Mesozoic and Tertiary Anura of Laurasia Zbyněk

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Article: Mesozoic and Tertiary Anura of Laurasia

Article in Palaeobiodiversity and Palaeoenvironments · November 2013 DOI: 10.1007/s12549-013-0131-y

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Palaeobiodiversity and Palaeoenvironments

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Palaeobio Palaeoenv (2013) 93:397-439 DOI 10.1007/s12549-013-0131-y

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Palaeobio Palaeoenv (2013) 93:397–439 DOI 10.1007/s12549-013-0131-y

REVIEW

Mesozoic and Tertiary Anura of Laurasia

ZbyněkRoček

Received: 4 April 2013 /Revised: 27 August 2013 /Accepted: 16 September 2013 /Published online: 6 November 2013 # Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2013

Abstract Anurans of Laurasia have a long history that begins when Eurasian and American herpetofaunas reached their peak with the earliest known anuran, Prosalirus, from the Early taxonomic diversities. In the Pliocene, some extant anuran spe- Jurassic of Arizona, USA. At that time, western Laurasia cies appeared, but at the same time taxa that had been dominant (North America) was still connected with Gondwana, so throughout the Oligocene and Miocene (e.g. Eopelobates, faunal interchange was still possible between those parts of palaeobatrachids) became extinct during this interval or during the former Pangean supercontinent. The anuran fossil record the subsequent Pleistocene glaciation. The brief biochronological from the Jurassic and Cretaceous of Laurasia is mainly repre- synopsis presented here is followed by a systematic review of sented by disarticulated skeletal elements similar to those of taxa with their diagnoses and published data on their stratigraphic Prosalirus (e.g. amphicoelous vertebrae indicating the pres- and geographic distributions. ence of continuous notochord; ilia without dorsal crest and dorsal tubercle; small body size). Because the morphology of Keywords Anura . Laurasia . Taxonomic diagnoses . the ilium, the most commonly preserved element of Mesozoic Systematic review . Stratigraphy . Distribution anurans, superficially recalls that of Recent Alytes, Bombina or Pelobates, Mesozoic anurans often were assigned to discoglossids and pelobatids. The Cretaceous portion of the Introduction Laurasian anuran record is marked by the appearance of procoelous and opisthocoelous vertebrae, ilia bearing a dorsal The aim of this paper is to summarise the Mesozoic and Tertiary crest and dorsal tubercle (although such ilia may rarely be record of anurans from Europe, Asia and North America, the found as early as in the Jurassic) and larger body sizes. three modern land masses in the Northern Hemisphere that were Cretaceous anuran assemblages include a mix of generalised once part of the single supercontinent of Laurasia. This paper taxa that are comparable to Recent basal anurans and more updates and complements earlier reviews (e.g. Holman 2003; specialised taxa lacking clear affinities with any extant an- Rage and Roček 2003;Roček 2000;Roček and Rage 2000a; urans. Some of these forms survived into the Paleocene, but in Sanchiz 1998a) by documenting new occurrences and taxa and general anuran faunas on all Laurasian continents were mark- presenting new interpretations. In the first part of this paper, I edly depleted in the Paleocene. Major groups of anurans provide an annotated, biochronological synopsis of the appeared in the Eocene. The early Miocene is the interval Laurasian anuran record, beginning with the Early Jurassic when anurans are first recorded, through to the end of Pliocene when the Recent herpetofauna was basically established. In the second part of this paper, I provide brief This article is a contribution to the special issue “Mesozoic and Cenozoic systematic accounts for all anuran taxa currently recognised lissamphibian and squamate assemblages of Laurasia” from the Mesozoic and Tertiary of Laurasia. These accounts Z. Roček (*) include a diagnosis (mainly as they were published, only slightly Department of Palaeobiology, Geological Institute, Academy of re-arranged for the sake of brevity), details on stratigraphic and Sciences of the Czech Republic, Rozvojová 135, 165 00 Prague 6, Suchdol, Czech Republic geographic occurrences and additional information or comments e-mail: [email protected] as appropriate. Ideally, a diagnosis should be a brief list of Author's personal copy

398 Palaeobio Palaeoenv (2013) 93:397–439 apomorphies, by which a given taxon can be distinguished from Biochronological synopsis other taxa of the same group (Bell et al. 2010). The intention of these systematic accounts is not to provide a comprehensive Prosalirus bitis, the earliest known anuran, was recovered revision of Mesozoic and Tertiary anurans of Laurasia, but to from the Early Jurassic age Kayenta Formation in Arizona, provide an up-to-date, encyclopedic treatment of currently USA (Jenkins and Shubin 1998; Shubin and Jenkins 1995). recognised taxa. The material consists of conglomerates of disarticulated Because this paper deals with anurans, neither of the Early bones, of which the presence of amphicoelous vertebrae with Triassic proanurans (i.e. Czatkobatrachus from southern a broad notochordal canal, sacral vertebra with narrow and Poland or Triadobatrachus from Madagascar; see Evans and slightly posteriorly inclined diapophyses and ilia with the Borsuk-Białynicka 2009;Roček and Rage 2000b) are consid- shaft rounded dorsally and lacking any crest or tubercle ered further, except to note that the presence of those (Fig. 1)suggeststhatProsalirus was a poor jumper. As the proanurans in widely separate places suggests that the ances- notochord obviously passed through the sacro-urostylar artic- tors of anurans were widely distributed on the Triassic super- ulation into the urostyle, the joint could not be bicondylar, a continent Pangaea. condition that allows for rotation of the pelvis in the vertical Although India is Gondwanan by origin and had merged plane and is typical of anurans that are good jumpers with Laurasia by the end of Cretaceous or in the earliest (Emerson 1979). Palaeogeographical reconstructions for the Cainozoic (the date of the earliest terrestrial contact between Pliensbachian suggest that Prosalirus was not geographically Indian and Eurasia is still matter of discussion), its anurans are isolated from the slightly younger, Early Jurassic anurans included in this review just to demonstrate the probability or Notobatrachus and Vieraella in South America because improbability of the Gondwanan origin of some taxa (e.g. Laurasia had not yet separated from Gondwana. A fragmen- Eopelobates). tary iliac shaft has been reported from the Early Jurassic age (Hettangian through Pliensbachian) Kota Formation of India and attributed to pelobatids (Yadagiri 1986). The Kota spec- Anatomical abbreviations imen indicates that a Prosalirus-like grade anuran occurred in the Indian portion of Gondwana. This formerly Gondwanan F, length of femur; LC, length of head; LtC, width of head; anuran subsequently became part of the Laurasian assemblage – SVL, snout vent length, measured in fossil anurans between when the Indian subcontinent collided with the rest of Asia. anterior end of premaxillary symphysis to posterior end of Because all of the above-mentioned Early Jurassic anurans ischia; TF, length of tibiofibula; V2,V3, etc., presacral verte- and, indeed, most other Mesozoic anurans lacked a dorsal brae, numbered anteriorly to posteriorly. crest on their iliac shafts (a situation analogous with some Recent taxa, such as Pelobates), the assumption is that those Institutional abbreviations anurans were poor jumpers. The next youngest Laurasian anuran is Eodiscoglossus GPIB, Steinmann-Institut für Geologie, Mineralogie und oxoniensis, which is based on isolated skeletal elements from Paläontologie of the Friedrich-Wilhelms-Universität, Bonn, the Middle Jurassic of Kirtlington, UK (Evans et al. 1990). Germany; MCZ, Museum of Comparative Zoology, These bones represent the earliest known anuran in Europe. Cambridge, Massachusetts, USA; MNA, Museum of Following the Pliensbachian, there is a gap of about 40 Ma Northern Arizona, Flagstaff, Arizona, USA; MNCN, Museo in the North American anuran record, until isolated bones and Nacional de Ciencias Naturales, Madrid, Spain; NHMUK, associated skeletons appear in the Late Jurassic (Kimmeridgian Natural History Museum, London, UK; NMP, National and Tithonian) age Morrison Formation in Wyoming, Utah, Museum, Prague, Czech Republic; PIN, Paleontological and Colorado, USA. An isolated anuran humerus from Quarry Institute, Moscow, Russia. 9, Wyoming, was described by Marsh (1887)asEobatrachus agilis and interpreted by Hecht and Estes (1960) as a possible pipoid. Another humerus from the same locality was described Chronological abbreviations by Hecht and Estes (1960)asComobatrachus aenigmatis. Both of those humeri are now considered indeterminate. Also ECret, Early Cretaceous; EEoc, Early Eocene; EJur, Early from Quarry 9, Evans and Milner (1993) described an isolated Jurassic; EMio, Early Miocene; EOlig, Early Oligocene; ilium as the holotype for the discoglossid Enneabatrachus EPaleo, Early Paleocene; LCret, Late Cretaceous; LEoc, Late hechti; this species also has been reported from the Morrison Eocene; LJur, Late Jurassic; LMio, Late Miocene; LOlig, Late FormationinUtah(Henrici1998a, b). Evans and Milner (1993) Oligocene; LPaleo, Late Paleocene; MEoc, Middle Eocene; described a second ilium from Quarry 9 as an indeterminate LPlio, Late Pliocene; MJur, Middle Jurassic; MMio, Middle pelobatid on the basis of its lacking a dorsal tubercle and having Miocene; MPaleo, Middle Paleocene; Plio, Pliocene. an oblique groove on the inner surface of the bone. Considering Author's personal copy

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as a possible ancestral palaeobatrachid (Wuttke et al. 2012). Ancestral palaeobatrachids, or early Palaeobatrachidae, may also be represented by a poorly preserved fragment of maxilla with peculiar lingual knobs between the alveoli, recovered from the late Barremian of Spain (Buscalioni et al. 2008). Recently, another palaeobatrachid-like anuran, Gracilibatrachus, has been described from the late Barremian of Las Hoyas (Báez 2013); it differs from true Palaeobatrachidae by having presacral vertebrae that are free from the sacral vertebra and by having posteriorly inclined and comparatively narrow sacral diapophyses. A slightly younger and rather uniform anuran assemblage (Dong et al. 2013) is contained in the endemic Jehol Biota of north-east China, which is known from several localities of Barremian/Aptian age. This assemblage consists of primitive anurans (amphicoelous vertebrae, free ribs with uncinate processes, monocondylar sacro-urostylar articulation), all of which are now assigned to the genus Liaobatrachus and ex- hibit little interspecific variation (Dong et al. 2013). Because these frogs are the earliest three-dimensionally preserved, articulated skeletons, they provide important information on the Fig. 1 Prosalirus bitis, the earliest known anuran, from the Early Juras- anatomical structure of Mesozoic anurans and their growth sic of Arizona, USA. a Left ilium (MNA V 8725); acetabular portion of (Roček et al. 2012). This assemblage is related to Late another ilium in medial view marked by arrow. b Reconstructed right Cretaceous Gobiates from central Asia (see below). It is not ilium (MCZ 9324A), reversed for comparison. Note absence of both a excluded that the combination in these Chinese anurans of an dorsal tubercle and a dorsal crest, as in the majority of Mesozoic anurans iliac shaft without a dorsal crest and dorsal tubercle, monocondylar sacro-urostylar articulation and strongly dilated how ubiquitous that pair of features is among Mesozoic an- sacral diapophyses indicate swimming as a predominant mode urans (e.g. characteristic for many ilia recovered from Late of locomotion, similar to extant Barbourula, Bombina and Cretaceous localities in Utah; Roček et al. 2010), they may Leiopelma. Approximately of the same age are disarticulated not be reliably diagnostic for the Pelobatidae. In contrast, the anuran bones from the Transbaikal region of Russia attributed presence of pipoids and, possibly, rhinophrynids, in the Late to indeterminate Discoglossidae (Skutschas 2003). Jurassic of North America is confirmed by partially articulated In North America, the Early Cretaceous (Comanchean, i.e. skeletons of Rhadinosteus parvus in the Morrison Formation late Aptian) Anura have a poor fossil record, represented by a of Utah (Henrici 1998a). discoglossid-like humerus and fragmentary urostyle from cen- The earliest, fully articulated anuran skeletons in Laurasia tral Texas, USA; in addition, this anuran assemblage also were recovered (Villalta 1954) from Early Cretaceous (late includes maxillae with pitted sculpture and amphicoelous and Berriasian or early Valanginian) age strata at Santa Maria de procoelous vertebrae (Winkler et al. 1990). These specimens Meià, Spain, and named Eodiscoglossus santonjae (Fig. 2). are consistent with the suggestion that in anurans, procoelous These skeletons, together with some disarticulated bones (e.g. and opisthocoelous vertebrae evolved from amphicoelous ver- atlas, sacral vertebra, humeri and ilia), recall the Discoglossidae. tebrae in the Early Cretaceous (Roček et al. 2010). Discoglossid-like anurans are also represented in Europe by a In contrast to the Early Cretaceous, Late Cretaceous anurans slightly younger, disarticulated skeleton of Wealdenbatrachus in North America are much better documented, although still jucarensis from the late Barremian of Spain which, although almost exclusively by disarticulated bones. In southern Utah, a characterised by amphicoelous vertebrae, was also tentatively relatively continuous anuran fossil record spans almost 25 referred to the Discoglossidae (Fey 1988). Iberobatrachus from million years, from the early Cenomanian to late Campanian the late Barremian of Spain (Báez 2013) also displays (Roček et al. 2010). Although the disarticulated and fragmen- discoglossid features, mainly in the postcranial skeleton. tary material from the Utah sequence provides only a limited Indeterminate discoglossid humeri have been reported from basis for taxonomic determinations, it does allow some trends the Early Cretaceous (Hauterivian/Barremian) of Spain in anatomical evolution to be recognised. For instance, small- (Canudo et al. 2010). However, another articulated skeleton bodied anurans prevailed until the early Campanian, then be- from Santa Maria de Meià, Neusibatrachus wilferti, was recent- ginning in the late Campanian larger-bodied anurans appeared. ly referred to Pipoidea (Báez and Sanchiz 2007)andsuggested Similarly, ilia in the majority of early anurans (including the Author's personal copy

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orbital margin of the maxilla there is a posterior orifice of a canal that runs anteriorly and opens at the base of the frontal process. Another, probably parallel canal enters the maxilla on the inner surface of the zygomaticomaxillar process. Maxillae with such canals are unknown in extant anurans. Ilia referrable to Scotiophryne were also recovered from the Santonian and CampanianofUtah(Roček et al. 2010) and maxillae referrable to Scotiophryne are known from the Campanian of Utah (Roček et al. 2010) and New Mexico (Armstrong-Ziegler 1980). Theatonius lancensis also is heavily ossified with beaded ornamentation on its cranial roofing bones; moreover, its maxilla is edentate and the pterygoid process is lacking (Fox 1976; Gardner 2008). This too is not paralleled among extant anurans. Maxillae having canals similar to those in Scotiophryne have been tentatively assigned to Eopelobates (Estes and Sanchíz 1982a; Gardner 2008), but they obviously have closer relations to Scotiophryne. Another anuran described from the late Maastrichtian is Paradiscoglossus americanus, known only by ilia that bear a dorsal crest (Estes and Sanchíz 1982a; Gardner 2008). It should be added that Estes and Sanchíz (1982a)assigned some ilia with an extensive acetabulum from the late Maastrichtian age Lance Formation to Palaeobatrachus (as P. occidentalis Estes et Sanchíz, 1982). This generic assignment was later brought into doubt by Sanchiz (1998a)and Wuttke et al. (2012) (but see also Gardner and DeMar 2013, this issue). However, pipoids were no doubt present in the latest

Cretaceous of North America, as evidenced by a fused V1 and V2 with opisthocoelous vertebral centra from Bug Creek Anthills, Montana (Gardner 2008;Wuttkeetal.2012). The taxonomic diversity of the Late Cretaceous anuran fauna in North America appears to be high, as indicated by different morphs of disarticulated ilia (Roček et al. 2013)and Fig. 2 Eodiscoglossus santonjae, the earliest articulated discoglossoid some other bones. However, it is difficult to decide whether frog (MNCN 4723; holotype), from the Early Cretaceous of Spain. Note variation of the ilia really reflects taxonomic differences or is the size. Photograph courtesy of the Museo Nacional de Ciencias Naturales Madrid the result of individual or developmental variation. It is possible that Cretaceous anuran assemblages include taxa that are anatomically different from those living today (e.g. Theatonius, earliest anuran Prosalirus) lacked both a dorsal tubercle and Scotiophryne). It should also be added that although a vast dorsal crest, and some had an oblique groove that traverses the majority of fossil anurans of North America are preserved as dorsal margin of the bone and extends onto the inner surface of disarticulated bones (see also Rowe et al. 1992), there are also the iliac shaft—a similar groove occurs in ilia of Pelobates, rare articulated skeletons, such as an unidentified articulated which was the reason why similar fossil ilia sometimes were frog specimen from the late Maastrichtian part of the North interpreted as pelobatid-like. Through the Late Cretaceous, Horn Formation (Cifelli et al. 1999). there is a trend towards an increasing majority of ilia having a Late Cretaceous anurans in Asia are represented by well-developed dorsal tubercle and dorsal crest, both of which discoglossoids. In terms of their basic features (e.g. frontal could be associated with changes in locomotor behaviour. fontanelle present between frontoparietals; amphicoelous Among these North American Late Cretaceous anurans, verterbal centra perforated by notochordal canal; imbricate some described taxa have morphologies profoundly different neural arches; ribs on V2–V4; fan-like sacral diapophyses) from those in extant anurans. Scotiophryne pustulosa from the these anurans are similar to Liaobatrachus, but they are late Maastrichtian and earliest Paleocene has its maxillae, smaller and, surprisingly, better ossified, the latter of which frontoparietals and squamosals covered externally with pustu- is expressed in the robust build and pitted sculpture of their lar sculpture (Estes 1969; Gardner 2008). Moreover, in the cranial bones. Fossils recovered from Albian through Author's personal copy

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Campanian strata in Mongolia and central Asia pertain to two genera: Gobiates, which contains multiple species and is known from incomplete articulated skeletons and disarticulated bones (Roček 2008;Roček and Nessov 1993; Špinar and Tatarinov 1986), and the monospecific Cretasalia, known by a three-dimensional skeleton (Gubin 1999; this study: Fig. 3b–d). In addition to gobiatoid-like anurans, the Late Cretaceous Asian assemblages also contain true Discoglossidae (e.g. Kizylkuma and some others), as can be judged from the shape of maxillae, opisthocoelous vertebrae, bicondylar sacro-urostylar articulation and morphology of the urostyle (see also Kordikova et al. 2001), as well as peculiar anurans such as Altanulia (Fig. 3a). Although some of these taxa have been described based on disarticulated bones, it is obvious that the anuran assemblages from Asia were highly diversified at the species and generic levels but, similar to North American assemblages, uniform at higher taxonomic levels. Anuran remains from the Maastrichtian intertrappean beds of Naskal, India, represent an interesting problem. The ilia have been attributed to indeterminate discoglossids by Prasad and Rage (1991, 1995), and a fragmentary ilium and a humerus display features of hylid morphology. If these frogs were autochthonous, as seems to be suggested by the geographic location and stratigraphic position of the fossils, they would be of Gondwanan origin. However, if the ilia represent true relations to Laurasian discoglossids, these frogs should be considered early (i.e. Maastrichtian) immigrants from Laurasia after the Indian subcontinent became attached to the rest of Asia. Late Cretaceous anuran assemblages from Europe were still mainly composed of small, discoglossid-like forms, as suggested by disarticulated bones attributed to Eodiscoglossus and Paradiscoglossus (Blain et al. 2010; Folie and Codrea 2005), or described as Hatzegobatrachus and Paralatonia from the Maastrichtian of Romania (Venczel and Csiki 2003), respectively, and to indeterminate discoglossids from Spain (Company and Szentesi 2012). In addition, Bakonybatrachus and Hungarobatrachus have been described from the Santonian of Hungary (Szentesi and Venczel 2010, 2012). Some of these, however, are too fragmentary to be attributed at a familial level. Scarce, and mostly indeterminate anuran material has also been Fig. 3 a Altanulia alifanovi, left maxilla in inner view (PIN 553/300; holotype), from the Late Cretaceous of Mongolia. b–d Cretasalia reported from the Cenomanian of France (Vullo et al. 2011)and tsybini, a Late Cretaceous articulated gobiatid (PIN 3142/399; holotype) latest Maastrichtian of Spain (Blain et al. 2010). from Mongolia: skull in dorsal view (b), postcranial skeleton in left In contrast to dubious discoglossid-like anurans, palaeo- dorso-lateral (c) and ventral views (d). Note large frontoparietal fonta- batrachids were already well established by the end of the nelle and amphicoelous vertebrae with imbricate neural arches Cretaceous (early Campanian), as evidenced by a fragmentary, but doubtlessly palaeobatrachid frontoparietal from the early occur in an uninterrupted sequence of strata that spans the latest Campanian of Villeveyrac in southern France (Buffetaut et al. Cretaceous to Paleocene. Records of anuran faunas just below 1996) and by incomplete maxillae and some postcranial ele- the K–T boundary present vital information on how these ments from the latest Maastrichtian of Spain (Blain et al. 2010). faunas may have been affected by K–T extinction. An excep- Although anurans from the latest Cretaceous (Maastrichtian) tion is the Sheep Pass Formation in east-central Nevada, where were recorded in various Laurasian localities, they only rarely an anuran of uncertain affinities, Eorubeta nevadensis,and Author's personal copy

402 Palaeobio Palaeoenv (2013) 93:397–439 recently also complete and partial skeletons of articulated frogs, 1984). However, the most surprising discovery is which have not yet been taxonomically identified, were recov- Thaumastosaurus in the late Eocene of southwestern France ered from Paleocene strata (AC Henrici, personal communica- and southern UK. It is assigned here to the Leptodactylidae tion). Other localities from the late Maatrichtian–Paleocene (an alternative determination was suggested by Laloy et al. transition are from the Lance and Hell Creek formations of 2013), which is a group poorly defined taxonomically and Wyoming and Montana, USA (Gardner 2008), North Horn most probably not monophyletic, but in spite of that it is Formation of Utah (Cifelli et al. 1999), Sânpetru Formation obvious that the genus undoubtedly has Gondwanan affinities. of Romania (Folie and Codrea 2005), northern Spain (Blain The genus vanished from Europe at the end of the Eocene, et al. 2010) and Naskal near Hyderabad city, India (Prasad and during the “grande coupure”. It can also be added that, judging Rage 1995). Unfortunately, except for specimens from the by their comparatively short anterior limbs, Eocene Sheep Pass Formation, all others are represented by palaeobatrachids were not so closely bound to dwelling in disarticulated skeletal elements. water as they were in the Oligocene and later times (Wuttke Nevertheless, it is apparent that at least some anurans et al. 2012). (Scotiophryne) survived from the Cretaceous up to the middle In North America, the taxonomic poverty of anurans also Paleocene (Estes 1976) in North America. It is not probable did not last long beyond the Paleocene. Two rhinophrynids are that among these North American survivors were true known: Eorhinophrynus from the late Paleocene of Montana discoglossids, as Estes (1975) believed based on his material (Estes 1975) and the middle Eocene of Wyoming (Hecht 1959) from the Big Horn Basin, Wyoming; rather, they display and Chelomophrynus from the middle Eocene of Wyoming “discoglossoid” or “pelobatoid” iliac morphologies, probably (Henrici 1991; Henrici and Fiorillo 1993). Scaphiopus guthriei reflecting non-saltatorial locomotion common to the majority from the early Eocene of Wyoming is the earliest record of the of Mesozoic anurans. However, in Europe, true discoglossids, Scaphiopodidae in North America (Henrici 2000). Eopelobates and also palaeobatrachid frogs may have survived the K–T grandis fromthelateEoceneofSouthDakotadiffersinsome boundary, as seems to be documented by taxa summarised by diagnostic characters from the Pelobatidae (see below) and may Rage and Roček (2003) from Hainin (middle Paleocene, be either related to the Scaphiopodidae or, based on the sculp- Belgium) and Cernay (late Paleocene, France). turing, to Scotiophryne (AC Henrici, personal communica- Apart from these rare Cretaceous survivors, some new tion), an anuran of uncertain systematic position. Finally, the anuran taxa appeared, mostly in the late Paleocene. In North rather enigmatic Elkobatrachus, unfortunately represented ex- America, Estes (1975)reportedEorhinophrynus sp. from the clusively by not fully ossified individuals, also may be assigned late Paleocene of Wyoming and in Europe, Rage (2003) to the American pelobatoids (Henrici and Haynes 2006). Also recovered the earliest Laurasian Bufonidae from the late during this epoch, the Pelodytidae appeared in North America, Paleocene of France. Data on the fossil record of Paleocene as suggested by Miopelodytes gilmorei from the middle Anura from Asia are lacking. It can be summarised that Eocene of Nevada (Henrici and Haynes 2006;Taylor1941). although anurans survived the K–T boundary, their abundance Eorubeta nevadensis from the early Eocene of Nevada was and taxonomic diversity strongly decreased. originally assigned to the leptodactylids (Hecht 1960), but is This situation changed only in the early Eocene. In Europe, now regarded as an anuran of unknown relations. Therefore, as mentioned above, some discoglossid-like forms and pipoid and pelobatoid anurans were dominant components of palaeobatrachids survived, but both were restricted to the North American herpetofauna in the Eocene. It should also be western part (Belgium, UK) of the present-day continent, added that some frogs have been recovered from the Eocene perhaps because the central and eastern parts were still cov- Green River Formation (Grande 1984), among which is a ered by sea. The Pelobatidae also appeared in Europe (Rage skeleton that is diagnostically pelobatid (not scaphiopodid) in and Roček 2003). Complete skeletons from the middle having tripartite frontoparietals. This finding seems to suggest Eocene of Messel and Geiseltal that include cranial elements common origins of European and North American pelobatoids, (especially the diagnostically critical frontoparietals) can un- which has been dated by molecular biology to the Cretaceous equivocally be attributed to Eopelobates (Wuttke 2012b), (Sage et al. 1982). Finally, a hylid ilium was recovered from the rather than Pelobates. These skeletons do not provide any late Eocene of Canada (Hyla swanstoni)byHolman(1968). evidence that Eopelobates was capable of burrowing. Another IntheEoceneofAsia,theanuranfossilrecordislimitedtoat important member of the European herpetofauna, the least five taxa from India. The three named species are: a Pelodytidae, appeared in the late middle Eocene (Duffaud discoglossid, Eobarbourula delfinoi, which seems to be related and Rage 1997) and late Eocene of Europe (Rage 1988), to extant Barbourula (Folie et al. 2013)andwaspublished and quickly spread over the western part of the continent. earlier as an indeterminate Discoglossidae by Bajpai and Kapur They were osteologically close to extant Pelodytes.Inaddi- (2008); the possible rhacophorid Indorana prasadi (Folie et al. tion, the earliest Ranidae were recovered from the late Eocene 2013); and the supposed myobatrachid Indobatrachus (Špinar of France and the UK (Holman and Harrison 1999;Rage and Hodrová 1985; this study: Fig. 4). Folie et al. (2013)also Author's personal copy

Palaeobio Palaeoenv (2013) 93:397–439 403 reported an ilium reminiscent of the Pelobatidae and assigned this Indian specimen to the European genus Eopelobates.This assignment is somewhat doubtful because isolated “pelobatoid” ilia are similar to those of many Mesozoic anurans that were clearly not related to the Pelobatidae; moreover, it would be hard to explain relations between European Eocene Eopelobates and presumed Eopelobates distributed at about the same time on the Indian subcontinent. Folie et al. (2013) also described isolated ilia, vertebrae and urostyles that they identified as “‘Ranidae’ indet.” In summary, on all Laurasian continents, the Eocene was an epoch of high taxonomic diversification of anurans, which contrasts with the poor diversity seen in the preceding Paleocene. In the Oligocene of Europe, diversification continued. Within the Discoglossidae, forms closely related to Discoglossus can be recognised beginning in the early Oligocene (Rage and Roček 2003) and are easily distinguishable from Latonia.Bothgenera probably originated from a common ancestor by heterochrony, which seems to be evidenced by the similarity of adult Discoglossus with juveniles of Latonia. In contrast, species diversity within these genera seems to be restricted; for example, D. troscheli and Opisthocoellelus hessi (Fig. 5), both from the late Oligocene, seem to be conspecific, and Latonia vertaizoni from the late Oligocene differs only in the degree of its cranial sculpturing from Miocene species of Latonia. Palaeobatrachids also are well represented in the Oligocene of Europe, but their taxonomy is poorly understood. Only recently was it determined that their tadpoles had a tendency to gigantism (Roček et al. 2006) and that metamorphosed individuals apparently attained various osteogenetic degrees when they became adults (Wuttke et al. 2012). An implication of the latter finding is that it is Fig. 4 Indobatrachus pusillus, enigmatic tiny frog (specimen “No 16” difficult to discern whether any given metamorphosed individual illustrated in Špinar and Hodrová 1985, 1986, now deposited in the is a fully grown adult or at an incompletely developed stage. It is National Museum Prague as NMP Pb Tv 1004) from the early Eocene of India. Note the small size also difficult to explain why hypertrophied palaeobatrachids (Albionbatrachus) disappeared from the UK at the beginning of the Oligocene. Regarding the Pelobatidae, the genus in the late Oligocene (Böhme 2008). These frogs are Pelobates appeared in European anuran assemblages in the osteologically very uniform, so it is difficult to decide whether early/middle Oligocene, often represented by forms largely cov- forms described as Rana aquensis, R. meriani, and others rep- ered with dermal bones (frontoparietal connected with the squa- resent distinct species. mosal), such as in Pelobates decheni. It should be noted that Compared with Europe, the Oligocene anuran record in although Pelobates and Eopelobates (E. bayeri;Fig.6)simul- North America is poor. The Rhinophrynidae continued to exist, taneously existed in Europe throughout the Oligocene, they as evidenced by Rhinophrynus canadensis from the early never occurred in the same localities. It also is worth emphasising Oligocene of Canada (Holman 1963). Scaphiopus skinneri is that both genera are easily distinguishable by their cranial oste- documented by a nearly complete skull from the middle ology, whereas their postcranial skeletons are uniform. The Oligocene of North Dakota (Estes 1970), and Scaphiopus sp. Oligocene P. decheni was not yet capable of burrowing, as can has also been found in the late Oligocene of Florida (Patton be inferred by the proportions of its hind limbs, absence of a 1969). In addition, an unidentified Bufo was recovered from spade and its articulated sacro-urostylar connection; the same the late Oligocene of Florida (Patton 1969). The more limited conditions also occur in Eopelobates (Roček and Wuttke 2010). Oligocene assemblage of Rhinophrynidae, Scaphiopodidae Finally, the earliest water frogs (“green frogs”), referred to as and Bufonidae in North America differs from that of Europe, Rana ridibunda,appearedintheearlyOligoceneofEurope where discoglossids, palaeobatrachids, pelobatids and ranids (Sanchiz et al. 1993); their existence has also been confirmed were comparatively well diversified. Author's personal copy

404 Palaeobio Palaeoenv (2013) 93:397–439

is from the middle Miocene of Hungary and Romania (Venczel 2004; Venczel and Sţiucă 2008), but unidentifiable Hyla was common in central and eastern Europe throughout the whole Miocene (Hír and Venczel 2005; Hodrová 1987, 1988; Sanchiz 1998b; Venczel et al. 2005). Also, the Bufonidae, including those identical or closely related to extant Bufo bufo, have been recorded from central Europe. Bufo close to B. viridis was recorded from the early and middle Miocene of both western and eastern Europe (Böhme 2010;Hodrová1988; Venczel 2004; Venczel and Sţiucă 2008). Several other fossils have also been identified for Bufonidae; for example, Palaeophrynos (now Bufo gessneri) from the middle Miocene locality of Öhningen (Sanchiz 1998a). With respect to Ranidae, the earliest “brown frogs” related to Rana temporaria were reported from the early Miocene of Germany (Böhme 2001). The Miocene was also an epoch when Eopelobates begantobe gradually replaced by Pelobates. However, evidence of this process should be treated with caution because the only reliable diagnostic characters differentiating the two genera are in the skull, whereas the postcranial skeleton is fairly uniform. In the late Miocene, well-ossified Pelobates (similar to extant P. syriacus, P. varaldii and P. cultripes) was joined by a less ossified form close to extant P. fuscus in which the postorbital connection of the frontoparietal with the squamosal is interrupted Fig. 5 Opisthocoelellus hessi, the single specimen documenting the even in full-grown individuals (e.g. late Miocene of Polgárdi, occurence of discoglossids in the late Oligocene of Bechlejovice, Czech Hungary; Venczel 1997). Miocene Pelobates probably was Republic (NMP Pb 1112, holotype; dorsal aspect). Absence of ossified epiphyses on femora indicates this is not a fully grown individual more diverse than it is today, covering a whole range of forms from those with a broad postorbital connection (P. fahlbuschi)to In Asia, Macropelobates appeared in the early Oligocene those with the postorbital ridge narrow (P. sanchizi). of Mongolia (M. osborni). This was a pelobatoid perhaps In North America, the Scaphiopodidae diversified, as related to the Scaphiopodidae (Roček 1982). Another evidenced by late Miocene occurrences of Scaphiopus pelobatoid from the early Oligocene of Mongolia is alexanderi from Nevada (Zweifel 1956), S. hardeni from Uldzinia (Böhme 2007;Gubin1996). Identification is based Kansas (Holman 1975), S. wardorum from Nebraska (Estes onasinglemaxilla,whichdiffersfromthatofMacropelobates and Tihen 1964) and unidentified Scaphiopus from the middle osborni (similar in size and area where it was found) by its pit- Miocene of Texas (Williams 2002). Simultaneously, less ossi- and-ridge sculpture (Gubin 1996). fied Spea appeared, as evidenced by S. neuter from the late At the beginning of the Miocene, the European herpetofauna Oligocene of South Dakota (Kluge 1966;Holman2003)andin reached the peak of its diversity. The Discoglossidae were still the early to late Miocene of Nebraska, Montana and Oklahoma represented by Discoglossus and Latonia (L. gigantea, L. (Henrici 2009; Parmley et al. 2010)andS. studeri from the late ragei, L. seyfriedi), but unidentifiable Bombina was also pres- Miocene of Kansas (Holman 1975). Miocene Pelodytidae in ent (Sanchíz and Schleich 1986). Another extant genus, Alytes, North America were represented by Tephrodytes brassicarvalis is known from the late Miocene of Spain (Rage and Roček from the early Miocene of Montana (Henrici 1994). Miocene 2003). Also, the palaeobatrachids remained diverse (although Hylidae were represented by the genus Acris recovered in their species diversity should be revised), and their main area of Miocene deposits of Florida, Oklahoma and Texas (Holman distribution shifted east of the Rhine Graben (Wuttke et al. 1967; Parmley et al. 2010;Rogers1976; Williams 2002), 2012). Another distinct group of anurans in the European Proacris (P. mintoni) from Florida (Holman 1961), Miocene were the Pelodytidae, which apparently were distrib- Pseudacris (P. nordensis) from Nebraska (Chantell 1964), uted across the whole of western and central Europe: Pelodytes and Hyla (H. miocenica and H. miofloridana) from Texas arevacus in the early and middle Miocene of Spain (Sanchiz and Florida (Holman 1966, 1967). Miocene bufonids are mostly 1978), and unidentified Pelodytes in the early and middle represented by the ilia. They include the extinct Bufo alienus, B. Miocene of France and Austria (Rage and Hossini 2000; pliocompactilis and B. spongifrons from Kansas (Tihen 1962; Sanchiz 1998b) and the late Miocene of Rudabánya, Hungary Wilson 1968), B. campi (based on a single, possibly anomalous (Roček 2005). The earliest Hyla referable to extant H. arborea tibiofibula) from Mexico (Brattstrom 1955a), B. holmani and Author's personal copy

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Fig. 6 Eopelobates bayeri, a European species of pelobatid from the late unfused sacro-urostylar joint. b Premetamorphic tadpole (NMP Pb Oligocene of Bechlejovice, Czech Republic. a Adult (NMP Pb 1694). 1716). Arrow marks unpaired median ossification, which takes part in Arrows mark diagnostic characters—moderately sinusoid anterolateral formation of ultimate frontoparietal margin of nasal, continuous posterior margin of frontoparietal and

B. kuhrei from Nebraska (Holman 1973; Parmley 1992), B. basaltica and by other, medium-sized members of the group. praevius and B. tiheni from Florida (Auffenberg 1957;Tihen Their development is documented both by premetamorphic 1951), B. hibbardi and B. valentinensis from Nebraska and tadpoles and individuals in various stages of metamorphosis. Kansas (Estes and Tihen 1964;Taylor1936;Voorhiesetal. The Pelobatoidea are represented by a large and robust spe- 1987) and extant forms identical with, or close to, B. cognatus cies, Macropelobates linquensis, which obviously was a good and B. woodhousii (Holman 2003). The microhylid genus burrower, as indicated by its large spade and comparatively Gastrophryne is represented by the extant G. carolinensis from short hindlimbs. Bufo shandongensis is represented by one of the early Miocene of Florida (Meylan 2005). Finally, Miocene the oldest known articulated skeletons of Bufo in Eurasia. ranids are represented by the extinct Rana pliocenica from the Rather surprising, when viewed from today’s perspective, is late Miocene of California, the extant species R. catesbeiana the absence of discoglossids at the Shanwang locality because andaformclosetotheextantR. clamitans. In addition, an at that time Bombina was already present in Europe. anuran ilium having a large acetabulum, dorsal tubercle and Eobarbourula, which appears to be closely related to the slender iliac shaft was described by Holman (1975)fromthe extant Asian discoglossid Barbourula, has also recently been late Miocene of Kansas as Tregobatrachus hibbardi; its family reported from the early Eocene of India (Folie et al. 2013). assignment remains obscure. In the Pliocene of Europe, Discoglossus was pushed south- The anuran fauna of Asia during the Miocene also was wards, as indicated by its late Pliocene occurrences in Spain similar to that of today, as evidenced by the faunal composi- (Bailon 1991), and only one form of Bombina continued to tion of the Shanwang locality in China (Roček et al. 2011). exist. Affinities of the latter to B. bombina is based on the fact The most frequent were the Ranidae, represented by Rana that it was better ossified that B . variegata (personal Author's personal copy

406 Palaeobio Palaeoenv (2013) 93:397–439 observation). The earliest B. bombina is recorded from the The fossil record of Pliocene anurans in North America late Pliocene (MN 15) of Wąże 1, Poland (Sanchíz and mainly consists of taxa related to extant species, few of which Młynarski 1979) and the earliest B. variegata from the late are known from the preceding Miocene. Evidence of extant Pliocene or early Pleistocene of Brasso, Hungary (Bolkay species in Pliocene deposits include Scaphiopus holbrooki 1913); now Braşov, Romania. Similar to Discoglossus, from Florida (Meylan 2005)andSpea hammondii from Latonia also was pushed to the southern part of the Idaho (Chantell 1970). Hylids are represented by Acris European continent, as indicated by its Pliocene occurrences crepitans from Texas (Rogers 1976), Hyla cinerea from in Sète (France), Polgárdi and Osztrámos 1 (both Hungary), Florida (Meylan 2005) and by a form closely resembling H. Ivanovce (Slovakia), Kuchurgan (Ukraine) and Arondelli regilla from Idaho (Chantell 1970). In general, hylids that were (Italy) (Bailon 1991; Chkhikvadze 1981;Roček 1994; comparatively abundant in the Miocene are not found in Pliocene Špinar 1978; Venczel 2001; Vergnaud-Grazzini 1970). Until localities. Other Pliocene anurans closely related to extant forms recently it was believed that the last European occurrences of are Bufo cf. B. alvarius from Arizona (Gehlbach 1965;Tihen Latonia were in the Pleistocene (latest Villafranchian) locali- 1962), Bufo cf. B. cognatus from Texas, Nebraska and Kansas ties of Pietrafitta and Triversa, Italy (Esu et al. 1993). (Eshelman 1975; Gehlbach 1965;Rogers1976, 1984;Tihen However, recent morphological and molecular studies have 1962), B. valliceps from Texas (Rogers 1976)andB . revealed that it survives as an extant species Latonia woodhousii from Texas, Arizona and Kansas (Brattstrom nigriventer (formerly Discoglossus nigriventer) in a small 1955b; Eshelman 1975; Rogers 1976). The only Pliocene refugial area in northern Israel (Biton et al. 2013). During bufonids having no relations to extant species are B. defensor the Pliocene, the Palaeo- batrachidae disappeared from west- from Florida, which might be, however, compared with Bufo ern Europe, but they were still present in the eastern part of the terrestris (Meylan 2005), B. rexroadensis and B. suspectus from continent (Slovakia, Hungary, Romania, Ukraine; see Wuttke Kansas (Holman 2003)andB. spongifrons from Nebraska et al. 2012). The last recorded occurrences of palaeobatrachids (Rogers 1984;Tihen1962). Among the Ranidae, Pliocene forms are from the middle Pleistocene (about 500 thousand years identical or close to extant species are Rana areolata and R. ago) of Russia (Wuttke et al. 2012). Eopelobates also palustris from Texas (Rogers 1976), R. aurora and R. pipiens disappeared (the last doubtless occurrence seems to be Idaho (Chantell 1970), R. capito from Florida (Meylan 2005), R. Osztramos 1, MN 14) in Hungary (Venczel 2001) and was catesbeiana from Nebraska, Kansas and Florida (Eshelman ultimately replaced by a form of Pelobates with affinities to 1975; Holman 1972; Meylan 2005; Rogers 1984) and R. the extant species P. fuscus. It should be emphasised, howev- sylvatica from Nebraska (Rogers 1984). Thus, it seems that the er, that Pliocene records of Eopelobates based exclusively on contemporary anuran fauna was shaped in North America only postcranial elements should be treated with caution because in the Pliocene.. postcranial elements (including ilia) of Pelobates and In Asia, Pliocene anurans were rarely recorded, and the Eopelobates are fairly similar. Concerning the Pelodytidae, fossils are not very informative. Rana yushensis (Rana whereas the extinct Miocene species P. arevacus disappeared, vushensis in the Chinese part of the text, published before it was replaced in the early Pliocene by a form that closely the English part; Liu 1961) from the locality of Yushe, recalls the extant P. punctatus (Bailon 1991). European hylids Wuxiang, Shanxi, may be diagnosed as a ranid only by the were represented by a form closely related to the extant H. proportions of its hindlimbs. Rana hipparionum from the late arborea as early as the middle Miocene in Hungary and Miocene or early Pliocene of the locality of Ertemte, Huade, Romania (Venczel 2004; Venczel and Sţiucă 2008), and this Inner Mongolia, is represented by a number of disarticulated, situation did not change until the end of the Pliocene, as not catalogued skeletal elements. These species may be con- evidenced by records of this hylid from Ivanovce (Hodrová sidered nomen nudum and nomen dubium, respectively 1981). The same holds true for Bufo bufo (Hodrová 1980, (Sanchiz 1998a). 1981). The earliest doubtless B. viridis was recorded in the It is obvious from the above review that Mesozoic anurans early Pliocene of Hungary (Venczel 2001), although forms are, with few exceptions, mostly represented by disarticulated tentatively referrable to B. viridis have been reported from the bones, of which the ilia are similar to those of contemporary early Miocene (Böhme 2010). The earliest form close to Rana discoglossoids not capable of strong jumping (e.g. leiopelmatids, arvalis is from the late Pliocene of Slovakia (Hodrová 1981). Bombina and Alytes) and of pelobatoids. However, these super- The earliest Rana perezi was also recorded from the Pliocene ficial resemblances do not reflect real relationships with contem- (Gómez de Soler et al. 2012). The relationships of well- porary representatives. Rather, they reflect an anatomical condi- preserved Rana strausi from the late Pliocene of Germany tion in early anurans, namely, poor jumping capability as inferred to extant species are still unknown. Among “brown frogs”, by the absence of a dorsal crest on the iliac shaft and less rigid only a form related to R. temporaria has been recorded from articulation between their amphicoelous vertebrae and at the the Miocene (Böhme 2001); other “brown frogs” are known sacro-urostylar joint. Another general feature of Mesozoic an- only from the Pliocene (see R. arvalis above). urans was that they were surprisingly small, sometimes no Author's personal copy

Palaeobio Palaeoenv (2013) 93:397–439 407 greater than 20 mm in SVL. Larger and well-ossified forms (such transverse processes; iliac shaft without dorsal crest (crest as Liaobatrachus;Roček et al. 2012), as well as those with only in saltatory forms like Discoglossus); sternum triradiate. dorsal crest on the iliac shaft and bicondylar sacro-urostylar articulation (both features indicate stronger jumping capability), Prosaliridae Shubin et Jenkins, 1995 appeared only in the Early Cretaceous. Mesozoic anurans includ- Prosalirus Shubin et Jenkins, 1995 ed some taxa that were profoundly different anatomically from Prosalirus bitis Shubin et Jenkins, 1995 (Fig. 1) extant forms (e.g. Scotiophryne characterised by distinctive canals in its maxillae). Some of those survived the K–Textinction Diagnosis (Shubin and Jenkins 1995): Tibiale and fibulare event; however, relatively soon thereafter (by the end of the unfused; roughly 60 maxillary teeth; well-ossified Paleocene and in the Eocene) they were replaced by taxa with sphenethmoid with moderately broad, bony nasal septum modern affinities. and ossified orbitonasal canal, sphenethmoid extends posteri- A major anuran faunistic shift in Laurasia occurred in orly to include the anterior margin of the optic foramen; ilium the Paleocene, when the majority of Mesozoic survivors lacking dorsal crest and supraacetabular fossa, with only a disappeared (European Discoglossidae and Palaeobatrachidae slight rugosity in place of a dorsal tubercle; atlas with a large, may be an exception) and new lineages (such as the Bufonidae funnel-shaped notochordal fossa between the cotyles; verte- and Rhinophrynidae) were established. Appearances of new brae notochordal; sacro-urostylar articulation cartilaginous lineages continued in the Eocene—pelobatoids as evidenced (non-condylar); four (possibly five) paires of ribs; uncinate by Eopelobates in Europe and by closely related forms in the processes project anteriorly; third rib trifurcate distally. Green River Formation and Elkobatrachus in North America. Stratigraphy and distribution: EJur (Pliensbachian) Gold In addition, pipoids represented by Rhinophrynidae spread Spring, Arizona USA (Shubin and Jenkins 1995). throughout North America, whereas the Pelodytidae and Ranidae did the same in Europe. Curiously, the European Gobiatidae Roček et Nessov, 1993 batrachofauna was enriched by temporary immigrants from Gondwana (Thaumastosaurus). True Discoglossidae and Diagnosis (after Roček and Nessov 1993;Roček 2008): Skull Palaeobatrachidae remained restricted to the western part of broader than long (LC:LtC 0.56–0.88); anterior end of maxilla Eurasia, but pelobatoids (Macropelobates) and bufonids deep; quadratojugals present; frontal portions of spread to the eastern part of the continent. Diversification of frontoparietals separated by a fontanelle, parietal portions herpetofaunas in Eurasia reached its highest point in the connected by median suture; lateral wings of parasphenoid Miocene, then during the Pliocene some forms disappeared. short and rounded; widely rounded median process on poste- In North America, the situation was different in that only a rior margin of parasphenoid; nine (exceptionally ten) presacral few Miocene taxa survived into the Pliocene. Nevertheless, vertebrae; vertebral centra amphicoelous, with notochordal the Pliocene was the epoch during which anurans closely canal entering urostyle; neural arches imbricate; sacro- related to extant species appeared both in Europe and North urostylar articulation monocondylar; urostyle with one pair America. of transverse processes; sacral diapophyses extended anteroposteriorly (“fan-like”), with slender basis and convex lateral margin; urostyle with pair of short diapophyses; acetabular Systematic review portion of ilium inclined ventrally, so dorsal margin of iliac shaft at level of anterior acetabular margin is markedly Discoglossoidea Günther, 1858 convex. Remarks: Tiny maxillae described by Roček and Nessov Diagnosis (Modified from Duellman 1975, see also Evans (1993)asGobiatoides and which are reminiscent in their and Manabe 1998; only characters identifiable in fossils are overall morphology to those of larger Gobiates might be- listed): Frontoparietal fontanelle present; frontoparietal long to juveniles of the latter genus. Liaobatrachus,as incrassation divided into frontal and parietal portions (but recently revised by Dong et al. (2013), fits into the scope not in left and right moieties); parahyoid V-shaped; vertebral of diagnostic features of Gobiatidae (see below, diagnosis centra amphicoelous or opisthocoelous (the latter evolved of Liaobatrachus). from the former in the Early Cretaceous); free ribs with uncinate processes or flared epiphyses on V2–V4;sacral Cretasalia Gubin, 1999 diapophyses dilated to varying degrees (those that are fan- Cretasalia tsybini Gubin, 1999 (Fig. 3b–d) like could be associated with swimming, whereas narrower and posteriorly declined ones with jumping); monocondylar Diagnosis (after Gubin 1999): Skull tapered anteriorly; der- or bicondylar sacro-urostylar articulation (bicondylar associ- mal bones smooth except for posterior surface of frontal ated with saltatory locomotion); urostyle with one or two process of maxilla which is covered by vestigial sculpture; Author's personal copy

408 Palaeobio Palaeoenv (2013) 93:397–439 frontal process of maxilla prominent; zygomaticomaxillar Stratigraphy and distribution: LCret (Coniacian) process of maxilla absent; orbital margin straight; posterior Dzhyrakuduk, Uzbekistan (Roček 2008). part of maxilla shallow; deep groove on anterior half of labial surface of maxilla. Gobiates dzhyrakudukensis Roček et Nessov, 1993 Stratigraphy and distribution: LCret Khermeen-Tsav, Mongolia (Gubin 1999). Diagnosis (Roček 2008): Processus zygomaticomaxillaris maxillae prominent above the level of margo orbitalis, its Gobiates Špinar et Tatarinov, 1986 posterior margin vertical (hence posterior part of bone which articulates with quadratojugal is slender); tooth row terminates Diagnosis (after Roček 2008): Frontoparietal table concave, at level of posterior end of the lamina horizontalis maxillae; sculptured around the margins but smooth in depressed area; margo orbitalis maxillae paralleled on inner surface by a dermal bones of skull roof and maxilla covered by irregular rounded ridge delimiting dorsally the groove for pit-and-ridge sculpture; frontoparietals paired, their posterior palatoquadrate bar. surface pierced by foramen for occipital artery; anterior end of Stratigraphy and distribution: LCret (Coniacian) maxilla not bifurcated; nasals barely in contact with tips of Dzhyrakuduk, Uzbekistan (Roček 2008). frontoparietals; palatines free in juvenile individuals, coalesced to palatine process of the maxilla in adults; Gobiates fritschi Roček et Nessov, 1993 squamoso-maxillary contact extensive; maxillary-pterygoid contact without participation of pterygoid process on the Diagnosis (Roček 2008): Tooth row of maxilla exceeds the horizontal lamina of maxilla; parasphenoid with distinct keel level where lamina horizontalis turns posterodorsally; edge for in the midline; ribs not expanded at their ends; ilium bears articulation with ramus maxillaris pterygoidei very distinct. variously defined tuber superius, iliac shaft rounded on its Stratigraphy and distribution: LCret (Coniacian) dorsal margin, with shallow groove which crosses dorsal Dzhyrakuduk, Uzbekistan (Roček 2008). margin of ilium obliquely immediately anterior to the tuber. Remarks: Presumed species diversity is inferred from morpho- Gobiates furcatus Roček et Nessov, 1993 logical variation of lamella alaris squamosi and posterior part of maxilla. Although it is admitted that some species may be Diagnosis (Roček 2008): Lamina horizontalis maxillae thin synonymous (e.g. G. kermeentsavi and G. sosedkoi), morpho- and narrow, its posterior end bifurcates in two ridges—upper logical variation of maxilla and squamosal suggests taxonomic ridge continues posterodorsally, the lower delimits end of differences, rather than individual or developmental variation. tooth row; tooth row exceeds posteriorly level of the end of As mentioned above, elements attributed to Gobiatoides by lamina horizontalis; inner surface of margo orbitalis and Roček and Nessov (1993) may represent juvenile Gobiates. processus zygomaticomaxillaris wrinkled. Stratigraphy and distribution: LCret (Coniacian) Gobiates asiaticus Roček et Nessov, 1993 Dzhyrakuduk, Uzbekistan (Roček 2008).

Diagnosis (Roček 2008): Lamella alaris squamosi slender, Gobiates khermeentsavi Špinar et Tatarinov, 1986 sometimes without processus dorsalis, declined posteriorly so that a perpendicular line constructed from the posterior Diagnosis (Roček 2008): Sculpture consisting of irregular, end of squamoso-maxillary suture cuts the orbital margin of sometimes widely spaced pits separated by ridges; quadratojugal the lamella anteriorly. long; processus zygomaticus (i.e. anterior process of the lamella Stratigraphy and distribution: LCret (Coniacian) alaris) slender and pointed; processus zygomaticomaxillaris Dzhyrakuduk, Uzbekistan (Roček 2008). maxillae absent. Stratigraphy and distribution: LCret (Santonian- Gobiates bogatchovi Roček et Nessov, 1993 Campanian) Khermeen Tsav, Mongolia (Špinar and Tatarinov 1986). Diagnosis (Roček 2008): Processus zygomaticomaxillaris maxillae prominent above level of margo orbitalis, confluent Gobiates kizylkumensis Roček et Nessov, 1993 with posterior, tapering part of maxilla; tooth row exceeds the level of the posterior end of lamina horizontalis maxillae; Diagnosis (Roček 2008): Processus dorsalis on the lamella sculpture on maxilla consisting of shallow, indistinct but regu- alaris squamosi prominent; posterior part of the lamella larly distributed depressions; quadratojugal fused to quadrate. slender. Author's personal copy

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Stratigraphy and distribution: LCret (Coniacian) ramus that has no contact with maxilla; postchoanal process Dzhyrakuduk, Uzbekistan (Roček 2008). of vomer as long as prechoanal process, or longer; angular with long, widely rounded coronoid process prominent Gobiates leptocolaptus (Borsuk-Białynicka, 1978) dorsomedially; mentomandibular fused to dentary; columella

present; parahyoid V-shaped; ribs on V2 with uncinate process Diagnosis (Roček 2008): Sculpture consisting of irregular at their end (“hatchet-like” rib may extend also anteriorly); extensive depressions and small shallow pits; quadratojugal ribs on V3 with uncinate process in middle of its length; ribs short; processus zygomaticomaxillaris maxillae absent. on V4 without uncinate process; ribs on V3 and V4 markedly Stratigraphy and distribution: LCret (Campanian or expanded at their medial and lateral ends; scapula short with Campanian-Maastrichtian) Khermeen Tsav II, Mongolia straight anterior margin and anteromedial margin overlapped (Roček 2008). by clavicle; ilium without dorsal protuberance or dorsal crest; 1 or 2 prepollex elements; F of about same length as TF; Gobiates sosedkoi (Nessov, 1981) carpus and distal tarsals ossified. Remarks: Originally, five monotypic genera were formally Diagnosis (Roček 2008): Sculpture on frontoparietals (and reported: Liaobatrachus Ji et Ji, 1998, Callobatrachus Wang supposedly also on lamella alaris squamosi and on maxilla) et Gao, 1999, Mesophryne Gao et Wang, 2001, consisting of numerous and comparatively small pits separated Dalianbatrachus Gao et Liu, 2004 and Yizhoubatrachus by broad ridges. Gao et Chen, 2004. Because each genus was based on a single Stratigraphy and distribution: LCret (Coniacian) specimen, some anatomical characters were ambiguous or Dzhyrakuduk, Uzbekistan (Roček and Nessov 1993). even incorrectly interpreted. A thorough analysis revealed that all, including juvenile Yizhoubatrachus , fit into the variation Gobiates spinari (Roček et Nessov, 1993) range of Liaobatrachus (Dong et al. 2013).

Diagnosis (Roček 2008): Squamoso-maxillary suture long; Liaobatrachus beipiaoensis (Gao et Wang, 2001) posterior margin of lamella alaris squamosi approximately perpendicular to squamoso-maxillary suture; perpendicular Diagnosis (Dong et al. 2013): Relatively long hind limbs; line constructed from the posterior end of squamoso- femur slightly longer than the tibiofibula; ilium with rounded maxillary suture cuts dorsal margin of the lamella alaris acetabular margin; two prepollex elements. squamosi posterior to processus dorsalis. Stratigraphy and distribution: ECret (Barremian/Aptian) Stratigraphy and distribution: LCret (Coniacian) Heitzigou, Liaoning China; ECret (Barremian/Aptian) Dzhyrakuduk, Uzbekistan (Roček 2008). Huangbanjigou, Liaoning China; ECret (Barremian/Aptian) Sihetun, Liaoning China (Dong et al. 2013). Gobiates tatarinovi (Nessov, 1981) Remarks: Originally described as Mesophryne beipiaoensis. Includes also Dalianbatrachus mengi as a junior synonym. Diagnosis (Roček 2008): Edge for articulation with ramus maxillaris pterygoidei gradually lowering and ultimately Liaobatrachus grabaui Ji et Ji, 1998 disappearing; margo orbitalis maxillae delimited medially by an edge; processus zygomaticomaxillaris low and widely Diagnosis (Dong et al. 2013): Relatively long presacral ver- convex. tebral column; relatively short hind limbs; TF nearly the same Stratigraphy and distribution: LCret (Coniacian) length as F; palatine process on maxilla present; one prepollex Dzhyrakuduk, Uzbekistan (Roček 2008). element. Stratigraphy and distribution: ECret (Barremian/Aptian) Liaobatrachus Ji et Ji, 1998 Sihetun, Liaoning China (Dong et al. 2013). Remarks:IncludesCallobatrachus sanyanensis as a junior Diagnosis (after Dong et al. 2013, slightly modified): Dermal synonym. bones of skull roof smooth; anterior end of maxilla bifurcated, only ventral branch articulates with premaxilla; frontal process Liaobatrachus macilentus (Gao et Chen, 2004) of maxilla prominent and slightly inclined anteriorly; frontoparietals with orbital margin straight and inclined ven- Diagnosis (Dong et al. 2013): Maxilla with palatine pro- trally; nasals with a long paraorbital process and extensive cess; cultriform process of the parasphenoid relatively contact in the midline; squamosal with a short zygomatic short (tapering abruptly one-third the length of the skull Author's personal copy

410 Palaeobio Palaeoenv (2013) 93:397–439 from the rostrum) and ends in a slender needle-like tip; Stratigraphy and distribution: LCret Altan-Ula II, ilium with an acetabular margin that points anteriorly; Mongolia (Gubin 1993). two prepollex elements present; F slightly longer than TF. Bakonybatrachus Szentesi et Venczel, 2012 Stratigraphy and distribution: ECret (Aptian) Hejiaxin, Bakonybatrachus fedori Szentesi et Venczel, 2012 Liaoning China; ECret (Aptian) WangJiagou, Liaoning China (Dong et al. 2013). Diagnosis (Szentesi and Venczel 2012): Small size (SVL 25– Remarks: Holotype is a subadult, not a fully-grown individ- 30 mm); iliac shaft with dorsal crest, dorsal protuberance ual. Originally described as Yizhoubatrachus macilentus. being lower than that of the iliac crest and provided with a thickened and flattened dorsolateral surface circumscribed by Liaobatrachus zhaoi Dong et al., 2013 shallow groove; dorsal acetabular expansion robustly built and ventral acetabular expansion small. Diagnosis (after Dong et al. 2013;Roček et al. 2012): Stratigraphy and distribution: LCret (Santonian) Iharkút, Relatively long hind limbs; parasphenoid with cultriform pro- Hungary (Szentesi and Venczel 2012). cess that tapers gradually from its midpoint; ilium with rounded acetabular margin; maxilla without palatine process; pubis Bombina Oken, 1816 ossified in full-grown adults. Stratigraphy and distribution: ECret (Barremian) Diagnosis: Living genus, osteological diagnosis not Qianyangou, Liaoning China (Dong et al. 2013). available.

Gobiatidae indet. Bombina cf. B. bombina (Linnaeus, 1761)

Stratigraphy and distribution: ECret (Aptian) Xierhuqiao, NE Stratigraphy and distribution: LPlio (Csarnotan) Ivanovce, China (Wang et al. 2007; Dong et al. 2013); LCret, Naskal, India Slovakia (Hodrová 1981); LPlio (Villanyian) Včeláre, (Prasad and Sahni 2009). Slovakia (Hodrová 1985). Remarks: The specimen from Xierhuqiao is the articulated skeleton of a juvenile individual, presumably related to Bombina sp. Liaobatrachus. Stratigraphy and distribution: EMio (Agenian, MN2) Discoglossidae Günther, 1854 Weissenburg 6, Germany (Sanchíz and Schleich 1986); EMio (Orleanian, MN3) Stubersheim 3, Germany (Sanchíz Diagnosis (after Duellman and Trueb 1994;Rageand and Schleich 1986); LMio (Vallesian, MN9) Rudabánya, Hossini 2000, partly modified): Eight presacral vertebrae Hungary (Roček 2005); Plio (MN 14) Osztramos 1, with opisthocoelous centra and imbricated neural arches; Hungary (Venczel 2001). sacro-urostylar articulation mono- or bicondylar; urostyle with one pair of transverse processes that can be Discoglossus Otth, 1837 rarely accompanied with an additional, vestigial pair; scapula short, partly overlain by clavicle; fossa cubitalis Diagnosis: Living genus, osteological diagnosis not humeri shallow; proximal part of tibiofibula on cross- available. section consisting of two elliptic bones fused with one another, the long axis of both being parallel. Discoglossus troscheli (Meyer, 1852)

Altanulia Gubin, 1993 Diagnosis (based on features preserved in GPIB Ro4056 Altanulia alifanovi Gubin, 1993 (Fig. 3a) a+b, holotype of Rana troschelii Meyer, 1852, and NHMUK 35657): Frontoparietals and maxillae smooth,

Diagnosis (after Gubin 1993, modified): Maxilla shallow at with no ornamentation; free ribs on V2–V4, only that on level of orbital margin; dorsal margin posterior to orbital V3 with uncinate process; diapophyses on posterior margin is horizontal, so processus zygomaticomaxillaris is presacrals are perpendicular to body axis; sacral not developed; labial surface smooth; processus pterygoideus diapophyses moderately dilated; sacro-urostylar articula- rounded in dorsal aspect, not prominent posteriorly; tooth row tion bicondylar; urostyle with a pair of triangular exceeds processus pterygoideus posteriorly. diapophyses; iliac shaft with dorsal crest. Author's personal copy

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Stratigraphy and distribution: LOlig (MP 30) Rott, Eodiscoglossus oxoniensis Evans, Milner et Mussett, 1990 Germany. Diagnosis (after Evans et al. 1990): Premaxilla with low alary Discoglossus cf. D. troscheli process; iliac shaft flattened and broad but narrow in cross- section with lateral ridge, little waisting at the junction of the Stratigraphy and distribution: LOlig (MP30) shaft and the acetabular region, dorsal tubercle poorly devel- Oberleichtersbach, Germany (Böhme 2008). oped, shallow and flush with the surface of the iliac shaft, supraacetabular fossa deep. Discoglossus sp. or cf. Discoglossus Stratigraphy and distribution: MJur (late Bathonian) Kirtlington, England (Evans et al. 1990). Stratigraphy and distribution: MMio (Badenian, MN6) Mátraszőlős 1, Hungary (Venczel 2004); MMio (Badenian, Eodiscoglossus santonjae Villalta, 1954 (Fig. 2) MN6) Mátraszőlős 2, Hungary (Venczel 2004); LMio (Vallesian, MN9) Rudabánya, Hungary (Roček 2005); LPlio Diagnosis (after Sanchiz 1998a): Iliac shaft rounded in cross- (MN16) Iles Medas, Spain (Bailon 1991); LPlio (MN16) section, without lateral ridge; waisting at junction of the shaft Galera 2, Spain (Bailon 1991). and acetabular region; tuber superius well developed; supraacetabular fossa shallow; premaxilla with elongate alary Enneabatrachus Evans et Milner, 1993 process. Enneabatrachus hechti Evans et Milner, 1993 Stratigraphy and distribution: ECret (Barremian) Galve, Spain (Estes and Sanchíz 1982b); ECret (Berriasian or Diagnosis (Evans and Milner 1993): Iliac shaft narrow Valanginian) Santa Maria de Meià, Spain (Sanchiz 1998a). and showing little development of a dorsal crest; dorsal tubercle prominent; distinct but moderately prominent cf. Eodiscoglossus sp. supraacetabular and subacetabular processes; acetabular rim expanded. Stratigraphy and distribution: ECret (Aptian-Albian) Stratigraphy and distribution: LJur (Kimmeridgian) Höovör, Övörkhangai Aimag, Mongolia (Gubin 1993); Rainbow Park Microsite, Utah USA (Henrici 1998a, b); LCret (Maastrichtian) Pui, Romania (Folie and Codrea LJur (?Tithonian) Quarry 9, Wyoming USA (Evans and 2005); Haţeg Basin, Romania (Grigorescu et al. 1999). Milner 1993). Iberobatrachus Báez, 2013 Eobarbourula delfinoi Folie et al., 2013 Iberobatrachus angelae Báez, 2013

Diagnosis (Folie et al. 2013): Reminiscent of Recent Diagnosis (after Báez 2013): Skull as long as wide; Barbourula from which it differs in smaller (about 1.5-fold) neurocranium narrow; frontoparietals fused at least along the body size; sacral diapophyses less laterally expanded (antero- posterior half of the antero-posterior orbital diameter, forming posterior/lateral length ratio, 1.1); tuber superius more devel- a flat dorsal table; maxilla bearing palatine process; sacral oped dorsally, more asymmetric in lateral view and placed diapophyses distally expanded; urostyle without dorsal crest. more anteriorly compared with the acetabulum. Stratigraphy and distribution: ECret (Barremian) Las Stratigraphy and distribution: EEoc (Ypresian) Vastan Hoyas, Spain (Báez 2013). Lignite Mine, Gujarat, India (Folie et al. 2013). Kizylkuma Nessov, 1981 Eodiscoglossus Villalta, 1954 Kizylkuma antiqua Nessov, 1981

Diagnosis (after Evans et al. 1990): 15–18 tooth positions on Diagnosis (after Roček and Nessov 1993): Maxilla low and premaxilla; about 50 tooth positions on maxilla, pterygoid elongated, with no sculpture on its outer surface, its postorbital process on maxilla poorly developed; coronoid process on section only slightly exceeds horizontal level of the orbital angular smooth and convex with no notches; atlantal cotyles margin; margo orbitalis slightly prominent medially, thus no elongate and flattened; neural arches lacking upturned flared roofing dorsally of the palatoquadrate bar; lamina horizontalis posterior margins; anterior vertebrae bearing free ribs; ilium maxillae terminates abruptly, at about a right angle; tooth row with dorsal crest and dorsal tubercle, iliac synchondrosis ends at the level of posterior end of lamina; caput humeri large, absent. so the epicondyles do not reach beyond distal level of the caput; Author's personal copy

412 Palaeobio Palaeoenv (2013) 93:397–439 ilium with long tuber superius; pars ascendens ilii long, pars (Venczel 2004); MMio (Badenian, MN6) Mátraszőlős2, descendens small; no dorsal crest on iliac shaft. Hungary (Gál et al. 2000); MMio (Astaracian, MN7-8) Opole, Stratigraphy and distribution: LCret (Turonian to Santonian) Poland (Młynarski 1984); MMio (Astaracian, MN7-8) Dzhyrakuduk, Uzbekistan (Roček and Nessov 1993). PrzewornoIandII,Poland(Młynarski 1976); MMio Sámsonháza 3, Hungary (Venczel 2004); MMio (Sarmatian, Latonia Meyer, 1843 MN7-8) Felsőtárkány “Güdör-kert”, Hungary (Hír et al. 2001); MMio (Astaracian, MN8) La Grive St.Alban,France(Hossini Diagnosis (Roček 1994): SVL up to 200 mm; skull moder- 1992); MMio Subpiatră, Romania (Hír and Venczel 2005; ately wider than long; frontoparietals fused both in adults and Venczel et al. 2005); MMio (Sarmatian) Tauţ,Romania juveniles (trace of median suture visible on inner surface of (Venczel and Sţiucă 2008); MMio (Orleanian, MN6) anterior portion of the bone); parietal foramen sometimes Belomechetskaya, Russia (Chkhikvadze 1988); LMio present; foramina for occipital arteries absent; frontoparietal (Vallesian, MN9) Gritsev, Ukraine (Roček 1994); LMio incrassation consists of two parts (elongated anterior and (Vallesian, MN9) Rudabánya, Hungary (Roček 2005); LMio circular posterior one); two coronoid processes on angular; (Turolian, MN13) Polgárdi, Hungary (Roček 1994); Plio (MN distinct recess on bottom of sulcus for Meckel’scartilageat 14) Osztramos 1, Hungary (Venczel 2001); Plio (MN14) the level of posterior coronoid (paracoronoid) process; ptery- Kuchurgan, Ukraine (Chkhikvadze 1981); Plio (MN15) Sète, goid process on maxilla slender and long; large depression on France (Bailon 1991); Plio (MN15) Ivanovce, Slovakia (Špinar inner posterior part of maxilla; distinct supracondylar depres- 1978); Plio (MN16) Arondelli, Italy (Vergnaud-Grazzini 1970). sion on prooticooccipital; posterior semicircular canal prominent as a distinct ridge; contact facets on frontoparietal, Latonia ragei Hossini, 1993 sphenethmoid, parasphenoid and prooticooccipitals irregularly striated; atlas with median keel on its ventral surface; Diagnosis (Hossini 1993;Roček 1994): Maxilla devoid of additional crest present parallel and lateral to ventral crista sculpture, tooth row exceeding posteriorly the basis of on humerus; caput humeri shifted laterally; dorsal crest on processus pterygoideus; scapula elongated; sacral iliac shaft thin with sharp edge declined dorsomedially; pars diapophyses only slightly dilated distally, declined posteriorly. ascendens of ilium prominent; F shorter than TF; tibiale and Stratigraphy and distribution: EMio (Agenian, MN 2) fibulare not fused and markedly different in size. Laugnac, France (Hossini 1993); EMio (MN 4) Oberdorf, Remarks: The genus Latonia was widespread in Europe Austria (Sanchiz 1998b). from the late Oligocene, and it was originally believed that it became extinct in the Pleistocene (Esu et al. 1993). Recently, Latonia cf. L. ragei however, it was discovered using a combination of morphological and molecular data that it survives today in a remark- Stratigraphy and distribution: EMio (Agenian, MN2b) ably small range in the Middle East (Biton et al. 2013). Navarette del Rio, Spain (Sanchiz 1998a); EMio (MN4) Béon 1, France (Rage and Bailon 2005). Latonia gigantea (Lartet, 1851) Latonia seyfriedi Meyer, 1843 Diagnosis (compiled from Rage and Hossini 2000;Roček 1994): Sculpture on frontoparietal of adult consisting of small, Diagnosis (Roček 1994): Frontoparietal with both orbital mar- densely scattered tubercles on the posterior part of the bone, gins converging towards the anterior; anterior margin of sacral tubercles tend to fuse into irregular ridges divergent anteriorly diapophyses perpendicular to main body axis; F/TF about 0.9. in its anterior part; pit-and-ridge sculpture in juveniles, Stratigraphy and distribution: LMio Öhningen, Germany frontoparietal without foramina piercing posterior surface, but (Roček 1994). parietal foramen present; zygomaticomaxillar process covered with sculpture consisting of ridges and tubercles arranged in Latonia vertaizoni (Friant, 1944) parallel arches open posterodorsally; sacral diapophyses may be dilated, so they can be perpendicular to main body axis. Diagnosis (Roček 1994): Maxilla smooth; processus Stratigraphy and distribution: EMio (Orleanian, MN4) coronoideus on angular less prominent than in other species Dolnice, Czech Republic (Hodrová 1987); EMio of Latonia; sacral diapophyses only moderately dilated dis- Sandelzhausen, Germany (Böhme 2010); MMio (Astaracian, tally, approximately perpendicular to main body axis. MN5) La Grive St Alban, France (Roček 1994); MMio Stratigraphy and distribution: LOlig (Stampien, MP30) (Astaracian, MN6) Sansan, France (Rage and Hossini 2000); Vertaizon, France (Roček 1994); EMio (Agenian, MN1, MMio (Astaracian, MN6) Devínska Nová Ves, Slovakia (Roček MN2) sites collectively termed St-Gérand-le-Puy, France 1994); MMio (Badenian, MN6) Mátraszőlős 1, Hungary (Roček 1994). Author's personal copy

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Latonia cf. L. vertaizoni Paralatonia Venczel et Cziki, 2003 Paralatonia transylvanica Venczel et Cziki, 2003 Stratigraphy and distribution: LOlig (MP30) Coderet, France (Roček 1994). Diagnosis (Venczel and Cziki, 2003): Iliac shaft with dorsal crest; iliac synchondrosis present. Latonia sp. Stratigraphy and distribution: LCret (Maastrichtian) Vălioara–Budurone, Romania (Venczel and Csiki 2003). Stratigraphy and distribution: MMio (Astaracian, MN7) Beni Mellal, Morocco (Roček 1994); MMio (Astaracian, Discoglossidae indet. MN7-8) Cerro del Otero, Spain (Sanchiz 1998a); Plio (Ruscinian, MN15) Sète, France (Bailon 1991); Plio Stratigraphy and distribution: ECret (Hauterivian- (Ruscinian, MN15) Serrat d’en Vacquer, France (Roček 1994). Barremian) La Cantalera, Spain (Canudo et al. 2010); ECret (Barremian-Aptian) Krasnyi Yar, Russia (Skutschas 2003); ? Latonia juvenile or Discoglossus adult ECret (Comanchean, =Albian according to Mancini and Scott 2006) Paluxy Church, Texas, USA (Winkler et al. Stratigraphy and distribution: MMio (Astaracian, MN6) 1990); LCret (Santonian) Shakh-Shakh, Kazakhstan Sansan, France (Rage and Hossini 2000). (Kordikova et al. 2001); LCret (Campanian) Ellisdale, New Remarks: Some bones in localities with Latonia (e.g. Sansan) Jersey, USA (Denton and O’Neill 1998); LCret (Campanian/ recall those of Recent adult Discoglossus. Therefore, it is difficult Maastrichtian) Chera Basin, Spain (Company and Szentesi to decide whether they belong to juvenile Latonia or to 2012); LCret (Maastrichtian) Pui, Romania (Folie and Discoglossus. Codrea 2005), LPaleo, Fort Union Formation, Big Horn Basin, Wyoming, USA (Estes 1975); EEoc (Ypresian) Opisthocoellelus Kuhn, 1941 Vastan Lignite Mine, Gujarat, India (Bajpai and Kapur 2008).

Diagnosis (after Špinar 1976, abbreviated): Large skull up to ?Discoglossidae. one-third of the SVL, broader than long; large fontanelle Wealdenbatrachus Fey, 1988 between anterior portions of the frontoparietals, and smaller Wealdenbatrachus jucarensis Fey, 1988 posterior fontanelle; large, strongly dentate vomers; transverse processes of V5–V8 robust and perpendicular to body axis; Diagnosis (after Fey 1988, modified): Frontoparietals sacro-urostylar joint bicondylar; prominent tubercle on the unfused; possibly amphicoelous vertebrae with imbricate neu- ilium; F strongly sigmoid, shorter than TF. ral arches; V2–V4 vertebrae with free ribs; third pair of ribs Remarks: The genus was based on a specimen from the with uncinate process; leiopelmatid-like scapula; ilium with middle Eocene of Geiseltal, described by Kuhn (1941) two (anterior and posterior) separated tubercles; iliac shaft as Opisthocoelellus weigelti. Although the individual is with rounded dorsal crest, rimmed by a broad groove medial- poorly preserved, the opisthocoelous vertebrae and sa- ly; tibiale and fibulare fused at their ends. cral diapophyses suggest that it may be related to the Stratigraphy and distribution: ECret (late Barremian-Aptian; Discoglossidae. Hauterivian after Sanchiz 1998a) Uña, Spain (Fey 1988).

Opisthocoellelus hessi Špinar, 1976 (Fig. 5) Discoglossoidea indet.

Diagnosis (based on data from Špinar 1976): Urostyle with a Stratigraphy and distribution: ECret Shokawa village, pair of slender and posteriorly inclined diapophyses. Japan (Evans and Manabe 1998). Stratigraphy and distribution: LOlig Bechlejovice, Czech Republic (Špinar 1976). Pipoidea Fitzinger, 1843 Remarks: Probably a species of Discoglossus close to D. troscheli, from which it differs only by shape of urostylar Diagnosis (based on data from Frost et al. 2006): diapophyses (pers. observ.). Sanchiz (1998a) considered it a Frontoparietals fused into an azygous element; lateral alae of juvenile Latonia; most probably it is not fully-grown individ- parasphenoid absent. ual, as suggested by the lack of epiphyses on the femur and the lack of terminal portions of the posterior presacral diapophyses. Palaeobatrachidae Cope, 1865 Assignment to Latonia is problematic, because an individual of that body-size should have its frontoparietals completely fused Diagnosis (from Wuttke et al. 2012): Maxilla without or with (see Roček 1994,fig.7). strongly reduced, zygomatico-maxillary process; Author's personal copy

414 Palaeobio Palaeoenv (2013) 93:397–439 quadratojugal absent; nasals narrow and crescent-like; Albionbatrachus oligocenicus Venczel, Codrea et Fărcaş, frontoparietal with dorsal surface flat or domed; parasphenoid 2012 extremely long and slender (“dagger-shaped”), with anterior tip extending beyond anterior border of sphenethmoid; Diagnosis (Venczel et al. 2012): Azygous, hourglass-shaped angular with coronoid process either smooth or bearing frontoparietal, covered by a weak pit-and-ridge dermal sculpture, muscle scars on dorsal surface; sphenethmoid long, with posterolaterally directing paraoccipital processes; dorsal with frontoparietal fenestra more than half the total anterior depression absent; impression of olfactory lobes length of the bone with two parallel ridges delimiting and cerebral hemispheres undivided; posterior portion of articulation area of parasphenoid on ventral surface, Meckel’s cartilage (i.e. articular) in angular is ossified; a and with septum nasi and lateral processes (i.e. ossi- small tubercle present on the preacetabular portion of ilium. fied parts of postnasal walls) very short; mentoman- Stratigraphy and distribution: EOlig (Rupelian, MP23-24) dibulars ossified, fused to dentary; vertebral centra Suceag, Romania (Venczel et al. 2012). procoelous, strongly compressed dorsoventrally, with condyle and cotyle typically crescentic; V1 and V2 Albionbatrachus wightensis Meszoely, Špinar et Ford, 1984 fusedinadults;V7,V8 and sacral vertebra (V9)fused to form synsacrum; V2–V5 bear free ribs in juveniles, Diagnosis (Meszoely et al. 1984): Posterolateral corners (the but ribs are coalesced to diapophyses in adults (con- prootic processes) of the frontoparietal drawn out into prom- sequently, diapophyses appear long); urostyle without inent spurlike processes. diapophyses, with dorsal crest poorly developed or Stratigraphy and distribution: LEoc (Headonian), HH2 entirely absent; clavicle with long, straight scapular process 1ocality near Alum Bay, Isle of Wight, England (Meszoely on lateral end; coracoid with beak-like rostriform process on et al. 1984). anterior margin of lateral end; rostriform process probably Remarks: As shape and size of the holotype and of similar absent in Eocene taxa; scapula short, with anterior margin frontoparietals correspond to those of large, fully grown straight, and acromial and glenoidal portions not separated Palaeobatrachus, it is not excluded that this form should be by incisure; humerus without cubital fossa (except for Eocene re-assigned to Palaeobatrachus,asP. wightensis (Wuttke taxa), with caput humeri comparatively small and located on et al. 2012). or near long axis of bone, and with epicondyles similar in size; metacarpals markedly elongated; ilium with massive ascend- Palaeobatrachus Tschudi, 1838 ing part, large acetabulum extending anteroventrally beyond margin of main body of bone (concealing descending part), Diagnosis: At present, same as the family (Wuttke et al. 2012). and symphysial area strongly protruding; dorsal tubercle pro- Albionbatrachus is based on frontoparietals of adult, fully trudes only slightly in dorsal direction, but is laterally prom- grown individuals that do not differ substantially from fully inent and bears muscle scars on lateral surface; distinct hori- grown Palaeobatrachus. Pliobatrachus, another genus con- zontal depression on inner surface of iliac shaft, ventral mar- ventionally distinguished from Palaeobatrachus, was original- gin of which continues onto dorsal margin of ascending part; ly erected on the basis of a synsacrum, which is a common pubes ossified; tibiale and fibulare not fused. feature for all palaeobatrachids. Attempts were then made to Remarks: It is obvious from the list of palaeobatrachid associate such synsacra (presumably belonging to taxa below that their diagnoses often involve features Pliobatrachus) with other disarticulated palaeobatrachid ele- that apply to all of them and should be thus considered ments in belief that these elements also must belong to to be diagnostic at the family or generic level. Because Pliobatrachus. It is astonishing that until the 1970s, no one the main goal of this paper is not a taxonomic revision, noted the fact that Pliobatrachus was erected on the basis of a the original diagnoses are maintained, but from their general palaeobatrachid feature. Nevertheless, that notion was comparisons with the diagnosis of the family above, it so deeply rooted that any unusual disarticulated palaeobatrachid is clear that some diagnostic characters differentiating elements from the Pliocene and younger strata were referred to among taxa at generic and even species levels overlap Pliobatrachus. To solve the problem of generic diversity within with diagnostic characters at the family level. the Palaeobatrachidae a comparison of individuals of corre- sponding age, preferably fully grown, is necessary. Albionbatrachus Meszoely, Špinar et Ford, 1984 Remarks: Palaeobatrachus occidentalis from the late Maastrichtian and early Paleocene was the only Diagnosis (Meszoely et al. 1984): Distinct, hourglass- palaeobatrachid reported from North America (Estes and shaped frontoparietal bone with a prominently sculptured Sanchíz 1982a;Gardner2008). It was based on the ilia with dorsal surface. a large acetabulum, the anterior margin of which extends Author's personal copy

Palaeobio Palaeoenv (2013) 93:397–439 415 over the pars descendens. However, they differ from those Stratigraphy and distribution: MMio (Badenian) of typical Palaeobatrachus by the nearly straight dorsal Mátraszőlős 2, Hungary (Venczel 2004); MMio Sámsonháza margin of the bone (the pars ascendens is markedly prom- 3, Hungary (Venczel 2004). inent postero-dorsally; compare Estes and Sanchíz 1982a, fig. 4B,E). Later, Sanchiz (1998a) doubted its generic as- Palaeobatrachus cf. P. hiri signment. Stratigraphy and distribution: MMio (Sarmatian) Tauţ, Palaeobatrachus aff. Albionbatrachus wightensis Meszoely, Romania (Venczel and Sţiucă 2008). Špinar et Ford, 1984 Palaeobatrachus langhae (Fejérváry, 1917) and Stratigraphy and distribution: LOlig (MP30) Palaeobatrachus cf. langhae Oberleichtersbach, Germany (Böhme 2008). Diagnosis (based on data from Venczel 2004): Frontoparietal Palaeobatrachus diluvianus (Goldfuss, 1831) with parasagittal crests. Stratigraphy and distribution: Plio (MN 14) Osztramos 1, Diagnosis (based on data from Venczel 2004): Hungary (Venczel 2001); Plio (MN 15) Ivanovce, Slovakia Sphenethmoid not co-ossified with prootics; diapophyses (Hodrová 1981, 1982); Plio (MN 15) Csarnóta, Hungary of V1 and V2 fused with each other; coracoid with (Hodrová 1982); Plio (MN 15) Wąże I, Poland (Młynarski hook-like processus rostriformis; synsacrum formed by 1962; Sanchíz and Młynarski 1979); Plio (MN 16) Rębielice diapophyses of V7,V8 and V9; cubital fossa of humerus Królewskie I and II, Poland (Młynarski 1962, 1977; Sanchíz and relatively shallow; distal part of forelimb relatively long, Młynarski 1979); Plio (MN 16) Hajnáčka, Slovakia (Hodrová so metacarpal IV is of the same length as ulna; TF 1982); Plio (MN 16) Apastovo, Russia (Ratnikov 1997, 2001); shorter than F. Plio (MN 16) Veretie (= Verkhnii Ol’shan), Russia (Ratnikov 1997, 2001); Plio (MN 16) Korotoyak, Russia (Ratnikov 1997, Palaeobatrachus grandipes (Giebel, 1851) 2001); Plio (MN 16) Uryv I, Russia (Ratnikov 1997, 2001); Plio (MN 17) Villány 6, Hungary (Hodrová 1982); Plio Diagnosis (based on data from Venczel 2004): Sphenethmoid (MN 17) Kisláng, Hungary (Sanchiz 1998a); Plio (MN not co-ossified with prootics; frontoparietal broader anteriorly 17) Liventsovka 5, Russia (Ratnikov 1997, 2001). than posteriorly; diapophyses of V1 and V2 fused with each Remarks: Because it was based on a synsacrum, which is a other; coracoid with hook-like processus rostriformis; general feature of all Palaeobatrachidae, Pliobatrachus synsacrum formed by diapophyses of V7,V8 and V9;cubital (monotypic genus consisting solely of Pliobatrachus fossa of humerus relatively shallow; distal part of forelimb langhae) was synonymized with Palaeobatrachus by relatively long, so metacarpal IV is of the same length as ulna; Wuttke et al. (2012). All of the occurrences listed above are TF shorter than F. founded on specimens that originally were identified as Stratigraphy and distribution: EOlig (MP 21) Hoogbutsel, “Pliobatrachus” langhae. The taxonomic identities of those Hoeleden, and Boutersem TGV sites, Belgium (Smith 2003). specimens deserve to be re-evaluated, but for the time being they are sufficient to establish the presence of palaeo- Palaeobatrachus cf. grandipes batrachids in those Pliocene localities.

Stratigraphy and distribution: LOlig Enspel, Germany Palaeobatrachus laubei Bieber, 1881 (Roček and Wuttke 2010). Diagnosis (based on data from Venczel 2004): Sphenethmoid

Palaeobatrachus hiri Venczel, 2004 not co-ossified with prootics; diapophyses of V1 and V2 fused with each other; coracoid with hook-like processus

Diagnosis (Venczel 2004): Frontoparietal unsculptured, rostriformis; synsacrum formed by diapophyses of V7,V8 frontoparietal table widened posteriorly, parasagittal crests and V9; distal part of forelimb relatively long, so metacarpal present, frontoparietal incrassation unbranched posteriorly; IV is of the same length as ulna; TF shorter than F. synsacrum consisting of V7,V8, and V9; nasals crescent- shaped; coracoid with hook-like processus rostriformis; area Palaeobatrachus aff. P. laubei of attachment of mm. glutaeus maximus separated from that of iliofemoralis and iliofibularis muscles; anterior part of Stratigraphy and distribution: LOlig (MP30) parasphenoid co-ossified with sphenethmoid. Oberleichtersbach, Germany (Böhme 2008). Author's personal copy

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Palaeobatrachus robustus Hossini et Rage, 2000 2010); EMio (MN 4) Oberdorf, Austria (Sanchiz 1998b); EMio (Agenian) Poncenat, France (Hossini and Rage 2000); Diagnosis (Hossini and Rage 2000): Large species; MMio (Astaracian, MN6) Sansan, France (Rage and Hossini frontoparietal table well-delimited, narrow, constricted in its 2000); MMio Subpiatră, Romania (Hír and Venczel 2005; medial part, not extending onto paraoccipital processes, Venczel et al. 2005); LMio (Vallesian, MN9) Rudabánya, paraoccipital processes short, pit-and-ridge sculpture lack- Hungary (Roček 2005); Plio (MN16) Ivanovce, Slovakia ing, longitudinal ridges on frontoparietal absent, margo (Hodrová 1981). occipitalis posteriorly convex; ventral surface of vertebrae smooth; coracoid elongated, with sharp ridge on anterior Palaeobatrachidae indet. border. Stratigraphy and distribution: EMio (Agenian, MN 2) Stratigraphy and distribution: LCret (Campanian) Laugnac, France (Hossini and Rage 2000). Villeveyrac, France (Buffetaut et al. 1996); MEoc (MP13) Geiseltal near Halle, Germany (Wuttke et al. 2012); LEoc Palaeobatrachus aff. P. robustus Kučlín near Bílina, Czech Republic (Wuttke et al. 2012).

Stratigraphy and distribution: LOlig (MP30) Rhinophrynidae Günther, 1858 Oberleichtersbach, Germany (Böhme 2008). Diagnosis (after Henrici 1991, slightly modified): Palaeobatrachus tobieni (Wuttke in Sanchiz, 1998a) Supraorbital flange present; premaxilla and maxilla edentate; palatine process of premaxilla absent; palatines absent; Diagnosis (Wuttke et al. 2012): Sphenethmoid partially parahyoid bone present; eight opisthocoelous presacral co-ossified with prootics; small process on prootic artic- vertebrae with imbricate neural arches; atlantal cotyles closely ulated with medial ramus of pterygoid; frontoparietal juxtaposed; atlantal neural arch elongate; transverse process of unsculptured, broader posteriorly than anteriorly, not V2 hook shaped; free ribs absent; sacral diapophyses expand- divided into left and right frontoparietal, dorsal surface ed; sacro-urostylar articulation bicondylar; pectoral girdle without parasagittal crests; vertebral complex V1+V2 arciferal; clavicle overlying anterior margin of scapula; tibiale bears diapophyses only on V2; coracoid without hook- and fibulare fused; prehallux and distal phalanx of first digit like processus rostriformis; diapophyses on V7 and V8 modified as bony digging spades. vestigial or absent, so diapophyses of synsacrum are formed only by diapophyses of sacral vertebra pro- Chelomophrynus Henrici, 1991 per (V9); cubital fossa of humerus relatively deep; Chelomophrynus bayi Henrici, 1991 distal part of forelimb relatively short, so metacarpal IV is about two-thirds of ulna; F and TF about equal in Diagnosis (Henrici 1991): Alary process of premaxilla broad length. and triangular; maxilla with anterior notch; vomer with elon- Stratigraphy and distribution: MEoc (Lower Geiseltalium) gate posterior process and elongate postchoanal ramus; “Grube Messel” near Darmstadt, southern Hessen, Germany diapophyses of last three presacrals laterally-oriented; caput (Wuttke et al. 2012). humeri round; atlantal neural arch tapers posteriorly; well- Remarks: This species was originally assigned to the developed lateral crest on humerus. monotypic genus Messelobatrachus, which was infor- Stratigraphy and distribution: MEoc (Uintan) Hot Springs mally erected in a PhD thesis by Wuttke (1988). Direct County, Wyoming, USA (Henrici 1991; Henrici and Fiorillo comparisons with the Eocene Palaeobatrachidae from 1993). Geiseltal and Kučlín revealed that all fit well into the range of variation seen in the Oligocene palaeobatrachids Eorhinophrynus Hecht, 1959 andeveninthegenusPalaeobatrachus (Wuttke et al. Eorhinophrynus septentrionalis Hecht, 1959 2012). Diagnosis (after Henrici 1991): Differs from all Palaeobatrachus sp. rhinophrynids in having atlantal neural arch laterally broad with square posterior margin; differs from Eorhinophrynus Stratigraphy and distribution: LPaleo (Thanetian, MP6) sp. in having caput humeri round; differs from Rhinophrynus Cernay, France (Vergnaud-Grazzini and Hoffstetter 1972); in having well-developed lateral crest on humerus. EOlig Möhren 13 near Treuchtlingen, Germany (Sanchiz Stratigraphy and distribution: MEoc (Bridgerian) et al. 1993); LOlig Enspel, Germany (Roček and Wuttke Tabernacle Butte, Wyoming, USA (Hecht 1959). Author's personal copy

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Remarks: The taxon was based on a single atlas vertebra. sacrum monovertebral, with sacral diapophyses slightly expanded; sacro-urostylar articulation bicondylar; meta- Eorhinophrynus sp. carpals exceeding 75 % the length of the radioulna; parahyoid bone; scapular cleft tiny; coracoid lacking a Stratigraphy and distribution: LPaleo Big Horn Basin, conspicuous rostral process and having a slightly ex- Wyoming, USA (Estes 1975). panded sternal end. Stratigraphy and distribution: ECret (late Berriasian– Rhinophrynus Duméril et Bibron, 1841 early Valanginian) Santa Maria de Meià, Spain (Báez and Sanchiz 2007). Diagnosis (after Henrici 1991): Lateral crest on humerus poorly developed; shaft of femur strongly bowed and narrow Pipoidea indet. directly proximal to distal condyle; distal condyle of femur longer than wide. Stratigraphy and distribution: LCret (Maastrichtian) or EPaleoc Bug Creek Anthills, Montana USA (Gardner 2008).

Rhinophrynus canadensis Holman, 1963 Remarks: The fused atlas and V2 was originally described by Gardner (2008) as belonging to Palaeobatrachus? Diagnosis (Henrici 1991;Holman2003): Ilium with en- occidentalis, but Wuttke et al. (2012) suggested the specimen tire dorsal acetabular region rotated forward so that it is be relocated to the Pipidae or another pipoid family on the almost perpendicular to the iliac shaft; dorsal prominence basis of the opisthocoelous shape of its centrum. of ilium well developed, almost dorsoventrally located and produced laterally rather than dorsally from the Pelobatoidea Bonaparte, 1850 shaft; caput humeri round rather than oval; space between tibiale and fibulare forming proximodistally elon- Diagnosis (mainly after Duellman and Trueb 1994): gate oval. Procoelous vertebrae with imbricate neural arches; atlantal Stratigraphy and distribution: EOlig, north branch of Calf cotyles closely juxtaposed; sacral diapophyses dilated; sacro- Creek near Eastend, Saskatchewan, Canada (Henrici 1991; urostylar articulation bicondylar; first postsacral vertebra

Holman 1963). (V10) bears rod-like or dilated transverse processes similar to those of the sacral diapophyses, which fuse to sacral ?Rhinophrynidae diapophyses by means of bony webbing in fully grown adults Rhadinosteus Henrici, 1998 (not known in Megophryidae and Pelodytidae); pectoral gir- Rhadinosteus parvus Henrici, 1998 dle arciferal; omosternum cartilaginous; anterior end of scapula not overlain by clavicle; two tarsalia. Diagnosis (Henrici 1998a): Frontoparietal azygous; parasphenoid without lateral alae; vertebrae ectochordal, Pelobatidae Bonaparte, 1850 amphicoelous, pierced by persistent notochord; teeth bicuspid, pedicellate; hind limb bones slender; femur Diagnosis (partly after Rage and Hossini 2000): Dermal sigmoid. roofing bones always covered with sculpture, which is an Stratigraphy and distribution: LJur (Kimmeridgian) integral part of the bone, not a secondary exostosis like in, Dinosaur National Monument DNM 96, Utah USA for example, Latonia or tropical hylids; in metamorphosed (Henrici 1998a). individuals, frontoparietal is azygous (at least in posterior part) with convex posterior margin, and in larvae it is tripartite Pipoidea indet. (consisting of a pair of frontoparietals plus one unpaired element inserted posteriorly between them; Roček 1982,figs. Neusibatrachus Seiffert, 1972 42, 43, 44, 50; Maus and Wuttke 2004, figs. 3, 4); Neusibatrachus wilferti Seiffert, 1972 quadratojugal is present; parasphenoid has a median keel and posterior median convexity; palatine process of maxilla Diagnosis (Báez and Sanchiz 2007): Frontoparietal azy- is elongate (=palatine fused to maxilla; Roček 1981,fig.59a– gous, forming a flat dorsal table; maxillary arch com- b; Lebedkina 2004, fig. 95a), postchoanal ramus of vomer plete; teeth pedicellate; squamosal T-shaped; articulation absent; pterygoid with moderately developed ventral flange for the lower jaw at the midlevel of the otic capsules; on its lateral margin; sternum ossified, with elongated stylus; parasphenoid lacking lateral alae; nine presacral verte- caput humeri shifted laterally, fossa cubitalis open laterally; brae with procoelous centra; V1 and V2 fused together; ilium without dorsal tubercle, iliac shaft without dorsal crest. Author's personal copy

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Eopelobates Parker, 1929 MMio (Badenian) Devínska Nová Ves–Bonanza, Slovakia (Hodrová 1988). Diagnosis (partly after Estes 1970; Špinar and Roček 1984): Sculpture on dermal roofing bones is exclusively of pit-and- Eopelobates cf. E. bayeri ridge type; anterolateral margin of nasals straight along the whole of its length; sphenethmoid exposed between posterior Stratigraphy and distribution: Plio (Csarnotan) Ivanovce, part of nasals and anterior ends of frontoparietals; Slovakia (Hodrová 1981); Plio Gorishnaya Vygnanka, frontoparietal not in contact with squamosal; urostyle long, Ukraine (K.A. Tatarinov in Chkhikvadze 1981, 1984). may reach the level of acetabular portion of pelvis; strong posterior projection of the ischium (difficult to quantify, but it Eopelobates grandis Zweifel, 1956 seems that the ischium is more prominent posteriorly than in Pelobates); tibiale and fibulare not fused; spade absent (no Diagnosis (Henrici 2002): Dermal sculpture highly tubercu- fossoriality). TF longer than or equal to F. lated; frontoparietal broad, in which the width is about 84 % of the length; otic process of the squamosal rounded. Eopelobates anthracinus Parker, 1929 Stratigraphy and distribution: LEoc, Ahern Member of the Chadron Formation, Pennington County, South Dakota, USA Diagnosis (after Estes 1970; Špinar and Roček 1984;personal (Zweifel 1956). observation): Pars facialis of the premaxilla almost in the Remarks: Assignment to Eopelobates remains doubtful. The midlength of the horizontal part of the bone, medial margin urostyle with a pair of transverse processes and strongly of the pars facialis widely convex; nasals are separated from arched ilium with the dorsal protuberance and dorsal crest each other, only their posterolateral part is covered with sculp- exclude E. grandis from the Pelobatidae and the tubercular ture; sculpture on frontoparietal is better defined only along sculpture from Eopelobates. the lateral edges of the bone; lamella alaris squamosi tapers anteriorly in a point, its orbital margin is shallowly concave, Eopelobates hinschei (Kuhn, 1941) contact margin with the zygomaticomaxillar process of the maxilla is not well defined, anterior and posterior parts of the Diagnosis (after Estes 1970; Wuttke 1988): Nasals in contact alar lamella are smooth; urostyle is articulated with the sacral with each other by a median suture at least in anterior third of vertebra; carpus and tarsus not ossified. their medial margin (based on GMH 6692; Estes 1970,fig. Stratigraphy and distribution: LOlig (MP 30) Rott, 12A), covered with pit-and-ridge sculpture except for a nar- Germany (Koenigswald et al. 1992). row area rimming the orbital margin; frontoparietals covered Remarks: Špinar and Roček (1984) suggested that E . with sculpture, pit-and-ridge posteriorly, consisting of grooves anthracinus is a juvenile of E. bayeri, but recent examination oriented anteriorly in its anterior part; posterior part of lamella of the type material of both species revealed that they repre- alaris squamosi thin and rather bent ventrally (GMH 6753; sent adults distinguishable by several diagnostic characters Estes 1970, fig. 20b); posterior margin of the scapula deeply (see diagnoses). concave, anterior margin only slightly concave in its dorsolateral section (GMH 6692; Estes 1970,fig.9e). Eopelobates bayeri Špinar, 1952 (Fig. 6) Stratigraphy and distribution: MEoc (Lower Geiseltalium) Geiseltal near Halle, Germany (Franzen and Haubold 1986). Diagnosis (based on personal observation): Pars facialis close Remarks: Pelobatids from the middle Eocene of Geiseltal to the medial end of the horizontal part of the bone; medial were reviewed by Estes (1970) and mostly referred to margin of the pars facialis premaxillae forms a rounded out- Eopelobates as E. hinschei (Kuhn, 1941). It is difficult to growth; nasals are in a long contact by median suture, whole understand why Estes (1970) choose Halleobatrachus nasals are covered with sculpture; sculpture on frontoparietal hinschei as the holotype of E. hinschei, a poorly preserved is well developed, except along the midline; lamella alaris specimen in which most diagnostic characters of the genus squamosi rounded anteriorly, its orbital margin straight, con- (including those listed by Estes 1970: p. 295) are not tact margin with the zygomaticomaxillar process of the max- recognisable, whereas he did not consider better preserved illa is well defined, whole lamella covered with sculpture; specimens referred by him to E. hinschei,suchasholotype urostyle is coalesced to sacral vertebra. of Archaeopelobates eusculptus.Kuhn(1941:p.371)provid- Stratigraphy and distribution: EOlig (MP 21) Hoogbutsel, ed only a very vague diagnosis of Halleobatrachus hinschei, Hoeleden, and Boutersem TGV sites, Belgium (Smith 2003); which does not even allow the decision to be made on whether LOlig Bechlejovice Czech Republic (Bellon et al. 1998; the specimen belongs to the Pelobatidae or Palaeobatrachidae. Špinar 1972); EMio Hrabák mine near Most and Nástup and Estes (1970) based his reassignment of Halleobatrachus to Merkur mines near Kadaň, Czech Republic (Špinar 1972); Eopelobates on general similarities with E. bayeri,stating Author's personal copy

Palaeobio Palaeoenv (2013) 93:397–439 419 that only the squamosals of E. hinschei and E. bayeri are Stratigraphy and distribution: MEoc (Lower Geiseltalium) different and that there are proportional differences of their Grube Messel near Darmstadt, Hessen, Germany (Wuttke nasals. At that time, E. wagneri was still considered a 2012a). pelodytid (see below), so no other Eocene Eopelobates was Remarks: It was considered a member of an indeterminate available for comparison. Similar to E. wagneri from the family, probably a palaeobatrachid (Sanchiz 1998a). Judging middle Eocene of Messel. Relations of these two species need by separate quadratojugal and quadrate, signs of segmented to be clarified. urostyle and by the absence of epiphyses of the humeri and radioulnae, it is obviously a juvenile. The structure of the Eopelobates cf. E. hinschei frontoparietal (with a posterior, unpaired element) suggests relations to the Pelobatidae. Stratigraphy and distribution:LEocHordleCliff,England (Milner et al. 1982). Pelobates Wagler, 1830

Eopelobates wagneri (Weitzel, 1938) Diagnosis (after Rage and Hossini 2000): Sculpture on dermal roofing bones tubercular; anterolateral margin of Diagnosis (based on data from Wuttke 2012b): Nasals widely nasals concave in its antero-medial section and convex in separated; frontoparietal covered with sculpture posteriorly, its postero-lateral section; presacral vertebrae with long smooth anteriorly; its posterior part is azygous, whereas both neural arches, and condyle and cotyle circular in shape; frontal portions are separated by median suture; palatines urostyle without dorsal crest; ilium without dorsal tubercle, either free or fused to maxillae; margin between ramus iliac shaft without dorsal crest, supra- and preacetabular maxillaris and ramus posterior of pterygoid is extended fossae absent; caput humeri shifted laterally, fossa cubitalis ventrally (“ventral flange”); scapula concave both anteriorly open laterally. and posteriorly; urostyle articulated (monocondylar articulation), not ankylosed, to sacrum. Pelobates decheni Troschel, 1861 Stratigraphy and distribution: MEoc (Lower Geiseltalium) Grube Messel near Darmstadt, Hessen, Germany (Wuttke Diagnosis (Böhme et al. 1982;Roček and Wuttke 2010): 2012b). Postorbital bridge wide, squamoso-frontoparietal suture may Remarks:OriginallyassignedtogenusPropelodytes attain up to 50 % of the frontoparietal length; quadratojugal (Weitzel 1938). Transferred to Eopelobates by Wuttke (1988). sculptured, not coalesced to the quadrate; urostyle articulated, not firmly fused, to sacrum; praehallux not modified to form a Eopelobates sp. spade for digging. Stratigraphy and distribution: LOlig (MP 30) Rott, Stratigraphy and distribution: LOlig (MP30) Germany (Koenigswald et al. 1992). Oberleichtersbach, Germany (Böhme 2008); EMio Sandelzhausen, Germany (Böhme 2010); EMio (Orleanian, Pelobates cf. decheni MN4) Dolnice, Czech Republic (Hodrová 1987); Plio (MN 14) Osztramos 1, Hungary (Venczel 2001); Plio (Csarnotan) Stratigraphy and distribution: LOlig Enspel, Germany Ivanovce, Slovakia (Hodrová 1981). (Roček and Wuttke 2010).

Lutetiobatrachus Wuttke in Sanchiz, 1998a Pelobates fahlbuschi Böhme, 2010 Lutetiobatrachus gracilis Wuttke in Sanchiz, 1998a Diagnosis (Böhme 2010): Highly prominent pit-and-ridge Diagnosis (after Wuttke 2012a, simplified): Frontoparietal sculpture on frontoparietal, squamosal and maxilla; tectum not sculptured, broader anteriorly than posteriorly, with its supraorbitale on frontoparietal broad; deep groove for frontal portions separated by medial suture, azygous posteri- arteria orbitonasalis on ventral side of the tectum orly; nasals large, triangular, separated by a long median supraorbitale; posterior margin of the frontoparietal suture; quadratojugal present; eight presacrals with imbricate incrassation deeply concave; foramen arteriae occipitalis neural arches; transverse processes of V5 short and perpendic- lying medial to the paraoccipital process and visible from ular to body axis, those of V6–V8 long and inclined anteriorly; the dorsal side; lamella alaris squamosi broad, sacral diapophyses broadly dilated; urostyle long, with possi- posteroventral process of squamosal prominent; ventral bly vestigial V10 and V11; ilia strongly bowed, without dorsal border of zygomatic process and lamellae alaris nearly tubercle and not coalesced to the ischium; femur shorter than perpendicular to each other; zygomatic process slender; the tibiofibula. squamoso-frontoparietal suture being one-fifth to one- Author's personal copy

420 Palaeobio Palaeoenv (2013) 93:397–439 quarter of the inferred frontoparietal length; anterior border Scaphiopodidae Cope, 1865 of pars descendens of ilium straight; shallow groove on dorsolateral edge of pars ascendens ilii; tuber superius Diagnosis (based on data in Roček 1981): Frontoparietals between this groove and the spiral groove rough, weak paired (thus the frontoparietal is not of tripartite origin, as is and oval; spinal nerve foramen in presacral vertebrae pres- the case with Pelobates and Eopelobates). Scaphiopus-Spea ent; neural spine missing in posterior presacral vertebrae. (= Scaphiopodidae) clade recognised by Henrici (1994)is Stratigraphy and distribution: EMio Sandelzhausen, diagnosed as follows (from Holman 2003): quadratojugal Germany (Böhme 2010). absent; postchoanal ramus of vomer elongate (= palatine fused to vomer; see Roček 1981, fig. 59); medial end of coracoid not Pelobates fuscus (Laurenti, 1768) expanded; sternum cartilaginous; ischium not extended posteriorly; sartorius muscle and associated tendon concealed, in Diagnosis (based on data from Roček 1981): Postorbital part, by gracilis major. Further characters are a well-developed bridge absent, no squamoso-frontoparietal contact. columella and ossified operculum. Stratigraphy and distribution: Plio (Ruscinian) Europe (Sanchiz 1998a). Macropelobates Noble, 1924

Pelobates sanchizi Venczel, 2004 Diagnosis (Noble 1924;Roček et al. 2011): Pit-and-ridge ornamentation on skull roof; frontoparietals paired; vertebral Diagnosis (after Venczel 2004): Large size; highly prominent centra procoelous; sacral diapophyses widely expanded, with pit-and-ridge sculpture; thin postorbital arch. a greater length than width; urostyle not ankylosed to sacral Stratigraphy and distribution: EMio (MN4) Mokrá- vertebra, articulated by a single condyle; tibiale and fibulare Western Quarry, Czech Republic (Ivanov 2008); MMio not fused with one another; prehallux modified as bony spade. (Badenian, MN6) Mátraszőlős 1, Hungary (Venczel 2004); Remarks: Comparative analysis suggested closer relations of MMio (Badenian, MN6) Mátraszőlős 2, Hungary (Venczel Macropelobates to Scaphiopus, rather than to Eopelobates 2004); MMio (Badenian, MN6) Sámsonháza 3, Hungary and Pelobates (Roček 1982). (Venczel 2004). Macropelobates linquensis (Yang, 1977) Pelobates sp. Diagnosis (Roček et al. 2011): Skull short and broad; robust Stratigraphy and distribution: LOlig (MP30) spade present; TF shorter than F; tibiale and fibulare as well as Oberleichtersbach, Germany (Böhme 2008); MMio (Astaracian, whole hindlimbs are short. MN6) Sansan, France (Rage and Hossini 2000); MMio Stratigraphy and distribution: MMio Shanwang, Shandong (Sarmatian) Tauţ, Romania (Venczel and Sţiucă 2008); Plio Province, China (Roček et al. 2011). (Villanyian) Včeláre, Slovakia (Hodrová 1985). Remarks: Originally described as Macropelobates cratus by Gao (1986); for details of nomenclatural history see Roček Pelobatidae indet. Larvae et al. (2011).

Tadpoles: Large size (up to 145 mm); tripartite frontoparietal Macropelobates osborni Noble, 1924 complex; frontoparietals sculptured; median ridge on ventral surface of cultriform process of parasphenoid. Diagnosis (compiled from Noble 1924,Roček 1982): Stratigraphy and distribution: LOlig Enspel (Maus and Frontoparietal with a small postero-lateral processes; trans-

Wuttke 2002, 2004;Roček and Wuttke 2010). verse processes of V7 and V8 moderately directed anteriorly; scapula long and having anterior crista; coracoids widened Pelobatidae indet. adults distally; ischium extending posteriorly greatly beyond the iliac pars ascendens; hindlimbs relatively long; pubis ossified. Stratigraphy and distribution: LCret, Asifabad, India Stratigraphy and distribution: EOlig Loh campsite, (Prasad and Sahni 1987, 2009); LCret (Campanian) Ellisdale, Mongolia (Noble 1924). New Jersey, USA (Denton and O’Neill 1998); EEoc (MP 7) Le Remarks: The otic ramus of the squamosal is not observable Quesnoy, France (Nel et al. 1999); EEoc (MP7) Silveirinha, in M. osborni; its listing among diagnostic characters of the Portugal (Rage and Auge 2003); MMio (Sarmatian) Karpov species by Sanchiz (1998a) was probably caused by confusion Yar, Moldova (Skutschas and Bannikov 2009); LMio with exposed posterolateral process of the squamosal (see (Vallesian, MN9) Rudabánya, Hungary (Roček 2005). Roček 1982). Author's personal copy

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Scaphiopus Holbrook, 1836 pterygoid process; prootic foramen widely emarginated; length of sacral diapophyses at their ends equal to the Diagnosis (personal observation): Scaphiopus contains well- length of three presacral vertebrae. ossified species, so some diagnostic characters are opposites Stratigraphy and distribution: MOlig (Orellan) Leo Fitterer to those listed in the diagnosis (see below) for Spea. Ranch, North Dakota, USA (Estes 1970).

Scaphiopus alexanderi Zweifel, 1956 Scaphiopus studeri Taylor, 1938

Diagnosis (Zweifel 1956;Sanchiz1998a): V9 (sacral) fused Diagnosis (Taylor 1938): Broad fontanelle between to V8 (presacral); preacetabular foramen on the ilium absent. frontoparietals; sphenethmoid with a rounded anterior Stratigraphy and distribution: Mio (Hemphillian?) Fish knoblike termination; sacral cotyle oval shaped; prehallux Lake, Nevada, USA (Sanchiz 1998a). spade relatively large. Stratigraphy and distribution: Mio (Hemphillian) Rhino Scaphiopus guthriei (Estes, 1970) Hill, Kansas, USA (Sanchiz 1998a).

Diagnosis (based on data from Estes 1970): Skull relatively Scaphiopus wardorum Estes et Tihen, 1964 short; median suture between frontoparietals runs up to posterior edge of the bones because there is no posterior median Diagnosis (Estes and Tihen 1964): Maxillary ornamentation element in the larval development (one of principal diagnostic of numerous separate denticles; ilium with an extremely nar- features of the Pelobatidae). row pars descendens anterior to the acetabulum, with a small Stratigraphy and distribution: EEoc (Lysitean) Fremont ridge-like tuber superius above the acetabulum. County, Wyoming, USA (Henrici 2000). Stratigraphy and distribution:MMio(Barstovian)Norden Remarks: Henrici (2000), and subsequently also Holman Bridge Quarry, Nebraska, USA (Sanchiz 1998a). (2003), removed this species from Eopelobates and assigned it to Scaphiopus. Scaphiopus sp.

Scaphiopus hardeni Holman, 1975 Stratigraphy and distribution: LOlig (Whitneyan) I-75 near Gainsville, Florida, USA (Patton 1969); MMio (Barstovian) Diagnosis (Holman 1975, 2003): Sphenethmoid with a strong Fort Polk, Texas, USA (Williams 2002). posterior median tubercle and with lateral processes at right angles to the longitudinal axis; ilium with well-developed Spea Cope, 1866 rather than an obsolete ventral acetabular expansion, dorsal prominence well-developed and rugged; dorsal part of the Diagnosis (based on data in Roček 1981): Due to overall ventral acetabular area less extensive; ventral acetabular ex- hypoossification, nasals and frontoparietals are reduced pansion narrow and deflected posteriorly, acetabulum rounded in size, thus sphenethmoid is exposed in frontoparietal rather than ovoid; orbital border of maxilla shallowly concave fontanelle in average-sized individuals; for the same and lacking extruded orbital rim. reason, frontoparietals lack ornamentation, lamella Stratigraphy and distribution: LMio (Clarendonian) alaris squamosi does not contact maxilla, and contact Wakeeney, Kansas, USA (Holman 2003). between the ramus interior of pterygoid and the prootic is weak. Scaphiopus holbrooki (Harlan, 1835) cf. Spea hammondii (Baird, 1859) Diagnosis: Living species, osteological diagnosis not available. Stratigraphy and distribution: Plio (Blancan) Inglis 1A, Stratigraphy and distribution: Plio (Blancan) Hagerman Florida, USA (Meylan 2005). area, Idaho, USA (Chantell 1970).

Scaphiopus skinneri Estes, 1970 Spea neuter (Kluge, 1966)

Diagnosis (after Sanchiz 1998a): Extensive skull orna- Diagnosis (after Henrici 2009; see also Sanchiz 1998a): mentation; absence of an open frontoparietal fontanelle; Sculpturing present on frontoparietal and maxilla; large large rounded tympanic process of the squamosal; pres- frontoparietal fontanelle; maxilla without pterygoid process; ence of squamoso-maxillary contact; maxilla with quadratojugal absent; urostyle fused to sacrum; maxilla and Author's personal copy

422 Palaeobio Palaeoenv (2013) 93:397–439 squamosal widely separated; dorsal protuberance of ilium Pelodytidae Bonaparte, 1850 large and prominent. Stratigraphy and distribution: LOlig (Arikareean) Diagnosis (after Henrici 1994; Rage and Hossini 2000; Wounded Knee area, South Dakota, USA (Holman 2003); Sanchiz et al. 2002): Eight presacral vertebrae with EMio (Arikareean) McCann Canyon, Nebraska, USA; condyle and cotyle compressed dorsoventrally; neural

MMio (early Barstovian) Anceney, Montana, USA (Henrici arches imbricate; atlas and V2 often fused with each 2009). other; atlantal cotyles closely juxtaposed; sacral diapophyses widely expanded anteroposteriorly; urostyle Spea indet. without dorsal crest, but with a pair of convexities anteriorly; iliac shaft without dorsal crest and dorsal Stratigraphy and distribution: LMio (Clarendonian), tubercle; tibiale and fibulare fused along whole their Whisenhunt Quarry, Oklahoma, USA (Parmley et al. 2010). length.

Elkobatrachus Henrici et Haynes, 2006 Miopelodytes Taylor, 1941 Elkobatrachus brocki Henrici et Haynes, 2006 Miopelodytes gilmorei Taylor, 1941

Diagnosis (Henrici and Haynes 2006): Premaxilla with Diagnosis (based on data from Henrici 1994): broad-based alary process that distally forms a laterally Frontoparietal paired, widely separated (especially in deflected, straight blade whose transverse width lies in a their frontal portion); alar lamella of squamosal bearing near parasagittal plane; frontoparietal fontanelle exposed sculpture; transverse processes of the last two presacrals dorsally; occipital canal roofed by bone and exits onto strongly inclined anteriorly; sacral diapophyses less ex- dorsal surface of frontoparietal; pars acromialis of scap- panded than in Pelodytes; scapula short; ischium does ula triangular, its anteriorly directed apex is positioned not extend posteriorly beyond the dorsal acetabular ex- at the level of the dorsal rim of the glenoid fossa; pansion of the ilium. urostyle longer than the remainder of vertebral column; Stratigraphy and distribution: MEoc Elko Formation, posteromedial process of parasphenoid has concave dis- Nevada, USA (Henrici and Haynes 2006). tal margin; nasal is triangular in dorsal aspect; clavicle anteriorly overlaps scapula; sculpture on dermal bones Pelodytes Bonaparte, 1838 absent; supraorbital flange absent. Stratigraphy and distribution:MEocElkoHills,Nevada, Diagnosis (Sanchiz et al. 2002): Columella present and ossi- USA (Henrici and Haynes 2006). fied; sacro-urostylar articulation monocondylar. Remarks: Elkobatrachus specimens do not represent fully ossified adults (Henrici and Haynes 2006). Pelodytes arevacus Sanchiz, 1978

Uldzinia Gubin, 1996 Diagnosis (Sanchiz 1978, 1998a): Atlas not fused with V2; Uldzinia kurochkini Gubin, 1996 sacral diapophyses strongly dilated anteroposteriorly; sacro-urostylar articulation intermediate between mono- Diagnosis (modified after Gubin 1996): Maxilla covered by and bicondylar; hypochordal process that contacts the sculpture posteriorly from the middle of the antero-posterior sacral vertebra persisting at least up to the end of diameter of the orbital margin (anterior part of the bone metamorphosis; short scapula with anterior crista and smooth); processus zygomaticomaxillaris slightly less promi- tuberosity of moderate size; tibiale and fibulare robust nent than processus frontalis; posterior margin of processus and fused. zygomaticomaxillaris vertical; processus posterior short; tooth Stratigraphy and distribution:MMio(Astaracian,MN7-8) row terminates anterior to processus zygomaticomaxillaris. Escobosa de Calatañazor, Spain (Sanchiz 1978). Stratigraphy and distribution:EOligErgiliyn-Dzo, Mongolia (Gubin 1996). Pelodytes cf. P. arevacus aff. Uldzinia Stratigraphy and distribution: EMio (Agenian, MN2b) Navarette del Rio, Spain; MMio (Astaracian, MN7-8) Cerro Stratigraphy and distribution:EOligTatalGol,Mongolia del Otero, Spain; MMio (Vallesian, MN9) Autilla 1, Spain (all (Böhme 2007). from Sanchiz 1998a). Author's personal copy

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Pelodytes cf. P. punctatus Daudin, 1802 anteriorly; sacro-urostylar articulation bicondylar; praepollex and praehallux present; F and TF of about same length; tibiale Stratigraphy and distribution: Plio (MN7-8) Sète, France and fibulare not coalesced. (Bailon 1991). Stratigraphy and distribution:EocWorliHillnearBombay, India (Špinar and Hodrová 1985; Prasad and Sahni 2009). Pelodytes sp. or cf. Pelodytes Leptodactylidae Werner, 1896 Stratigraphy and distribution: LEoc (MP 16, Bartonian) Bretou, France (Rage 1988; Duffaud 2000); EMio (MN 4) Diagnosis (after Duellman and Trueb 1994): Palatines pres- Oberdorf, Austria (Sanchiz 1998b); MMio (Astaracian, MN6) ent; parahyoid absent; eight presacral vertebrae with Sansan, France (Rage and Hossini 2000); LMio (Vallesian, procoelous centra; sacrum with rounded diapophyses; sacro- MN9) Rudabánya, Hungary (Roček 2005). urostylar articulation bicondylar; urostyle without transverse processes; pectoral girdle arciferal; omosternum and sternum Pelodytidae indet. cartilaginous; anterior end of scapula not overlain by clavicle.

Stratigraphy and distribution: MEoc (MP13) Saint Thaumastosaurus De Stefano, 1903 Maximin, France (Duffaud and Rage 1997). Diagnosis (based on data from Roček and Lamaud 1995): Tephrodytes Henrici, 1994 Skull hyperossifed, dermal bones (frontoparietals, nasals, Tephrodytes brassicarvalis Henrici, 1994 parasphenoid) fused not only with each other, but also with braincase. Diagnosis (Henrici 1994): Frontoparietals are in contact Remarks: A new cladistic analysis including postcranial along the whole of their length; both frontoparietals characters reveals ranoid affinities (Laloy et al. 2013). produce posteriorly a moderate median tip; otic plate of squamosal present; atlas not fused with V2;neuralarches Thaumastosaurus bottii De Stefano, 1903 of presacral vertebrae elongate; anterior lamina of scapula absent; sacral diapophyses strongly expanded. Diagnosis (Roček and Lamaud 1995): Dermal roofing Stratigraphy and distribution: EMio (Arikareean) Flint bones heavily sculptured with pit-and-ridge sculpture; na- Creek Basin, Montana, USA (Henrici 1994). sals and frontoparietals of the left and right side co-ossified in large individuals; frontoparietals and nasals in contact, Hyloidea Stannius, 1856 sphenethmoid exposed even in large individuals; frontoparietals and nasals co-ossified with sphenethmoid Diagnosis: Arciferal pectoral girdles (coracoids not fused), and prooticooccipitals in large individuals; one foramen in procoelous vertebrae (Frost et al. 2006). posterior wall of orbit; frontoparietal with deep vertical marginal cristae forming pars contacta; frontoparietal Myobatrachidae Schlegel, 1850 incrassation paired, its posterior part may be completely enclosed into ossified tectum synoticum; foramina pro Diagnosis (Frost et al. 2006): Notochordal (ectochordal) ver- arteriae occipitales present; frontoparietal not in contact tebrae with intervertebral discs; vomers reduced with concom- with squamosal; spine on either side of posterior surface itant reduction of vomerine teeth. of prooticooccipital, medial to prominentiae ducti semicircularis posterior; the latter compressed laterally Indobatrachus Noble 1930 and confluent with processus paraoccipitales of Indobatrachus pusillus (Owen, 1847) (Fig. 4) frontoparietals; maxillae toothed, not fused with quadratojugals; palatines present, anterior tip of Diagnosis (slightly modified after Noble 1930,fromŠpinar parasphenoid not extending between them. and Hodrová 1985): Very small frog (SVL about 20 mm); Stratigraphy and distribution: LEoc (Lower Ludian, MP skull broader than long; sphenethmoid largely cartilaginous; 17) La Bouffie, Lot, France; LEoc (Marinesian through broad anterior fontanelle between frontoparietals; palatines Upper Ludian) Perrière, Sainte-Néboule, Coânac 1, possibly present; parasphenoid with narrow lateral alae direct- Escamps, Rosières 2, Sindou D, perhaps also Aubrelong 2 ed posterolaterally; eight procoelous presacral vertebrae; sa- and Malperie, Quercy, France (all from Roček and Lamaud cral diapophyses dilated towards their ends, slightly inclined 1995). Author's personal copy

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Thaumastosaurus gezei Rage et Roček, 2007 Hylidae Rafinesque, 1815

Diagnosis (Rage and Roček 2007): Anterior end of Diagnosis (simplified from Holman 2003): Eight procoelous lamella alaris of squamosal extends anteriorly along presacrals with non-imbricate neural arches; atlantal cotyles dorsal edge of maxilla, thereby separating maxilla from widely separated; sacral diapophyses dilated; sacro-urostylar orbit and nasal; dermal sculpture consists of pits and articulation bicondylar; urostyle without transverse processes; ridges, without any tubercles; two foramina in posterior omosternum and sternum cartilaginous; clavicle does not over- wall of orbit; medial face of lamina horizontalis of lap scapula anteriorly. maxilla convex (instead of flat); base of ramus paroticus of squamosal straight, not curved. Acris Duméril et Bibron, 1841 Stratigraphy and distribution: MEoc or LEoc Quercy, France (Rage and Roček 2007; Laloy et al. 2013). Diagnosis: Living genus, osteological diagnosis not available.

Thaumastosaurus sulcatus Holman et Harrison, 2003 Acris barbouri Holman, 1967

Diagnosis (Holman and Harrison 2003): Sculpture of maxilla Diagnosis (Holman 2003): Iliac shaft almost straight, with and alar lamella of squamosal composed of elongate, ridge- low dorsal crest gently arising from anterior border of dorsal bordered sulci; lamina horizontalis of maxilla thin and sharply prominence; dorsal tubercle rounded and relatively smooth. bowed upward through its length; contact area for premaxilla Stratigraphy and distribution: Mio (Hemingfordian) extensive; tooth crowns relatively narrow and pointed and Thomas Farm, Gilchrist County, Florida, USA (Holman 2003). with auburn pigmentation; dorsal portion of ramus paroticus of lamella alaris squamosi arising more posteriorly than in T. Acris crepitans Baird, 1854 bottii and T. wardi; ventral border of lamella alaris squamosi less concave. Diagnosis: Living species, osteological diagnosis not Stratigraphy and distribution: LEoc (Headonian, available. MP 17) Hordle Cliff near Mildford-on-Sea, UK (Holman Stratigraphy and distribution: Plio (Blancan) Beck Ranch, and Harrison 2003). Scurry County, Texas (Rogers 1976).

Thaumastosaurus wardi Holman et Harrison, 2002 Acris sp. Duméril et Bibron, 1841

Diagnosis (Holman and Harrison 2002): Lamina horizontalis Stratigraphy and distribution: MMio (Barstovian) Fort maxillae flat rather than rounded in medial view; ridge Polk, Texas, USA (Williams 2002); LMio (Clarendonian), separating fossa maxillaris from suborbital part of maxilla Whisenhunt Quarry, Oklahoma, USA (Parmley et al. 2010). directed anteriorly rather than posteriorly; teeth larger and fewer in number (if compared with T. bottii); left squa- Hyla Laurenti, 1768 mosal with much larger and deeply excavated pits especially on the processus zygomaticus; orbital margin of Diagnosis: Living genus, osteological diagnosis not squamosal evenly curved and hemispherical; squamosal available. in medial view with ramus paroticus hemispherical rather than sinusoidal. Hyla cf. arborea Laurenti, 1768 Stratigraphy and distribution: LEoc (MP 17) Hatherwood Point, Isle of Wight, England (Holman and Harrison Stratigraphy and distribution: MMio (Badenian, MN6) 2002). Mátraszőlős 1, Hungary (Venczel 2004); MMio (Badenian, MN6) Mátraszőlős 2, Hungary (Venczel 2004); MMio Leptodactylidae indet. (Badenian, MN7-8) Felsötárkány 1, Hungary (Venczel 2004); MMio (Sarmatian) Tauţ, Romania (Venczel and Stratigraphy and distribution: EEoc (Ypresian) Vastan Sţiucă 2008); LPlio (Csarnotan) Ivanovce, Slovakia Lignite Mine, Gujarat, India (Bajpai and Kapur 2008). (Hodrová 1981).

?Leptodactylidae or Hemisotidae Hyla cinerea (Schneider, 1799)

Stratigraphy and distribution: LCret Naskal, India (Prasad Diagnosis: Living species, osteological diagnosis not and Sahni 2009). available. Author's personal copy

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Stratigraphy and distribution: LPlio (Blancan) Inglis 1A, (Hodrová 1988); MMio (Barstovian) Fort Polk, Texas USA Florida, USA (Meylan 2005). (Williams 2002); LMio (Vallesian, MN9) Rudabánya, Hungary (Roček 2005); LMio (Clarendonian), Whisenhunt Hyla miocenica Holman, 1966 Quarry, Oklahoma, USA (Parmley et al. 2010).

Diagnosis (after Holman 2003): Ilium similar to those Proacris Holman, 1961 of Hyla chrysoscelis and H. versicolor, but differing in Proacris mintoni Holman, 1961 that the dorsal tubercle is more dorsally produced from the shaft. Diagnosis (Holman 2003): Acetabulum very large; ventral Stratigraphy and distribution: MMio (Barstovian) Fort acetabular expansion almost completely ventral to the dorsal Polk, Texas, USA (Williams 2002); Mio (Barstovian) Trinity acetabular expansion; dorsal tubercle very large and ball-like, River, Texas, USA (Holman 2003). supported by relatively small dorsal prominence; iliac shaft very high along posterior two-thirds of its extent and then Hyla miofloridana Holman, 1967 becoming abruptly lower anteriorly. Stratigraphy and distribution: Mio (Hemingfordian) Diagnosis (after Holman 2003): Dorsal tubercle roughly Thomas Farm, Gilchrist County; Florida, USA (Holman rounded in shape, rather weakly produced laterally from the 2003). iliac shaft, not highly distinct from the prominence; anterior edge of dorsal prominence lies slightly posterior to anterior Pseudacris Fitzinger, 1843 edge of acetabular fossa; acetabulum only moderately excavated, and its border is rather weak; ventral acetabular expan- Diagnosis: Living genus, osteological diagnosis not sion well developed and wide; anterior edge of ventral ace- available. tabular expansion makes an angle of much greater than 90° with the iliac shaft; iliac shaft compressed, without dorsal Pseudacris nordensis Chantell, 1964 crest; long deep groove extends anteriorly on the ventral part of the iliac shaft. Diagnosis (Holman 2003): Dorsal acetabular expansion Stratigraphy and distribution: Mio (Hemingfordian) very small and narrow; dorsal tubercle very robust and Thomas Farm, Gilchrist County; Florida, USA (Holman elevated, far from acetabular fossa; ventral acetabular ex- 2003). pansion very narrow dorsally, making a wide angle with the long axis of the iliac shaft; well-developed dorsal crest Hyla cf. H. regilla Baird et Girard, 1852b on ilium present. Stratigraphy and distribution:MMio(Barstovian)Norden Stratigraphy and distribution: LPlio (Blancan) Hagerman Bridge Quarry, Brown County; Nebraska, USA (Holman area, Idaho, USA (Chantell 1970). 2003).

Hyla swanstoni Holman, 1968 Hylidae indet.

Diagnosis: Similar to Hyla miofloridana, but dorsal tubercle Stratigraphy and distribution: LCret, Asifabad, India of ilium more rounded, dorsal iliac prominence less extensive, (Prasad and Sahni 1987, 2009). acetabular fossa smaller with its anterior part hemispherical rather than oval, and iliac groove much less well-developed Bufonidae Gray, 1825 (Holman 2003). Stratigraphy and distribution: LEoc (Chadronian) Calf Diagnosis (from Duellman and Trueb 1994): Five to eigth

Creek near East End; Saskatchewan, Canada (Holman procoelous presacrals (V1 and V2 may be fused), with imbri- 1968). cate neural arches; atlantal cotyles juxtaposed; sacral diapophyses dilated, sacro-urostylar articulation bicondylar; Hyla sp. urostyle without transverse processes; sternum ossified; scapula is not overlain by clavicle; premaxilla and maxilla edentate. Stratigraphy and distribution: EMio (Orleanian, MN4) Dolnice, Czech Republic (Hodrová 1987); EMio (MN 4) Bufo Laurenti, 1768 Oberdorf, Austria (Sanchiz 1998b); MMio Subpiatră, Romania (Hír and Venczel 2005;Venczeletal.2005); Diagnosis: Living genus, osteological diagnosis not MMio (Badenian) Devínska Nová Ves—Bonanza, Slovakia available. Author's personal copy

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Bufo alienus Tihen, 1962 the genus. Although these crests are the same shape as those of Bufo terrestris, they are larger and end in a tall hemisphere Diagnosis (Holman 2003): Dorsal prominence of ilium trian- posteriorly. gular in lateral view, posterior slope of the prominence is Stratigraphy and distribution: LPlio (Blancan) Inglis 1A, steeper than the anterior; the prominence projecting somewhat Florida, USA (Meylan 2005). medially from the axis of the shaft. Stratigraphy and distribution: LMio (Hemphillian) Quarry Bufo gessneri (Tschudi, 1838) E near Long Island, Kansas, USA (Holman 2003). Diagnosis (after Sanchiz 1998a): Vertebral transverse pro- Bufo cf. B. alvarius Girard, 1859 cesses long and strong; femur and ilium of subequal length; F slightly longer than TF. Stratigraphy and distribution: Plio (Blancan) Benson local- Stratigraphy and distribution: Mio (Astaracian, MN7-8) ity, Cochise County, Arizona, USA (Gehlbach 1965). Öhningen, Germany (Sanchiz 1998a).

Bufo bufo (Linnaeus, 1758) or Bufo cf. B. bufo B. hibbardi Taylor, 1936

Diagnosis: Living species, osteological diagnosis not Diagnosis (from Holman 2003): Heavy, porous to granular available. supraorbital and postorbital crests; almost all the posterior Stratigraphy and distribution: MMio Devínska Nová Ves, portion of the frontoparietal, including the otic plate, is Slovakia (Hodrová 1980, 1988; Špinar et al. 1993); EPlio covered by the material of the crests; dorsal iliac prominence (MN 14) Osztramos 1, Hungary (Venczel 2001); LPlio high and about 45 % of the length of its base, with the anterior (MN16) Ivanovce, Slovakia (Hodrová 1981). slope of the prominence being much steeper than the posterior slope; sacral centrum relatively long and narrow. Bufo campi Brattstrom, 1955a Stratigraphy and distribution: LMio (Hemphillian) Edson beds, Sherman County, Kansas, USA (Holman 2003). Diagnosis (Holman 2003): Tibiofibula with a thin and wide longitudinal ridge along the central portion of the postaxial Bufo cf. B. hibbardi surface. Stratigraphy and distribution: LMio (Hemphillian) Stratigraphy and distribution: MMio (Barstovian) Fort Patterson Field locality; Arroyo de los Burros, Chihuahua, Polk, Texas, USA (Williams 2002); MMio (Barstovian) Mexico (Holman 2003). Egelhoff, Keya Paha County, Nebraska, USA (Chantell Remarks:Holman(2003) suggested that the ridge used to 1971); MMio (Barstovian) Hottell Ranch rhino quarries, diagnose this species is pathological. Banner County, Nebraska, USA (Voorhies et al. 1987).

Bufo cf. B. cognatus Say, 1822 or Bufo aff. B. cognatus Bufo holmani Parmley, 1992

Stratigraphy and distribution: Mio (Clarendonian) Diagnosis (Parmley 1992,fromHolman2003): Ilium large WaKeeney, Trego County, Kansas, USA (Wilson 1968); and extremely robust, with dorsal prominence long, moder- Mio (Hemphillian) Driftwood Creek, Hitchcock County, ately high and smooth; acetabulum large and moderately Nebraska, USA; Mio Quarry E, near Long Island, Phillips shallow; iliac shaft compressed along upper two-thirds, but County, Kansas, USA (Tihen 1962; Gehlbach 1965); Plio wider and ridge-like along the lower one-third. (Blancan) Beck Ranch, Scurry County, Texas, USA (Rogers Stratigraphy and distribution: LMio (Hemphillian) Devils 1976); Plio Big Springs, Antelope County, Nebraska, USA Nest Airstrip site, Knox County, Nebraska, USA; Santee, (Rogers 1984); Plio Hornet’s Nest Quarry, Knox County, Knox County, Nebraska USA (Holman 2003). Nebraska, USA (Rogers 1984); Plio Rexroad, Meade County, Kansas, USA; Plio White Rock, Republic County, Bufo kuhrei Holman, 1973 Kansas, USA (Tihen 1962; Gehlbach 1965); Plio White Rock Fauna, Republic County, Kansas, USA (Eshelman 1975). Diagnosis (after Holman 2003): Iliac prominence long and low, with a height only 17 % of the length of its base; two tubercles Bufo defensor Meylan, 2005 occurring on the summit of theiliacprominence,asmaller anterior and a larger posterior one, neither produced laterally; Diagnosis (Meylan 2005): Supraorbital crest of the dorsal prominence with small web anterior to the most anterior frontoparietal larger than any other New World member of tubercle; acetabular fossa extending almost to the anterior edge Author's personal copy

Palaeobio Palaeoenv (2013) 93:397–439 427 of the ventral acetabular expansion; acetabulum moderately thickened at this point of juncture, but not covering the excavated, with a pronounced lip occurring on approximately entire portion of the bone; ilium with a very high promi- its ventral half; a large, elongate, roughened tubercular bridge nence that has subequal anterior and posterior slopes or has produced from the posterodorsal margin of the acetabular cup a slightly steeper anterior slope; centrum of sacral vertebra and extending posterodorsally; a large interiliac tubercle on the short, and the crests of sacral neural arch meet in a broad, medial surface of the acetabular region. subangular arc. Stratigraphy and distribution:MMio(Barstovian)Norden Stratigraphy and distribution:Plio(Blancan)FoxCanyon, Bridge, Brown County, Nebraska, USA (Holman 2003). Meade County, Kansas, USA (Holman 2003).

Bufo marinus (Linnaeus, 1758) Bufo shandongensis Roček et al., 2011

Diagnosis: Living species, osteological diagnosis not Diagnosis (Gao 1986;Roček et al. 2011): Size large; head available. short and widely rounded; parasphenoid less than half as long Stratigraphy and distribution: Mio (Clarendonian) as wide, with lateral alae perpendicular to its longitudinal axis; WaKeeney, Trego County, Kansas, USA (Holman 2003). anterior margins of sacral diapophyses perpendicular to longitudinal body axis, posterior margins slightly inclined poste- Bufo pliocompactilis Wilson, 1968 riorly, so that diapophyses are moderately dilated towards their widely rounded ends; articulation between urostyle and Diagnosis (Holman 2003): Frontoparietals with supra- and sacral by double condyle; TF shorter than F. postorbital ridges absent and with dermal sculpturing most Stratigraphy and distribution: MMio Shanwang, Shandong pronounced toward the lateral border; orbital shelf with its Province, China (Roček et al. 2011). greatest overlap at the posterior corner of the orbit and Remarks: This species was originally described as Bufo covering the anterior opening to the occipital canal; linquensis by Yang (1977). However, the holotype specimen frontoparietal incrassation delimited by a low ridge; iliac was later recognised by Gao (1986)asMacropelobates.The prominence very high, with its central portion being an second specimen, designated as the paratype of B. linquensis elongated tubercle that is straight up or slightly reflected by Yang (1977), thus became the holotype of B. linquensis. anteriorly and with webbed portions forming its anterior As the species name linquensis was transferred to and posterior slopes; anterior margin of ventral acetabular Macropelobates, the new species name shandongensis was expansion extends well beyond the anterior margin of the established for this species of Bufo. For details of nomencla- acetabulum; iliac shaft slightly ventrally curved to almost tural history of Bufo shandongensis see Roček et al. (2011). straight. Stratigraphy and distribution: LMio (Clarendonian) Bufo (?) speciosus Girard, 1854 WaKeeney, Trego County, Kansas, USA (Holman 2003). Stratigraphy and distribution: Mio (Hemphillian) Santee Bufo cf. B. pliocompactilis Wilson, 1968 site, Knox County, Nebraska, USA (Holman 2003).

Stratigraphy and distribution: LMio (Clarendonian) Bufo spongifrons Tihen, 1962 Whisenhunt Quarry, Oklahoma, USA (Parmley et al. 2010). Diagnosis (Holman 2003): Cranial crests high but not Bufo praevius Tihen, 1951 especially heavy; the supraorbital, postorbital, and to a lesser extent the frontoparietal surfaces have a spongy Diagnosis (Holman 2003): Skull with high, granular supraor- texture; tuber superius ilii is high, between 45 and 50 % bital crests, oriented almost vertically and forming a sharp of the length of its base, with the posterior slope steeper angle with the postorbital crests; frontoparietals fused with than the anterior one. prootics; ilium with tuber superius with a height about 20 % of Stratigraphy and distribution: Mio (Hemphillian) Quarry E its base length. near Long Island, Phillips County, Kansas, USA; Plio Stratigraphy and distribution: Mio (Hemingfordian) (Blancan) Big Springs, Antelope County, Nebraska, USA Thomas Farm, Gilchrist County, Florida, USA (Holman 2003). (Rogers 1984; Tihen 1962).

Bufo rexroadensis Tihen, 1962 Bufo suspectus Tihen, 1962

Diagnosis (Holman 2003): Heavy supraorbital and postorbit- Diagnosis (Holman 2003): Iliac prominence low and long, al crests meeting each other at an obtuse angle, crests greatly about 17–18 % of the length of its base; anterior edge of Author's personal copy

428 Palaeobio Palaeoenv (2013) 93:397–439 acetabulum extends well behind the anterior edge of the iliac Stratigraphy and distribution: EPlio (MN 14) Osztramos 1, prominence. Hungary (Venczel 2001). Stratigraphy and distribution:Plio(Blancan)FoxCanyon, Meade County, Kansas, USA (Holman 2003). Bufo cf. B. viridis Laurenti, 1768

Bufo tiheni Auffenberg, 1957 Stratigraphy and distribution: EMio (MN 4) Vieux- Collonges, France (Bailon and Hossini 1990); EMio (MN 4) Diagnosis (Holman 2003): Sacral vertebra having a relatively EMio Sandelzhausen, Germany (Böhme 2010); MMio (MN 7 long, dorsoventrally depressed centrum with a shallow ventral or 8) Escobosa de Calatañazor, Soria, Spain (Sanchíz 1977); groove between the condyles, a horizontally oval glenoid MMio (MN 7) La Grive-Saint-Alban, France (Bailon and cavity and closely juxtaposed condyles; ilium with a high Hossini 1990); MMio (Badenian) Devínska Nová Ves – dorsal prominence, with a height 33–42 % of the length of Bonanza, Slovakia (Hodrová 1988); MMio (Badenian, its base and with pronounced webbing anterior and posterior MN6) Sámsonháza 3, Hungary (Venczel 2004); MMio to the elongated tubercular portion; an anteriorly–posteriorly (Sarmatian) Tauţ, Romania (Venczel and Sţiucă 2008). oriented and anteriorly deflected knob capping the elongated tubercular portion of the ilium. Bufo woodhousii Girard, 1854 Stratigraphy and distribution: Mio (Hemphillian) Haile locality, Alachua County, Florida, USA (Holman 2003). Diagnosis: Living species, osteological diagnosis not available. Bufo valentinensis Estes et Tihen, 1964 Stratigraphy and distribution: Mio (Hemphillian) White Cone, Navajo County, Arizona, USA (Parmley and Peck Diagnosis (Holman 2003): Frontoparietal somewhat thick- 2002); Plio (Blancan) Beck Ranch, Scurry County, Texas, ened laterally, without prominent crests but with finely USA (Rogers 1976); Plio (Blancan) Benson, Cochise County, granular ornamentation over the entire surface; iliac prom- Arizona, USA (Brattstrom 1955b); Plio (Blancan) White Rock, inence high and short (about 25–30 % of the length of its Republic County, Kansas, USA (Eshelman 1975). base); anterior edge of iliac prominence sloping steeply into the iliac shaft; anterior edge of acetabulum extending ante- Bufo sp. or ?Bufo sp. riorly from slightly beyond to slightly behind the anterior edge of the iliac prominence. Stratigraphy and distribution:MMio(MN6–8) Hasznos, Stratigraphy and distribution: Mio (Barstovian) Norden Hungary (Venczel 2004); LOlig (Whitneyan) I-75 near bridge Quarry, Brown County, Nebraska, USA; Mio Gainsville, Florida, USA (Patton 1969); EMio (MN4) (Barstovian) Egelhoff, Keya Paha County, Nebraska, USA; Mokrá-Western Quarry, Czech Republic (Ivanov 2008); Mio (Barstovian) Hottell Ranch rhino quarries, Banner EMioc Sandelzhausen, Germany (Böhme 2010); MMio County, Nebraska, USA; Mio (Barstovian) Kleinfelder Farm (Barstovian) Fort Polk, Texas, USA (Williams 2002); LMio near Rockglen, Saskatchewan, Canada; Mio (Barstovian) (Clarendonian), Whisenhunt Quarry, Oklahoma, USA Railway Quarry Cherry County, Nebraska, USA; Mio (Parmley et al. 2010); Plio (Villanyian) Včeláre, Slovakia (Barstovian) Lemoyne Quarry, Keith County, Nebraska, (Hodrová 1985); Plio (Blancan) Hagerman area, Idaho USA USA; Mio (Clarendonian) WaKeeney, Trego County, (Chantell 1970). Kansas, USA (all occurrences from Holman 2003). Remarks:Sanchiz(1998a) synonymised this species with B. Bufonidae indet. suspectus. Stratigraphy and distribution: LPaleo (Thanetian, MP6) Bufo valliceps Wiegmann, 1833 Cernay, France (Rage 2003).

Diagnosis: Living species, osteological diagnosis not Ranoidea Rafinesque-Schmaltz, 1814 available. Stratigraphy and distribution: Plio (Blancan) Beck Ranch, Diagnosis (Frost et al. 2006): Pectoral girdle firmisternal. Scurry County, Texas, USA (Rogers 1976). Microhylidae Günther, 1858 Bufo viridis Laurenti, 1768 Diagnosis (Holman 2003): Eight procoelous, holochordal Diagnosis: Living species, osteological diagnosis not presacral vertebrae, either imbricate or non-imbricate; atlantal available. condyles widely separated; sacral diapophyses broadly Author's personal copy

Palaeobio Palaeoenv (2013) 93:397–439 429 dilated; sacro-urostylar articulation bicondylar; urostyle lacks Rana basaltica Young, 1936 transverse processes; sternum cartilaginous. Diagnosis (Roček et al. 2011): A small ranid frog (SVL 29– Gastrophryne carolinensis (Holbrook, 1836) or Gastrophryne cf. 46 mm, depending on degree of development); LC somewhat G. carolinensis less than maximum LtC; comparatively long hind limbs (SVL/F+TF 0.85–1.08); TF longer than F (F/TF 0.71–0.90), Stratigraphy and distribution: Mio (Hemingfordian) Thomas suggesting that this frog was a relatively good jumper. Farm, Gilchrist County, Florida, USA (Holman 2003); LPlio Stratigraphy and distribution: MMio Shanwang, Shandong (Blancan) Inglis 1A, Florida, USA (Meylan 2005). Province, China (Roček et al. 2011). Remarks: Developmental series of tadpoles up to nearly Ranidae Rafinesque-Schmaltz, 1814 completely metamorphosed individuals also is known from the Shanwang locality (Roček et al. 2011). Diagnosis (after Duellman and Trueb 1994): Eight presacral vertebrae with non-imbricate neural arches; atlantal cotyles Rana capito LeConte, 1855 widely separated; V1–V7 are procoelous, V8 biconcave, sacrum biconvex; sacral diapophyses cylindrical and inclined Diagnosis: Living species, osteological diagnosis not available. posteriorly; sacro-urostylar articulation bicondylar; urostyle Stratigraphy and distribution: LPlio (Blancan) Inglis 1A, lacking transverse processes; sternum ossified; astragalus Florida, USA (Meylan 2005). and calcaneus fused at their ends; frontoparietal incrassation of typical ranid form; TF longer than F. Rana catesbeiana Shaw, 1802 or Rana cf. R. catesbeiana

Rana Linnaeus, 1758 Stratigraphy and distribution: LMio (Hemphillian) Mailbox Prospect site, Antelope County, Nebraska, USA; LMio Diagnosis (Špinar 1980, modified): Frontoparietal (Hemphillian) Pipe Creek Sinkhole Biota, Grant County, incrassation consists of an unpaired anterior part and a Indiana, USA; LMio (Hemphillian) Santee site, Knox paired posterior part; both frontoparietals meet in a median County, Nebraska, (Farlow et al. 2001; Parmley 1992); Plio suture; vomer dentigerous; vertebral column consists of (Blancan) Hornet’s Nest Quarry, Knox County, Nebraska, nine vertebrae (V1–V7 are procoelous, V8 is amphicoelous, USA (Rogers 1984); Plio (Blancan) Sand Draw, Brown V9, sacral, is opisthocoelous); sacro-urostylar articulation County, Nebraska, USA (Holman 1972); Plio (Blancan) bicondylar; sacral diapophyses slender, inclined postero- White Rock, Republic County, Kansas, USA (Eshelman laterally; scapula deeply incised; ossified part of sternum 1975); Plio (Blancan) Inglis 1A, Florida, USA (Meylan 2005). widened to the same extent both anteriorly and posteriorly; carpus of fully grown adults consists only of five elements Rana cf. clamitans Latreille, 1801 (2 in proximal row, 2 in central zone and 1 distal carpal near the base of metacarpal 1). Stratigraphy and distribution: Mio (Barstovian) Hottell Ranch rhino quarries, Banner County, Nebraska, USA Rana cf. R. areolata Baird et Girard, 1852a (Holman 2003).

Stratigraphy and distribution: Mio (Clarendonian) Rana esculenta Linnaeus, 1758 WaKeeney, Trego County, Kansas, USA (Wilson 1968); Plio (Blancan) Beck Ranch, Scurry County, Diagnosis: Living species, osteological diagnosis not Texas, USA (Rogers 1976). available. Stratigraphy and distribution: MMio (Astaracian, MN6) Rana cf. R. arvalis Nilsson, 1842 Sansan, France (Rage and Hossini 2000); MMio (Badenian, MN6) Sámsonháza 3, Hungary (Venczel 2004); MMio Stratigraphy and distribution: LPlio (MN16) Hajnáčka, (Badenian, MN6) Mátraszőlős 1, Hungary (Venczel 2004); Slovakia (Hodrová 1981). MMio (Badenian, MN6) Mátraszőlős 2, Hungary (Venczel 2004); MMio Hasznos, Hungary (Venczel 2004); MMio Rana cf. R. aurora Baird et Girard, 1852b Szentendre, Hungary (Venczel 2004).

Stratigraphy and distribution: Plio (Blancan) Hagerman Rana palustris LeConte, 1825 area, Elmore, Owyhee and Twin Falls counties, Idaho, USA (Chantell 1970). Diagnosis: Living species, osteological diagnosis not available. Author's personal copy

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Stratigraphy and distribution: Plio (Blancan) Beck Ranch, strongest and inclined posterolaterally; sacral diapophyses Scurry County, Texas, USA (Rogers 1976). slender and cylindrical, declines 35–42° from transversal axis of sacral vertebra; sacro-urostylar joint bicondylar; Rana cf. perezi (López-Seoane, 1885) pectoral girdle firmisternal; ilium with a dorsal crest well differentiated laterally from tuber superius; descending part Stratigraphy and distribution: Plio, Camp dels Ninots, of acetabular portion of ilium only moderately exceeding Spain (Gómez de Soler et al. 2012). acetabulum, ascending part prominent; F shorter than TF; length of F plus TF equal to SVL. Rana cf. R. pipiens Schreber, 1782 Stratigraphy and distribution: LPlio Willershausen, Germany (Špinar 1980). Stratigraphy and distribution: LPlio (Blancan) Hagerman area, Idaho, USA (Chantell 1970). Rana sylvatica LeConte, 1825

Rana pliocenica Zweifel, 1954 Diagnosis: Living species, osteological diagnosis not available. Diagnosis (Holman 2003): Sacral vertebra with relatively Stratigraphy and distribution: Plio (Blancan) Hornet’sNest deep medial groove on the ventral surface; iliac blade rela- Quarry, Knox County, Nebraska, USA (Rogers 1984). tively deep. Stratigraphy and distribution: LMio (Hemphillian) Rodeo Rana cf. temporaria Linnaeus, 1758 locality, Contra Costa County, California, USA (Holman 2003). Stratigraphy and distribution: EMio (MN 3) Dietrichsberg. Germany (Böhme 2001); LPlio (MN16) Ivanovce, Slovakia Rana pueyoi Navás, 1922 (Hodrová 1981).

Diagnosis (Špinar 1980): Head larger than in R. strausi. Rana sp. Stratigraphy and distribution:EMio(AgenianMN2) Navarette del Rio, Spain; LMio (Turolian MN 13) Arquillo Stratigraphy and distribution: LEoc (MP17a) Hordle, UK 1, Valdecerebro, Spain (Martín and Sanchiz 2013). (Holman and Harrison 1999); LOlig Enspel (Roček and Remarks: Although this species of European water frogs is Wuttke 2010); EMio (Orleanian, MN4) Dolnice, Czech documented by more than 60 articulated and comparatively Republic (Hodrová 1987); EMio (MN 4) Oberdorf, Austria well-preserved skeletons (both adults and associated (Sanchiz 1998b); EMio (MN4) Béon 1, France (Rage and tadpoles), mainly from the type locality Libros (Spain), it Bailon 2005); EMio (MN4) Mokrá-Western Quarry, Czech is poorly diagnosed (Navás 1922; see also comment in Republic (Ivanov 2008); EMioc Sandelzhausen, Germany Sanchiz 1998a). This is why Sanchiz (1998a) considered (Böhme 2010); MMio (Barstovian) Fort Polk, Texas, USA it nomen dubium, but admitted that it is a valid distinct (Williams 2002); MMio Subpiatră, Romania (Hír and species. Venczel 2005; Venczel et al. 2005); MMio (Sarmatian) Tauţ, Romania (Venczel and Sţiucă 2008); LMio (Vallesian, Rana ridibunda Pallas, 1771 or Rana (ridibunda)sp. MN9) Rudabánya, Hungary (Roček 2005); LMio (Clarendonian), Whisenhunt Quarry, Oklahoma USA Stratigraphy and distribution: EOlig (MP 22) Möhren 13 (Parmley et al. 2010); EPlio (MN 14) Osztramos 1, near Treuchtlingen, Germany (Sanchiz et al. 1993); LOlig Hungary (Venczel 2001); LPlio (Villanyian) Včeláre, (MP30) Oberleichtersbach, Germany (Böhme 2008). Slovakia (Hodrová 1985).

Rana strausi Špinar, 1980 Ranidae indet.

Diagnosis (after Špinar 1980, slightly modified): Length of Stratigraphy and distribution: EEoc (Ypresian), Vastan skull less than 20 % of SVL; frontoparietals with sagittal Lignite Mine, Gujarat, India (Folie et al. 2013); LEoc crest on the dorsal surface; medial portion of nasals large, (Rhenanian) Grisolles, northeastern France; LEoc irregularly triangular in shape, with lateral process of the (Headonian) Perrière, Malpérié, Lavergne, Lébratières 1, bone slender; tooth row covers about two-thirds of the Aubrelong 2, La Bouffie, Escamps, Rosières 1, Coanac 1, length of maxilla; frontal process of maxilla short and Sindou D, Ste Néboule, all Phosphorites du Quercy, south- robust, not in contact with lateral process of nasal; eight western central France (Rage 1984); EMio Sandelzhausen, procoelous presacral vertebrae; diapophyses on V3 are Germany (Böhme 2010). Author's personal copy

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?Ranidae Remarks: Maxillae with pit-and-ridge sculpture, associated with similarly ornamented squamosals and fragmentary Stratigraphy and distribution: LPlio, Dhammi and Balli, frontoparietals, have the same canal within anterior half of Jammu, India (Rage et al. 2001). orbital margin as Scotiophryne (Gardner 2008, compare with his fig. 13.1O and 13.3B), but not like any pelobatid. Thus, Ranoidea indet. rather than to Pelobatidae, these dermal bones belong to a representative of another family, probably the same as Stratigraphy and distribution: LCret Naskal, India (Prasad Scotiophryne. and Sahni 2009); EEoc (Ypresian) Vastan Lignite Mine, Gujarat, India (Bajpai and Kapur 2008). Anura incertae sedis Gracilibatrachus Báez, 2013 ?Rhacophoridae Hoffman 1932 Gracilibatrachus avallei Báez, 2013 Indorana prasadi Folieetal.,2013 Diagnosis (after Báez 2013): Quadratojugal absent; Diagnosis (after Folie et al. 2013): Differs from Rhacophorus mentomeckelian bones present; eight presacral vertebrae; and Polypedates in having weak dorsal crest on ilium; differs presacral vertebrae not fused with sacral; sacral diapophyses from Buergeria buergeria in having thicker iliac shaft, wider narrow; sacro-urostylar articulation bicondylar; inferred five acetabulum, and ossified pubis fused with ilium and ischium. pairs of free ribs in metamorphosing individuals and juveniles; Stratigraphy and distribution: EEoc (Ypresian), Vastan sternal end of coracoid moderately expanded; F longer than TF. Lignite Mine, Gujarat, India (Folie et al. 2013). Stratigraphy and distribution: ECret (Barremian) Las Hoyas, Spain (Báez 2013). Anura incertae sedis Eorubeta Hecht, 1960 Anura incertae sedis Eorubeta nevadensis Hecht, 1960 Hatzegobatrachus Venczel et Cziki, 2003 Hatzegobatrachus grigorescui Venczel et Cziki, 2003 Diagnosis (after Hecht 1960): Maxilla toothed; seven presacral vertebrae; sacral diapophysis expanded distally; Diagnosis (after Venczel and Csiki 2003): Dorsal protuber- diapophyses of presacral vertebrae of equal or subequal width ance on ilium well-developed and undivided; iliac shaft with- and flattened distally. out dorsal crest; pre- and supraacetabular regions small. Stratigraphy and distribution: EEoc, Sheep Pass Stratigraphy and distribution: LCret (Maastrichtian) Formation, White Pine County, Nevada, USA (Hecht 1960). Vălioara—Fântânele, Romania (Venczel and Csiki 2003).

Anura incertae sedis Anura incertae sedis Eopelobates sp. Hungarobatrachus Szentesi et Venczel, 2010 Hungarobatrachus szukacsi Szentesi et Venczel, 2010 Stratigraphy and distribution: EEoc (Ypresian), Vastan Lignite Mine, Gujarat, India (Folie et al. 2013). Diagnosis (modified after Szentesi and Venczel 2010): Remarks: Diagnostic characters of pelobatid (Pelobates, Middle-sized anuran (estimated SVL 50–60 mm); iliac shaft Eopelobates) ilia share also other anurans (e.g. those from with prominent dorsal crest; longitudinal ledge runs on lateral Mesozoic); therefore, assignment to Eopelobates should be surface of iliac shaft along its ventral margin, posteriorly supported by evidence from cranial bones (see also Rage and terminating in an elevation anterior to acetabulum and Augé 2003). delimiting a depression sculptured by longitudinal grooves and ridges that anastomose posteriorly; interiliac synchondrosis Anura incertae sedis prominent. cf. “Eopelobates” sp. Stratigraphy and distribution: LCret (Santonian) Iharkút, Hungary (Szentesi and Venczel 2010). Stratigraphy and distribution: LCret (Campanian) W of Farmington, New Mexico, USA (Armstrong-Ziegler 1980); Anura incertae sedis LCret (Maastrichtian) Bushy Tailed Blowout, Wyoming, Paradiscoglossus Estes et Sanchiz, 1982 USA (Gardner 2008); LCret (Maastrichtian) or EPaleo, Bug Paradiscoglossus americanus Estes et Sanchiz, 1982 Creek Anthills, Montana, USA (Gardner 2008); Eoc, Mission Valley Formation, California, USA (Golz and Lillegraven Diagnosis (after Gardner 2008): Ilium with well-developed 1977). dorsal crest inclined medially so it forms a broad groove Author's personal copy

432 Palaeobio Palaeoenv (2013) 93:397–439 parallel to dorsal margin of the shaft; dorsal tubercle is a low, Scotiophryne sp. elliptical elongate knob placed dorsolaterally on posterior end of dorsal crest; dorsal acetabular expansion prominent; Stratigraphy and distribution: MPaleo, Tongue River supraacetabular fossa is a deep pit open posterodorsally. Formation, Montana, USA (Estes 1976). Stratigraphy and distribution: LCret (Maastrichtian) Bushy Tailed Blowout, Wyoming, USA (Gardner 2008). Anura incertae sedis. Theatonius Fox, 1976 cf. Paradiscoglossus sp. Theatonius lancensis Fox, 1976

Stratigraphy and distribution: LCret (Maastrichtian) Pui, Diagnosis (Fox 1976; Gardner 2008): Inferred small body Romania (Folie and Codrea 2005), LCret (Maastrichtian) size; maxilla, frontoparietal and squamosal sculptured with Blasi 2, Spain (Blain et al. 2010). moderate-sized, tightly packed pustules; maxilla edentulous, with lamina horizontalis an indistinct ridge; processus Anura incertae sedis palatinus maxillae massive, with prominent facet for su- Scotiophryne Estes, 1969 tured contact with nasal, groove for ductus nasolacrimalis Scotiophryne pustulosa Estes, 1969 deep and extending across base of processus palatinus; processus pterygoideus short but broad, with prominent Diagnosis (partly after Gardner 2008): Ilium lacks dorsal crest articular facet that wraps posterodorsally onto dorsal edge and has dorsal tubercle poorly developed; preacetabular of processus zygomaticomaxillaris; posterior end of region anteriorly short, dorsal acetabular expansion low, processus posterior bluntly pointed; squamosal having la- and ventral acetabular expansion moderate, with depth mella alaris as a broad plate, with tiny spines along greater than height of dorsal acetabular expansion; maxilla, posteroventral margin; frontoparietal relatively short and nasal, squamosal and frontoparietal ornamented with fine broad, with nearly transverse anterior margin and processus beadlike tubercles that rarely contact one another; lamina lateralis broadly expanded laterally, with extensive facet for horizontalis of maxilla thick, with flat lingual surface; inner contact with squamosal; incrassatio frontoparietalis surface of zygomaticomaxillar process and orbital margin of undivided. maxilla are pierced by orifices of canals that run anteriorly Stratigraphy and distribution: LCret V-5711 near Lance to open on the posterior surface of the frontal process; Creek, Wyoming, USA (Fox 1976; Gardner 2008). groove for nasolacrimal duct runs horizontally along sculptured margin of the bone, crossing the base of the frontal Anura incertae sedis process; alar lamella of squamosal dorsally elongate, in- Tregobatrachus Holman, 1975 clined slightly posteriorly, with obtuse bend midway along Tregobatrachus hibbardi Holman, 1975 its height; the zygomatic process of squamosal broadly sutured with maxilla. Diagnosis (Holman 2003): Ilium with dorsal acetabular ex- Stratigraphy and distribution: LCret (Santonian) North pansion reduced in size, its dorsal border extending almost Side Pasture Wash, Utah, USA (Roček et al. 2010); LCret straight back from the dorsal prominence; iliac shaft without (Campanian) White Flats Road, Campbell Canyon, Websters dorsal crest; dorsal prominence well developed (about twice Flat, Howards Blues, Utah, USA (Roček et al. 2010); LCret as long as high) but lacking a tubercle, with one half of its (Campanian) W of Farmington, New Mexico, USA extent anterior to acetabulum, with a smoothly rising pos- (Armstrong-Ziegler 1980); LCret–EPaleo (Lancian-Puercan) terior slope and more abruptly rising anterior slope, and Bug Creek Anthills, Montana, USA; LCret (Lancian) with a tip of prominence deflected medially; iliac shaft Wyoming, USA (Estes 1969;Gardner2008). compressed, its medial surface with an anterodorsally di- Remarks:Estes(1969) designated the holotype of rected wide groove and entire shaft abruptly constricted Scotiophryne as an ilium from Bug Creek Anthills of anteriorly; large foramen present on shaft just anterior to Montana, USA, and associated it with 14 fragmentary maxil- anterior edge of ventral acetabular expansion; acetabulum lae and distal ends of humeri from the same locality—how- large, with pitted surface. ever, without giving any evidence supporting this association. Stratigraphy and distribution: Mio (Clarendonian) He probably only inferred that all of these skeletal elements WaKeeney, Trego County, Kansas, USA (Holman 2003). from the single locality should belong to the same species. However, if this is ultimately proved not to be true, then the Anura incertae sedis holotype ilium will keep the name Scotiophryne pustulosa, and a new name will be needed to accomodate the distinctive Stratigraphy and distribution: ECret (Comanchean) Paluxy cranial bones currently contained in S. pustulosa. Church, Texas, USA (Winkler et al. 1990). Author's personal copy

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Anura incertae sedis Bieber V (1881) Über zwei neue Batrachier der böhmischen Braunkohleformation. Sitzber Akad Wiss Math-Naturwiss Kl 82: 100–124 Stratigraphy and distribution: MEoc, Green River Biton R, Geffen E, Vences M, Cohen O, Bailon S, Rabinovich R, Malka Formation, fossil “Lake Uinta”, Utah, USA (Gardner 1999). Y, Oron T, Boistel R, Brumfeld V, Gafny S (2013) The rediscovered Remarks: Cast of a fossil tadpole. Hula painted frog is a living fossil. Nat Commun 4:1959. doi:10. 1038/ncomms2959 Blain H-A, Canudo J-I, Cuenca-Bescós G, López-Martínez N (2010) Amphibians and squamate reptiles from the latest Maastrichtian (Upper Cretaceous) of Blasi 2 (Huesca, Spain). Cretac Res 31: Acknowledgements I would like to thank Jim Gardner and Jean- 433–446 Claude Rage for reviewing the manuscript and making helpful sug- Böhme M (2001) The oldest representative of a brown frog (Ranidae) from gestions for its improvement, and Sinje Weber for her patience with the Early Miocene of Germany. Acta Palaeontol Pol 46:119–124 my tardy submission of this manuscript. Thanks are also due to the Böhme M (2007) Herpetofauna (Anura, Squamata) and palaeoclimatic late Farish Jenkins Jr. who kindly allowed me to study material of implications: preliminary results. In: Daxner-Höck G (ed) Prosalirus under his care. Yuri Gubin from the Paleontological Insti- Oligocene-Miocene vertebrates from the Valley of Lakes (Central tute Moscow allowed me to take photographs of Altanulia and Mongolia): Morphology, phylogenetic and stratigraphic implica- Cretasalia. The Museo Nacional de Ciencias Naturales Madrid pro- tions. Ann Naturhist Mus Wien 108A:43–52 vided the photograph of the Eodiscoglossus santonjae holotype. 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