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 nori- ca (Triassìco superiore) nella località di Endenna nelle prealpi Lom- barde. 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 lom- bardo, 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 Ger- many. 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 Zor- zino, deposited in intraplatform basins some hundreds of meters deep and several kilomerres wide, surrounded by the huge Dolomia Principale (Hauptdoiomit) carbo- nate 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 inverte- brate life. Fossil findings consist mainly of fishes and are characterised by thousands of superbly preserved speci- mens, representing a well differentiated communiry, (Tintori, 1992), with also some previously unknown ge- nera (Tintori & Lombardo, 1996). Reptiles are much ra- rer, 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 fau- nal 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 Aetosdu- rus (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 ranystro- pheids and belonging ro rhe new genus Langobard.isau- ',4s (Renesto, 1994c) were found along with a tiny speci- men of the sphenodontid Diphydontosaurus (Renesto, 1995a). Most of these reptiles are of small or medium size, excluding the large specimen of psephoderma (Rene- sto & TintorL, 1995), nearly t.wo meters long, and a newly found complere phytosaur, currently under pre- paration, that reaches nearly four merers in length (Tin- tori 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 for- med 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, main- iy in the nofthern hemisphere, Europe, USA and India (I4cGregor, 1906; Gregory 1962; Chatterjee, 1978, Bal- lew; 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 fin- dings 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 be- longing to the genus Mystriosuchws and no description was provided. Buffetaut (1993) also mentioned the speci- men 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. Furt- her preparation by Mr. M. Pandolfi, on staff at the Mu- seum of Natural History of Bergamo, revealed the dorsal side, allowing a complete description and a positive ge- neric attribution, despite the fact that many of the bo- nes 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. Fun- hermore, 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", Ber- gamo (lombardy, Nonhern Italy). Curator Dr. Anna Paganoni. Cata- logue number MBSN 2. Provenance. IJppermost level of the Calcare di Zorzino (Zor- zino 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 nar- row, 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 rid- ges. Both show two medial rounded ridges on the ven- tral 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 spa- ced, 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 visi- ble, 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 premaxil- Iae are very extended posteriorly, nearly reaching the anterior border of the external nares. The maxillae are extremely difficult to recon- struct, because their dorsai region is crushed and split, and the dorsal outline m y not be observed; the antor- bital fenestra also is obliterated. About 20 rounded, spa- ced 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 an- terodorsally 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 mediola- terally. A ridge divides rhe squamosal into lateral and dorsal portions. The posterior process of the squamosal is short and not pointed as in Pseudopalatus (Hunger- bùhler, 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 quadratoju- gal. 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 nar- row; 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 ec- topterygoid 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 oblirera- ted but it can be reconstructed from the pattern of the surrounding bones. A small triangular supraoccipital is surrounded dorsaily and laterally by the parietals, for- ming an inverted IJ strucrure (Ballew; 1989; Hunt & Lu- cas, 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 occi- pital condyle. They continue lareroposreriorly into the long and stour paroccipital processes of the opisthotic, but no suture is visible between these bones. The paroc- cipital processes of the opisthotic are surrounded distal- ly by the squamosals and support dorsally the greatly depressed parieto-squamosal arcade. The posttemporal fenestra is reduced to a smail slit. The balllike basiocci- pital 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 Angi- storhinus, 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 be- longs to the genus Mystriosucbu.s: unserrated and proba- bly homodonr teeth that decrease in size posteriorly, heavily sculptured skull roof region and greatly depres- sed, compressed, supratemporal fenestra, inverted U sha- ped parietal-supraoccipital complex, short and not poin- ted 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 Mystrio- suchus 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 si- gnificantly more depressed than in M. planiro.srds. F{ow- ever, 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 pla- nirostris, 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 conrinen- tal, 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 car- casses coming from distant life environmenrs, as hypot- hesised by Renesto (1995b). FIowever, the recent finding of a new, Iarge (4 merer in length) phytosaur (Tintori et al., 1996), possibly another Mystriosucbus, requires chan- ging 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 clo- ser 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 en- demic 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 pro- vided 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 consi- dered 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 mari- ne environments, like some modern crocodilians. Ac- tually, Crocodylws porosus inhabits both freshwater and marine environments and can swim for several kilome- ters in the open sea (Ross & Magnusson, 1989); Mystrio- suchus 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. (1989) - Genetic analysis of the Phytosauria from the Late Triassic of the Western United States, pp. 309- 339 in S. G. Lucas Er A. P. Hunt (eds)- Dawn of the Age of Dinosaurs in the American Southwest. New Mexico Museum of Natural History, Albuquerque. Buffetaut E. (1993) - Phytosaurs in time and space. pp. j9-44 in Mazin J. M. 8{ Pinna G. (eds): Evolution, ecology and biogeography of the Triassic Reptiles. Paleont. Lom- bardaN. S. v. 2, Milano. Buffetaut E., Martin M. & Monod O. (1988) - Phltosaur re- mains from the Cenger formation of the Lycian Taurus (lfestern Turkey): Stratigraphical implications. Geobios, v. 21, pp. 237-243, Lyot Cal.zavara M., Muscio G. 6 \lild R. (1980) - Megalancosaurus preonensis, n. g.n. sp., a new reptile from the Norian of Frrrli. Gortania, v.2, pp. 59-64, Udine. Chatterjee S. (1978) - A primitive parasuchid (phytosaur) rep- tile from the Upper Triassic Maleri Formarion of India. Palaeontology, v. 27, pp. 83-127, London. Doyle K. D. & Sues H. D. (1995) - Phytosaurs (Reptilia: Ar- chosauria) from the Upper Triassic New Oxford forma- tion of York County, Pennsylvania. fourn. Wrt. Pa- leont., v. 15, pp. 545-553, Lawrence. Fraas E. (1896) - Die schwàbischen Trias-Saurier nach dem Material der Kgl Naturalien-Sammlung in Stuttgan zu- sammengestellt. Stuttgaft (Schweizerbart) Festgabe des Kónigl. Nat. Cabinets irr Stuttgart, 42. Versamml. der Dt. geol. Ges in Stuttgart, pp. 1-18, Stuttgart. Gregory I. T. (L962) - The genera of phytosaurs- Am. Journ. of Science, v 260, pp. 652-690, New llaven. FIuene F. von (1911) - Beitràge zur Kenntnis und Beurteilung der Parasuchier. Geol Palàont. Abh., N. F., v. 10, pp. 67-121, lena. FIuene F. von (1922) - Neue Beitràge zur Kenntnis der Parasu- chier. Jb. preuss. geol. Landesanst. v. 42, pp.59-160, Ber- lin. Hunt A. P. (1989) - Cranial morphology and ecology among phytosaurs- pp. 349-354 in S. G. Lucas Ec A. P. Hunt (eds)- Dawn of the Age of Diaosaurs in the American Southwest. Ne-w Mexico Museum of Natural Historv. Albuquerque. Hunt A. P. & Lucas S. G. (1989) - New Genotype designa- tions {or the phytosaurs Mystriosucbus anà Rutiodon with a discussion of the taxonomic status of Mystriosu- chus, Clepsysaurus and Rutiodon. pp. 340-348 in S. G. Lucas & A. P. Hunt (ed$- Dawn of the Age of Dino- saurs in the American Southwest. Ne-w Mexico Mu- seum of Natural History, Albuquerque. Jadoul F., Berra F. Er Frisia S. (1992) - Stratigraphic and paleo- geographic evolution of a carbonate platform Ír an ex- tensional tectonic regime: the example of the Dolomia Principale in Lombardy (italy). Rlz. It. Paleont. Strat., v. 98, pp. 29-44,Mitano. Long R. & Murry P. A. (1995) - Late Triassic (Carnian and Norian) tetrapods from the southwestern United States. Bull. Neu. Mexico. Mus. Natur. Hist. Sci., v. 4, 254 pp Albuquerque. McGregor J. H. (1906) - The Phytosauria with especial refer- erì.ce to Mystriosuchus and Rbytidodon. Mem. Am. Mus. Nat. Hist.,.r. 9, pp. 30-101, New York. Meyer H. von (1863) - Der Schàdel des Belodon aus dem Stu- bensandstein des oberen Keupers. Palaeontographica, v. 10, pp. 227 -246, Stuttgart. Pinna G. (1979) - Il cranio di un giovane Placochelide (Ae- pboderma alpinum Meyer, 1858) del Norico Superiore di Endenna (Bergamo). Atti Soc. It. Sc. Nat., v. 120, pp. D5-2A2, Milano. Pinna G. (1980) - Drepanosaurus unguicaudatus, nuovo genere e nuova specie di lepidosauro del Trias alpino. Atti Soc. It. Sc. Nat., v. 721, pp. 181.-192, Milano. Pinna G. (1984) - Osteologia di Drepanosaurus unguicaudatus, Lepidosauro triassico del sottordine Lacerttlía. Mern. Soc. It. Sc. Nat., v. 24, pp. 7-28, Milano. Pinna G. (1987) - I nuovi Lagerstàtten fossili del Triassico Ita- hano. Le Scienze, v. 224, pp. 62-7A,Mìlano. Pinna G. 6c Nosotti S. (1989) - Anatomia, morfoiogia funzio- nale e paleoecologia del rettile Placodonre Psephoderma alpinum Meyer, 1858. Mem. Soc" It. Sc. Nat, v. 25, pp. l./-fu- lvlrlano. Renesto S. (1984) - A new Lepidosaur (Reptilia) from the No- rian beds of the Bergamo Prealps. Rio. It. Paleont. Strat. v. 90, pp. 756-167, Milano. Renesto S. (1992) - The anatomy and relationships oÍ Enden- nasaurus acutirostris (Reptilia, Neodiapsida) from the Norian (Late tiassic) o{ Lombardy. Riv. It. Paleont. Strat. v. 97 (1991), pp. 4A9-430, Milano. Renesto S. (1993) - An isolated sternum oî Eudimorpbodon (R.eptilia, Pterosauria) from the Norian (Late Triassic) of the Bergamo Preaips (Lombardy, Northern Italy) Ria. It. Paleont. Strat., v. 99, pp. 415-422, Milano. Renesto S. (1,99a1 - Megalancosaurus, a possibly arboreai ar- chosauromorph (Reptilia) from the Upper Triassic of Northern kaly. Journ. Wrt. Paleont., v. 14, pp. 38-52, Lawrence. Renesto S. (1994b) - The shoulder girdle and anterior limb of Drepanosaurus unguicaud.atus (Reptiiia, Neodiapsida) from the Upper Triassic (Norian) of Northern Italy. Zool. Journ. Linn. Soc., v. 111, pp. 247-264, London. Renesto S. (1994c) - A new prolacertiform reptile from the Late Triassic of Northern ltaly. Riv. It. Paleont. Strat., v. 100, pp. 285-306, Milaro. Renesto S. (1995a) - A sphenodontid from the Norian (Late Triassic) of Lombardy (Northern Italy): A preliminary note. Modern Geology, v. 20, 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 Litographic Limestones, pp. 127-129, Madrid. Renesto S. & Tintori A. (1995) - Functional morphology and mode of life of the Late Triassic placodont Psephoderma alpinum Meyer from the Calcare di Zorzrno (Lombar- dy, N. Italy). Riv. It. Paleont. Strat., v. 101, pp. 37-48, Milano. Ross C. A. & Magnusson \L E. (1989) - Living Crocodilians, pp. 58-73 in Ross C. A. (ed.) Crocodiles and alligators, Golden Press, Silverwater. A pbytosaur skull from the Norian of Lombardy 121 Tintori L. (1992) - Fish taphonomy and Triassic anoxic Ba- sins from the Alps: a case history. Rio. It. Paleont. Strat. v.97 (1991), pp. 393-408, Milano. Tintori A. Muscio G. & Nardon S. (1985) - The Triassic fos- sil Fishes localities in Italy. Rip It. Paleont. Strat. v.91, pp. 127-210, Milano. Tintori A. & Lombardo C. (1996) - Gabanellia agilis gen. n. sp. n. (Actinopterygii, perleidiformes) from the Calcare di Zorzlno of l-ombardy (North Italy). Rio. It. Paleont. Strat. v. 102, pp. 227-236, Milano. Tintori A., Renesto S., Iombardo C., Manarolla G., Manenti M. Er Vendico M. (1996) - An exceptional reptile find in the Norian (Late Triassic) Lagerstàtte of Endenna (Zogno, Bergamo, Italy). Riz,. It. PaLeont. Strat. v. 702, pp. 131,-134, Milano. l7estphal F. (1976) - Phytosauria. pp. 99J2A In O. Kuhn (Ed.): Handbùch der Palàoherpetologie, v. 13. Gustav Fischer Verlag, Stuttgart. \íild R. (1978) - Die Flugsaurier (Reptilia, Pterosauria) aus der Oberen trias von Cene bei Bergamo, Italien,. Boll. Soc. Paleont. It., v. 17, pp. 176-256, Modena. \fild R. (1991) - Aetosaurus (Reptilia, Thecodontia) from theUpper Triassic (norian) of Cene near Bergamo, Italy, with a revision of the genus. Rio. Mus. Cio. Sc. Nat. Bergamo, v. 14, pp. 1-24,Bergamo.