Taxonomic Ciarification of Diplocynodon POMEL, 1847 (Crocodilia) from the Miocene of Styria, Austria

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N. Jb. Geol. Paläont. Abh. 261/2, 177—193 Article ~ published onhine April 2011

Taxonomic ciarification of Diplocynodon POMEL, 1847 (Crocodilia) from the Miocene of Styria, Austria

Jeremy E. Martin and Martin Gross With 8 figures and 1 table

MARTIN, J. & GRoss, M. (2011): Taxonomic ciarification of Diplocynodon POMEL, 1847 (Crocodilia)

from the Miocene of Styria, Austria. — N. Jb. Geol. Paläont. Abh., 261: 177—193; Stuttgart.

Abstract: A re-examination of the original type series of PRANGNER (1845) and HOFMANN (1887a) of the primitive alligatoroids from the middle Miocene of Styria, led to a reappraisal of the taxonomy of the following species of Diplocynodon: D. steinen and D. styriacus. Of unsettled affinities, En neodon ungeni was also re-examined. lt is here demonstrated that it belongs to the same taxon of the specimens described by HOFMANN (1887a). These taxa are in fact junior synonyms of the previously erected Enneodon ungeri. Moreover, comparison with other European alligatoroids reveals that the Austrian specimens described by PRANGNER (1845) and HOFMANN (1887a) belong to the same genus: Diplocynodon POMEL, 1847. According to the rules of the International Commission on Zoological Nomenciature (ICZN), Diplocynodon has priority over Enneodon. Under the principle of priority, it is therefore proposed to rename all the Miocene remains of alligatoroids from Styria as Diplocyno don ungeri PRANGNER, 1845. Comparison of almost complete skulls from various Miocene contempo raneous localities reveals that there is no reason to erect another taxon for the French specimens of D. “styriacus“ described in GINSBURG & BULOT (1997). However, these specimens need to be redefined as D. ungeri as weil. D. ungeri was coded and included in a character matrix to cladistically test its affinities with other alligatoroids. A total of six species of Diplocynodon were analysed including: D. ratelii, D. hantoniensis, D. tormis, D. inuelleri, D. darwini and D. ungeni. The resuits are consistent with previous studies and favour a monophyletic diplocynodontid clade. D. ungeni is the first species of the genus to be recognized from distant coeval European deposits, namely the Paris and the Pan nonian Basins.

Key words: Diplocynodon, Alligatoroidea, Styria, Miocene, taxonomy.

1. Introduction partly a much more blunt or much more acute snout, this certainly does not enable a comparison with Cr. Among the oldest described fossil crocodilians from Alligator styriacus nov. spec.; the same applies for the Europe is the enigmatic Enneodon ungeri of PRANG gavialiform Crocodilus Ungeri Prangn.“]. In the same NER (1845). Enneodon has been recovered from the work, HOFMANN (1887a) described and erected two middle Miocene lacustrine deposits of Schönegg in new species from geographically dose sites of approx Styria, Austria. Soon after its description, it was at imately the same age: Schönegg and Vordersdorf, the tributed to the genus Crocodylus (FITZINGER 1846) and former corresponding to the locality from which En HOFMANN (1887a) considered it to be a gavialiform neodon was previously recovered. HOFMANN (1887a) without providing further explanation [see HOFMANN erected Crocodilus (Alligator) styriacus and Croco p. 35: “What concerns the species described by OwEN dilus steinen for these two new species. In a short note from the Tertiary-Formation of England, these have HOFMANN (1887b) mentioned two additional findings

l3~E 300km 47 N 4 Sarmatian fl MiocenePho-/Pleistocenevolcanics& A Badenian ~ Basenient * Styrian Karpatian Basin

“Upper Eibiswald Beds“

“Middle Eibiswald Beds“

Crystalline basement

Fig. 1. A — Simplified geological map and position of the Styrian Neogene Basin. B — Geological map of the Wies-Eibiswald coal mining district (after BECK-MANNAGETTA & STINGL 2002).

of C. steinen from Wies (Schönegg and Brunn). These don, the choice regarding the taxonomy of the Aus Austrian crocodilian remains have been reviewed sub trian Diplocynodon at the species level remained un sequently by several workers who debated affinities clear. The latest work invoking this problem is that of with the genus Diplocynodon, previously erected by BERG (1966). Re favoured the use of D. styriacus over POMEL (1847) from exquisite remains from the Mi D. steinen for the reason that the mandible in D. styri ocene of Allier, Central France. But despite that the acus provided a better diagnosis than any remains of general consensus favoured affinities with Diplocyno D. steinen. He therefore considered D. steinen to be a Taxonomic clarification of Diplocynodon POMEL, 1847 (Crocodilia) 179

Standard Central MN >. Epoch stages Paratethys Zones W: Wies coal seam 5 with type specimens of 0 Enneodon ungeriand 0) 5 Pontian Crocodylus styriacus Messinian MN13 deltaic-coastal 7- EN: EibiswaldNordersdorf — 7.2 — coal seam with type specimens MN12 of Crocody!us steinen 8- 0) CO MN11 -J T: radiometric dated 9- Pannonian cS M 5 tuffitic Iayers Tortonian MN1O 0 W lo~lying mire ~ 10- coal MN9 11- 0 0 0 sand — 11.6 >~ Co 12- Sarmatian MN fluvial (5 t3) Serravallian 2 .2 (braided river) ~ LlJ 7/8 0 0) EZ~1 pelite 13- LL 0 () —13.7- 14- MN6 CO ~ gravel .2 :~ . Badenian Co ~ Lartghian .0 15- w 5

16- — 16.0 — MN5 0 ~~_ETow~lying, 15.22 myr Karpatian (+/- 0.17) 17- 7fr~hwater mire -cl Ottnangian ~ 0 0 18- Burdigalian Co 51 0 0 0 0 19- MN3 afluvial fan (5 CO (proximal, Radl Fm.) 5 OD CO Eggen 2 0 and 0 c 20- uJ burgian 0 (5 —20.4— LC~ fan-delta (distal, 0) “Lower Eibiswald Beds“) 0 21- 0 MN2 CO 0 Aquitanian c-. 22- Egerian MN1 A Crystalline basement B

Fig. 2. A — Correlation of Central Paratethys stages, Standard stages and Neogene Mammal-Zones (after HARZHAUSER

& PJLLER 2007; PILLER et al. 2007). B — Lithostratigraphy, facies interpretation and position of the Wies and Eibiswald/ Vordersdorf coal seams (after NEBERT 1983; GRUBER et al. 2003; HANDLER et al. 2006).

junior synonym of D. styriacus. He briefly argued that and C. steinen are proposed as paratypes ofD. ungeri. E. ungeri should be placed under D. styriacus but did Together with Austrian material, recent description not finalize its claim. This, added to the fact that the of fairly complete material of the same species from synonymy of D. styriacus over D. steinen or the other France but referred as D. styriacus (GINSBURG & way around has never been substantiated, shows that BULOT 1997) is used to code and test the affinities of some ciarification is needed for the Austrian Diplo Diplocynodon with other alligatoroids. cynodon taxa: D. styriacus and D. steinen. Also, the status of E. ungeri needs to be assessed. 2. Geology and stratigraphy As a consequence, the holotype of PRANGNER (1845) previousiy referred as Enneodon ungeni as The herein discussed crocodilian remains originate weil as the type series of HOFMANN (1887a) previously from middle Miocene coal-bearing strata of the south referred as Crocodilus styriacus or C. Steinen are here eastern part of the Styrian Neogene Basin (Fig. 1). The re-examined. PRANGNER‘S generic name Enneodon Styrian Basin is an around 100 km long and 60 km is here recognized to be replaced by Diplocynodon. wide subbasin of the Pannonian back-arc basin System Considering the principle of priority, the species name at the eastern margin of the Alps (EBNER & SACHSEN Diplocynodon ungeri is then applied to all the Austrian HOFER 1991; see GROSS et al. 2007 for a compilation). Diplocynodon. The described and more complete From a palaeogeographical view, the Styrian Basin specimens listed in HOFMANN (1887a) as C. styniacus is part of the Central Paratethys, for which regional 180 J. E. Martin and M. Gross

A ltJ////~y

Fig. 3. The rostrum OfPRANGNER‘S original Enneodon ungeri from the middle Miocene of Schönegg, Austria (mv. No. 1774,

UMJ = holotype of Diplocynodon ungeri) in ventral view (A — photograph of the specimen; B — scan OfPRANGNER‘S original plate dating of 1845). The large arrow points to the premaxillary-maxillary notch. The small arrows point to the region of confluent fourth and fifth alveoli. Scale bar = 20 mm.

chronostratigraphical stages were established (for re biswald Fm. is divided informally into the “Middle cent reviews see HARZHAUSER & PILLER 2007; PILLER Eibiswald Beds“ and the “Upper Eibiswald Beds“. Re et al. 2007; Fig. 2). cent radiometric dating of tuffitic layers just above the Internally, the N—S striking Middle Styrian Swell EibiswaldlVordersdorf coal seam indicate an age of (a Paleozoic basement high) divides the Styrian Ba 15.22 (+1-0.17) myr (Early Badenian, MN5; HANDLER sin into the Western and the Eastern Styrian Basin. et al. 2006), which is in contrast to the earlier proposed Three depocentres are distinguished within the West Karpatian age based on mammal faunas (MOTTL 1961; ern Styrian Basin: the embayments of Stalihofen, K0LLMANN 1965; DAXNER-HÖCK 2003). Groß-St. Florian and Eibiswald. The most southern Probably linked with the ongoing eustatic sea level one, the embayment of Eibiswald, is filled with ap rise in early Langhian/Badenian times (T.B. 2.3 cy proximately 1000 m thick, siliciciastic sediments of cle of HAQ et al. 1988) the water table rises and the the Radl and Eibiswald Formation ranging in age from low-lying, freshwater mire facies of the Eibiswald/ early to middle Miocene (WINKLER 1927; STINGL 1994; Vordersdorf coal seam became replaced by limnic and GRUBER et al. 2003; Figs. 1-2). Above ?Ottnangian to deltaic sedimentary environments. Coarse gravels of Badenian, mainly coarse grained, subaeric alluvial fan a braided river system (“Pitschgau Conglomerate“) (Radl Fm.) and subaquatic fan-delta deposits (“Lower above might correspond to a regressive period. Lim Eibiswald Beds“; the more distal complement of the nic-deltaic sediments up section and again the growth Radl Fm.) coal-bearing layers are deposited. During of a low-lying mire (Wies coal seam) with rather high coal mining from the l8th to the 2Oth century (WE suiphur contents point to the following transgressive BER & WEiss 1983) many vertebrate (for a compilation phase and some influence of brackish (ground) waters see M0TTL 1970) and plant remains (v. ETTINGSHAUSEN from the lagoonal area at the northern Groß-St. Flo 1890, 1891) were found. Nevertheless, a modern re rian embayment (GRUBER et al. 2003). vision of those fossils is missing up to now. Due to NEBERT (1983) calculated for the sediments between the development of two main coal seams (Eibiswaldl the Eibiswald/Vordersdorf and the Wies coal seam Vordersdorf and Wies coal seam) this part of the Ei- (mean thickness c. 470 m) a duration of deposition of Taxonomicclarification of Diplocynodon POMEL, 1847 (Crocodilia) 181

pfr hl €~fr J~ ~>/ pt ~lr ‚r

c D

Fig. 4. Photographs of the skull of Diplocynodon ungeri (mv. No. 21, MUL) in A, dorsal and B, ventral views and the associated original drawings of HOFMANN (1887a) in C, dorsal and D, ventral views. Scale bar 20 mm. Abbreviations: an, angular; art, articular; den, dentary; f, frontal; j, jugal; LTF, lower temporal fenestra; mx, maxilla; oc, occipital condyle; Or, orbit; pa, palatine; pfr, prefrontal; pmx, premaxilla; pt, pterygoid; q, quadrate; qj, quadratojugal; san, surangular; soc, supraoccipital condyle; sp, splenial; STF, supratemporal fenestra. Scale bar = 20 mm. 182 J. E. Martin and M. Gross

around 670 kyrs. The type specimen of C. steinen was 3. Material found in the footwali of the coal seam at Vordersdorf Institutional abbreviations: UMJ, Department of Geology (Fig. 2). That means the type specimen of E. ungeri & Palaeontology, Universalmuseum Joanneum, Graz, Aus and C. styriacus, which were found in the hanging tria; MUL, Montanuniversität Leoben, Austria. wall of the Wies coal seam, are about less than 1 myr Other abbreviations: MN foliowed by a number: Mammal younger than the specimens mentioned before. Neogene zone of the Buropean Land Mammal Age (LMA) according to BIOCHRO‘M 97. The description is based on the following material: Most A of the material is housed in the Universalmuseum Joan neum, Graz and comprises the holotype of Enneodon un gen PRANONER. 1845, UMJ 1774 consisting of the rostrum on siab (Fig. 3A) as weil as a siab containing several osteo derms (UMJ 1775, Fig. 3B). All the elements of the type series for Crocodilus (Al ligator) styriacus as figured in HOFMANN (1887a) are also housed in the Universalmuseum Joanneum, Graz (for more information concerning the history of those specimens see GROSS & MARTIN 2008). Key specimens used for the fol lowing description are the mandible (UMJ 203838, Fig. 4A- B B), a left posterior mandibular portion (UMJ 203839, Fig. 4C-D) and a series of articulated osteoderms (UMJ 203848, Fig. 3C). The type series on Crocodilus steinen consists of sev eral postcranial elements as weil as an almost compiete but somehow crushed skull. The skull of the original type se ries of HOFMANN (1887a) on C. steinen is currently on loan at the Staaliches Museum für Naturkunde in Karlsruhe (Germany). lt is housed at the Montanuniversität Leoben (MUL) under the access code MUL 21 (HOFMANN, Fig. 5). The skuli is no longer in the state of HOFMANN‘S original de scription, some of the original plaster having been removed. lt is now available in two pieces. Its original shape can be examined from a plaster cast (Mus~e d‘Histoire Naturelle de Genäve-2416). All the surface of the skull is covered with brownish plasticine rendering suture relationship difficult to discern. Original inscriptions that may have been superim posed by Hofmann are still visible at the surface of the skull and their presence prevent further cleaning (the following inscription are written in red: te, ty, d, j?, fo.te, fp, or, pa, pt). Finaliy, two blocks recorded as UMJ 5775 and UMJ 5780 found in the Eibiswaid coal seam and stored in the collections of the Universalmuseum Joanneum, Graz (UMJ) were also included in the description. They consist of fragmentary skull table and rostrum for UMJ 5775 (Fig. 6) and of the occipital region in ventral view for UMJ 5780.

4. Systematic palaeontology

Crocodilia GMELIN, 1789 Fig. 5. Details of the skull of Diplocynodon ungeri (Inv. Eusuchia HUXLEY, 1875 No. 21, MUL): A, rostrum from the left anterior view with Alligatoroidea GRAY, 1844 arrow pointing at the double caniniform dentition from Diplocynodontidae HUA, 2004 right to left in the maxilla and dentary; B, detail on the right posteroventral region showing the cont act between Diplocynodon P0ME1, 1847 ectopterygoid and maxilla. The arrow points to the last two alveoli, scale bar 20 mm; C, left quadrate in posterodorsal Note: Here, the taxonomy foliows that of MARTIN & BENTON view with the arrow pointing to the medially shifted (2008). For an alternate point of view, refer to BROCHU et al. foramen a~reum. Abbreviations: ec, ectopterygoid; mx, (2009). maxilla; pmx, premaxiila scale bar = 10 mm. Taxonomic ciarification of Diplocynodon POMEL, 1847 (Crocodilia) 183

Fig. 6. Photographs of the mandible of Diplocynodon ungeri. A, mandible in connection (mv. No. 203838, UMJ); A, detail of the anterior region of (A) in left lateral view with suture organization of the spienial (sp); posterior mandibular portion of mv. No. 203839 (UMJ) in C, left lateral and D, occlusal views. Scale bar = 20 mm.

Differential diagnosis: The following combination of traits and maxilla; ectopterygoid adjacent to at least the last two is unique to the genus. maxillary alveoli; quadrate separates parietal from squa mosal ventral to the orbitotemporal foramen; dorsal margin Diplocynodon: enlarged fourth and fifth maxillary alveoli of lower temporal fenestra formed by quadratojugal, pre of roughly identical diameter; confluent third and fourth venting quadrate from reaching the fenestra; tip of spienial dentary alveoli containing caniniform teeth; ontogenetic excluded from the symphysis; overbite of the premaxilla and development of a pit into a notch between the premaxilla anterior portion of the maxilla. 184 J. E. Martin and M. Gross

Diplocynodon ungeri (PRANGNER, 1845) comb. nov. Nomenciatural remarks: Under the pninciple of pnior Figs. 3-7 ity, the species name Enneodon ungeni PRANGNER, 1845: 1) should apply to all the Styrian remains of Diplocynodon. v. 1845 Enneodon Ungeri; PRANGNER, p. 114, p1. 1, fig. a-r. Therefore, D. Steinen and D. styniacus should be consid

1846 [Crocodilusj Enneodon ungeri. — FITZINGER, p. 190. ered junior synonyms of D. ungeri. 2) Moreover, the genus

1847 [Crocodilus] Cr. Ungeri. — GIEBEL, p. 123. name Enneodon PRANGNER, 1845 is older than the genus

1854 Crocodilus ungeri Prangner. — V. MEYER, p. 580. name Diplocynodon POMEL, 1847. Therefore, Diplocyno v. 1887a Crocodilus Steinen nov spec. — HOFMANN, p. 27, p1. don should be considered junior synonym of Enneodon. 11, figs. 1-8; p1. 12, figs. 1-6; p1. 13, figs. 1-14. However, statement (2) cannot be fulfilled because Diplo v. 1887a Crocodilus (Alligator) styriacus nov spec. — HOF cynodon POMEL, 1847 repnesents a case of nonien pnotectum MANN, p. 33, p1. 14, figs. 1-9; p1. 15, figs. 1-6. (article 3.9. of ICZN) as it fulfils the following conditions:

1887a Crocodilus Ungeri PRANGN. — HOFMANN, p. 35. an older synonym or homonym of Enneodon PRANGNER,

1887b Crocodilus Steinen. — HOFMANN, p. 219. 1845 was not used as a valid name after 1899 (23.9.1.1.), and

1898 Diplocynodon Steinen HOFM. — ROGER, p. 386. its younger synonym (here, Diplocynodon POMEL, 1847) or 1902 Diplocynodon Steinen. — ROGER, p. 58. homonym was used for a particular taxon, as presumably

1936 Crocodilus ungeri PRANG. 1845. — KÜHN, p. 126. valid, in at least 25 publications, published by at least 10

1936 Crocodilus steinen HOFMANN 1886. — KÜHN, p. 128. authors during the last 50 years immediately preceding, and 1936 Crocodilus styriacus HOFMANN 1886. — KÜHN, p. spanning a peniod of at least 10 years (23.9.1.2.). The old 129. est name of Enneodon PRANGNER, 1845 is here considered a

1955 Diplocynodon styriacus (HOFMANN). — THENIUS, p. nomen oblitum. Nomina oblita remain available names and 187. in the present case, thene is no restriction against the use of 1966 D. styriacus (HOFMANN 1885) = cf. syn. D. steinen the species name “ungeni“. Therefore, Diplocynodon ungeni

(HOFMANN 1885). — BERG, p. 38, p1. 3, figs. 17-18. is preferred to designate all the remains from the Miocene 1966 “Enneodon Ungeri“ PRANGNER 1845, ?syn. D. Styri of Stynia as well as the remains from MN4 of France (see GINSBURG acais. — BERG, p. 38. & BULOT 1997). That D. steinen or D. styniacus were randomly used in the literature for designating taxa Holotype: UMJ 1774 consists of a rostrum in ventral view from Stynia or France without justification appears confus encased on siab (Fig. 3A). ing. In order to get clear from such confusion, a different and older name is pneferned, here D. ungeni. Locallty and age: Schönegg and Vordersdorf, middle Miocene (early Badenian, MN5) from the coal seams and associated sediments of the Eibiswald Formation, Stynia, 5. Description Austnia. Rostrum: As shown in MUL 21, the premaxilla com pletely surrounds the external nares and therefore ex Referred specimens: All the following specimens reported from the same area (Wies/Eibiswald) share morphological cludes the nasals from these openings, at least from correspondence with the holotype UMJ 1774 and are consid the dorsal surface. The tip of the nasals seems to be ered to belong to a single taxon: UMJ 1775: siab with osteo located quite far from the postenior narial margin. The derms; UMJ 5775: a fragmentary skull table and rostrum; premaxillae are longer than wide as are the external UMJ 5780: occipital region in ventral view; UMJ 203838: a nares. Anteriorly, both premaxillae are pierced by mandible; UMJ 203839: a left posterior mandibular portion; UMJ 203848: a series of articulated osteoderms; MUL 21: a an occlusal foramen for reception of the first dentary partial skull with mandible in occlusion. tooth. The premaxillae send a posterior dorsal proc esses that meet medially at the level of the fourth den Diagnosis: Member of the genus Diplocynodon character tary tooth. The ventral side is observed in UMJ 1774. ized by the following combination of characters: elongate The maxillae do not reach the nanial contour hut the snout representing 65 percent of the skull length (tip of snout to front of orbits / tip of snout to postenior margin of incisive foramen is broken. The alveolar and tooth the skull table), anterior process of the palatine thin (lami succession of the premaxilla and maxilla are as fol nar), dentary symphysis projecting to the posterior level of lows (dimensions are provided in Appendix). As seen the fourth dentary alveolus. in MUL 21, the premaxilla bears five alveoli. A large interalveolar space is located between the first and the Remarks: When mentioning Diplocynodon from the second alveoli, and then a pit for reception of a dentary Miocene of Styria, KÜHN (1936) referred to HOFMANN (1886) and BERG (1966) to HOFMANN (1885). Both authors point to tooth lies between the third and the fourth alveoli. The the same publication of A. Hofmann, which was printed largest alveolus is the fourth and is followed by a smail (special reprints) at the end of the year 1885 (the plates) alveolus almost as small as the first one. A weil formed and dated (the special reprint) as February, 1886. However, notch marks, in dorsal view, the premaxillary-maxil the whole volume was distributed in the year 1887. For lary junction and is well visible in MUL 21 and UMJ that reason we use this date as the valid date of publication (HOFMANN 1887a~. 1774. The left maxilla is more complete than the right Taxonomic clarification of Diplocynodon POMEL, 1847 (Crocodilia) 185 one and shows several teeth occluding labially to the not straight but form a shelf. In that same specimen, dentary. A first tooth of small size testifies of the pres the choanae are incomplete hut provide some infor ence of the first alveolus. The alveoli rapidly increase mation. Given that the pterygoids are fused anteriorly, in diameter until the fourth and fifth alveoli, which are the choanae must open posteriorly and fully within the of same diameter. More posteriorly, the preservation pterygoids. The right pterygoid shows a well devel in occlusion hides the alveoli and it is only possible to oped posteromedial notch as in MUL 21 (Fig. 4) and establish the number of alveoli: the maxilla bears at UMJ 5780. MUL 21 clearly shows that the right ectop least thirteen alveoli. terygoid borders the posterior portion of the toothrow As for the frontal and prefrontal, which are best (Fig. 5D). The maxilla does not send any outgrowth observed in UMJ 5775, a preorbital ridge in front of and the ectopterygoid suture lies along the level of the the orbits seems absent. This specimen shows that last two maxillary alveoli. the frontal divides the posterior tip of the nasals and From MUL 21, the exoccipital does not descend projects further anteriorly than the prefrontal. The along the basioccipital plate hut stops just below the anterior extension of the lacrimal is not visible hut level of the occipital condyle. The basisphenoid expo reaches further anteriorly than the prefrontal. These sure is large posteriorly on the braincase wall. How configurations are obscured in MUL 21. ever, the region is crushed and shades Organisation of the bones around it. Skull table: In MUL 21 and UMJ 5775 (Fig. 4), the skull table is wider than long. The parietal is medially Mandible: UMJ 203838 is represented by a mandible depressed and shallow ridges mark the contour of with articulated dentaries. Alveolar diameters are re the medial sides of the well-rounded supratemporal ported in Appendix. The left dentary bears 19 alveoli. fenestra. UMJ 5775 shows that the frontoparietal The posterior portion of the left ramus is broken off. suture is concavoconvex. However, the frontoparietal The symphysis is longer than in Diplocynodon ratelii, suture lies at the anterior limit of the fenestra. The reaching the level of the fifth dentary tooth. All other frontal prevents the postorbital and parietal from characters are common to Diplocynodon ratelii. The meeting and thus enters the anteriormost corner of this spienial is excluded from the symphysis, the third and fenestra. A shallow fossa is present at its anteromedial fourth alveoli are enlarged and confluent and the tooth corner. The supraoccipital is well visible and can be row shows a medial constriction at the level of the described as large, but does not prevent the parietal eighth and ninth alveoli for reception of the large dou from participating in the posterior wall of the skull ble caniniform fourth and fifth maxillary teeth. All table. The squamosal region of UMJ 5775 is broken the alveoli are rounded. The pattern of dentition is un hut MUL 21 presents a notch at the squamosal-parietal known in the mandible. As expressed in UMJ 203839, suture on the posterior wall of the skull table. In MUL the dorsal surface of the surangular is large for recep 21, the jugal is raised anterior to the postorbital bar. tion ofM. adductor rnandibulae externus superficialis The postorbital bar is inset below the dorsal surface and inedius. The posteriormost region of the dorsal of the jugal. The quadratojugal-jugal suture lies at the surangular surface is marked hy a deep groove. posterior corner of the lower temporal fenestra. The quadratojugal ascends along the dorsal margin of Osteoderms: The specimen of Enneodon ungeri the lower temporal fenestra and may have prevented (UMJ 1774) is associated with two osteoderms. They the quadrate to meet the squamosal. The otic notch both present their external surface only. One of them is clearly inset hut the squamoso-quadrate suture is is complete: it is square and do not show any median unclear. The foramen adrum lies on the mediodorsal keel. The second, a dorsal osteoderm, is broken hut the surface of the quadrate in both MUL 21 and UMJ median keel is well visible. 5775. A large siab (UMJ 1775) belongs most probably to the same finding mentioned by PRANGNER. A large Palatal and occipital regions: As visible in UMJ number of square osteoderrns lie at the surface of this 1774 (Fig. 3), the anterior process of the palatine is siab, which is composed of the same sediment as the thin. In MUL 21, the palatine-pterygoid suture reaches plate of UMJ 1774. Many osteoderms present their vis the posteriormost level of the suborbital fenestra. No ceral finely striated surface. Some of them show a me notch is present at this level and the dorsal and lat dian keel on the dorsal surface. All these osteoderms eral surfaces of the posterior part of the palatine are are square in shape. In some of them, it is possible to ob- 186 J. E. Martin and M. Gross serve thefacies articularis on their rostral margin (Fig. sible presence of the confluent fourth and fifth alveoli 3). The lateral and medial sutures are also observed. and the narrow spaced alveohi, unlike those in gavial. The specimen UMJ 203848 is a siab presenting a The phylogenetic diagnosis of Diplocynodon as re collection of osteoderms more or less in connection. cently redefined in PIRAS & BUSCALIONI (2006) is based Two ribs, connected to two fragmented vertebrae run on a data matrix used for the ciadistic analysis of Alli between the rows of osteoderms (Fig. 3B). Three sagit gatoroidea (BROCHU 1999, 2004) and is limited to only tal rows of osteoderms and two longitudinal rows are two characters: largest dentary alveolus is 10, 11 or 12 preserved. Their external surface presents no median (167-2) and medial jugal foramen is large (120-1). Nev keel and suggests that they belonged to the ventral ar ertheless, focusing on the formal taxonomic diagnosis mour as shown by articulated skeletons of Diplocyno of Diplocynodon provides sufficient evidence to sup don. A second set of connected osteoderms lies be port the affinities of PRANGNER‘S Enneodon to this ge neath and present their internal finely striated surface. nus. First, the presence of a notch at the premaxillary Because these osteoderms are obscured by the above maxillary suture has never been observed in any longi described series, it is not possible to give a proper rostrine taxa. lt is however commoniy present among count. Most of the mentioned osteoderms are sutured several species of the genus Diplocynodon including together. Their external surface present ovoid pits of D. darwini, D. ratelii, D. tormis and D. hantoniensis uniform diameter. as weil as in Baryphracta deponiae and is observed in the skull of Enneodon ungeri PRANGNER, 1845 as weil 6. Discussiori as in Crocodilus steinen HOFMANN, 1887a. Also, the premaxilla is very long in longirostrine taxa, which is not the case here and among aihigatoroids; and as 6.1. The status ofEnneodon ungeri PRANGNER, 1845 noted by BERG (1966) while deteriorated in PRANGNER‘S The holotype of Enneodon is a fragmentary rostrum, (1845) Enneodon ungeri, the premaxillary and max and for this reason did not provide a clear basis for iilary alveoli are dose to each other unhike in longi comparison with other crocodilian remains. Neverthe rostrines where alveohi are widely spaced. Also don less, several evidences offer the opportunity to reassess cordant with the observations of BERG (1966), the tooth its taxonomic identity. Enneodon of PRANGNER (1845) count corresponds to the double caniniform, part of and Crocodilus (Alligator) styriacus HOFMANN, 1887a the diagnosis of Diplocynodon where fourth and fifth were both recovered from the Miocene freshwater maxihiary alveoli are of same size (Fig. 3a). The naris sediments of the Eibiswald Formation. Since the Oh is large, as in other specimens reported by HOFMANN gocene, freshwater habitats of Europe rarely include (1887a) as weil as in the French specimens referred to more than one crocodilian taxon. Despite one men Diplocynodon styriacus (GINSBURG & BULOT 1997). tion from freshwater deposits (BUFFETAUT et al. 1984), The osteoderms reported in UMJ 1775 and UMJ Miocene longirostrine tomistomines, represented in 203848 as well as the two osteoderms present on the Europe by Gavialosuchus are mainly recovered from slab of the holotype mv. No. 1774 (PRANGNER‘S Enneo marine or coastal deposits (TAPLIN & GRIGG 1989). don) have the same morphology. They all bear Uni Longirostrine taxa may have been able to invade fluvi form pits, they are square and sutures are visible. Os al habitats. On the other hand, Diplocynodon is mostiy teoderms presenting an anteroposteriorly long median recovered from lacustrine deposits as is the case for the keel are considered dorsal. This suite of traits charac localities of Wies and Eibiswald. Therefore, the chance terizing the osteoderms, while not unique to Diplocy of recovering Diplocynodon and a longirostrine taxon nodon points in favour of a similarity with this genus. in a same deposit is reduced. So far, such combina The morphology of osteoderms of other contempora tion has been reported from the Falun de Touraine and neous crocodihians such as Gavialosuchus or Tomis Aquitaine but the deposits represent a coastal mixture toma are different: their keels are short and restricted of continental remains transported to proximal marine to the posterior region (BROCHU et ah. 2007). Therefore, settings (LECUYER et al. 1996). the morphology of the osteoderms points in favour of a BERG (1966) raised doubts on the proposed gaviali Diplocynodon affinity for Enneodon. form affinities of Enneodon ungeri as stated by HOF As a consequence of all the above-mentioned MANN (1887a). In addition, BERG (1966) wrote that the reasons, wie regard Enneodon as a representative of rostrum fragment of E. ungeri was probably similar to Diplocynodon (see Nomenciatural remarks above for that of Diplocynodon styriacus, highlighting the pos a detaihed expianation). Taxonomic ciarification of Diplocynodon POMEL, 1847 (Crocodilia) 187

Table 1. Historical account for the synonymy of Diplocynodon ungeri (PRANONER, 1845)“

Name Author Status Provenance Diplocynodon ungeri this study Diplocynodon ungeri Austria, France Diplocynodon styriacus Ginsburg & Bulot, 1997 Diplocynodon ungeri France Diplocynodon styriacus Berg, 1966 Diplocynodon ungeri Austria Diplocynodon styriacus Thenius, 1955 Diplocynodon ungeri Austria Diplocynodon Steinen Thenius, 1955 Diplocynodon ungeri Austria Crocodilus Kühn, 1936 Diplocynodon ungeri Austria Diplocynodon steinen Roger, 1898; 1902 Diplocynodon ungeri Austria Crocodilus steinen Hofmann, 1887b Diplocynodon ungeri Schönegg & Brunn, Austria Crocodilus styriacus Hofmann, 1887a Diplocynodon ungeri Schönegg, Austria Crocodilus steinen Hofmann, 1 887a Diplocynodon ungeri Vordersdorf, Austria Crocodilus Fitzinger, 1846 Diplocynodon ungeri Schönegg, Austria Enneodon ungeri Prangner, 1845 Diplocynodon ungeri Schönegg, Austria

6~2. Synonymy of Diplocynodon ungeri (ROGER 1902) just mentioning that HOFMANN (1887a) (PRANGNEF~ 1845) reported two species of Diplocynodon. ROGER did not Recent studies on or referring to Austrian Diplocyno declare either why its preference went for D. steinen over D. styriacus. For some reason, KÜHN (1936) re don applied the species Diplocynodon styriacus HOF grouped all species erected so far (i.e., ungeri, steinen MANN, 1887a (BERG 1966; GINSBURG & BULOT 1997) for the middle Miocene material. An historical account of and styriacus) under the same genus Crocodilus. One the names used so far is provided (Table 1). In Aus more time, the generic name Crocodilus has been re tria, the first Neogene fossil crocodilian was recovered placed with Diplocynodon by THENIUS (1955). His mo tivations were that dentition, mandible morphology and from Schönegg. PRANGNER (1845) described and fig ured this material under the name Enneodon ungeri, ventral osteoderms were referable to Diplocynodon consisting of a rostrum in ventral view (UMJ 1774). and he stated that he confirmed the thoughts of ROGER This material was transferred to Crocodilus by FITz (1902). THENTUS (1955) considered D. steinen and D. styriacus to be two separate species but no compari INGER (1846) later and mentioned by HOFMANN (1887a). son was provided and most of its article dealt with the HOFMANN (1887a) erected two new species: Crocodilus steinen, whose holotype consists of a complete skull status ofD. styriacus. The status ofD. steinen was not and jaws in connection (MUL 21) and Crocodilus (Al explained, neither was discussed the fact that ROGER ligator) styriacus. This distinction was made because (1902) almost dismissed D. styniacus. He also noted of some differences in tooth morphology, tooth count differences between ungeni and styriacus, however he also mentioned he did not see the original material and shapes of the mandible. HOFMANN (1887a) attrib uted the finds from Vordersdorf to C. steinen, and the of ungeni (at this time PRANGNER‘s specimen was lost; finds from Schönegg to C. styniacus. Both localities see KÜHN 1952). Finally, BERG (1966) in his review of are middle Miocene in age (MN5, Early Badenian). Diplocynodon stated that the species name styniacus was preferable over Steinen because the mandible as Anyway, HOFMANN (1887b) attributed findings from Schönegg and Brunn to C. steinen too. sociated with this name provided a better diagnosis. He considered that D. steinen was a junior synonym ROGER (1898) was the first to apply the name Diplo cynodon to crocodilian remains from Austria. He at of D. styniacus. BERG (1966) also questioned whether tributed a jaw fragment and isolated postcranial ele the holotype of Enneodon ungeni should be regarded ments to Diplocynodon steinen. This author did not or not as D. styniacus. Given that E. ungeni can now present any explanation for such a generic choice and be considered a Diplocynodon and that all specimens latter, continued to refer isolated teeth to D. steinen reported from the same area (Wies!Eibiswald) belong 188 J. E. Martin and M. Gross

to a single taxon, it is presently considered that PRANG the short length of the dentary symphysis of the speci NER‘S material has priority. Diplocynodon styriacus men from C~reste do not match the diagnostic condi and Diplocynodon steinen are then considered junior tion of D. ungeri, suggesting that there is no reason to synonyms of Diplocynodon ungeni (PRANGNER, 1845). synonymize these species. Several specimens were described and attributed to D. styriacus based on complete skulls from the Mi ocene of South and Central France (GINSBURG & BULOT 6.4. Cladistic analysis and the diagnosis of 1997). Examination of the original type material from Diplocynodon Austria allows confirming this attribution. However, Diplocynodon ungeri was coded in the matrix of BR0- as discussed above and for purpose of clarity these CHU (1999, 2004) based on the description and plates French specimens should be referred to D. ungeri. of GINSBURG & BUL0T (1997) as weil as from observa tions of the above-described specimens from the col lections of Leoben and Graz (Appendix). The matrix 6.3. Comparison of Diplocynodon ungeri with of BROCHU (1999,2004) was chosen because it includes other Diplocynodon a maximum of ailigatoroid taxa (37). Moreover, other Beside that cranial proportions were taken as evidence Diplocynodon species were included in the matrix to refer the French remains to the Austrian Diplocyno from the work OfPIRAS & BUSCALIONI (2006) and MAR don styniacus ~GINSBERG & BULOT 1997), both French TIN (2010). Therefore, the genus Diplocynodon con and Austrian specimens share a thin anterior process sidered in this analysis inciudes D. ungeni, D. natelii, of the palatine, a long symphysis and a relatively long D. hantoniensis, D. danwini, D. nuielleni, D. tonmis, snout. The anterior palatine process is short and blunt and D. elavenicus. The foilowing analysis is identical

in D. ratelii, D. muelleri ‚ and D. hantoniensis. lt is long to that conducted in MARTIN (2010) but with the addi and wide in D. tormis. The dentary symphysis is long tion of D. ungeni. The software TNT (GOLOBOFF et al. in D. ungeri and projects behind the fourth dentary 2003) was used to compute a matrix of 167 characters alveolus in the French specimens (GINSBURG & BULOT for 51 taxa inciuding Hylaeochampsa as the outgroup 1997) and to the posterior level of the fifth dentary al taxon. The matrix was analysed using a traditional veolus in the Styrian specimens. On the other hand, it search (1000 random seeds and 1000 replicates). PIRAS is short and projects near the level of the third dentary & BUSCALIONI (2006) coded D. rnuelleni as having no alveolus in D. ratelii and D. gervaisi (pers. obs.), to the keei on the dorsal osteoderms but the description men third in D. ‚nuelleri (PIRAS & BUSCALIONI 2006) and is tions otherwise. This was corrected for character 35. described as short in D. toninis (BuscALIoNI et al. 1992). The resuits are provided as a strict consensus tree The proportion of the snout in D. ungeni can be esti (Fig. 8) with the following characteristics: the strict mated in MUL 21 and in more compiete skulis from consensus is built of 54 trees; the best tree length has France. The snout represents 65 percent of the skull 409 steps; CI = 0.474 and RT = 0.78. The topology of length, which is substantially longer than in any other the tree is characterized by the presence of Leidyo known complete skulis of Diplocynodon except D. suchtis as the basalmost ailigatoroid. The ciade com tormis. The sutural contact between the ectopterygoid prising the genus Diplocynodon iies in the next more and maxilla in Diplocynodon was discussed in MARTIN derived position to Leidyosuchus, the basalmost al (2010). Here, it is confirmed that D. ungeri shows the ligatoroid taxon. The interreiationships of the clade same condition as in D. hantoniensis, D. elavenicus, Diplocynodon arc solved with Baryphnacta then D. D. ratelii, D. gervaisi and D. tormis: an ectopterygoid darwini lying at the base of the clade as obtained in contacting the posteriormost portion of the tooth row. previous studies (BU5cALI0NI et al. 1992; BROCHU 1999; BRINKMANN & RAUHE (1998) synonymised the Styrian PIRAS & BUSCALIONI 2006; MARTIN 2010). The present Diplocynodon with D. ratelii on the basis of a skull analysis does not provide new characters supporting from the Oligocene of C~reste, southeastern France the monophyly of Diplocynodon as already obtained but did not detail their choice. These authors did not in PIRAS & BUSCALIONI (2006). D. ratelii and D. han compietely describe their referred D. ratelii skuil, thus toniensis arc polytomic and sit in a basal Position rela the diagnostic characters for D. ungeni cannot be com tive to the two sister ciades formed by D. ungeni + D. pared with the present work. Nevertheless, according elavenicus and D. muelleni + D. tormis respectively. to the figures of BRINKMANN & RAUHE (1998) the skull However, character support is weak for most of the proportions (snout is 57 percent of skull length) and nodes within Diplocynodon as demonstrated by the Taxonomic ciarification of Diplocynodon POMEL, 1847 (Crocodilia) 189

1~ ‘1, ‘S‘w / ~ ~‘ ~%—_ 1

Fig. 7. Postcranial elements of Diplocynodon ungeri. A — The osteoderm siab (mv. No. 1775, UMJ), which was signalled in PRANGNER‘S description (1845). A dose up on a dorsal keeled osteoderm is provided with the arrow pointing to the facies articularis, scale bar = 40 mm; B — postcranial elements in connection (mv. No. 203848, UMJ) including two ribs and two vertebrae (v). The arrow points to the limit between two longitudinal rows of ventral osteoderms (v.o.). Dorsal osteoderms (d.o.) are also aligned but are mostly hidden by the ventral series. Scale bar = 20 mm.

calculated Bremer values superimposed on Fig. 7. This Dipiocynodontidae, they should not be included in the demonstrates that the relationships of the genus are genus. Therefore, D. darwini should be applied a dif poorly supported; however the consensus of its relation ferent generic name and D. nwelleri could be renamed to other alligatoroids fits at the base of Alligatoroidea Hispanocharnpsa nutelleri. However, the present anal and is fairly weil supported. ysis clearly does not place D. rnuelleri at the stern of The first ciadistic hypothesis involving the relation the Diplocynodontinae but in a derived position with ships of Diplocynodon was that of BUSCALIONI et al. D. tormis. Thus, the monophyly of Diplocynodon is (1992) in which the authors argued for a polyphylet here confirmed. ic genus. However, their resuits were very similar to those of subsequent analyses (see BROCHU 1999, 2004) hut discussion of the mono- or polyphyletic nature of 6.5. Spatial distribution of Diplocynodon ungeri Diplocynodon clearly suffered frorn a lack of taxa in The Diplocynodon lineage is recognized as endemic cluded in the analysis (5 species of Diplocynodon for a to Europe and evolved in this region for more than 40 total of 7 taxa in BuscALI0NI et al. 1992). More recently, myrs. The youngest species of the genus Diplocyno PIRAS & BuscALIoNI (2006) recovered a monophyletic don is D. ungeri (late Marnmal Neogene zone MN5), Diplocynodon but pinpointed that if Baryphracta, D. although much younger remains referred at genus darwini and D. muelleri were forming the stern of the rank have been reported in the literature. In particu 190 J. E. Martin and M. Gross

Hylaeocharnpsa vectiana Gavialis gangeticus 2 Thoracosaurus rnacrorhynchus Borealosuchusforinidabilis Borealosuchus wilsoni 3? 3? Lj______Borealosuchus acutidentatus Borealosuchus sternbergii Asiatosuchus gernianicus Prodiplocynodon langi 1 1 Crocodilus ‘affinis‘ Pristichampsus spp 1 Leidyosuchus canadensis Baryphracta deponiae Diplocvnodon darwini 2 Diplocynodon hantoniensis 3?/ Diplocynodon rate/ii Diplocynodon elavericus 3 1 Alligatoroidea Dz~plocyiiodor, wigeri Diplocynodon tormis Diplocynodon rnuelleri basal globidontans A lligatorinae 3 Caimaninae 9

Fig. 8. The position of Diplocynodon ungeri within Alligatoroidea. Strict consensus of 54 optimal trees obtained in this study using a maximum parsimony analysis. Numbers at nodes represent Bremer decay values.

lar, another species, D. levantinicum was erected from localities within a narrow window of time. This high younger deposits (MN9-10) of Bulgaria (v. HUENE & lights the patchy nature of our current knowledge on NiKoLov 1963) but the diagnostic material consists of a the distribution of Diplocynodon in space and time, a mandibular ramus and can at best be attributed to the genus with a remarkably long time range (Eocene to genus level. Moreover, according to the figured den late Miocene; see MARTIN 2009). The occurrence of tary, the symphysis does not extend beyond the fourth two species of Diplocynodon at a same locality has alveolus, unlike D. ungeri. One interesting outcome never been reported, and one observes that the time of D. ungeri lies in its wide longitudinal distribution range of the various species do not overlap, neither in from Austria to Central France. Although one must the Paleogene (MARTIN 2010) and neither in the Neo keep in mmd the spatial gaps of specimen sampling, gene with D. ratelii (MN1-2) occurring weil before D. it is noticeable that, however, other species of this ge ungeri (MN5): each species is reported from separate nus have often been recognized from ciosely spaced European Land Mammai Ages (LMA, BIOCHRO‘M Taxonomic clarification of Diplocynodon POMEL, 1847 (Crocodilia) 191

1997). The only exception could be Diplocynodon Acknowledgements cf. rateli from the Oligocene of Cdreste (BRINKMANN The authors would like to thank M. RABI for reviewing an & RAUHE 1998) but the attribution to that species re earlier version of this manuscript. The reviewers, A.D. Bus mains uncertain. The absence of premaxilla-maxillary CALIONI and M. DELFINO provided helpful comments that notching combined to the small size of the specimen helped to improve the quahity of the manuscript. suggests a juvenile and renders comparison with other Diplocynodon species difficult. This apparent suc References cession of species of Diplocynodon through time led GINSBURG & BULOT (1997) to view Diplocynodon as BECK-MANNAGETTA, P. & STINGL, K. (2002): Geologische Karte 1:50.000. Blatt 206. Eibiswald; Wien (Geolo representing an anagenetic lineage. But the present gische Bundesanstalt). phylogenetic results contradict this view in showing a BERG, D. (1966): Die Krokodile, insbesondere Asiatosuchus discontinuity between a phyletic change and the age und aff. Sebecus?, aus dem Eozän von Messel bei of the various species involved (see present study and Darmstadt/Hessen. — Abhandlungen des Hessischen MARTIN 2010). Moreover, the record of Diplocynodon Landesamtes für Bodenforschung, 52: 1-105. BIOCHRO‘M 97 (1997): Synthäses et tableaux de corr~la is sparse and the numerous non-diagnostic crocodil tions. — In: AGUILAR, J.-P., LEGENDRE, 5. & MICHAUX, ian fragments bridging the gaps between the various J. (Eds.): Actes du Congräs BiochroM‘97, 769-805; Land Mammal Ages where species of Diplocynodon Montpellier. have been recognized are to be regarded as potentially BÖHME, M. (2003): The Miocene climatic Optimum: evi informative for improving the time resolution of each dence from ectothermic vertebrates of Central Europe. — Palaeogeography, Palaeoclimatology, Palaeoecology, species. For now, the presence of two or more species 195: 389-401. of the same age in different catehment basins has not BRINKMANN, W. & RAUHE, M. (1998): Diplocynodon ratelii been reported yet but there is no reason to exclude this POMEL, 1847 (Crocodylia, Leidyosuchidae) aus dem possibility. Here, the extensive geographie distribution Unter-Oligozän von C~reste (Südfrankreich). — Neues of D. ungeri during the middle Miocene (MN5, 16-14 Jahrbuch für Geologie und Paläontologie, Abhandlun Ma) highlights that this species occupied at least two gen, 209: 295-321. BROCHU, C.A. (1999): Phylogenetics, taxonomy, and histori distant basins of similar latitude, the Pannonian Basin cal biogeography of Alhigatoroidea. — Society of Verte and the Paris Basin. The distribution of Diplocynodon brate Paleontology, Memoir, 6: 9-100. could be more continuous because unrecorded occur BROCHU, CA. 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