lssuedzz February2oo8 of SYstemoticPolaeontology 6 (r)' 4t-Sg lournol iOThe Natural History Museum doi:to.tot7lSt4772orgo7oo228) Printed in the UnitedKingdom A UEWSALAMANDRID AMPHIBIAN FROM THEMIODLE MIOCENE OF HUUGARY ANDITS PHYLOGEN ETIC RELATIONSHIPS M6rtonVenczel Romania l6rii CrigurilorMuseum, B-dut Dacia r'3, R0-4ro464 0radea, centraland eastern Paratethys region, surviving up to lateAstaracian times (MN 7 + 8). Taphonomic settingof theMdtraszdt6s localities indicates that Corpathotriton matraensis was closely associated withaquatic environments. KEYWORDS Astaracian, Paratethys, Pataeobiogeography, Palaeoenvironment, Salamandridae, Taxonomy Contents lntroduction 4t Materiat,osteologicaI terms and conventions 42 Systematicdescri Ptions 43 LissamphibiaHaecke[, 1855 43 CaudataScoPoli,t777 43 UrodelaDumdrit, 18o6 43 SalamandridaeGotdfuss, r8zo 43 Carpathotritongen' nov. 43 Carpathotritonmatraensis gen. et sp' nov' 43 Phytogeneticrelationships of Carpothotriton 52 PalaeoenvironmentaIandpataeogeographicat imptications 55 Acknowledgements 55 References 56 Appendix:List of characters 5E and Recentrepresentatives found within Europe.Thus one lnrnooucrloN may concludethat the Europeancontinent and adjacentter- evolutionaryhistory of this The family Salamandridae,comprising 20 generaand more ritories were important in the than 60 living species,represents one of the most diverse salamanderfamily. two majorgroups groups of caudateamphibians distributed acrossEurope' Traditionally,within Salamandridae 'true' salamanders(Chioglossa, fto.t-tl Rftitu, Asia and North America but with most fossil are distinguished, the E-mail: [email protected] 42 M. Venczel Mertensiellaand Salamandra), and the newts(Cynops, Ech- cene),Salamandra Laurenti, 1768 (Late Eocene-Recent)' inotriton,Euproctus, Neure rgus, Notophtalmus, Pachytriton' SalamandrinaFitzinger,I 826 (Early Miocene-Recent),Trit- Paramesotriton, Pleurode le s, Salamandrina,Taricha, Trit- urus Rafinesquel8l5 (?Eocene-Recent)and Tylototriton and Tylototriton).This arrangementis supportedby Anderson.1871 (Middle Eocene-Recent). The fossilrecord urus 'true' numerousstudies that indicate the monophyly of sala- of salamandridsalso containsseveral genera and species excludingSalamandrina (e.9. Titus & with uncertaintaxonomic status(taxon name usually pre- mandersand newts, 'cf 'aff Larson 1995;Larson et al. 2003; Montori & Henero 2004; cededwith or ) while a numberof taxonnames are Weisrockel a/.2006). The latter genus was seen as a newtby synonymsof other taxa or are consideredtobe nominadu- Naylor(1978), but it wasplaced in theSalamandrc-group by bia (e.g. Salamandragoussardiana, Triturus sansaniense, Wake& Ozeti(1969). Estes (1981) grouped Salamandrina T. minimus,T. lacasianum:Estes 198l; Rage & Hossini in his mostprimitive salamandrid group ('group I') together 2000).Most salamandridfossils are knownfrom squashed w ith Chi o g lo s s a, M ert en s i ella, M eg a I o t rito n and Sa Laman- skeletonsor from dissociatedthree-dimensional material, dra.Nevertheless, several recent morphological and molecu- which usuallyconsists of vertebrae,appendicular skeleton lar contributionshave considerably changed the taxonomy of and,very rarely, of cranialbones. salamandrids.The generic nameLyciasalamandra Veith & Until now only a few salamandertaxa have been repor- (MN Steinfartz,2004 is usedto accommodateL. (= Mertensiella) ted from the Middle Miocene 6-8) of the Carpath- (: luchaniand a further six relatedspecies (Veith & Steinfartz ian Basin, mostly from DdvinskaNov6 Ves Neudod 2004).The genusCabtriton Gray,1858 was resurrected to D6v6nyrijfalu)(Bratislava district), Slovakia' The fossil sala- includethe Pyreneanbrook newts,c' (: Euproctus)asper mandersdescribed from this locality include:Bargmannia and C. arnoldi, while the name Euproctusis retainedfor wettsteini,Salamandra broilii, Voigtiella ludwigi and lls- theTynhenian brook newts only (E montanusand E. platy- sotriton(:Triturus) roehrsi(Herre 1955).The taxonomic cephalus)(Cananza & Amat 2005).Furthermore, the large positionof Bargmanniawettsteini has not yet beenprecisely Europeangenus Triturus appearsnot to be monophyletic established,since it is consideredeither a fossil dicamp- (Milner (Cacconeet al. 1994;Titus & Larson 1995;Larson et al. todontidGstes l98l) or a caudateincertae sedls 2003;Montori & Henero2004; Litvinchuk et al' 2005).The 2000), while Salamandrabroilii and higtiella ludwigi are (Estes first personto addressthe taxonomicsubdivision of Triturus synonymsof Salamandrasansaniensis 1981). wasBolkay (1928), who establishedthree groups within the In the last three decadesa number of Astaracian genusTriturus based on cranial osteologicalcharacters: the vertebratelocalities in Hungary (Szentendre,Hasznos, subgeneraPalaeotiton (Triturus boscai, T. italicus,T. hel- Siimsonhilza,M6trasz6l6s, Fels6t6rkiiny, Feln6met), have & veticus,T. montandoni,T. vulgaris andT' vittatus),Mesotri' beendescribed (Kordos 1981, 1986; Hir 2003,2004a;Hft ton (7. alpestris)and Neotriton (actuallya synonymof Trit- K6kay 2004; Hir et al. 1998,2001; G6l et al. 1999a'b, urus) with T. cristatus and T' marmoratus.However, more 2000). Reportson fossil vertebrates,including salaman- al. recently,the large-bodiedT. vittatushas been placed in the drids,were made by Hir et al. (1998,2001) andG6l et thesecontri- subgenusTriturus (see Arntzen & Olgun 2000 and refer- (1999a,b,2000).The taxa described bricfly in (M6trasz616s2),Lisso- encestherein). Montori & Herrero(2004),beside Triturus butionsconsist of Archaeotritonsp. cf. (= Neotritonof Bolkay),resurrected the generic names Lfs- triton (:Triturus) roehrsi(Fels6t6rkdny 3/2), Lissotiton (M6trasz6l6sl) and sotriton(: Palaeotritonof Bolkay,excluding T. vittatus)and roehrsi(Mr{trasz6l5s 2), Lissotritonsp. (Siimsonhi{za)' Mesotriton.Finally, Litvinchuk et al. (2005) split Triturus Salamandridaeindet. In fact,Archaeotriton frontalbone into four monophyleticgenera: Triturus Rafinesque, l8l5; sp.was identified enoneously on the basisof a due LophinusRafinesque in Gray, 1850 (due to priority, this andtrunk vertebrae (Venczel, p. 45 in Gfi et a|.2000)' Oligocenenewt nameis a synonymof LissotritonBell' 1839);Mesotriton to theirapparent similarity with thoseof the (Bcihme Bcihme& Riissler Bolkay,1928 and OmmatotitonGray, 1850, to accommod- Archaeotritonbasalticus 1998; materialfrom ateO. ophryticusand O. vittatus. 2002). Amore detailedevaluation of the fossil to rep- Theearliest known fossil salamandrid is KoaLliellagen- Miitrasz6lSsshows that the caudateremains aPpear one a new salamandridamphibian zeliHene, 1950,from theUpper Palaeocene(MP 6) of Wal- resenttwo distincttaxa: post-cranialskeletal features, beck,Germany and Cernay, France (Estes et aL.1967;Estes with someunique cranial and Lissolritonroehrsi, 1981).Koalliella genzeli has also beenrecorded from the theother, less frequent, closely related to vertebraland appen- LowerEocene (MP 7) of Dormaal,Belgium (Godinot el al' asindicated by a combinationofskull, 1978)and probably from the Lower Eocene(MP 7) of Le dicularskeletal features. This paper will (l) provide a detailed morpho!9- Quesnoy,France (Nel et al. 1999) Furtherfossil repres- from M6trasz6l6s, entativesof salamandrids,have been described (e.g. Estes & gicaldescription of thenew salamandrid its phylogenetic Hoffstetter1976; Sanchiz & Mlynarski1979;Westphal 1980; (2) assessits taxonomicstatus and ascertain (3) discussthe pa- Estes1981; Hodrova 1984; Bailon 1989, 1991; Rodek 1988' relationshipswithin Salamandridaeand implications 1994,1996a,b,2005;Bbhme 1998; Sanchiz 1998; Rage & laeoenvironmentaland palaeobiogeographical Hossini2000;Bbhme & Rdssler2002; Bdhme & Ilg 2003) of thenew taxon. including:Archaeotriton von Meyer, 1860(Early-Late Oli- gocene),Brachycormus von Meyer,1860 (Late Oligocene)' Material,osteological terms and conventions ChrlotritonPomel, 1853(Middle Eocene-LatePliocene)' repeatedintensive washing ChioglossaBocage, I 864 (LateEocene-Recent), M egalotri- The fossils were collectedafter using screenswith meshes ton iittel, 1890(Upper Eocene or Lower Oligocene-Early and sievingof the sediments (Hit 2004b).The materialincludes a Miocene),M ertensiellnWolterstorff, 1 925 (Early Miocene- of 0.8 and 0.5mm bones(premaxilla, frontal, Recent),Oligosemia Navils, 1922 (Upper Miocene)' Pa- seriesof disarticulatedcranial parasphenoid,squamosal, quadrate, Iaeopleurodeles Herre, I 94I (UpperOl i gocene-Middle Mio- parietal,orbitosphenoid, NEwsALAMANDRTD FRoM Mtoolr MtocrHe or Hulcnnv 43 alternatingwith sandsand clays cover the fossiliferous layers. The whole non-marinelimnic seriesis situatedbetween the Riikos Leithakalk (Early Badenian)and Kozi{rdformations (Early Sarmatian);the microvertebrate bearing sediments are datedto theAstaracian European Mammal age (MN 7) (Hir & parativematerial used in this study(disarticulated dried skel- K6kay 2004). etonsand alcianblue and alizarin red colouredsalamandrid REFERREDsPECIMENS. M6trasz6l6s 1: frontals (n=2): specimens)came from the Museo Nacional CienciasNat- MMP. 2005.366,MMP. 2005.369;parietal (n: 1): MMP. urales,Madrid and the Crigurilor Museum, Oradea. Jlrii 2005.370;squamosals (n:3): MMP. 2005.371ll-3;paras- CommonEnglish terms and the standard anatomical orienta- phenoids(n:2): MMP. 2005.362,MMP. 2005.3'12;dentar- tion systemare usedthroughout this paper;measurements ies (n:4): MMP.2005.373114:vertebrae (n:5): MMP' of salamandervertebrae follow Haller-Probst& Schleich 2005.363114,MMP 2005.374;humeri (n= 19): (MMP. (1994);the classification and nomenclature