2�) "1(% "1#!� #4 )�$�%" 3#3'�(" #%! 6�"#)$1 (,3��� !#�("�+ "��"1� 3()"! #4 �#%2 -#%�!� $('+(� 0�)"�-)(�� ,�"(3#+ (� -#%�! (%+ 31(�(%2�! ()� )("1�) $#,,#%� 91 �� #"1�) . %+! #4 0�)"�-)(�� !.'�� -#%�! (%+ ,#!" !1#)" -#%�!� )��� /��������"� ���"�� 0!�������� ����"���1 � �� ��� &� �"��� ���� "�� �"-�*2���� 03���� " �� ��1 �7 e 2/ � 5 �����$ �������" 6�#)+,(%%� @7?8/ 6(8 � +�>"/� 6-8 �� +�>"/� 6$8 �� +�>"/ n min. max. mean sd m 1 length 4 1.02 1.08 1.05 0.02 width 1.05 1.23 1.12 0.08 m 2 length 2 1.14 1.14 1.14 width 1.14 1.17 1.15 m 3 length 3 0.81 0.84 0.82 0.02 width 1.05 1.11 1.07 0.03 m1 length 2 1.08 1.17 1.12 width 0.96 1.05 1.00 m3 length 1 0.84 width 1.02 �(-/ g 5 � ,�%! #%! #4 ,#�()! #4 � ����� "�������% e 2/ � 5 � ����� "������� 6�(��(!� ff@8/ 6(8 �� +�>"/� 6-8 �� ! %/ n min. max. mean sd m 1 length 5 1.68 1.92 1.81 0.10 width 1.17 1.35 1.27 0.07 m1 length 3 1.68 1.89 1.76 0.11 width 1.20 1.29 1.23 0.05 �(-/ f 5 � ,�%! #%! #4 ,#�()! #4 !���� ��"�� ������"� ���/ e 2/ 7 5 !���� ��"�� ������"� ��� 6� %%(�'!� f=@8/ 6(8 � +�>"/� 6-8 � ! %/� 6$8 � ! %/� 6+8 � ! %/� 6�8 � +�>"/ -�$('!� #4 "1� ) 4)(2 �� !")'$"')� (%+ !,(�� ! ;�� ()� ,'$1 ,#)� )()��* 4#'%+/ �#!" #4 "1� "��"1 ()� !#�("�+� -'" "1�)� ()� (�!# !#,� ,(> ��(�l,(%+ -��! 9 "1 "��"1/ �#%2 -#%�! 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(%+ 4#)�!"5!"�33� 6!�� ���� � ����� (%+ !% � ������ !���� �# ����� "����"�� �����$ �������"� ��� �������� $4/ �����# ������8h �� !3�$ �! #4 !"�33� (%+ !�, 5() + )�2 #%! 6� ���# ��� � ������� � �������� �� ��� ���� ������� �� ������ )�# � �� �� � ��� !�� ���� � ����8/ �% "�),! #4 "1� %',-�) #4 )�,( %!� "1� %1("(%"! #4 #3�%� )("1�) +)* (%+ !"�33� 1("("! 6�i�8 9�)� 3)�+#, %(%" % -#"1 �(*�)! � (%+ 7� 91 �� "1#!� � 0 %2 % ,#)� 1', +� 9(), #) 4#)�!" 1("("! 9�)� ,'$1 ��!! $#,,#% 6e 2/ =8/ �1� 3�)$�%"(2�! #4 "1� 2)#'3! � � (%+ � % �(*�)! � (%+ 7 ()� !") . %2�* ! , �()h "1�)� ()� #%�* !� 21" + 44�)�%$�! % "1� 4)�b'�%$* #4 4#)�!" (%+ ,�(+#9 ���,�%"!� ,#)� 4#)�!" (%+ ��!! ,�(+#9 !3�$ �! &� ��'����' �%� (�"����' �%� !���������' �%� )�*� ����' +% , !�����-����' .%�7� order species lagomorpha lepus sp. ochotona pusilla (pallas, 1769) carnivora canis lupus linnaeus, 1758 vulpes vulpes (linnaeus, 1758) ursus spelaeus rossenmüller & heinroth, 1794 gulo gulo (linnaeus, 1758) martes martes linnaeus, 1758 meles meles (linnaeus, 1758) mustela nivalis linnaeus, 1766 crocuta spelaea (goldfuss, 1823) panthera spelaea (goldfuss, 1810) panthera pardus (linnaeus, 1758) felix silvestris schreber, 1777 proboscidea mammuthus primigenius (blumenbach, 1799) perissodactyla coelodonta antiquitatis (blumenbach, 1799) equus ferus boddaert, 1785 equus hydruntinus regalia, 1907 artiodactyla megaloceros giganteus (blumenbach, 1799) cervus elaphus linnaeus, 1758 bos primigenius bojanus, 1827 bison priscus (bojanus, 1827) capra ibex linnaeus, 1758 rupicapra rupicapra (linnaeus, 1758) rodentia spermophilus cf. citelloides (kormos, 1916) castor fiber linnaeus, 1758 sicista subtilis (pallas, 1773) cricetulus migratorius (pallas, 1773) cricetus cricetus linnaeus, 1758 mesocricetus newtoni nehring, 1898 arvicola terrestris (linnaeus, 1758) chionomys nivalis (martino, 1842) microtus (m.) arvalis (pallas, 1778) & m. (m.) agrestis (linnaeus, 1761) microtus (stenocranius) gregalis (pallas, 1779) microtus (terricola) subterraneus (de sélys-longschamps, 1836) clethrionomys glareolus (schreber, 1780) lagurus lagurus (pallas, 1773) apodemus ex gr. sylvaticus-flavicollis (linnaeus, 1758)/(melchior, 1834) spalax leucodon (nordmann, 1840) dryomys nitedula (pallas, 1778) muscardinus avellanarius (linnaeus, 1758) insectivora indet. chiroptera indet. �(-/ @ 5 � !" #4 ,(,,(� "(>( 4#'%+ % "1� (0�! �()(% $( � (%+ �� 6(4"�) �)2(%" � � , ") :�0 $ �??f (%+ �#2 $�0 $ �" (�/ % 3)�3/8/ % �(*�) 7/ �1 ! %+ $("�! 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()� "*3 $(� #4 "1� �(!" ��($ (� #4 "1 ! 3()" #4 "1� �(�.(% ��% %!'�( 6�#3#0 �???h �#3#0 � �() %!.( �??f8/ �! �#9(�!. 6 <@�8 1(! (�)�(+* !"("�+� % "1�!� . %+ #4 �#$(� " �!� "1� 4)�b'�%$* #4 !"�33� !3�5 $ �! ! 1 21�) "1(% % $�%")(� �')#3�� (%+ 4#)�!" !3�$ �! ()� (�,#!" (�9(*! 3)�!�%" 6(�"1#'21 )�3)�!�%"�+ -* )()� !3�$ ,�%!8� �0�% +') %2 "1� $#�+�!" 3�) #+!/ �#9�0�)� "1�)� ()� (�!# !#,� + 44�)�%$�!� ������� ! ,'$1 ,#)� $#,,#% % �()(% $( "1(% % (%* #4 "1� ,�%" #%�+ �#$(� " �!� 91 �� )� � �� (%+ ����"���� "�# ����� ()� %#" (! 4)�b'�%" (! % "1� "1)�� �'�2() (% $(0�!/ �% �'�2() (% $(0�! +#), $� ()� � "1�) (-!�%" #) 4#'%+ 0�)* )()��* 6(! % (0� g8� 91 �� % �(*�) 7 % �()(% $(� "9# +#), $� !3�$ �! 9�)� 4#'%+/ �1 ! $#'�+ ,�(% "1(" "1� $� ,("� #4 �()(% $( 6(" ��(!" #4 �(*�) 78 9(! ( � ""�� 9(),�) (%+ ,#)� 1', +� 9 "1 ,#)� 4#)�!" (%+ ��!! !"�33� (%+ ,#'%"( % !3�$ �!h -'" "1� + 44�)�%$�! % 4('5 %(� $#,3#! " #% $#'�+ (�!# -� +'� "# "(31#%#, $(� )�(5 !#%!/ �$$#)+ %2 "# "1� (%(�*! ! #4 "1� )#+�%" 4('%( 4)#, �()(% $(� "1� �%0 )#%,�%" #4 "1 ! 3()" #4 ��)( +') %2 "1� �(!" ��($ (� $#'�+ -� %4�))�+� " 9(! 3)#-5 (-�* #3�%� +)* (%+ )("1�) $#�+� 9 "1 "1� 3)�!�%$� #4 !#,� 4#)�!"�+ ()�(!/ )��� /��������"� ���"�� 0!�������� ����"���1 � �� ��� &� �"��� ���� "�� �"-�*2���� 03���� " �� ��1 �7� e 2/ = 5 � !") -'" #% #4 )#+�%"! 9 "1 + 44�)�%" �$#�#2 $(� 3)�4�)5 �%$�! % �(*�)! � (%+ 7 % "1� �()(% $( (0�/ � d 9##+5 �(%+ !3�$ �!� � d !3�$ �! %1(" %2 ,# !" ,�(+#9! (%+ ) 0�) -(%.!� d ,#'%"( % +9����)! (%+ 3�")#31 �#'! !3�$ �!� � d %1("(%"! #4 >�)#5,�!#31 �#'! �%0 )#%5 ,�%"!� � d %1("(%"! #4 !"�33� (%+ !�, 5() + )�2 #%!/ species relative abundance in % layer 2 layer 3 layer 4 spermophilus cf. citelloides 1.59 1.5 sicista subtilis 1.59 12 cricetulus migratorius 3.17 3 cricetus cricetus 4.76 7.5 mesocricetus newtoni 9.52 9.1 12.5 arvicola terrestris 1.59 9.1 2 clethrionomys glareolus 4.76 5 microtus subterraneus 11.11 27.3 7.5 microtus arvalis & m. agrestis 33.33 27.3 30.5 chionomys nivalis 7.94 7 microtus gregalis 6.35 lagurus lagurus 11.11 18.2 2.5 apodemus ex gr. sylvaticus-flavicollis 1.59 4 spalax leucodon 1.59 9.1 1 dryomys nitedula 2 muscardinus avellanarius 2 �(-/ < 5 �1� )��(" 0� (-'%+(%$�! #4 "1� )#+�%" !3�$ �! 6-(!�+ #% ���8 % + 44�)�%" �(*�)! % �()(% $(/ �2� #4 "1� 4('%( �0�% +') %2 "1� 3)�� , %()* %0�!" 2(" #%!� " 9(! �!"(-� !1�+ "1(" "1� +�3#! 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'�!� )��!�� $� ,� ������ $� $'� &���$�������� ��� ����$�$'��� ���"�#� ,*$ ��"� $/� �� +�*� #)����# -*+ � ��������� �+ ������� ��� )��,�,"� �+ �+( ��� �������� � ��� �+ �+( ��$���'. )��!�� $� ,� /���#)���� �"#� �� $'� ����$�$'�#( �'� �$'��# ��� $�)���" "�$�#$ &�##����� ��6 �����# -�+ ����� � ���� �+ � ����� �&���� �+ $�!���� � ���� �+ ������&�� �+ ����������. 0��/� �:�"*#�!�"� +��� ��+6 +����$ #�$�# �+ $'� &���$�������� ���"� -�+ ���������� ������� �3 #��� "���� �� #�$ ������� �� /4�������0 3��$ ����$� � /56 7����0 ?a? *� )+ , ����� -+.+?a@ taxon specimens l h h/l la la/l c c/h h h/h ribs e. subodessae mfea.g-1619 7.8 5.5 0.72 2.3 0.30 9+6 e. subodessae mfea.g-1617a rv 10.0 7.2 0.72 3.5 0.35 8+6 p. bollenensis bs.132.02.009/05 lv 26.2 22.5 0.86 8.2 0.31 7.5 0.33 3.2 0.14 20+7 p. bollenensis bs.132.02.009/01 lv 10.1 7.6 0.75 4.1 0.41 3.3 0.43 1.8 0.24 20+6 p. bollenensis bs.132.02.009 rv fragm.* 12.6 9.5 0.75 4.4 0.35 4.1 0.43 23+8 p. bollenensis mfea.g-1617b rv 12.6 ~9 0.71 4.2 0.33 20+7 p. incerta chiae bs.132.02.001 lv fragm.* ~9 ~6 0.7 ~3.3 0.37 14+7 p. cf. magdalenensis bs.132.01/03-1 lv ~7 5.6 0.80 2.5 0.36 2.0 0.36 17+6 p. cf. magdalenensis bs.132.01/03-2 rv* 7.2 5.0 0.69 3.2 0.44 20+ ? p. cf. partschi bs.132.02.006 cf. pontalmyra (?subgen. nov.) sp. bs.132.01/01 rv* 7.3 4.9 0.67 2.5 0.35 19+6 p. nevesskayae mpur7-3425 rv 6.2 4.7 0.76 1.9 0.31 1.4 0.30 0.7 0.15 23+7 p. nevesskayae bs.132.02.002-1 lv 6.0 4.0 0.67 1.8 0.30 1.2 0.30 24+6 p. nevesskayae bs.132.02.002-2 rv* 7.5 4.7 0.63 3.0 0.40 1.6 0.34 22+7 p. cf. pseudocatillus mfea.g-1620 lv 13.0 9.0 0.76 3.5 0.26 28+?5 p. cf. simplex bs.132.02.003 lv ~5 3.7 0.74 ~2 0.40 1.3 0.35 22+11 p. cf. submedius bs.132. 02.005 rv 10.5 10.0 0.95 ~5 0.47 3.0 0.30 22+? cf. pseudocatillus sp. bs.132.02.005/01-1 rv bs.132.02.005/01-2 rv ~10 11.3 7.8 8.0 0.78 0.71 ~4 0.40 3.0 0.38 1.7 0.22 23+~7 21+7 p. sabbae bs.132.03.001 lv bs.132.03.001 lv* 18.5 17.0 16.0 13.7 0.86 0.81 6.0 4.5 0.32 0.26 ~7 6.5 0.44 0.47 4.1 4.0 0.26 0.29 19+7 p. sabbae bs.132.01.001/01 lv 19.4 17.2 0.89 7.2 0.37 5.5 0.32 20+5 cf. prosodacnomya sp. mfea.g-1618 rv mfea.g-1615 rv 11.0 24.2 ~8 18.9 0.73 0.78 4.8 6.7 0.44 0.28 19+5 17+6 cf. pachydacna sp.1 mfea.g-1616 6.4 ~5 0.8 2.6 0.41 12+?5 cf. pachydacna sp.2 mfea.g-1614 21.9 7.9 0.36 4.4 0.20 7+? ��,( � 6 &��#*��� #'�"" �����#���#( �� !�"!� "��1$'= �� !�"!� '��1'$= "�� "��1$' �+ $'� ��$����� )��$ �+ $'� #'�"" +��� ,��0= �� !�"!� ���!�:6 �$�= '� ,��0 '��1'$( �,#� �*�,�� �+ ��,# �+ ��$����� o )�#$����� ���� �+ $'� !�"!�= � � "�+$ !�"!�= � ��1'$ !�"!�= �#$���#0 ������$�# ���#*�����$ �+ +��1���6 $��� #'�""# ,�#�� �� 1��/$' "���#( �"" �����#���# �� ��""���$��( lymnocardiinae in turin museum lymnocardiinae in alba museum sacco (1899) or name in collection revised name cavallo & repetto (1992) or name in collection revised name pontalmyra? bollenensis 02.007, 02.007/01; pontalmyra? spratti 02.009, 02.009/01/05, 02.008 pontalmyra bollenensis pl. 2, figs 4-9 limnocardium (s. l.) sp. g-1617a, g-1619 euxinicardium subodessae pl. 1, figs 8a, 9 pontalmyra carinata 02.001 pontalmyra incerta chiae pl. 2, fig. 10 limnocardium (s. l.) spratti g-1617b pontalmyra bollenensis pl. 1, fig. 8b limnocardium sp. 01../03 pontalmyra cf. p. magdalenensis pl. 1, figs 3, 4 limnocardium (s. l.) carinatum pontalmyra incerta chiae pontalmyra partschi 02.006 pontalmyra cf. p. partschi pl. 2, fig. 13 without name g-1620 pseudocatillus cf. p. pseudocatillus pl. 1, fig. 12 limnocardium sp. 01..01 cf. pontalmyra (?subgen. nov.) sp. pl. 1, fig. 1 without name g-1615, g-1618 cf. prosodacnomya sp. pl. 1, figs 13, 14 pontalmyra solitaria 02.002; pontalmyra castellinensis 02.004 pseudocatillus nevesskayae pl. 2, figs 17-19 without name g-1616 cf. pachydacna sp. 1 pl. 1, fig. 10 pontalmyra simplex 02.003 pseudocatillus cf. p. simplex pl. 2, fig. 14 without name g-1614 cf. pachydacna sp. 2 pl. 1, fig. 11 pontalmyra novarossica 02.005 pseudocatillus cf. p. submedius pl. 1, fig. 7 pontalmyra 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��� �� ����" $��"�*�"��� �"��' � $ �&�!���� �! ��##�!� +�;<>,� � $ �������� �"� � "�� &������7 $ ���"��� !� #�����7 � $ ������ !� -������� )��� ���"���� ��&�"��7 � $ ����. &����� !� �������7 � $ ���� ������� �"� ��&��� "��7 # $ ��������7 � $ �� ����" ��&��� "��7 � $ ������ ������88�� ��&��� "��7 � $ ��������7 9 $ ����� ��&��� "��7 . $ "�� &������7 � $ ������ ��&��� "��7 & $ ��. �� �"� �������' � $ ���!# +�;<>,� $ ��������7 � $ ��. ���7 ��� $ ��&��� "�� �"� ������ /�"���&�������*7 ��� $ "�� &������ /�"���&�������*7 �� $ �� ����" ��&��� "��7 !.�� $ ����� ������ !� � "�� &������7 !.& $ ����� ������ !� &���� )��� 1#��� �������7 !���� $ ����� ���� ��"� � "�� &������ �"� ��&��� "��7 ��� $ � ��"� � "�� &������7 �& $ � ��"� &����7 �# $ � ��"� #�����' $ ����� ���� # ��� �� ����" $����"� !"�� +g �� ����" $��"�*�"��� �"��, !��� ���"� �" ��� �� ����" ���� ��� ���"� � :!��"�$:���� +�;<@,� <$ %��"���� -�������7 <� $��. ��� #�����7 <� $��&��� "� �"� ������7 ; $ �� ����" ��&��� "��7 �> $ "�� &������ �"� &���� ��&��� "�� )��� 1#��� �������7 �� $ "�� &������ �"� ��&��� "��7 �� $ "�� &������7 ��� $ � "�� %�����' : $ ����� ���� # ��� �� ����" $����"� !"�� +g �� ����" $��"�*�"��� �"��, !��� ���"� �" ��� �� ����" ���� ��� ���"� � � ��� �� ��' +�;;@,� 9&@ $ %��"���� -�������7 9&? $ ��� "�� ��. ���7 9&c $ ������ !� ��&��� "��7 9� $ �� ����" ��&��� "��7 �� $ "�� &������7 ��� $ 1����7 �� $ ��&��� "�� �"� � "�� &������7 �&$� $ ���"��� !� %�����' � g �����7 ����' g ������ !�7 � g � )��7 1 g 1�����7 2 g 2!������7 h g ������ !�7 �4# �' g �4# ����"7 h�&&�' g h�&&�������"' %������##� �"� %���##� (" ) ����##� �"@=c lithologies age fossil records authors l senonian (campanian) mg. gr. lapparenti tricarinata, g. gr. lapparenti bulloides, g. rosetta g. gr. stuartiformis, g. elevata and hedbergella j. magné in rieuf 1980; middle senonian (?santonian) jglobigerinae and globotruncana (globotruncana gr. sigali, g. gr. marginata, g. gr. fornicata ) j. sigal in amaudric du chaffaut 1980; l cretaceous globotruncana, globigerinae amaudric du chaffaut 1977; l cretaceous globotruncana de booy 1957; limasset 1958; durand-delga 1975, 1984; rossi et al. 1994; e berriasian calpionella alpina , tintinnopsella carpatica, crassicollaria cf. parvula rieuf 1980; durand-delga 1984; cretaceous htrocholinae amaudric du chaffaut 1980; berriasiankimmeridgian propeneroplis, aptychus, belemnites, saccomidae, calpionella cf. alpina amaudric du chaffaut 1980; kimmeridgianoxfordian helicerina, diceras, protopeneroplis rossi et al. 1994; kimmeridgianoxfordian protopeneroplis, trocholinae, clypeina, favreina* durand-delga 1984; * see beauvais in rieuf 1980; l oxfordian cladophyllia dichotoma, stylina micromammata, stylohelia coalescens, aplosmilia crassa* l. beauvais, in rieuf 1980; l jurassic clypeina, solenoporees limasset 1958; amaudric du chaffaut 1977; m-?l jurassic protopeneroplis, lenticulina cf. vidalina de booy 1957; m jurassic rossi et al. 1994; durand-delga 1984; jurassic trocholinae de booy 1957; limasset 1958; durand-delga 1975; amaudric du chaffaut 1977, 1980; rieuf 1980; dogger-e malm protopeneroplis striata, rossi et al. 1994; durand-delga 1984; jurassic trocholinae amaudric du chaffaut 1977; jurassic or m jurassic trocholinae de booy 1957; limasset 1958; durand-delga 1975; amaudric du chaffaut 1977; dogger-? malm protopeneroplis striata, planiinvolutina carinata caporalino limestones rossi et al. 1994; durand-delga 1984; siliciclastic, coarse-grained deposits marls and/or limestones conglomerates arkosic flysch limestones and cherts s favreina salavensis, protocoprolithus centripetus, *full list in beauvais & rieuf 1980; protopeneroplis striata ���" +�&�!���� �! ��##�!� �;<>,' �(� ��� �� ��$ ��" ��&��� "�� ��� � "�� &������ ���� # 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��� �� ����" $��"�*�"��� �"�� )���� ����$ ��"��� ��� ��� ���(� !� ��� ������ �!� ��� ���������$ ��� # ���� !"��' ����� " ��� ���� ������������� #���!��� �"��!��"� ��� � &� ���� " # ��� � ����$����"�� ��� �$ ��� # ��� &����� !"��� ��� �� ����" $��"�*�"��� �"�� �� � ����. �"� � ����$ � #�"�$����"�� ������������� �!������ " ��!##�� )��� 1�� 8 �� ���� ����� +�'�' �$ � ����" ��&��� "��, �"� � ����$����"�� ��� ���� ���� # ���"���� ������ +�'�' �!"�� ���88 � � "�� &������ �"� -�������, ��&���� � ��� 1�� 8 �� ���� �����' ��$ � ���"� � ��� ������� !� "�"" # ����� ��� �"���� �$ � ����" $��"�*�"��� �"�� �� � ���� "��" +���� 1����� � ��"�, ������� )����' "��3!�"��� ����� ���!��� ��$ � )�� !� � ��9��� ��� ��� ������ )���� � ��!����� # � ���� !"�� ��� �4����"�� # � � "�$��(�� +1����� 2!������$ 1����� � ��"�, �"� �&��� ����&�"���� -���" )��� �" �����!��� & ��� � ���� ��!��� -� � ��"����&�"���� ���$ � "�� ����(���' �!� "�) �"���������� " � �"����� ���$ ������ )��� ��� ��� ������ # 1����!�� � �� !�� +�;=?, )���� ����!����� ���� �� ����" $��"�*�"��� �"�� �� �" � ��"� ������� )���� ���!&!����� �" � ���$ ��"��(� -���" # ��� �!� ���" � "��"�"��� &����" �$ ���� -���!�� ��##���"��� #� & ����� �!�� �� ��� ��!���� � �� �*����� �+ �� ��������&$����� �����#���&��� ,��*�#� -�� . ������/ @<; %������##� �"� %���##� (" ) ����##� �"@;> int. units metres sinonimies metres missing in the studied sections metres authors cfr. jaspes et calcaires (h) 0-12 amaudric du chaffeaut 1980; tèrme calcaréo-siliceux 0-30/50 amaudric du chaffeaut 1980; cfr. calcaires de caporalino (g) 0-30/50 amaudric du chaffeaut 1980; cfr. calcaire de caporalino (cc) 0-50 rieuf 1980; cfr. calcaires de caporalino p.p. (malm calcaries 9) 0-100 durand-delga 1984; cfr. calcaires de caporalino p.p. (js) 0/20-100 rossi et al. 1994; cfr. brèches p.p. (f) 0-< 10 amaudric du chaffeaut 1980; cfr. les conglomérats intermediaries (cgi) 0-< 10 rieuf 1980; cfr. calcaires de caporalino p.p. (9 malm calcaries): conglomérats 0-20 durand-delga 1984; cfr. calcaires de caporalino p.p. (js): conglomérats 0-20 rossi et al. 1994; cfr. jaspes et calcaires (e) 0-12 amaudric du chaffeaut 1980; cfr. series calcaire et siliceuse intermédiaires (csi) 0-50 rieuf 1980; cfr. formation calcaréo-siliceuse 0-50 durand-delga 1984; cfr. calcaire à lit siliceuse (jm6) 0-50 rossi et al. 1994; cfr. pélites micacées (d) 0-12 amaudric du chaffeaut 1980; cfr. grés et brèches a ciment calcaire (c) p.p. 0-25 amaudric du chaffeaut 1980; cfr. flysch greséux (b) 0-100 amaudric du chaffeaut 1980; cfr. les arkoses (a) 0-200 rieuf 1980; cfr. flysch arkosic (8c dogger-malm inférieur p.p.) 0-200 durand-delga 1984; cfr. arkoses de setonia (jm5) 0-250 cfr. brèches et conglomérats (a) p.p 0-300 cfr. les brèches p.p. (b) 0->300 cfr. formation de setonia (8b dogger-malm inférieur p.p.) 0-300 durand-delga 1984; cfr. brèches de francardo (jm4) p.p. 0-300 rossi et al. 1994 c ap or al in o lim es to ne s m b. m id dl e cl as ti c in te rv al l ow er c la st ic in te rv al 0-100 0-250 c ap or al in o br ec ci as a nd co ng lo m er at es m b. m er lo ng o f m . se to ni a sa nd st on es f m . 0-250 0-350 amaudric du chaffaut 1980 rieuf 1980 rossi et al. 1994; ; ; ��-' �� $ ����� "��"�� -��)��" ��� �����"�!����� !"�� +��� %��' <, )��� ����&���� ����."��� )��� ��� ���� � ����� ����&-����� �"� ����."��� ��� ���� �" �������!��' "�' g "���(��� nterva units metres sinonimies metres missing in the studied sections metres authors cfr. arkose et conglomérats (level m) 0-200 aumadric du chaffaut 1980 cfr. calcaire (level l) 0-40 aumadric du chaffaut 1980 cfr. conglomérats et calcaires (level k) 0-60 aumadric du chaffaut 1980 cfr. les grès (ge) eocène 0-300 rieuf 1980 cfr. les marnes (me) eocène 0-20 rieuf 1980 cfr. les conglomérats (cg-e) eocène 0-10 rieuf 1980 cfr. flysch de tonda (12a eocène détritique p.p.) 0-300 durand-delga 1984 cfr. pelites et marnes (12 eocène détritique p.p.) 0-50 durand-delga 1984 cfr. conglomérats (12 eocène détritique p.p.) 0-20 durand-delga 1984 cfr. flysch gréseux (em-s) 0-300 rossi et al. 1994 cfr. flysch gréseux de tonda (em-s p.p.) 0-300 rossi et al. 1994 cfr. marnes pélitiques ou gréseuses (em-s p.p.) 0-20 rossi et al. 1994 cfr. conglomérats (em-s p.p.) 0-50 rossi et al. 1994 conglomérats et calcaries (upcgl) 0-50 rieuf 1980 calcaries gréseux (11 upper paleocene) 0-20 durand-delga 1984 calcaries et conglomérats (ei) 0-50 rossi et al. 1994 cfr. calcaire argileux (level j) 0-50 aumadric du chaffaut 1980 cfr. marnes (ukm) crétacé supérieur 0-30 rieuf 1980 cfr. marno-calcaires (10 p.p.) 0-50 durand-delga 1984 cfr. marnes (cs1) rossi et al. 1994 cfr. brèches (level i) p.p. 0-100 aumadric du chaffaut 1980 conglomérats (ukcg) crétacé supérieur 0-30 rieuf 1980 cfr. conglomérats néocrétacés (10 crétacé supérieur) 0-50 durand-delga 1984 la chapelle de sant'angelo olistolith 0-12 aumadric du chaffeaut 1980 cfr. conglomérats (cs) 0-40 rossi et al. 1994 u pp er c la st ic in te rv al m id dl e cl as ti c in te rv al 030 0-120 sa nt 'a ng el o sa nd st on es f m . c ap or al in o m ar ls m b. p un ta c ap iz zo lo co ng lo m er at es an d b re cc ia s m b. 0-250 ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ���� ��� ���� ���!&!����� �" � ��" � ��"�� � &������(� -�$ ��" �!��"� ��� "� �"� ��# �&��� " # ��� � &���4 // �����"** ������� "��� )����' ��7���#��*� �� �' ��� �!�� �� ��� �"��-��� � �� #' �' :��$ ��" # � ��� ���"��#����� " # ��� # ��&�"�#��� �"� ��� #�!��#!� ����!��� " )���� ��� ) !� � �&�� (� ��� #���� (���� " # ��� &�"!������' e� ��� ��� �����#!� � "� # ��� �" "�& !� ��#����� )���� ���� !� � ������ �&�� (� ��� #�"�� (���� " # ��� &�"!������' e� ) !�� ��.� � ��." )����� :�' :' ��""�"� # � ��� #�"�� ������� " # ��� #�$ �!���' � �� �*����� �+ �� ��������&$����� �����#���&��� ,��*�#� -�� . ������/ @;� � � % � � � � � � �&�!���� �! 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��� �,��"�+"� &��� ��"� �� ����� $%�% & ���'�� (%�%110 chapskii n m or n m or n m or absolute length 2 85.0; 89.0 4 148.2 143.0-153.3 3 153.1 152.0-154.0 length of deltoid crest 2 55.0; 56.0 4 81.0 78.0-83.9 3 83.3 82.6-83.4 height of head 4 20.8 19.5-23.0 4 30.4 28.9-31.5 3 30.7 28.7-34.5 height of trochlea 3 15.0 14.0-16.0 21.0 4 29.5 28.0-31.0 3 27.8 27.7-28.0 width of head 4 23.3 22.0-25.0 4 34.5 34.0-35.1 3 35.9 34.6-38.0 width of deltoid crest 2 19.5; 20.0 4 36.8 31.0-41.5 3 45.5 44.1-50.0 width of distal epiphysis 2 33.0; 34.5 4 57.3 55.0-60.0 3 57.2 56.7-63.0 width of proximal epiphysis 2 29.0; 32.0 4 53.8 49.5-58.0 3 56.0 53.5-60.0 width of trochlea below 3 18.0 17.0-19.0 4 31.3 30.6-32.0 3 33.1 30.6-38.0 width of trochlea, frontal view 3 21.3 20.0-23.0 49.5 4 31.7 28.0-34.0 3 35.9 35.0-37.7 transverse width of diaphysis 4 18.3 16.5-21.5 27.0 4 23.6 23.2-24.0 3 23.9 22.2-27.0 thickness of proximal epiphysis 2 29.0; 35.0 4 49.1 44.7-53.5 3 53.9 53.7-68.0 thickness of medial condyle 3 16.3 15.5-17.0 23.0 4 25.0 24.0-26.1 3 25.7 24.1-27.0 thickness of lateral condyle 3 15.2 15.0-5.5 20.0 4 28.2 26.4-30.0 3 27.8 27.0-31.0 diameter of diaphysis with deltoid crest 2 31.5; 33.0 4 53.8 53.0-55.0 3 58.4 53.2-66.0 characters pachyphoca cystophora cristata ukrainica female male ��+� 0 # ����% �&�!�� .&&2 �* �%&� �� �! 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�2 ��%��"d 32 � �# !��" �!� �2 � �$�&�" ,��-�� (2 �� �������� ��+�� .����#� 7?#>05� �� �&&��% �2 �! ��%��" ,��-� �� �������� *�+%"� .�����#� 7?#>0/� �� �&&��% �� * �& ��� ��&� �!��,��%�" �� 7?#>052 �! �2 ��%��" �!� 42 � �!��" ,��-�� chapskii n m or width of proximal epiphysis 3 33.2 32.0 35.0 40.0 height of proximal epiphysis 3 20.0 18.0 22.0 27.0 width of distal epiphysis 2 20.5; 21.5 31.5 characters pachyphoca ukrainica ��+� a # ����% �&�!�� .&&2 �* ��+���� "�!)� ,� � -�"" ��,�"���� ��)� .+�))� �!� �� �!)� ���! �! �% �� ������2� ��� �!* ����!�%� *���� �� ���� �!� -��� ���! ��� �%� ����!�%� *����� �! ��!� ��� �� �% �� ������� ��� �� ����� � ����� �� ����'� ��� �� �!� -���� ��� �����" +� �� �* ��� �%� ����!�%� *���� �!�� �� � �� ��)�� "�!� .�� �! �% �� ������2d ��� ��%��" �!)"� �� !�� � ��� ,��� ��� )"�!��� *���� �� ���""�.���� �! �% �� ������2� -����%� � � �!�%!��� "��� �!� -����%� � +� �� .�! ��!� ��� �� �% �� ������2� ��� ��&�"��� +�!� �� ,� � ����'� ���,�� �!� &%�� +�))� �!� ����'� ���! 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? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? pachyphoca chapskii ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? mirounga angustrirostis 1 1 2 0 0 1 0 0 0 0 1 1 2 2 2 2 0 2 0 0 1 0 1 0 monachus schauinslandi 1 1 2 0 0 1 1 1 1 2 1 1 1 1 2 2 1 1 1 1 1 1 0 1 callophoca obscura 0 0 2 0 1 1 0 0 0 1 0 1 1 0 2 0 0 1 0 1 0 0 0 1 pliophoca etrusca 0 0 2 0 2 1 0 1 0 1 2 0 0 0 1 0 0 1 1 1 0 1 0 1 pontophoca sarmatica ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? ? ? 1 0 0 ? 0 0 phoca vitulina 0 0 0 1 0 1 1 2 1 0 1 0 0 0 0 0 1 0 1 1 1 1 1 1 leptophoca lenis 0 0 0 1 0 1 0 1 1 1 1 1 1 0 0 0 0 0 1 1 1 1 0 1 erignathus barbatus 0 1 0 1 0 1 1 2 1 0 1 1 1 1 1 2 0 0 0 1 1 1 1 1 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 lutra canadensis 0 0 1 0 1 0 1 0 0 1 0 1 0 1 2 0 0 0 0 1 1 0 1 1 devinophoca claytoni 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? cystophora cristata 0 0 1 1 0 1 2 1 1 0 0 1 0 0 2 1 0 1 2 0 0 1 1 1 pachyphoca ukrainica ? ? ? 1 0 2 0 2 2 0 1 0 0 0 0 0 0 1 0 0 1 0 1 0 pachyphoca chapskii ? ? ? ? 0 ? ? ? ? ? 1 ? 1 0 0 0 0 1 ? 0 1 ? ? 0 mirounga angustrirostis 0 0 1 0 0 1 2 2 2 1 1 0 1 1 2 1 1 0 2 0 1 2 1 1 monachus schauinslandi 1 0 1 1 1 1 0 0 1 1 1 1 1 1 0 1 1 1 0 1 0 1 1 1 callophoca obscura 1 0 2 1 1 0 1 1 1 2 0 1 0 0 0 0 1 0 1 1 1 0 0 1 pliophoca etrusca 1 0 1 1 1 0 1 0 2 1 0 0 0 0 0 0 1 1 1 0 0 0 0 1 pontophoca sarmatica 0 ? ? 0 1 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 0 phoca vitulina 0 1 2 0 2 1 0 0 0 0 0 2 1 0 1 0 1 1 0 1 1 1 0 1 leptophoca lenis 0 0 0 0 2 0 0 0 0 1 1 1 0 0 1 0 0 1 0 0 1 1 0 0 erignathus barbatus 1 0 0 0 2 1 2 2 2 0 1 1 1 0 1 0 1 1 2 1 1 1 1 1 ��+� /5�� + # ��� �$ *� ��� ���� #����� ���� *� �������� ��$� �!� �%�) �%� �!�"�9��� 3��� !�������� ���� &����� �* ����� � �&�# ��,� ��� ���� �� 4� �$�&�"�� ��� �&�""� ������� 2�����# ����� �� ������2 ��� � +���� #��,�"���� "���� � ����!# �� �* ��� *�&% ���! ��� "� )� ������� .�% ��������2� -��"� ��� "� )� ��� � ����'� �!� +�))� �!�� � ����!# �� �� � ���� ��� "� )� .�% ��������2 ��� �!!�&�!��� +�!�� -��� � ����� ��!���" �����+%"%&� �!� ��� &� )�!� �* ��� �����+%"� *���� � � ����� ��)� �+�,� ��� �"�!� �% *��� �* ��� +�!�� �! ��!� ���� ��� �&�""� ������� ��� � �%+�� -��� � +�)� -�""#��,�"���� ��)� *� ������&�!� �* ��� ���� ��� &%��"�� .-���� ��%�� �%�-� � ������! �* ��� ��� g��!�2� � ����'� -��� �!� �+%�� ������" ���!� *� ������&�!� �* ��� ������ )�'� �� &%��"� .�! �$��!�� �* ��� ��� g��!�2� �!� � ���� *���� �! ��� &����" ������ �* ��� �"�%& *� ������&�!� �* ��� " ����� '����� &%��"� .�"�� �! �$��!�� �* ��� ��� g��!�2� ��� � ���!�� �* ����� �!���&���" � ����� �!� ���# ��*���""� ��� -�""#��,�"���� "���� � ����!�� �* ��� *�# &% � �&�"��� ���� ��� �&�""� ���������� �� ������ -�� &� � ������� �� �� ��� ��" "���&����! .&� � � �&���,�2 ���! ��� "� )� �"���,�� �! �������!� +��� !�������� � � &� � � �&���,� �!� +���� ������� *� �� ��� ��" "���# &����! ���! �!� "�,�!) �� ���!����,�� �* ��� %+*�&�"� �������� �!��� ����+"�� ��� ��$%�" ��&� ����& �%�# ����� �! ��� �-� !�������� �* ���������� �� &� � �+,��%� ���! ��� ��&�"� ��� ���� � �! &��� ! �����# ��� �!�� .��� ��+� 0� 72� ���� !�&��� ��" �* *����" ���"� * �& ��� ����"� �����!� . � &����!2 �������� �* ��� �' ��!� �""�-� �&�!��� ���)!����� ����� �����!�� �!� ������&�!� �* )��) ����� ���� �+%���!� �� ��"� �"� �*� ���"�)�!���� �# "����!����� -����! ��� %+*�&�"� �������� �!�� .����# ���""� ��&�� ���!� -��� ��� ��!� �,� ���" )�!%� !�# ���"�2� ��� *����" ��� � �* �������� �!�� * �& ��� �� ��� ! 3"��' �� �)��! .�� �������! 3���!2 �� �! �) ��&�!� -��� ��� �� "�� ���������� ���� ��� � �)�! �* �������� -�� �! ��� ��"�!��� ����! �!�"%��!) ������ .�� ���'� � 3� !�� 05572� (������ ��� �!��&�"��� *����" �,���!�� *� �!���!� ����� ��" ����� !�� �% ����������� +���� �! ���� !�&��� ��"� �!������� ���� �������� �# !�� � �)�!���� �! ��� �� �������! 3���!� � �+�+"� &�# ) ���� -���-� � �+�%� //�0 &�""��! ��� � �)�� �!� �%+# ��<%�!�"� ��,���� �!�� ��� �-� &��� ! )�!� � . ����# ��� � �!� !� ���"�2� 4% ��� � !� ���"� ����"* ��,���� �!�� ��� !� ��� ! �!� ��%��� ! �"����!� ���"�� �� ��� ! �"����!� ���"� .!� ���"� ��"���� ��� ��2 �� ��� �� �%)� ��� �� �� ��"�!��� ����! -��� ������� � � ������ �!�� ��� � ���� �)��!� �%��� ! �"����!� ���"� .!� ���"� ������2 ����� ��� �!�� ��� �!�� ���� �)��! ,�� ��� �%�� ��"�!��� ����!� !% ��"���� ��� �� &�) ���� �� �%)� ��� 3� �!) �� �!� "��� �! +���&� ��� �� �� ����*�� �"����!� ���"� -��"� % � ������ +���&� ��� ������ ���"� �* ��� � ���� �� �"� �!� �� ��� ! ����*��� ����� !�*�!��!)� ��"� �$�"��! ��� � �)�!� ����� ��"� �!� ���"�)�!���� �"����!� �&�!) ��� ��,� �" �%+*�&�# "��� �* ��� 4�&�"� ��������� �% ���"�)�!���� �!�"����� +���� �! � �!��" �!� ����� �!��" ��� ���� � �* *����" �!� ����!� ���"�� �%�# �� �� � &�!����"���� 4�&�"� �������� ���� �!�"%��� ��� �%+*�&�"��� (�,�!������!��� �����!��� ��!����# �� ����� $%�% & ���'�� (%�%1?? 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