Geo.Cro.2-3-61-KB.pdf 333 � Daniela Basso1, Davor Vrsaljko2 and Tonći Grgasović3 AB STRA CT The fossil coralline fl ora of the Badenian bioclastic limestone outcropping in Northern Croatia is known by the name “Litavac”, shortened from “Lithothamnium Limestone”. The name was given to indicate that unidentifi ed coralline algae are the major component. In this fi rst contribution to the knowledge of the coralline fl ora of the Litavac, Lithoth- amnion valens seems to be the most common species, with an unattached, branched growth-form. Small rhodoliths composed of Phymatolithon calcareum and Mesophyllum roveretoi also occur. The Badenian benthic association is dominated by melobesioid corallines, thus it can be compared with the modern maërl facies of the Atlantic Ocean and Mediterranean Sea. Since L. valens still survives in the present-day Mediterranean, an analogy between the Badenian Litavac and the living L. valens facies of the Mediterranean is suggested. Keywor ds: calcareous Rhodophyta, Corallinales, rhodoliths, maërl, Badenian, Croatia 1 Dipartamento di Scienze Geologiche e Geotecnologie, Università degli Studi di Milano – Bicocca, Piazza della Scienza 4, 20126 Milano, Italy; (daniela.basso@unimib.it) 2 Croatian Natural History Museum, Demetrova 1, HR-10000 Zagreb, Croatia; (davor.vrsaljko@hpm.hr) 3 Croatian Geological Survey, Sachsova 2, HR-10000 Zagreb, Croatia; (tgrgasovic@hgi-cgs.hr) The coralline fl ora of a Miocene maërl: the Croatian “Litavac” Geologia Croatica 61/2–3 333–340 1 Fig. 1 Pl. Zagreb 2008 Geologia CroaticaGeologia Croatica � 1. INTRODUCTION Since Roman times, a building stone named “Litavac” has been quarried at many Croatian localities, particularly on the SW slopes of the Medvednica Mts. (Fig. 1). The name derives from the shortening of the Croatian words litotamnijski vap- nenac meaning “Lithothamnium Limestone”. The Litavac corresponds to a facies of biolithites and bio- clastic limestones (e.g. biocalcirudites, biocalcarenites and biocalclutites) belonging to the Upper Badenian succession (Lower Serravallian of the standard chronostratigraphy; de- tails in PILLER et al., 2007) which are characterized by fully marine environments in Northern Croatia (KOCHANSKI, 1944; ŠIKIĆ, 1967, 1968; AVANIĆ et al., 1993; VRSALJKO et al., 2005, 2006). The Upper Badenian deposits unconform- ably overlie older rocks, and in the area of Mt. Medvednica over Upper Triassic dolomites. The Litavac facies is inter- posed between a facies of coarse-grained clastics (with clasts from underlying dolomites and limestones), at the base and an overlying facies of fi ne-graded clastics: fi ne-graded sands, marls, clayey limestones and calcsiltites (VRSALJKO et al., 2006, 2007a). Litavac is almost structureless, but cross-bed- ding or even fl at-bedding can occasionally be observed. A description of the stratigraphy and palaeogeography of Miocene deposits from the investigated area (SW Medved- nica) and neighbouring Samoborsko gorje – Žumberak Mts. is provided by VRSALJKO et al. (2005). The sedimentology and palaeoenvironmental evolution of Upper Badenian and Sarmatian deposits of Mt. Medvednica is discussed by VR- SALJKO et al. (2006). The Litavac is composed predominantly of corallines with fragments of bivalves including Lucinoma boreale (L.), Nuc- ula sp., Pectinidae, Ostrea sp., Arcidae, Glycymerididae, Car- diidae, Glossus humanus (L.), Corbula gibba (OLIVI), gas- tropods (Conus sp. Ficus sp.), echinoderms (Clypeaster sp.), marine benthic foraminifers (Textulariidae, Planostegina sp., Amphistegina sp., Brizalina dilatata (REUSS), Heterolepa dutemplei (D’ORBIGNY) with common Lobatula lobatula Geologia Croatica Geologia Croatica 61/2–3 334 (WALKER & JACOB), Elphidium fi chtelianum (D’OR BIG- NY) and rare planktonic forams), ostracods (i.e. Cytheretta tenuipunctata dentata BRESTENSKA), bryozoans, hydro- zoans and corals (Flabellum sp.), denoting a shelf palaeoen- vironment within the infralittoral to shallow circalittoral zones (VRSALJKO et al., 2007a, c,d). The skeletal fragments are cemented with calcite. The name “Lithothamnium limestones” refers to the dom- inance of coralline nodules and remains but has simply a litho- genetic sense, since several genera of non-geniculate coralline algae other than Lithothamnion can be identifi ed. The same is true for the Tortonian “Calcari a Briozoi e Litotamni” of the central-southern Apennines in Italy, (= Bryozoan and Lithoth- amnium limestones; SIMONE & CARANNANTE, 1985; BRANDANO, 2002), for the Badenian Leitha Limestone of the Vienna basin (=Nullipora, Lithothamnium or Lithotham- nion limestone; RIEGL & PILLER & 2000), for the Badenian Leithakalk units of Hungary (RANDAZZO et al., 1999) and for the Early Badenian Pinczow Limestones in Poland (STU- DENCKI, 1988). Despite the historical use of Litavac as a building stone (more than 2250 m3 were used to build the present Zagreb Ca- thedral; VRSALJKO & al., 2007a,b), and its importance as a reservoir rock for water and oil and gas (LUČIĆ et al., 2001), the taxonomy of the coralline algae composing the limestone has never been investigated in detail. The aim of this paper is to provide a fi rst account of the fossil coralline fl ora of the northern Croatian Litavac, and its possible palaeoenvironmen- tal interpretation. 2. MATERIAL AND METHODS Sampling has been performed at the quarries of Gornje Vrapče and Bizek (NW of Zagreb, Medvednica Mt.; Fig. 1), in the bioclastic limestones facies of Upper Badenian age. Figured specimens are from the Bizek quarry where the Litavac crops out and is about 30 m thick (see VRSALJKO et al., 2007d). Observations of coralline anatomy and biometry have been made on thin sections. Thallus nomenclature follows BASSO et al., (1997). In particular, the terms cortex and medulla, in the description of protuberances, are used with a merely topo- graphic sense, to distinguish the central cell fi laments running parallel to the main axis of the protuberance (= the medulla) from the derivatives diverging and bending toward the surface of a protuberance (= the cortex). The terms unattached branch- es and prâline are used according to BASSO (1998). Growth- form nomenclature follows WOELKERLING et al., (1993). Percentage quantifi cation of total coralline algal thalli is based on visual estimate. 3. RESULTS The Litavac from the studied localities is a calcareous rud- stone, mainly composed of fragments of fruticose corallines (VRSALJKO et al., 2007d, fi g. 2). The Litavac corallines are free-living, unattached branches or small rhodoliths with var- ious degrees of protuberance development. The Litavac un- derwent dolomitization and dissolution, preventing identifi ca- tion of a large part of the algal remains. However, despite Fi gu re 1: Map of the investigated Litavac quarries in the Zagreb area, Croatia. Geologia CroaticaDaniela Basso et al.: The coralline fl ora of a Miocene maërl: the Croatian “Litavac” 335 fossil diagenesis, numerous algal fragments still have their microscopical anatomy preserved, therefore allowing identi- fi cation. 3.1. The coralline fl ora Division Rhodophyta WETTSTEIN, 1901 Order Corallinales SILVA & JOHANSEN, 1968 Family Hapalidiaceae GRAY, 1864 emend. HARVEY, BROADWATER, WOELKERLING & MITROVSKI, 2003 Subfamily Melobesioideae BIZZOZERO, 1885 Genus Phymatolithon FOSLIE, 1898 Phymatolithon calcareum (PALLAS) ADEY & MCKIBBIN, 1970 (Pl. 1, Fig. 1) Neotype: BM Box Collection No. 1626 (WOELKERLING & IRVINE, 1986). Basionym: Millepora calcarea PALLAS 1766, p.265. 1943 Pomatophyllum operculatum CONTI, p. 52–54, fi g. 3, pl. VI/4, pl. VIII/5 1950 Lithothamnion operculatum CONTI, p. 117 1997 Phymatolithon calcareum – BASSO et al., p. 168–170, pl. 36 The lumpy growth-form of this species has been identi- fi ed in a single rhodolith, about 6 mm in diameter, among dominant branches of L. valens. The thallus shows a thin hy- pothallium, with cells L 6–11 x D 6–7 μm, a perithallium with cells L 8–11 x D 7–8 μm and rounded multiporate concepta- cles with diam. 125–150 x 75–90 μm in height. Although the epithallial cells are not preserved, the specimen is attributed to Phymatolithon calcareum on the basis of the correspond- ence of all the other known anatomical features (BASSO et al., 1997). P. calcareum is distributed from the Oligocene to Recent (BASSO et al., 1997). Genus Mesophyllum LEMOINE, 1928 Mesophyllum roveretoi CONTI, 1943 (Pl. 1, Figs. 2–3) Protologue: CONTI, 1943, p.55, pl. VII, fi g. 1a–c; pl. 8, fi g. 6. Rhodolith-forming plant with a lumpy growth-form shows stout protuberances reaching about 2 mm in length. Each thal- lus barely exceeds 450 μm in thickness, but several thalli are superimposed to give the total thickness. In thin sections, the protuberances appear to have originated from the superposi- tion of fertile and sterile layers of the thallus, which thicken up in correspondence with the conceptacle chambers. Hypoth- allium coaxial is 100–200 μm thick, with cells L 22–25 x D 12–14 μm. Perithallial cells are L 9–16 x D 8–9 μm. Multipo- rate conceptacles abundant has D 275–425 x H 155–205 μm with roof thickness of 50–75 μm. Several conceptacle cham- bers show a partial infi ll of irregularly-shaped, large “cells”. The stratigraphic distribution of M. roveretoi ranges from the Upper Eocene to the Miocene of the Tertiary Piedmont Basin (FRAVEGA et al., 1987) and in the Badenian of the Leitha Limestone (Vienna Basin; CONTI, 1946b). Genus Lithothamnion HEIDRICH, 1897 Lithothamnion valens FOSLIE, 1909 (Pl. 1, Figs. 4–6) Holotype: TRH, unnumbered; includes slides 1731–1733. Fig ur ed by BASSO et al. (1997, pl. 37, fi gs. 1–8; pl. 38, fi gs. 3, 5, 6). 1946a Lithothamnium ramosissimum (GÜMBEL non RE- USS) CONTI, p.18–22, pl. I/1 a–f; pl. VII/1–3 1997 Lithothamnion valens – BASSO et al. (1997), p. 170– 176 The species is common as branches and their fragments, with diameters ranging from 2 to 4–5 mm. Also some prâlines with fruticose growth-forms have been identifi ed. Single pro- tuberances (branches) can regularly exceed 2 cm in length. Fragments of branches containing the hypothallium have not been observed. The perithallium shows a sharp zonation, with large rectangular cells (L 17–24 x 9–12 μm) at the base of each growth zone, gradually decreasing in size toward the more square shaped cells at the top of the same zone (L 10–12 x 9–12 μm). This superimposed growth zone begins sharply with the large cells, without any transition. Multiporate con- ceptacle chambers D 260–380 x H 105–120 μm show roof thickness ranging from 32 to 45 μm. The stratigraphic distri- bution of L. valens ranges from the Priabonian to the Recent (BASSO et al., 1997). 3.2. The facies On the basis of the lithology and fossil components (relative abundance of prâlines, more or less fragmented coralline un- attached branches, small and large benthic/planktic foramin- ifera and molluscan remains) at least three different facies can be distinguished within the Bizek limestone. Branches dominate in facies A, small rhodoliths dominate in facies B, corallines are fragmented and probably were not autochthonous in facies C. Facies A and B were also recog- nized at Gornje Vrapče (Fig. 1), though their geometry and palaeoenvironmental / stratigraphic relationships need further investigation. Facies A (Pl. 1, Fig. 7) has been sampled at about 50 cm above the Triassic base. It represents a micro-breccia with an- gular and sub-rounded clasts of dolomite mixed with bioclasts. Coralline branches and their fragments are dominant in the limestone. Other components include centimetre to millime- tre-sized fragments of bryozoan colonies and small benthic foraminifera (Lobatula lobatula, Textularidae, Elphidiidae, Miliolidae). External moulds of large Arcidae and Glycymeris sp. randomly occur. Dolomitization of bioclasts is extensive, and was followed by dissolution, both of bioclasts and litho- clasts. The resulting mouldic porosity involves about 20–30% of the rock. The well preserved algal fragments (about 20–40% of the total algal fragments) are irregularly cylindrical, 1–5 mm in diameter and up to about 1 cm in length. Corallines with lumpy to fruticose growth-forms compose sparse, small prâlines (sen- su BASSO, 1998), 1–3 cm in diameter. Most algal remains belong to L. valens, with the contribution of P. calcareum and M. roveretoi. Geologia Croatica Geologia Croatica 61/2–3 336 PLATE 1 1 Phymatolithon calcareum (PALLAS) ADEY & MCKIBBIN, 1970. Two conceptacles with the characteristic roof. Note thallus zonation and abraded thallus surface. Thin section BIZ1, Bizek quarry, OM photograph. 2 Mesophyllum roveretoi CONTI, 1943. Several superimposed fertile thalli which compose a protuberance. The coaxial hypothallium is visible. Thin section CRO4/1C, Bizek quarry, OM photograph. 3 Mesophyllum roveretoi CONTI, 1943. Detail of the multiporate conceptacles. The improper thallus orientation hides the coaxial organization of the hypothallium. Thin section CRO4/1C, Bizek quarry, OM photograph. 4 Lithothamnion valens FOSLIE, 1909. An oblique section of a protuberance showing the typical thallus zonation and multiporate conceptacles. Note diagenetic fractures deforming the original conceptacle shape. Thin section CRO4/1A, Bizek quarry, OM photograph. 5 Lithothamnion valens FOSLIE, 1909. A longitudinal section of a protuberance showing the sharp zonation in the medulla and the multiporate conceptacles developing in the. cortex. Thin section BIZ6, Bizek quarry, OM photo- graph. 6 Lithothamnion valens FOSLIE, 1909. Detail of the multiporate conceptacles of Fig. 5. Note the preservation of some (?sporangial) remains hanging from the roof of the conceptacle. Thin section BIZ1, Bizek quarry, OM photograph. 7 Litavac facies A (Bizek quarry, sample BIZ 1). Note the preserved fossil algal thalli appearing as white, irregular cylindrical clasts in the limestone (thick arrows on branching growth-forms). Mouldic porosity due to dissolution of dolomitic clasts is indicated by thin arrows. 8 Litavac facies B (Bizek quarry, sample BIZ 6). Note the common occurrence of small rhodoliths (thick arrows), together with branching growth-forms and the abundant larger foraminifera (arrows). 9 Litavac facies C (Bizek quarry, sample BIZ 8). Most coralline algae that occur as micro-fragments are detectable only under O.M. Benthic and planktic foraminifera are the major components. Note also common Ditrupa (thick arrows) and molluscan (arrow) shell fragments. Geologia CroaticaDaniela Basso et al.: The coralline fl ora of a Miocene maërl: the Croatian “Litavac” 337 Geologia Croatica Geologia Croatica 61/2–3 338 Two other facies B and C (Pl. 1, Figs. 8, 9) appear as lat- eral variations at the same stratigraphic level, at about 15 m height within the Upper Badenian strata. In the algal biocal- carenite of facies B (Pl. 1, Fig. 8), which represents the most typical Litavac, the fragments of coralline branches become less abundant and smaller, while small rhodoliths become com- mon. The coralline fl ora is composed of Lithothamnion spp. and Mesophyllum spp. Large benthic foraminifera (Amphiste- gina and Planostegina) are the co-dominant components. In facies C (Pl. 1, Fig. 9), an evident orientation of the grains is observed. It represents a mud supported micro-con- glomerate with small sub-rounded dolomitic clasts in a matrix of fi ne grained crushed bioclasts. The dominant components are the large benthic foraminifera, mostly as fragments, to- gether with small benthic and common planktic foraminifera. Other components are small gastropods and fragments of bi- valves together with annelid tubes. Millimetre-sized fragments of uncommon corallines also occur. 4. DISCUSSION AND CONCLUSIONS The accumulation of prevalently loose-lying non-geniculate, branched corallines corresponds to the modern concept of maërl. Maërl is a Breton word defi ning a benthic association dominated by free-living, branching algal thalli. Along the north-western coast of France, at many localities in Ireland, and in Galicia, maërl covers large areas of the infralittoral sea- fl oor at < 20 m of water depth in a macrotidal environment, thus exposed to oscillating tidal currents (CABIOCH, 1969; BIOMAERL team, 2003). In the Mediterranean, maërl occurs infrequently at about 40 m of water depth, in current-swept, coastal bio-detritic, benthic environments of the upper circalit- toral zone (BASSO & BRUSONI, 2004). The available literature reports analogues of fossil algal facies in the Miocene of Europe. The “branching algae facies” reported by STUDENCKI (1988) occurs in the basal layer of the Pinczow Limestone outcropping west of Pinczow (NE of Cracow, Poland). It is composed of an accumulation of branch- ing thalli of Sporolithon sp. and Lithothamnion valens (re- ported as Archaeolithothamnium sp. and Lithothamnium ra- mosissimum (GÜMBEL) respectively in STUDENCKI, 1988) and subordinate encrusting growth-forms of Mesophyllum in- gestum CONTI, 1946b and Neogoniolithon contii (MASTRO- RILLI) QUARANTA, VANNUCCI & BASSO, 2007 (report- ed as Mesophyllum rigidum MASTRORILLI in STUDENCKI, 1988). This facies has been interpreted as the fi rst step of an ecological succession leading to a rhodolith pavement along a shallowing trend (STUDENCKI, 1988). Maërl occurs as a facies of the Leitha Limestone in the Vienna Basin (DULLO, 1983; RIEGL & PILLER, 2000). Sev- eral species of non geniculate corallines have been identifi ed in the Austrian Leitha Limestone, among which the most com- mon are Lithothamnion valens, Lithothamnion ramosissimum (REUSS) PILLER, 1994, Phymatolithon calcareum, Meso- phyllum roveretoi, Spongites albanensis (LEMOINE) BRA- GA, BOSENCE & STENECK, 1993 (for reference: CONTI, 1946b; PILLER, 1994; BASSO et al., 1997). Among these species, L. valens and L. ramosissimum commonly show a free-living, branching growth-form, while the others have been identifi ed as encrusting thalli or rhodolith builders. A cool-water, algal rudstone/fl oatstone microfacies com- posed of whole and fragmented red algae has been described for the Hungarian Leithakalk (RANDAZZO et al., 1999). Cor- allines appear in numerous growth-forms, can be referred to maerl or prâlines and are associated with benthic foraminifera (Amphistegina and Heterostegina) and worm tubes. They are reported to belong to several coralline genera, including Litho- thamnion, Sporolithon (quoted as Archaeolithothamnion), Meso phyllum and Lithophyllum. Unluckily, the absence of palaeontological descriptions and insuffi cient illustration pre- vent confi rmation and more detailed identifi cation of the re- ported taxa (RANDAZZO et al., 1999). Branch fragments of corallines in a marly matrix, pos- sibly comparable with a maërl, occur in the Badenian algal limeston es of the Transylvania Basin (Romania; BUCUR & FILI PES CU, 1994). The diversifi ed coralline fl ora of the Badenian algal limestones includes several species of the genera Sporolithon, Lithothamnion (among which L. ramo- sissimum, quoted as P. archaeotypum CONTI), Mesophyllum (among which M. roveretoi), Lithophyllum and Spongites albanensis (quoted as Lithophyllum?albanense LEMOINE), mostly composing rhodoliths (BUCUR & FILIPESCU, 1994). These observations support the conclusion that in Para- tethys, a Badenian maërl was predominantly composed of Lithothamnion (free-living branches) and Mesophyllum, with locally subordinate Sporolithon, Spongites and Phymatolithon (in rhodoliths). We can attempt a comparison of this Badenian fossil as- sociation with the present-day maërl. The coralline association of the Atlantic maërl (NW France, Ireland and Galicia) is dom- inated by Lithothamnion corallioides (P.L. & H.M. CROUAN) P.L. & H.M. CROUAN, 1867 and Phymatolithon calcareum. Other species contributing to the Atlantic maërl are Lithoth- amnion glaciale KJELLMAN, Lithothamnion tophiforme (ESP ER) UNGER, 1858, Lithophyllum dentatum (KÜTZ- ING) FOSLIE, 1900, Lithophyllum fasciculatum (LAMARCK) FOSLIE, 1900 and, more occasionally, Mesophyllum. The Mediterranean maërl has a diversifi ed algal fl ora, with the occurrence of several species of Lithothamnion, Phyma- tolithon, Lithophyllum and Mastophoroideae. Presently, L. valens is a Mediterranean endemic, infrequently occurring at considerable depth (50–70 m), mainly in the Western Medi- terranean (BASSO, 1996; BABBINI & BRESSAN, 1997). In the tropics, an analogous maërl (based on the algal growth- form) is represented by the accumulation of unattached branch- es of Neogoniolithon in seagrass meadows (BASSO et al., in press). The Badenian maërl has no obvious analogies with the tropical Neogoniolithon facies. The Atlantic and the Mediter- ranean maërl and their Badenian fossil counterparts are dom- inated by melobesioid corallines. However, one of the most characteristic and abundant species in the Badenian maërl, Lithothamnion valens, does not occur in the present-day maërl association living in the Atlantic. Since L. valens still occurs in relatively deep Mediterranean waters, we conclude that the Geologia CroaticaDaniela Basso et al.: The coralline fl ora of a Miocene maërl: the Croatian “Litavac” 339 temperate-water, modern Mediterranean maërl is fl oristically the most similar analogue of the Badenian Litavac. However, direct palaeodepth interpretations based on the Mediterranean occurrence of L. valens would be speculative, until further palaeoecological investigations based on the whole benthic association and sedimentary environment of the Litavac have hopefully clarifi ed the matter. REFERENCES ADEY, W.H. & MCKIBBIN, D. (1970): A revision of the Foslie crustose coralline herbarium. – K. Norske Videns. Selsk. Skr., 1, 1–46. AVANIĆ, R., KOVAČIĆ, M., PAVELIĆ, D., MIKNIĆ, M., VRSALJKO, D., BAKRAČ, K. & GALOVIĆ, I. (2003): The Middle and Upper Miocene Facies of Mt. Medvednica (Northern Croatia). – In: VLA- HOVIĆ, I. & TIŠLJAR, J. (eds.): Stages in Evolution of the Mio- cene North Croatian Basin (Pannonian Basin System). 22nd IAS Meeting of Sedimentology, Field Trip Guidebook, 167–172. BABBINI, L. & BRESSAN, G. (1997): Recensement des Corallinacées de la Mer Méditerranée et considérations phytogéographiques. – Bibliotheca Phycologica, 103, 421 p. BASSO, D. (1996): Soft bottom Mediterranean calcareous algae (non- geniculate Corallinaceae): distribution and ecology. – In: ALBER- TELLI, G., DE MAIO, A. & PICCAZZO, M. (eds.): Atti dell’11 Congresso A.I.O.L., Sorrento, 26–28 ottobre 1994, 225–234. BASSO, D. (1998): Deep rhodolith distribution in the Pontian Islands, Italy: a model for the paleoecology of a temperate sea. – Paleo. Paleo. Paleo., 137, 173–187. BASSO, D. & BRUSONI, F. (2004): The molluscan assemblage of a transitional environment: the Mediterranean maërl from off the Elba Island (Tuscan Archipelago, Tyrrhenian Sea). – Boll. Malacol., 40, 37–45. BASSO, D., FRAVEGA P. & VANNUCCI, G. (1997): The taxonomy of Lithothamnion ramosissimum (Gümbel non Reuss) Conti and Litho- thamnion operculatum (Conti) Conti (Rhodophyta, Corallinaceae). – Facies, 37, 167–182. BASSO, D., NALIN, R. & NELSON, C.S. (in press): Shallow water Spo- rolithon rhodoliths from North Island (New Zealand). – Palaios. BIOMAERL team (2003): Conservation and management of northeast Atlantic and Mediterranean maërl beds. – Aquatic Conservation: Marine and Freshwater Ecosystems, 13/1, S65–S76. BRAGA, J.C., BOSENCE, D.W.J. & STENECK, R.S. (1993): New an- atomical characters in fossil coralline algae and their taxonomic implications. – Palaeontology, 36, 535–547. BRANDANO, M. (2002): La Formazione dei “Calcari a Briozoi e Lito- tamni” nell’area di Tagliacozzo (Appennino centrale) e considera- zioni paleoambientali sulle facies rodalgali. – Boll. Soc. Geol. It., 121, 179–186. BUCUR, I.I. & FILIPESCU, S. (1994): Middle Miocene Red Algae from the Transylvanian Basin (Romania). – Beitr. Paläont., 19, 39–47. CABIOCH, J. (1969): Les fonds de maërl de la baie de Morlaix et leur peuplement végétal. – Cahiers de Biologie Marine, 10, 139–161. CONTI, S. (1943): Contributo allo studio delle Corallinacee del terziario italiano. II: Le Corallinacee del Miocene del Bacino Ligure-Pie- montese. – Palaeont. Ital., 41, 37–61. CONTI, S. (1946a): Revisione critica di Lithothamnion ramosissimum Reuss. – Pubbl. Ist. Geol. Univ. Genova, Serie A, 1, 3–29. CONTI, S. (1946b): Le Corallinaceae del calcare miocenico (Leithakalk) del Bacino di Vienna. – Pubbl. Ist. Geol. Univ. Genova, Serie A, 2, 31–68. CONTI, S. (1950): Alghe Corallinaceae fossili. – Pubblicazioni dell’Isti- tuto di Geologia dell’Universita di Genova 4 (ser A), 1–155. CROUAN, P.L. & CROUAN, H.M. (1867): Florule du Finistère. – Frie- drich Klincksieck & J.B. et A. Lefournier. DULLO, W.C., (1983): Fossildiagenese im miozänen Leitha-Kalk der Paratethys von Österreich: Ein Beispiel für Faunenverschiebung durch Diageneseunterschiede. – Facies, 8, 1–112. FOSLIE, M. (1900): New or critical calcareous algae. – Kongelige Nor- ske Videnskabers Selskabs Skrifter, 5, 1–34. FOSLIE, M. (1909): Algologische notiser. VI. – K. norske Vidensk. Sel- sk. Skr., 2, 1–63. FRAVEGA, P., GIAMMARINO, S., PIAZZA, M., RUSSO, A. & VAN- NUCCI, G. (1987): Signifi cato paleoecologico degli episodi coral- gali a Nord di Sassello. Nuovi dati per una ricostruzione paleogeo- grafi co-evolutiva del margine meridionale del Bacino Terziario del Piemonte. – Atti Soc. Tosc. Sc. Nat. Mem, s. A, 94, 19–76. KOCHANSKI, V. (1944): Fauna marinskog miocena južnog pobočja Medvednice (Zagrebačke gore) (Miozäne marine fauna des südli- chen Abhanges der Medvednica – Zagreber Gebirge). – Vjestnik Hrvatskog drž. geol. zavoda i Hrv. drž. geol. muz., 2–3, 171–280. LUČIĆ, D., SAFTIĆ, B., KRIZMANIĆ, K., PRELOGOVIĆ, E., BRITVIĆ, V., MESIĆ, I. & TADEJ, J. (2001): The Neogene evolution and hydrocarbon potential of the Pannonian Basin in Croatia. – Marine and Petroleum Geology, 18, 133–147. PALLAS, P.S. (1766): Elenchus zoophytorum sistens generum adum- brationes generaliores et specierum cognitarum succinctas descrip- tiones cum selectis auctorum synomymis. – P. van Cleef, Hague, 265, 28+451 p. PILLER, W.E. (1994): Nullipora ramosissima Reuss, 1847 – a redisco- very. – Beitr. Paläont., 19, 181–189. PILLER, W.E., HARZHAUSER, M. & MANDIĆ, O. (2007): Miocene Central Paratethys stratigraphy – current status and future direc- tions. – Stratigraphy, 4, 151–168. QUARANTA, F., VANNUCCI, G. & BASSO, D. (2007): Neogonioli- thon contii comb. nov. based on the taxonomic re-assessment of Mastrorilli’s original collections from the Oligocene of NW Italy (Tertiary Piedmont Basin). – Riv. Ital. Paleont. Strat., 113, 43–55. RANDAZZO, A.F., MÜLLER, P., LELKES, G., JUHÁSZ, E. & HAM- -OR, T. (1999): Cool-water limestones of the Pannonian basinal system, Middle Miocene, Hungary. – J. Sed. Res., 69, 283–293. RIEGL, B. & PILLER, W.E. (2000): Biostromal coral facies – a Miocene example from the Leitha Limestone (Austria) and its actualistic in- terpretation. – Palaios, 15, 399–413. ŠIKIĆ, L. (1967): Torton i sarmat jugozapadnog dijela Medvednice na osnovi faune foraminifera (Torton und Sarmat des südwestlichen Teils der Medvednica auf Grund der Foraminiferenfauna). – Geol. vjesnik, 20, 127–135. ŠIKIĆ, L. (1968): Stratigrafi ja miocena sjeveroistočnog dijela Medved- nice na osnovu faune foraminifera (Über di Miozänstratigraphie des nordöstlichen Teiles des Medvednica Gebirges auf Grund der Fo- raminifferenfaunen). – Geološki vjesnik, 21, 213–227. SIMONE, L. & CARANNANTE, G. (1985): Evolution of a Miocene Carbonate open shelf from interception to drowning: the case of the southern Apennines. – Rend. Acc. Sci. Fis. Mat., IV, LII/2, 1–43. STUDENCKI, W. (1988): Facies and sedimentary environment of the Pinczow limestones (Middle Miocene; Holy Cross Mountains, Cen- tral Poland). – Facies, 18, 1–26. UNGER, F. (1858): Beiträge zur näheren Kenntniss des Leithakalkes, namentlich der vegetabilischen Einschlüsse und der Bildungsge- schichte desselben. – Denkschr. Kais. Akad. Wiss., Math.-nat. Klas- se, 14, 13–35. Geologia Croatica Geologia Croatica 61/2–3 340 VRSALJKO, D., PAVELIĆ, D. & BAJRAKTAREVIĆ, Z. (2005): Stra- tigraphy and palaeogeography of Miocene deposits from the mar- ginal area of Žumberak Mt. and the Samoborsko gorje Mts. (nor- thwestern Croatia). – Geol. Croat., 58/2, 133–150. VRSALJKO, D., PAVELIĆ, D., MIKNIĆ, M., BRKIĆ, M., KOVAČIĆ, M., HEĆIMOVIĆ, I., HAJEK-TADESSE, V., AVANIĆ, R. & KUR- TANJEK, N. (2006): Middle Miocene (Upper Badeninan/Sarmati- an) palaeoecology and evolution of the environments in the area of Medvednica Mt. (North Croatia). – Geol. Croat., 59/1, 51–63. VRSALJKO, D., HEĆIMOVIĆ, I. & AVANIĆ, R. (2007a): Miocene de- posits of Northern Croatia. – In: GRGASOVIĆ, T. & VLAHOVIĆ, I. (eds.): 9th International Symposium on Fossil Algae, Field Trip Guidebook and Abstracts. Croatian Geological Survey, Zagreb, 143– 153. VRSALJKO, D., MARKOVIĆ, S. & GRGASOVIĆ T. (2007b): Zagreb Cathedral of the Assumption of the Blessed Virgin Mary. – In: GRGASOVIĆ, T. & VLAHOVIĆ, I. (eds.): 9th International Sym- posium on Fossil Algae, Field Trip Guidebook and Abstracts. Cro- atian Geological Survey, Zagreb, 155. VRSALJKO, D., MIKNIĆ, M., HAJEK-TADESSE, V., BAKRAČ, K., AVANIĆ, R., GRIZELJ, A. & KOCH, G. (2007c): Miocene deposits in Gornje Vrapče. – In: GRGASOVIĆ, T. & VLAHOVIĆ, I. (eds.): 9th International Symposium on Fossil Algae, Field Trip Guidebook and Abstracts. Croatian Geological Survey, Zagreb, 161–163. VRSALJKO, D., AVANIĆ, R., MIKNIĆ, M., GRIZELJ, A., HAJEK- TADESSE, V. & BAKRAČ, K. (2007d): The Badenian deposits of Bizek quarry. – In: GRGASOVIĆ, T. & VLAHOVIĆ, I. (eds.): 9th International Symposium on Fossil Algae, Field Trip Guidebook and Abstracts. Croatian Geological Survey, Zagreb, 165–167. WOELKERLING, W.J. & IRVINE, L.M. (1986): The typifi cation and status of Phymatolithon (Corallinaceae, Rhodophyta). – British Phy- col. J., 21, 55–80. WOELKERLING, W.J., IRVINE, L.M. & HARVEY, A.S. (1993): Growth -forms in non-geniculate coralline red algae. – Austral. Syst. Bot., 6, 277–293. Manuscript received May 25, 2008 Revised manuscript accepted July 31, 2008