Albanerpetontid amphibians from the Late ...

Canadian Journal of Earth Sciences

Albanerpetontid amphibians from the Late Cretaceous (Santonian) of Iharkút, Hungary, with remarks on regional differences in Late Cretaceous Laurasian amphibian assemblages

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Journal: / Journal :

Canadian Journal of Earth Sciences 2012-0049.R1

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Article / Article

Date Submitted by the Author: / Date de soumission de l'auteur :

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Szentesi, Zoltán; Eötvös University, Department of Paleontology Gardner, James; Royal Tyrrell Museum of Palaeontology, Venczel, Márton; łării Crişurilor Museum,

Keyword:

Albanerpetontidae, Cretaceous, Csehbánya Formation, Hungary, Lissamphibia

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Albanerpetontid amphibians from the Late Cretaceous (Santonian) of Iharkút, Hungary, with remarks on regional differences in Late Cretaceous Laurasian amphibian assemblages

Zoltán Szentesi, James D. Gardner, and Márton Venczel

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Z. Szentesi. Department of Paleontology, Eötvös University, MTA-MTM-ELTE Research Group for Palaeontology, P.O. Box 120, H-1518, Budapest, Hungary. J.D. Gardner.1 Royal Tyrrell Museum of Palaeontology, Box 7500, Drumheller, Alberta, T0J 0Y0, Canada.

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M. Venczel. łării Crişurilor Museum, B-dul Dacia 1-3, RO-410464 Oradea, Romania. 1

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Corresponding author ([email protected]).

Abstract: Since its discovery in 2000, the Iharkút fossil locality in the Upper Cretaceous

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(Santonian) Csehbánya Formation of western Hungary has yielded a taxonomically diverse

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assemblage of terrestrial and freshwater vertebrates that continue to provide insights into the diversity, paleobiogeography, and paleoecology of Late Cretaceous vertebrates in Europe. Albanerpetontidae, which are an extinct group of superficially salamander-like amphibians that were widespread across Laurasia during the latter part of the Mesozoic, are represented at Iharkút by 16 fragmentary jaws. Here we describe and figure these specimens as Albanerpetontidae genus and species indeterminate. Based on the age of the Iharkút locality, several premaxillary features, the known distribution (late Early Cretaceous–late Pliocene) of the

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type genus Albanerpeton, and an unusually large dentary specimen, we suggest that the Iharkút albanerpetontid may pertain to a previously unrecognized species of Albanerpeton, but verification of that must await the recovery of more diagnostically informative specimens, such as frontals and more nearly complete premaxillae. The Iharkút lissamphibian assemblage contains a mixture of taxa with Laurasian (the albanerpetontid and a discoglossid anuran) and Gondwanan (a neobatrachian anuran) affinities. Intriguing higher level differences are evident among Late Cretaceous Laurasian assemblages; for example, urodeles are scarce or absent (as at

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Iharkút) in Europe, whereas albanerpetontids are scarce in Middle Asia.

Introduction

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Albanerpetontidae Fox and Naylor, 1982 are a Middle Jurassic–Pliocene clade of

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salamander-like lissamphibians that are closely related to frogs (anurans), salamanders

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(urodeles), and, perhaps, caecilians (apodans) (e.g., Fox and Naylor 1982; McGowan and Evans 1995; Gardner 2001; Gardner and Böhme 2008). Albanerpetontids are known mostly by isolated and rare articulated bones from Europe, the North American Western Interior, Middle Asia, and

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northern Africa (see Gardner and Böhme 2008, table 2 and references therein) and are a

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characteristic component of non-marine vertebrate assemblages, particularly from the latter part of the Mesozoic in North America and Europe. The European record for the clade extends from the early Bathonian (Seiffert 1969; Evans and Milner 1994; Evans and Waldman 1996) to late Pliocene (Delfino and Sala 2007), encompasses 11 countries (summarized by Gardner and Böhme 2008, table 2), and includes all four of the currently recognized genera: Albanerpeton Estes and Hoffstetter, 1976 (middle or late Campanian–late Pliocene in Europe); Celtedens McGowan and Evans, 1995 (Kimmeridgian–early Albian); Anoualerpeton Gardner, Evans, and

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Sigogneau-Russell, 2003 (late Bathonian in Europe); and an unnamed genus (here informally called the “IoW genus”) from the Barremian of the Isle of Wight, southern England (Sweetman and Gardner in press). Although the European record spans about 165 million years, according to the time scale of Gradstein et al. (2004), it contains several lengthy temporal gaps that hamper attempts to interpret the evolutionary history of the clade. For example, until recently Late Cretaceous occurrences in Europe were limited to the Campanian and Maastrichtian of France, Spain, and Romania (Table 1). This differs from the situation in the North American Western

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Interior, where albanerpetontids are known from every stage of the Late Cretaceous (Estes 1981; Fox and Naylor 1982; Gardner 1999a, 2000a). Geologically older albanerpetontid fossils recently have been reported (e.g., Makádi et al. 2006; Szentesi 2008; İsi et al. in press) from the

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Santonian-age Iharkút locality in Hungary. Here we document and discuss the significance of these new fossils.

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Iharkút fossil locality

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The Iharkút fossil locality (İsi et al. 2003, in press) is situated between the villages of

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Iharkút and Németbánya (Fig. 1A), in the Bakony Mountains of western Hungary, within an

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open pit bauxite mine that exposes rocks of Mesozoic and Cenozoic age (Fig. 1B). The fossiliferous layers are within the Csehbánya Formation, a fluvial and floodplain sedimentary unit (İsi and Mindszenty 2009) that is reliably dated as Santonian based on palynological (Knauer and Siegl-Farkas 1992) and paleomagnetic (Mártonné 20051) studies. Vertebrate fossils 1

Mártonné, S.E. 2005. Paleomágneses vizsgálatok Iharkúton. [Paleomagnetic studies at Iharkút.] Department of Environmental Geology, Eötvös University, Budapest, Hungary. Unpublished manuscript. [In Hungarian.]

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documenting a taxonomically diverse assemblage of freshwater and terrestrial taxa (e.g., Makádi et al. 2006; İsi and Rabi 2006; Kocsis et al. 2009; İsi et al. in press) are found throughout the 50–60 metre thick exposures of the Csehbánya Formation at Iharkút, but mostly are restricted to fossiliferous pockets within about a four metre interval in the lower part of the exposures. The Iharkút vertebrate assemblage includes a mix of taxa with Laurasian and Gondwanan affinities, and contains many of the non-marine taxa that are typical for the Late Cretaceous of Laurasia, including: lepisosteids and pycnodontiform actinopterygians; albanerpetontid and anuran

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lissamphibians; scincomorph and mosasauroid lizards; bothremydid, dortokid, and Kallokibotion-like turtles; eusuchian and mesoeucrocodylian crocodiles; azhdarchid pterosaurs; nodosaurid ankylosaurs; rhabdodontid ornithopods; coronosaurian ceratopsians; abelisaurid,

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maniraptoran, and basal tetanuran non-avian theropods; and enantiornithine birds (e.g., Makádi

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et al. 2006; İsi and Rabi 2006; İsi, Apesteguía, et al. 2010; İsi, Butler, et al. 2010; İsi and

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Buffetaut 2011; Rabi et al. in press; İsi et al. in press). Notably absent from this otherwise typical Late Cretaceous assemblage are chondrichthyians, snakes, mammals, and—as highlighted in the Discussion portion of this paper—urodele amphibians. The same four metre

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interval in the lower part of the unit that contains most of the vertebrate fossils also has yielded an angiosperm-dominated assemblage of seeds and fruits (Bodor 2011; Bodor et al. 2012).

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Within the basal part of this fossiliferous interval are two localities that have produced all of the amphibian (anuran and albanerpetontid) fossils recovered to date (Fig. 1C). Thirteen of the albanerpetontid specimens come from the stratigraphically lower, fine-grained, amber-rich, microvertebrate bonebed at level SZ-7-8, which is interpreted as having been deposited in swamps or pools (İsi et al. in press). The remaining three specimens come from the stratigraphically higher, coarse-grained, macrovertebrate-dominated bonebed layer at level SZ-6,

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which is interpreted as having been deposited under higher energy conditions within a fluvial channel (İsi et al. in press). The two bonebeds are about 300 metres apart horizontally (İsi et al. in press) and appear to be within about one metre of each other stratigraphically, although the latter is not certain because of extensive post-depositional faulting within the walls of the mine.

Materials and methods Specimens reported in this study are fragmentary, three-dimensionally preserved jaws

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that were recovered by screenwashing of fossiliferous matrix from two levels (SZ-7-8 and SZ-6) at the Iharkút locality. Albanerpetontid premaxillae, maxillae, and dentaries are sufficiently

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distinctive (e.g., Estes and Hoffstetter 1976; Estes 1981; Fox and Naylor 1982; McGowan and Evans 1995; McGowan 1996, 2002; McGowan and Ensom 1997; Gardner 1999a–c, 2000a, b,

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2001, 2002; Venczel and Gardner 2005; Sweetman and Gardner in press) that even fragmentary

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specimens can be readily identified. The Iharkút jaws were compared directly with other albanerpetontid specimens and with published descriptions. Collectively we (especially JDG) have seen most of the relevant albanerpetontid collections, including examples of all named

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genera and species. In addition to taxa and specimens listed by Venczel and Gardner (2005, p.

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1274), we also have been able to examine material of the new genus and species from the Barremian of the Isle of Wight (Sweetman and Gardner in press) and unpublished material from the Cenomanian–Campanian of Utah, USA, and the Maastrichtian of north-central Romania. Osteological terms for albanerpetontid jaws follow Venczel and Gardner (2005); see also Fig. 2. Institutional abbreviations for specimens included in this report are: MÁFI, Magyar Állami Földtani Intézet (Geological Institute of Hungary), Budapest, Hungary; MNHN.LGA, Muséum National d’Histoire Naturelle, collection from La Grive-Saint-Alban, Paris, France;

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MTM, Magyar Természettudományi Múzeum (Hungarian Natural History Museum), Budapest, Hungary (note that the correct format for specimen numbers from this collection includes a period at the end of the number; e.g., MTM V2008.22.1.); UALVP, University of Alberta Laboratory for Vertebrate Paleontology, Edmonton, Alberta, Canada; and UCM, University of Colorado Museum, Boulder, Colorado, USA.

Systematic Palaeontology

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Subclass Lissamphibia Haeckel, 1866 Order Allocaudata Fox and Naylor, 1982

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Family Albanerpetontidae Fox and Naylor, 1982

Genus and species indeterminate

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(Figs. 2, 3A)

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SPECIMENS: Premaxillae (n=5): MTM V2008.22.1., V.2008.23.1., V2008.24.1., V2009.1.1.,

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and V2010.288.1.; maxillae (n=2): MTM V2008.28.1. and V2009.29.1.; dentaries (n=8): MTM

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V2008.25.1., V2008.26.1., V2008.29.1., V2009.8.1., V2009.9.1., V2009.26.1., V2009.32.1., and V2010.287.1.; and indeterminate jaw fragment (n=1): MTM V2009.10.1.

LOCALITY, UNIT, AND AGE: Iharkút fossil locality, western Hungary; Csehbánya Formation; Late Cretaceous (Santonian). Three specimens (MTM V2008.25.1., V2008.28.1., and V2009.32.1.) are from the stratigraphically higher, coarse-grained, macrovertebrate-dominated

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bonebed layer at level SZ-6, whereas the remaining specimens are from the stratigraphically lower, fine-grained, and amber-rich microvertebrate bonebed at level SZ-7-8.

Description General remarks None of the specimens is complete. Bone surfaces are well preserved and features are clearly visible in some specimens, but in other specimens broken edges, extensive cracking, and

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patches of matrix obscure many details. Due to the fragmentary nature of the specimens, the descriptions below are brief and focus on those features that are taxonomically informative; more

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detailed descriptions of albanerpetontid jaws can be found in papers cited in the “Materials and methods” section.

Teeth

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and V2008.29.1; Fig. 2J and K, respectively), and one jaw fragment (MTM V2009.10.1.; Fig

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2M), also preserve one or more intact or nearly intact teeth. In the latter specimens, the teeth are diagnostic for albanerpetontids in the following combination of features: highly pleurodont attachment (i.e., attached along at least half of their length to the lingual surface of the pars dentalis on upper jaws or of the dental parapet on lower jaws); closely spaced; non-pedicellate; shafts elongate and slightly expanded lingually; and crowns labiolingually compressed, chisel like, and, occasionally, have a lighter-colored, apical cap. In some specimens the tooth crowns are faintly tricuspid, consisting of a central cuspule that is flanked mesially and distally by a

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slightly smaller cuspule, whereas in others the three cuspules are less distinct and the apex has a more arcuate lingual or labial profile (Fig. 2J versus M). Even for specimens lacking any reasonably complete teeth, it is still evident from the broken tooth bases or empty tooth slots that the teeth were highly pleurodont, narrow, and closely spaced.

Premaxillae The three most nearly complete specimens (MTM V2008.23.1., V.2008.22.1., and V.

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2008.24.1.; Fig. 2A–C) preserve portions of the pars dorsalis and pars dentalis, but most of the shelf-like pars palatinum is broken away and only a broken face remains on the lingual surface. These specimens are characteristic for albanerpetontid premaxillae in the following non-dental

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features: pars dorsalis is relatively broad, tall, and projects dorsally; pars dentalis is deep; labial

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surface of bone is perforated by small, scattered nutritive foramina and ventrolaterally there is a

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smooth, cup-shaped facet for receipt of a complementary process from the maxilla; medial surface of bone bears a parallel series of alternating ridges and grooves for sutured contact across the midline with the opposite premaxilla; lingual surface of pars dorsalis has a prominent

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suprapalatal pit; and broken lingual face of pars palatinum exposes a canal extending through the shelf, between the ventral and dorsal openings of the palatal foramen. None of the premaxillary

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specimens has a dorsally complete pars dorsalis, which means that several features (e.g., proportions of the pars dorsalis; presence and relative size of dorsal boss on labial surface; ornament pattern on upper part of labial surface; pattern of contact with the nasal) useful for differentiating albanerpetontid species cannot be determined. The suprapalatal pit, which is best exposed in MTM V2008.22.1., is relatively moderate in size, elliptical in outline, faces lingually, and is located approximately midway across the pars dorsalis and slightly above the broken face

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of the pars palatinum. MTM V2008.22.1. and MTM V2008.23.1. both show that the canal connecting the dorsal and ventral openings of the palatal foramen extends vertically through the pars palatinum. Compared to the other premaxillae reported here, MTM V2008.24.1. has a more robust build and is from a larger individual, probably at least twice the size as that represented by MTM V2008.23.1. Only one of the five specimens (MTM V2009.1.1.; Fig. 2D) preserves a nearly intact tooth.

Maxillae

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The two maxillary specimens (MTM V2008.28.1. and V2009.29.1.; Fig. 2E and F, respectively) are broken both anteriorly and posteriorly, lack any intact teeth, and were from

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relatively small individuals. Both are characteristic for albanerpetontid maxillae in the following

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non-dental features: bone elongate; pars dorsalis low and becoming shallower posteriorly; and

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dorsal surface of pars dorsalis shallowly flattened for contact with lacrimal and jugal. The structure of the anteriorly incomplete pars palatinum (i.e., a lingually expanded shelf, indented by a concave internal narial margin, narrows posteriorly, and bears shallow grooves and ridges

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on its dorsal surface) on MTM V2008.28.1. and the triangular nasal process on MTM V2009.29.1. also are characteristic for albanerpetontids. As in most albanerpetontids, the ventral

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margin of the pars dentalis is relatively straight (versus ventrally convex in Anoualerpeton [Gardner et al 2003, figs. 1C] and Albanerpeton nexuosum [Gardner 2000a, fig. 3A]) and the labial surface is unornamented (versus ornamented with irregular ridges and pustules in large maxillae of Al. inexpectatum [Gardner 1999b, pl. 1J]). An unusual feature of MTM V2009.29.1. is the concave depression in the labial surface below the nasal process.

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Dentaries The eight specimens collectively document most of the tooth-bearing region (i.e., about the anterior two-thirds) of the dentary. Three specimens (MTM V2009.9.1., Fig. 2G; MTM V2008.26.1., Fig. 2H; and MTM V2009.26.1., not figured) preserve the anterior end of the bone, including the symphyseal prongs that are autapomorphic for albanerpetontids. Another three specimens (MTM V2009.8.1., V2010.287.1., and V2008.29.1.; Fig. 2I–K, respectively) preserve the median portion of the ramus, between the symphyseal region and the opening for the

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Meckelian canal. The two remaining specimens, MTM V2008.25.1. (Fig. 2L) and MTM V2009.32.1. (not figured), preserve the posterior portion of the tooth bearing-region and the opening for the Meckelian canal. Other characteristic, non-dental albanerpetontid features of

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these dentary specimens are: labial surface bears a horizontal row of external nutritive foramina

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and, more ventrally, a low ridge probably for attachment of intermandibularis muscles; dental

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parapet is relatively high; subdental shelf is shallow, moderately broad, and gutter-like anteriorly, becoming deeper and narrower posteriorly; and opening for Meckelian canal lies below the posterior portion of the tooth row. As in most albanerpetontids, the dorsal edge of the

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dental parapet along about the anterior half of the dentary is shallowly convex dorsally in labial or lingual profile (versus more sinuous in Anoualerpeton [Gardner et al. 2003, fig. 2D] and

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Albanerpeton nexuosum [Estes 1964, fig. 43e; Gardner 2000a, fig. 2A–E]), the labial surface is unornamented (versus ornamented with irregular ridges in large dentaries of Al. inexpectatum [Gardner 1999b, pl. 2D]), and the dorsal edge immediately behind the tooth row lacks a low, dorsally-projecting flange (versus flange present in Al. inexpectatum [Gardner 1999b, pl. 2A, E]). Most of the Iharkút dentary specimens are small (i.e., less than 2 mm in preserved length), and fall within the size range typical of other albanerpetontids. However, MTM V2008.29.1. is

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6.4 mm in preserved length and was from a large individual. When depths of the subdental shelf are compared, MTM V2008.29.1. is over twice the size of the next largest, comparable Iharkút specimen (MTM V2010.287.1.; Fig. 2J) and about 20 percent larger than the largest dentaries known for larger-bodied albanerpetontid species (Fig. 3).

Jaw fragment MTM V2009.10.1. (Fig. 2M) is a jaw fragment consisting of five teeth in sequence and

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the adjacent portion of bone to which those teeth are attached. Too little of the bone is preserved to determine which element and side the specimen is from. Nevertheless, the teeth are diagnostic for albanerpetontids in their structure, arrangement, and attachment. The teeth are comparable in

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size to those on the largest dentary (MTM V2008.29.1.; Fig. 2K).

Discussion

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vi Taxonomic identification of the Iharkút specimens

Although fragmentary, the 16 jaw specimens reported here can be assigned to

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Albanerpetontidae based on diagnostic features of the bones and teeth. Two albanerpetontid

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autapomorphies (Gardner 2001) are seen in seven of the specimens: symphyseal prongs are present on the three specimens that preserve the anterior end of the dentary and non-pedicellate teeth with labiolingually compressed crowns bearing three mesio-distally aligned cuspules are preserved in two other dentaries, one premaxilla, and the indeterminate jaw fragment. As detailed in the above descriptive accounts, numerous other features also are diagnostic for albanerpetontids, including the following: teeth are highly pleurodont, elongate, and closely spaced; premaxilla has relatively tall and broad pars dorsalis, deep pars dentalis, suprapalatal pit

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in lingual surface of pars dorsalis, parallel ridges and grooves along medial surface of bone, and nutritive foramina scattered across labial surface of bone; maxilla has deep pars dentalis and low pars dorsalis that becomes shallower posteriorly, triangular nasal process, and pars palatinum indented for internal narial margin; and dentary has high dental parapet, subdental shelf low and gutter like anteriorly and becoming deeper and narrower posteriorly, opening for Meckelian canal located posteriorly, and bears row of external nutritive foramina labially and ridge ventrolabially for muscle attachment.

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Generic assignment of the Iharkút specimens is problematic, for two reasons. First, no frontals are known from the locality, which is unfortunate because frontals are widely considered the most diagnostic bone for albanerpetontid genera (e.g., McGowan and Evans 1995; McGowan

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1998; Gardner 2000b, 2002; Gardner et al. 2003; Skutschas 2007; Sweetman and Gardner in

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press). Second, although jaws can be useful for differentiating albanerpetontid genera, the

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specimens from Iharkút do not preserve any feature or combination of features that can be used to assign them unambiguously to a particular genus. The following five jaw features are relevant in this context:

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(1) Although neither of the maxillary specimens and none of the eight dentary specimens from Iharkút preserve a complete occlusal margin, the labial or lingual profile of that margin

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appears to have been nearly straight to shallowly convex. If correct, that is similar to the condition in Celtedens, the IoW genus, and most species of Albanerpeton, but it differs from the more strongly convex profile seen in maxillae and dentaries of Anoualerpeton and Al. nexuosum. (2) In the two premaxillae (MTM V2008.22.1. and V2008.23.1.) for which the orientation of the suprapalatal pit can be determined, this opening faces lingually as it also does

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in Celtedens, the IoW genus, and Albanerpeton. By contrast, the suprapalatal pit faces more laterally in Anoualerpeton. (3) The same two Iharkút premaxillae also show that the suprapalatal pit lies at or just slightly above the pars palatinum. In Anoualerpeton, Celtedens, and one species of Albanerpeton (the basal Al. arthridion) the suprapalatal pit lies well dorsal to the pars palatinum, whereas in the remaining, more derived and geologically younger (all Late Cretaceous and Cenozoic) species of Albanerpeton the suprapalatal pit is located lower, as in the two Iharkút specimens.

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Both conditions occur in premaxillae of the monospecific IoW genus. (4) Based on the one Iharkút premaxilla (MTM V2008.22.1.) in which the suprapalatal pit is entirely free of matrix and comparisons with better preserved premaxillae from other

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albanerpetontid species, the suprapalatal pit appears to be moderate in size relative to the pars

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dorsalis. This admittedly is not certain, because the dorsal part of the pars dorsalis is missing

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from the Iharkút specimen. If correct, however, the moderate-sized suprapalatal pit inferred for MTM V2008.22.1. differs from the relatively smaller pit in Anoualerpeton, Celtedens, and Albanerpeton arthridion and from the substantially larger suprapalatal pit in Al. galaktion. The

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remaining species of Albanerpeton have a moderate-sized suprapalatal pit. As with the previous feature, the relative size of the suprapalatal pit also varies (from small to moderate) in the

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monospecific IoW genus.

(5) In the two premaxillae (MTM V2008.22.1. and V2008.23.1.) in which the canal connecting the dorsal and ventral openings of the palatal foramen is exposed in the broken face of the pars palatinum, the canal extends vertically. The orientation of this canal is unknown for Celtedens, but in the remaining genera the canal extends vertically in Albanerpeton, diagonally

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(dorsolaterally–ventromedially) in Anoualerpeton, and both conditions occur in the monospecific IoW genus. To summarize, the Iharkút jaws can be excluded from both Anoualerpeton (all five features) and Celtedens (features 3 and 4). However, none of the five jaw features is decisive for differentiating between Albanerpeton and the IoW genus. The Iharkút condition for each feature occurs in both genera, as follows: feature 2 occurs consistently in both genera; feature 1 occurs in most species of Albanerpeton and in the monospecific IoW genus; features 3 and 4 occur in

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most species of Albanerpeton and variably in the monospecific IoW genus; and feature 5 occurs in all species of Albanerpeton and variably in the monospecific IoWgenus. Although there is no morphological basis for assigning the Iharkút specimens to either

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the IoW genus or Albanerpeton, the latter assignment is more consistent with known

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stratigraphic and geographic occurrences for the two genera. The IoW genus is limited to the

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Early Cretaceous (Barremian) of southernmost England and, possibly, Spain (Sweetman and Gardner in press). Albanerpeton is the albanerpetontid genus currently recognized from the postBarremian (late Aptian–late Pliocene), and during the Late Cretaceous in Eurasia it has been

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reported from slightly younger (Campanian and Maastrichtian) localities in France (Duffaud 2000) and Romania (e.g., Grigorescu et al. 1999; Duffaud 2000; Folie and Codrea 2005) and also

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from an older (Cenomanian) locality in Uzbekistan (Skutschas 2007). Outside of Eurasia, Albanerpeton is known from the late Aptian/early Albian–late Paleocene of the North American Western Interior (e.g., Gardner and Böhme 2008, table 2). More diagnostically informative specimens—ideally frontals or premaxillae exhibiting features (such as medial fusion) that are synapomorphic for more derived species of Albanerpeton—will need to be recovered before the generic identity of the Iharkút albanerpetontid can be resolved.

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The Iharkút specimens are too fragmentary and non-distinct to be assigned to a recognized species. Based on the lack of convincing taxonomic differences among the 16 specimens, we conservatively assume these pertain to one species. Although a range of different sized premaxillae and dentaries is available, neither element shows any obvious ontogenetic variation like, for example, the development of labial ornament reported in larger dentaries of Albanerpeton inexpectatum (Gardner 1999b). Considering that the only other Santonian albanerpetontids (Albanerpeton galaktion and Al. nexuosum from Alberta, Canada [Gardner

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2000a] and undescribed specimens from Utah, USA [Gardner and Eaton, unpublished observation 2010]) are geographically far removed from Iharkút and that the temporally closest European albanerpetontid species, Al. inexpectatum (latest Oligocene–late Miocene; see Gardner

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and Böhme 2008, table 2; Böhme 2008), is about 60 million years younger (timescale of

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Gradstein et al. 2004), it is likely that the Iharkút albanerpetontid is a previously unrecognized

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species. The Iharkút material could be conspecific with albanerpetontid material known from Campanian and Maastrichtian localities elsewhere in Europe (see Table 1), but that possibility will be difficult to evaluate given the equally fragmentary nature of many specimens from those

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younger localities. In the context of discussing the potential specific affinities of the Iharkút albanerpetontid, two of the specimens each exhibit a feature that is worth noting. The depression

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in the labial surface of the maxilla MTM V2009.29.1. is an unusual feature that we have not seen in any other albanerpetontid maxilla. Because albanerpetontid maxillae tend to be fairly conservative in their structure, we interpret that depression as an anomalous feature of no taxonomic significance. The notably large size of the dentary MTM V2008.29.1. is more intriguing, because based on the depth of the subdental shelf that specimen is about 20 percent larger than the next largest albanerpetontid dentaries currently known (see Fig. 3). But due to the

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limited sample currently available from Iharkút, there is no way to judge whether MTM V2008.29.1. is typical of the size attained by the Iharkút albanerpetontid or is simply an aberrantly large specimen.

Late Cretaceous European and other Laurasian lissamphibian assemblages Compared to contemporaneous deposits in the North American Western Interior and Middle Asia, the Late Cretaceous record of non-marine vertebrates in Europe is sparse (see

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reviews by Le Loeuff 1991; Pereda-Suberbiola 2009). This is especially true for the lissamphibian record (Table 1, Fig. 4), particularly in the following aspects: occurrences are known only from France, Spain, Hungary, and Romania; the record is temporally restricted to

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the Santonian–Maastrichtian, except for a single early Cenomanian occurrence; many localities

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have yielded small numbers of specimens (i.e., several dozen or fewer); and specimens consist of

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isolated bones. This limited fossil record likely is the result of high sea levels during the Late Cretaceous (e.g., Golonka and Kiessling 2002), which fragmented the European continent into an archipelago of various-sized islands (e.g., Dercourt et al. 2000; Smith et al. 2001; Pereda-

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Suberbiola 2009). For the localities listed in Table 1, paleogeographic reconstructions (see Pereda-Suberbiola 2009, p. 61 and fig. 2) place localities in France and Spain on the Ibero-

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Armorican island, the Romanian localities on the Rhodope-Moesian platform, and the Iharkút locality on the Austroalpine island. Even though the Late Cretaceous record of European lissamphibians is limited, some interesting observations can be made. Based on specimens collected and identified to date, the Iharkút assemblage contains a low diversity of three lissamphibian taxa: the indeterminate albanerpetontid reported here; a discoglossid (sensu lato) anuran, Bakonybatrachus fedori Szentesi and Venczel, 2012; and a

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neobatrachian anuran Hungarobatrachus szukacsi Szentesi and Venczel, 2010. The cooccurrence of these taxa at Iharkút is consistent with the interpretation (İsi et al. in press) that the two fossiliferous layers were deposited in fluvial and marshy conditions, which are among the most favoured habitats for lissamphibians (e.g., Duellman and Trueb 1986; Wells 2007; Vences and Köhler 2008). Considering that the Iharkút layers sample prime lissamphibian habitats and that the known diversity is based on only about 30 bones (16 albanerpetontid, 4 Bakonybatrachus, and 12 Hungarobatrachus), we predict that additional lissamphibian

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specimens and taxa will be identified as matrix continues to be processed. The Iharkút vertebrate assemblage is biogeographically interesting, because it contains a mixture of taxa with Laurasian and Gondwanan affinities (see summary by İsi et al in press).

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The lissamphibians conform to this pattern (Szentesi and Venczel 2010, 2012): albanerpetontids

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and discoglossids are primarily Laurasian groups, whereas neobatrachians—although now

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having a near cosmopolitan distribution—are considered to have originated in Gondwana (e.g., Savage 1973; Duellman and Trueb 1986; Feller and Hedges 1998; Roelants and Bossuyt 2005; Bossuyt et al. 2006; Gardner and Böhme 2008). The presence of albanerpetontids and

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discoglossids at Iharkút is not unexpected, because both groups are known in Europe from the Middle Jurassic to the late Pliocene (albanerpetontids) or to the present (discoglossids) and

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during that interval their remains are commonly encountered in fossiliferous, non-marine deposits like those at Iharkút (e.g., Evans and Milner 1994; Roček 2000; Gardner and Böhme 2008, table 2). Hungarobatrachus is the oldest (Santonian) record of neobatrachians in Laurasia, and its presence at Iharkút may have been the result of island hopping or rafting across the Tethys from Africa (Szentesi and Venczel 2010). Interestingly, there is a gap of about 25 million

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years (according to the timescale of Gradstein et al. 2004) in the Laurasian neobatrachian record, before they re-appear in the late Paleocene of Cernay, France (Estes et al. 1967; Rage 2003). Although all three Laurasian Late Cretaceous lissamphibian groups are known from Europe (Table 1), the European localities are dominated by albanerpetontids and anurans, which are present at 10 and 15 localities, respectively. Urodeles are conspicuously uncommon, being present at just one locality in Spain and two or, perhaps, three localities in France. The scarcity or absence of identifiable urodele specimens at the European localities is curious. During the

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Early Cretaceous all three lissamphibian groups were broadly distributed across Laurasia, including in Western Europe (e.g., Estes 1981; Estes and Sanchíz 1982; Evans and Milner 1996; Milner 2000; Roček 2000; Evans and McGowan 2002; Sweetman and Evans 2011). As sea levels

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rose at the end of the epoch, presumably members of all three groups would have been marooned

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on islands of the European archipelago, yet by the latter part of the Late Cretaceous all three

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groups were not equally represented on the paleo-islands. Most of the localities listed in Table 1 contain a typical Late Cretaceous mixture of terrestrial and freshwater, small-bodied tetrapods, including other lissamphibians (albanerpetontids and anurans), which argues against a

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depositional or taphonomic explanation for urodeles not being preserved. Many of the Spanish and French localities that lack urodele fossils admittedly have produced only small numbers of

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other lissamphibian fossils, so at those localities the absence of urodele fossils could be an artefact of small sample size. However, that explanation is less compelling for the Santonian Iharkút locality in Hungary and the Romanian Maastrichtian localities, because those have produced moderate numbers of other lissamphibian specimens—about 30 from Iharkút and several times that amount collectively from the Romanian localities. Although it would be difficult to establish from the fossil and geological record, compositions of lissamphibian

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communities across the European archipelago conceivably could have been influenced by regional differences in weather patterns, disease, competition, predation, or other factors. If urodeles were present during the Late Cretaceous on the Austroalpine island (Iharkút locality) and the Rhodope-Moesian platform (Romanian localities), they evidently were even less common than on the larger Ibero-Armorican island (French and Spanish localities) or were living in areas outside of the depositional basins. The geologically oldest records of urodeles in presentday Hungary and Romania are from the middle Miocene (e.g., Venczel 2008; Venczel and Ştiucă 2008).

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Outside of Europe, other significant Laurasian occurrences of Late Cretaceous lissamphibians are known from the following four regions:

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(1) Southern Mongolia. This record is limited to the Campanian–Maastrichtian (possibly

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also Santonian) of the Gobi Desert and consists exclusively of anurans (e.g., Borsuk-Białynicka

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1978; Špinar and Tatarinov 1986; Gubin 1993, 1999; Shishkin 2000). (2) Middle Asia. Centered in the Kyzylkum Desert of Uzbekistan, but also including the neighbouring countries of Kazakhstan and Tadzikistan, this record extends from the

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Cenomanian–Campanian and consists of anurans, urodeles, and albanerpetontids (e.g., Nessov, 1981, 1988; Roček and Nessov 1993; Shishkin 2000; Skutschas 2007, 2009; Roček 2008).

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Although anuran and urodele fossils are locally abundant in many of the localities, albanerpetontids are rare, being known by only six jaws and two frontals (Gardner and Averianov 1998; Skutschas 2007). (3) Eastern North America. The only described lissamphibian assemblage from the North American eastern subcontinent is from the Campanian age Ellisdale locality, in New Jersey,

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USA, which has produced abundant urodele and lesser numbers of anuran bones (Denton and O’Neill 1998). (4) North American Western Interior. Dozens of localities that collectively span the Cenomanian–terminal Maastrichtian extend from southern Alberta and Saskatchewan, Canada, south through the USA, and into northern Mexico. Urodeles and anurans are relatively abundant at most localities throughout the epoch; albanerpetontids are similarly abundant, although they become scarce in the late Maastrichtian (e.g., Estes 1964; Sahni 1972; Armstrong-Zeigler 1980;

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Carpenter 1979; Rowe et al. 1992; Gardner 1999a, 2000a, 2005; DeMar and Breithaupt 2008; Gardner and Böhme 2008, table 2; Aguillon Martinez 2010; Gardner et al. in press). To summarize the above, there are intriguing regional differences among Laurasian Late

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Cretaceous lissamphibian assemblages: anurans always seem abundant; urodeles are abundant in

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North American and Middle Asia, but not in Europe; and albanerpetontids are abundant in the

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North America Western Interior until the late Maastrichtian and from the Santonian to Maastrichtian in Europe, yet they are rare in Middle Asia and unknown from eastern North America or Mongolia. The semi-arid, depositional environments of the Mongolian localities

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during the Late Cretaceous (e.g., Jerzykiewicz 1998) likely are a significant factor in the absence of non-anuran lissamphibians from those localities. The remaining three regions had broadly

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similar depositional environments during the Late Cretaceous (i.e., floodplains with abundant rivers and marshes), which suggests that differences in the taxonomic compositions of their lissamphibian assemblages may be attributable to other factors, such as the unique paleobiogeographic histories of taxa, variation (both temporal and areal) in local and regional conditions, and differences in the number of localities, sample sizes, and how samples were collected.

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Conclusions •

Sixteen fragmentary jaws from the Santonian age Iharkút vertebrate fossil locality (Csehbánya Formation), western Hungary, are diagnostic for Albanerpetontidae, but cannot decisively be assigned to genus or species. Details of the suprapalatal pit on the best preserved premaxillary specimens, coupled with the known geographic and temporal range of the type genus Albanerpeton, suggests that the Iharkút albanerpetontid likely

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pertains to that genus. However, more informative specimens, such as frontals, which are diagnostic for albanerpetontid genera, or premaxillae that have synapomorphies for less

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inclusive clades within Albanerpeton, need to be recovered to establish the generic identity of the Iharkút albanerpetontid. One dentary specimen is unusually large for an

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albanerpetontid. Large size potentially could be diagnostic for the Iharkút species, but

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additional specimens are needed to assess the reliability of that feature. •

As currently known, the Iharkút assemblage contains a low diversity of lissamphibians:

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one albanerpetontid and two anurans. As is typical for other components of the Iharkút vertebrate assemblage, the lissamphibians are a mix of taxa with Laurasian (the

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albanerpetontid and a discoglossid anuran) and Gondwanan (a neobatrachian anuran) affinities. •

The absence of urodeles from Iharkút and many of the European Late Cretaceous lissamphibian assemblages is notable. Intriguing differences in the higher level taxonomic composition of lissamphibian assemblages also are apparent across Laurasia: anurans always seem abundant; urodeles are abundant in North American and Middle Asia, but are rare in Europe; and albanerpetontids are abundant in the North America

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Western Interior until the late Maastrichtian and from the Santonian to Maastrichtian in Europe, but are rare in Middle Asia and are unknown from eastern North America or Mongolia. The reasons for these regional differences in Late Cretaceous lissamphibian assemblages are unknown.

Acknowledgements Atilla İsi (MTM and Hungarian Academy of Sciences, Budapest, Hungary) directed

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collecting operations at the Iharkút locality and permitted description of the albanerpetontid specimens from that locality. Field work at Iharkút is supported and facilitated by the Hungarian

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Natural History Museum, The Hungarian Scientific Research Fund (OTKA PD 75021 and NF 84193), Bakonyian Bauxite Mine Ltd., and Geovolan Ltd. Former Iharkút mine geologist Andrea

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Mindszenty provided information on the stratigraphy of the amphibian-bearing layers and Ágnes

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Görög (Department of Paleontology, Eötvös University, Budapest), Károly Bóka (Department of Plant Anatomy, Eötvös University, Budapest), and Péter Gulyás (Hungarian Dinosaur Research Group, Budapest) provided technical support. Access to comparative specimens depicted in

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Figure 4 was facilitated by: László Kordos (MÁFI); Daniel Goujet and Jean-Claude Rage

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(MNHN); Michael Caldwell, Richard Fox, and Mark Wilson (UALVP); Toni Culver and Jaelyn Eberle (UCM). George Braybrook (University of Alberta, Edmonton, Canada) took the scanning electron micrographs in Figures 2 and 3. Vlad Codrea (Universitatea Babeş-Bolyai, Cluj-Napoca, Romania), Julio Company (Universidad Politécnica de Valencia, Spain), Jean-Claude Rage (MNHN), and Zbyněk Roček (Academy of Sciences and Charles University, Prague, Czech Republic) shared information on Late Cretaceous European amphibian occurrences. Financial support for this project was provided by grants from the Jurassic Foundation, the Hantken

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Foundation, and the Támop 4.2.2./B-10/1-20110-0030. Venczel received funding from the CNCS (grant No. PN-II-ID-PCE-2011-3-0381) and costs for his research trip to Canada in 2007 were defrayed by the Royal Tyrrell Museum Cooperating Society. Jean-Claude Rage (MNHN) and Pavel Skutschas (St. Petersburg State University, Russia) provided constructive reviews, and Hans-Dieter Sues (National Museum of Natural History, Washington, D.C) served as the editor. This is MTA-MTM-ELTE Paleo contribution No. 162.

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Figure captions Fig. 1. Location and geological context of the Iharkút fossil locality. (A) Map showing geographic location of the Iharkút locality in western Hungary. (B) Generalized stratigraphic column showing rock units exposed at the Iharkút locality. (C) Detailed stratigraphic column showing positions of the two amphibian-bearing layers within exposures of the Upper Cretaceous (Santonian) Csehbánya Formation at the Iharkút locality. Adapted from Szentesi and Venczel (2010, fig. 1).

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Fig. 2. Incomplete jaws of Albanerpetontidae genus and species indeterminate from the Upper

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Cretaceous (Santonian) Csehbánya Formation, Iharkút, western Hungary. (A–D) Premaxillae: (A) MTM V2008.23.1., left premaxilla, in lingual and slightly ventromedial view; (B) MTM

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V2008.22.1., left premaxilla, in lingual (B1) and labial (B2) views; (C), MTM V2008.24.1., right

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premaxilla, in lingual (C1) and mediolingual and slightly ventral (C2) views; (D) MTM V2009.1.1., right premaxilla, lingual views of entire specimen (D1) and close up of nearly intact tooth (D2). (E, F) Maxillae: (E) MTM V2008.28.1., left maxilla, in labial and slightly dorsal (E1),

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dorsal (E2), and ventral (E3) views; (F) MTM V2009.29.1., right maxilla, in labial (F1) and

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lingual (F2) views. (G–L) Dentaries: (G) MTM V2009.9.1., right dentary, in lingual view; (H) MTM V2008.26.1., left dentary, in ventrolabial view; (I) MTM V2009.8.1., right dentary, in lingual (I1) and dorsal and slightly lingual (I2) views; (J), MTM V2010.287.1., left dentary, entire specimen in lingual (J1) and dorsal (J2) views, both with arrows pointing at teeth with intact crowns, and close up of same teeth in occlusal view (J3); (K) MTM V2008.29.1., right dentary, in lingual (K1) and labial (K2) views; (L), MTM V2008.25.1., left dentary, in lingual (L1) and labial (L2) views. (M) Jaw fragment (element and side uncertain), MTM V2009.10.1., oriented

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with tooth crowns pointing to top of page, entire specimen (M1) and close up of three tooth crowns showing lighter apical cap (M2). Most of the figured specimens are from the stratigraphically lower, fine-grained, and amber-rich microvertebrate bonebed at level SZ-7-8; however, two specimens (maxilla MTM V 2008.28.1 and dentary MTM V 2008.25.1) are from the stratigraphically higher, coarse-grained, macrovertebrate-dominated bonebed layer at level SZ-6. Images are at different magnifications.

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Fig. 3. Albanerpetontid dentaries, all in lingual view and at same scale, to show size difference between largest dentary from Iharkút and representative dentaries from four species of Albanerpeton. (A) Albanerpetontidae genus and species indeterminate, MTM V2008.29.1.,

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median portion of right dentary, from the Upper Cretaceous (Santonian) Csehbánya Formation,

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Iharkút, Hungary. (B) A. pannonicum (Venczel and Gardner, 2005), MÁFI V.11.133.1, left

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dentary missing most of area for attachment of post-dentary bones, from lower Pliocene fissure infill, Csarnóta 2 locality, Hungary. (C) A. nexuosum (Estes, 1981), UCM 38714, left dentary missing symphyseal end and posterior half of bone, from the Upper Cretaceous (upper

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Maastrichtian) Laramie Formation, UCM locality 77062, Colorado, USA. (D) A. inexpectatum Estes and Hoffstetter, 1976, MNHN.LGA 1248, nearly complete left dentary, from an

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unrecorded lower or middle Miocene fissure infill, La Grive-Saint-Alban, France. Lines above and below this specimen demarcate approximate portion preserved in the figured dentary (A) from Iharkút. (E) A. galaktion Fox and Naylor, 1982, UALVP 16217, right dentary missing posteriormost end of tooth row and area for attachment of postdentary bones, from the Upper Cretaceous (upper Santonian) Deadhorse Coulee Member, Milk River Formation, UAVLP locality MR-9B, Alberta, Canada. Specimens at same magnification.

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Fig. 4. Map showing locations of Upper Cretaceous lissamphibian localities in Europe (see also Table 1).

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Table 1. Late Cretaceous occurrences of albanerpetontids, anurans, and urodeles in Europe (see also Fig. 4). The Spanish locality of Molí Vell is problematic: Roček (2000, p. 1324) mentioned undescribed anuran bones from the locality in the collections of the Institut de Paleontología Miquel Crusafont (Sabadell, Spain), but according to Julio Company (personal communication, 2011 to Z.Sz.) this locality is unknown to him and other Spanish paleontologists currently working in the Upper Cretaceous of Spain. Not

Fo

listed is the upper Campanian–lower Maastrichtian Lo Hueco locality, in eastern Spain, from which unidentified lissamphibian

rR

postcranial bones have been reported (Ortega et al. 2009).

ev

Geological Age

Locality

Albanerpetontidae

Anura

Maastrichtian

Oarda de Jos,

X

X

(precise age

central Romania

uncertain) Maastrichtian

Molí Vell,

(precise age

northeastern Spain

iew X

Urodela

References Codrea et al. (2010)

On

Roček (2000)

ly

uncertain) late Maastrichtian

Blasi 2,

X

X

Blain et al. (2010)

X

X

Roček (2000); Company et al. (2009a)

northeastern Spain late Maastrichtian

La Solana (= “Carlet” in Roček 2000), eastern Spain

-1-


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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48


late Maastrichtian

Cassagnau,

X

Laurent et al. (2002)

southern France late Maastrichtian

Peyrecave,

X

?

Gheerbrant et al. (1997)

southern France early Maastrichtian

HaŃeg Basin

X

Fo

X

Grigorescu et al. (1999); Duffaud (2000); Folie

localities,

southwestern Romania

early Maastrichtian

et al. (2002); Smith et al. (2002); Codrea et al.

rR

Fontllonga,

(2002); Venczel and Csiki (2003); Folie and Codrea (2005); Csiki et al. (2008); Grigorescu

ev

X

X

X

northeastern Spain early Maastrichtian

Cruzy (=

(2010); Panaiotu and Panaiotu (2010); Bojar et

iew

“Massecaps” in Buffetaut 2005),

al. (2011) López-Martínez et al. (1999); Roček (2000)

On

Buffetaut et al. (1999); Buffetaut (2005)

ly

southern France late Campanian or

Laño,

early Maastrichtian

northern Spain

late Campanian or

Chera,

early Maastrichtian

eastern Spain

middle–late

La Neuve,

Campanian

southeastern France

-2-

X

X

X

X

X

X

X

Astibia et al. (1990); Duffaud and Rage (1999)

Company et al. (2009b)

X

Duffaud (2000); Garcia et al. (2000)



1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48

early Campanian

Villeveyrac,

X

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Buffetaut et al. (1996)

southern France Campanian

Champ-Garimond,

(precise age

southeastern France

uncertain) Santonian

Fo

Iharkút,

western Hungary

early Cenomanian

X

Font-de-Benon,

X

rR

X

Duffaud (1995, 2000); Sigé et al. (1997)

Szentesi (2006, 2008); Makádi et al. (2006); Szentesi and Venczel (2010, 2012); İsi et al. (in press)

ev

southwestern France

X

X

iew

Vullo and Néraudeau (2008); Vullo et al. (2011)

On

ly

-3-


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r Fo Re

Fig. 1. Location and geological context of the Iharkút fossil locality. (A) Map showing geographic location of the Iharkút locality in western Hungary. (B) Generalized stratigraphic column showing rock units exposed at the Iharkút locality. (C) Detailed stratigraphic column showing positions of the two amphibian-bearing layers within exposures of the Upper Cretaceous (Santonian) Csehbánya Formation at the Iharkút locality. Modified from Makádi et al. (2006, figs. 1, 2). 99x54mm (300 x 300 DPI)

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Fig. 2. Incomplete jaws of Albanerpetontidae genus and species indeterminate from the Upper Cretaceous (Santonian) Csehbánya Formation, Iharkút, western Hungary. (A–D) Premaxillae: (A) MTM V2008.23.1., left premaxilla, in lingual and slightly ventromedial view; (B) MTM V2008.22.1., left premaxilla, in lingual (B1) and labial (B2) views; (C), MTM V2008.24.1., right premaxilla, in lingual (C1) and mediolingual and slightly ventral (C2) views; (D) MTM V2009.1.1., right premaxilla, lingual views of entire specimen (D1) and close up of nearly intact tooth (D2). (E, F) Maxillae: (E) MTM V2008.28.1., left maxilla, in labial and slightly dorsal (E1), dorsal (E2), and ventral (E3) views; (F) MTM V2009.29.1., right maxilla, in labial (F1) and lingual (F2) views. (G–L) Dentaries: (G) MTM V2009.9.1., right dentary, in lingual view; (H) MTM V2008.26.1., left dentary, in ventrolingual view; (I) MTM V2009.8.1., right dentary, in lingual (I1) and dorsal and slightly lingual (I2) views; (J), MTM V2010.287.1., left dentary, entire specimen in lingual (J1) and dorsal (J2) views, both with arrows pointing at teeth with intact crowns, and close up of same teeth in occlusal view (J3); (K) MTM V2008.29.1., right dentary, in lingual (K1) and labial (K2) views; (L), MTM V2008.25.1., left dentary, in lingual (L1) and labial (L2) views. (M) Jaw fragment (element and side uncertain), MTM V2009.10.1., oriented with tooth crowns pointing to top of page, entire specimen (M1) and close up of three tooth crowns showing lighter apical cap (M2). Most of the figured specimens are from the stratigraphically


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lower, fine-grained, and amber-rich microvertebrate bonebed at level SZ-7-8; however, two specimens (maxilla MTM V 2008.28.1 and dentary MTM V 2008.25.1) are from the stratigraphically higher, coarsegrained, macrovertebrate-dominated bonebed layer at level SZ-6. Images are at different magnifications. 206x235mm (300 x 300 DPI)

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r Fo

Fig. 3. Albanerpetontid dentaries, all in lingual view and at same scale, to show size difference between largest dentary from Iharkút and representative dentaries from four species of Albanerpeton. (A) Albanerpetontidae genus and species indeterminate, MTM V2008.29.1., median portion of right dentary, from the Upper Cretaceous (Santonian) Csehbánya Formation, Iharkút, Hungary. (B) A. pannonicum (Venczel and Gardner, 2005), MÁFI V.11.133.1, left dentary missing most of area for attachment of postdentary bones, from lower Pliocene fissure infill, Csarnóta 2 locality, Hungary. (C) A. nexuosum (Estes, 1981), UCM 38714, left dentary missing symphyseal end and posterior half of bone, from the Upper Cretaceous (upper Maastrichtian) Laramie Formation, UCM locality 77062, Colorado, USA. (D) A. inexpectatum Estes and Hoffstetter, 1976, MNHN.LGA 48, nearly complete left denary, from an unrecorded lower or middle Miocene fissure infill, La Grive-Saint-Alban, France. Lines above and below this specimen demarcate approximate portion preserved in the figured dentary (A) from Iharkút. (E) A. galaktion Fox and Naylor, 1982, UALVP 16217, right dentary missing posteriormost end of tooth row and area for attachment of postdentary bones, from the Upper Cretaceous (upper Santonian) Deadhorse Coulee Member, Milk River Formation, UAVLP locality MR-9B, Alberta, Canada. Specimens at same magnification. 75x30mm (300 x 300 DPI)

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109x65mm (300 x 300 DPI)

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