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 of salamandrids 2005.36411-3,MMP. 2A05.367,MMP. 2005.3',7611-15; follows Montori & Herrero(2004), Litvinchuk et al. (2005) ilia h:3): MMP. 2005.37511-3;femurs (n:47): MMP. andWeisrock et al. (2O06t. 2005.365| | -l 4, MMP. 2005.368/ I -3, MMP. 2005.37 7 | | -30. Institutionalabbreviations used are as follows: Mdtrasz6l6s2: frontals(n:26)'. MMP' 2005.378/l- BMNH, The Natural History Museum,London; 10, MMP. 2005.388/l-16; premaxillae (n:6): MMP. (n:l): MMP. 2005.387;orbito- MMGD, MuseumfiirMineralogie und Geologie, Dresden; 2005,392114;parietal (n:2): MMP. 2005.38911-2;parasphenoids MMP, MunicipalMuseum Pdszt6, P6szt6, Hungary; sphenoids (n:4)'. MMP. 2005.380,MMP. 2005'39011-3;squamosals MNCN, MuseoNacional Ciencias Naturales, Madrid; (n: 10):MMP. 2005.436/1-3,MMP' 2005'39111-7;qoad- CriqurilorMuseum, Oradea, Romania; MTC, Tdrii rates(n : 10): MMP. 2005.408/1-10;prearticular+angular Bonn' PILJB,Paliiontologisches Institut, Universitdt (n:26)'.MMP.2005.379ll-15, MMP. 2005,39411'l l; dent- aries(n:28): MMP. 2005.43711-3,MMP' 2005.39311-25; ceratobranchial(,?: l): MMP. 2005.403;atlases (n=38): IPTlo NS MMP. 2005.381/l-8,MMP. 2005.395/l-30;trunk verteb- SvsrrmnncDEScR rae(n:128): MMP. 2005.38211-5,MMP' 20053831143, (n=4): MMP. 2005.4041l- BIAHaeckel, 1866 MMP. 2005.3961140;ribs LISSAMPHI 4; scapulocoracoids(n:10): MMP' 2005.386/1-5'MMP. CAUDATAScoPoli, 1777 2O05.39711-5humeri (n= 120): MMP. 2005.384/l-30' MMP. 2005.398/I -90; radiuses(n : 4)'.MMP. 2005.399/1- URODELADumEri[, 18o6 4; cubituses(n:2): MMP. 2005.40011-2;ilia (n=48): ischia(n=2): SALAMANDRIDAE Goldfuss, rSzo MMP' 2005'438/l-5'MMP' 2005'4011143; MMP. 2005.40211-2;femurs (n:300): MMP' 2005.385/1- CARPATHOTRITOIVgen. nov. 105, MMP. 2005.405/l-195; tibiae (n = 38): MMP. 2005.40611-38 ; fi bulae(n : I 0): MMP. 2005.407/I - I 0; mal- TYPESPECIES. Carpathotriton matraensir gen. et sp.nov' formedspecimens: I atlas(MMP.2005.409), I ilium (MMP. 2005.410),3 femurs (MMP. 2005.411/l-3). ErYMoLocY. Carpathian,geographical name of mountain chainfrom CentralEurope, + triton, Greek,a salamander. DIAGNOSIS.Carpathotriton nwtraensis differs from Meso- triton, Neurergus, Oligosemia, Lissotriton montandoni,L. DIAcNosls.As for thetype and only knownspecies. vulgaris and Triturus in having a large posterolateralpro- cessto its frontal.It differs from Calotriton andEuproctus in Corpothotrltonmatroensls gen. et sp.nov. (Figs r, 3, having a comparativelyshorter frontal and a more promin- 5-7) ent supraorbitalcrest. Cynops, Echinotriton, Notophtalmus, 2C/ll0Archaeotriton sp.;G{l et a|.,45' lig. 3. Pachy t r i ton, Parameso t ri to n, PI e u ro de le s and Taric ha have a completefronto-squamosal arch, but differ in having a Mdtra, Hungarian, geographical name of ETYMoLocY. more prominentsupraorbital crest to the frontal. Carpatho- rangein thevicinity of thelocality. mountain triton matraen^sisfurther differs from Cynops,Echinotriton sculptureon Holorvpe. MMP. 2005.361,a partialright frontalmissing andParamesotriton by its lack of pit and ridge L hel' its postero-medialborder and distal sectionof posterolateral the frontal surface.It differs ftom Lissotritonboscai, process. veticusand Ommatotriton in havinga stouterand posteriorly wideningposterolateral process. OccuRReNceAND AGE. M6trasz6l6s locality 2, a section belongedto an of roadbelow the R6k6cziChapel, situated at the northem DESCRIFTION.Frontal. The holotypefrontal posteromedialmargin and marginof the smallvillage of M6trasz6l6s,North Hungary adultand is incomplete,with the processbroken off (GdIet a|.2000;Hir 2004a;Hfr & K6kay2004). The Mio- the distal section of the posterolateral surfaceof the frontal cenelacustrine series with freshwaterclays and lignite beds (Fig. lA, B). In dorsalview, the outer secondarydermal belongto the Saj6Valley Formation (Hfr & K6kay 2004)' is sligtrtty convexdorsally and lacks any of the frontalthere Thefossiliferous horizons rich in microvertebratesstart with sculpture.Alongside the medialmargin that, in life, it was about20cm of greenclays with freshwatermolluscs and is a lesselevated bony surfacesuggesting sectionofthe gypsumconcretions and are followed by about lOcm of overlappedby its counterpart.The anterolateral situatedin a deep giey clays,with abundantfreshwater molluscs. Lignite beds frontal is penetratedby severalforamina 44 M. VENczEL

Flgurer Carpothotritonmatruensis Een. et sp.nov. from Mitraszdlds z. A, B,MMP.2oo5.361 (holotype) frontal; C, D, MMP.zoo5.378/r frontahE, MMP.zoo5.38Z parietah F, G, MMP.2oo5.39olr parasphenoid; H, l, MMP.zoo539zlrprefrontal; J, K, MMP.2oo5.395h atlas; [, MMp.2oo5.4o+/rrib; M-P, MMP.zoo5.38zl1 trunk vertebrai q MMP.zoo5.382/2trunk vertebra; R, MMP.zoo5'386/1 scaputa. A C,F, l, l{' R, ventralviews; B, D, E, G, H, M, dorsalviews; l, posteriorview; K, O, anterior views; L, P, (L lateralviews. Scate bar = 1mm. groove,which is closedposteriorly' On the anteriorsection the premaxillary-frontalcontact. This morphologysuggests of the tablethere is a prominentarea produced as two diver- that in the living animal the distal sectionof the premaxil- gent ridges,defining a hemi-cylindricalshaped groove for lary dorsalprocess was almostcompletely embedded in the NewsnrnmnNDRtD FRoM Mtoolr MtoceHE or HuHennv 45 anteromedialsection of the frontal. On the anterolateralsec- smallerindividual, the middle section of theparietal crest is tion of the frontal thereis a small laterally inclined surface, extremelythin and prominent. In MMP.2005.387the anterior which waspresumably overlapped by the prefrontalin life. marginof theparietal is slightlydamaged but imprintson its The posterolateralprocess is relativelywide and presum- dorsalsurface suggest that it was overlappedby the frontal. ably wasrather long. Its dorsalsurface is nearlyhorizontal, The anterolateralmargin is depressedand piercedby several piercedby severalpits; and the supraorbitalcrest is low. A smallforamina. The lateralprocess of theparietal is tapering low crestruns from the medial margin of the posterolateral with a laterodorsalorientation indicating possible contact processtoward the frontal'sposteromedial margin, delimit- with the squamosalover the otoccipital.An arch-likeridge ing a less elevatedposterolateral area. In ventral view, the extendsbetween the parietal's lateral process and its posterior medialsurface is concaveventrally and borderedlaterally by margin,delimiting a lesselevated posterolateral area pierced a prominentventrolateral ridge. The fronto-orbitosphenoidal by severalsmall foramina. The latterstructure is alsoseen in contactis markedby an imprint on the medialside of the MMP.2005.370. ventrolateralridge. The areabetween the lateralside of the Orbitosphenold.Specimen MPP. 2005.389/1 (Fig. 3F ventrolateralridge and the supraorbitalmargin is concave andsee Fig. 7A) in lateralor medialview is oblong-shaped laterallyand piercedby severalforamina of varyingsizes. andhas a rathersimple morphology. The posteriorthird of A bonycrest ventrally connects the posteriorsection of the the boneis concavelaterally and pierced by two foramina: ventrolateralridge and the base of theposterolateral process. an anteriorlarger one for the aditusof the optic nerveand a The morphologyof the remainingfrontals closely re- posteriorsmaller one for the exit of the occulomotornerve. semblesthat of theholotype, exhibiting a relativelyn.urow Theanterior section is concavemedially and lacks foramina. rangeof intraspeciticvariation. ln MMP.2005.378/1,which About two-thirdsof the orbitosphenoid'santerior section, belongedto a largerindividual (Fig. 1C,D), theanterolateral facingdorsolaterally, in life wasoverlapped from aboveby projectionmarking the contactterminus for the prefrontal thefrontal; the posterior section, facing dorsomedially, was is distinctlylarger than that in the holotype;the bonycrest overlappedby the parietal.The ventralmargin of the orbito- runningfrom the medialbase of the posterolateralprocess sphenoidcontacted the parasphenoidfrom above,while its and the posteromedialmargin is extremelyprominent and anteriormostsection overlapped, dorsally, the vomer.Both inclinedposterolaterally. A comparablecondition may also the dorsaland ventralmargins are slightly widenedante- be seenin someRecent salamandrids (e.g. Lissotriton, Para' riorly. rnesotriton:Fig. 28, C, D. In MMP.2005.378/2the medial and Parasphenoid.In all specimensthe cultriform process anteriorsection ofthe frontal tableis brokenoff, but thereis is brokenoff. In MMP.2005.390/1(Fig. lF, G) thebasal plate a well-preservedposterolateral process whose distal section isbroadly rectangular with someposterior waist; its posterior is distinctlywidened (Fig. 3A, B). In MMP.2005.378/3the sectionis slightlyconcave. The ventralsurface is finelyorna- groovefor thefronto-premaxillary contact is distinctlywider mentedby smallpits and ridges,excepting the post-central andlonger than in otherspecimens. Several fragmentary spe- area,which is penetratedby severalforamina of variable cimensbelongingto large individuals bearafi ne dorsal sculp- size.On both sidesnear the baseof the cultriformprocess ture,produced by extremelyshallow pits andridges. thereis a shallowconcavity. A steepbony laminaconnects Premaxilla.The premaxillaeare completelyfused; the the posteriorcrest and the posteriormargin. The lattertapers parsdentalis is relativelylong and somewhat arcuate poster- posteriorly.In dorsalview theendocranial imprint is exposed olaterally.The bestpreserved specimen is MMP.2005.392ll in the form of an elongatedfossa delimited laterally by sharp (Fig. lH, I) with 18 tooth positionspreserved, of which bonylamellae. The lattelis flankcdat thelevel of thewidest I I teethare on theright sideof thepremaxilla, demonstrat- extentof the basalplate by a sinuousgroove for the carotid ing that therewere at least22 teeth.The labial surfaceof duct.In the posteriorarea there is a smallerconcavity de- the premaxillais slightlyconvex and withoutsculpture. In limited by a transversebony ridge.The ventralsurfaces in MMP.2005.392/|only the broken base of thedorsal process MMP.2005.372.380 and 390/2 are flat with somesmall irreg- (= alaryprocess) is preserved.The dorsalprocess, based on ularembayments and a convexityat thebase of thecultriform specimenMMP.2005.392l2,bears a distinct proximal keel on process;the posterior crest projects slightly. MMP.2005.362 its labialand lingual surfaces. The ramiof thedorsal process resemblesother specimens,but its posteriorcrest is more areunited at theirbase and diverge slightly dorsally; the free prominentand thereis a low crestdetached from the ventral portionis subcylindricalin shape.On the lateralside of the lateralmargin running obliquely onto the ventralsurface of dorsalprocess there is a distinctrecess with a foramenpier- the cultriform process. - cing thelamina horizontalis.The distal section of thedorsal Quadrate.The avail able specimens (MMP.2005.408/1 processin all specimensis brokenoff. However,based on the l0) belongedto various-sizedindividuals. This boneis mod- morphologyof availablefrontals, I deducethat the premax- eratelylong androbust and provided with a relativelywide illary dorsalprocesses were ratherlong and contactedthe otic processthat facesdorsally. The quadratecrest is sinu- anteromedialpart of the frontal. The pars palatina is relat- ous,ending in a spur-likeprocess projected dorsally. On the ivelysmall when compared to othersalamandrids (Fig. 4). It posterolateralarea there is an imprint formed by the ventral hasa slightposterodorsal orientation and is connectedmedi- ramusof the squamosal,which in life would havepartially ally to thebase of thedorsal process. overlappedthe quadrate.A prominentprocess is observed Parietal.The dorsalsurface of the parietalis smooth on theanterolateral side of the quadrate,which presumably withoutany secondary sculpture. In MMP.2005.387(Fig. 1E) servedfor muscularattachment (Fig. 3H, I). themedial margin of thebone is slightlycurved dorsally to Squamosal.MMP.2005.371ll (Fig. 3C) is tri-radiate- contactits counterpart.Obviously in the living individual shapedand stronglyffattened medio-laterally. The ventral a parietalridge was presentwith contributionsfrom both ramusis relativelylong (its posterior margin is broken),over- parietals.In MMP.2005.370,which belongedto a somewhat lappingthe quadrateposterolaterally. The posteriorramus of 46 M. Veruczel

.\ *flll

o tl -{ ,r1 df

Flgure2 Rightfrontals of Recentsalamandrids (dorsal views).A, Pleurodeteswaltl(MNCN 19648). B, Lissotritonboscol(MNCN rgr8z). c,Lissotritonhelvefrcus(MNCN19z6).D,Ttiturusmarmorcfus(MNCN t9275).E,cynopspynhogaster(MNCN2J822).F,Tarichagronuloso (MNCNtt8:o)' G'ommototritonuitfatus (MNCN r+oq6z).H, Mesotriton potomesotriton alpestris(MTC2345o). a, hongkongensis(MNCNztssz). Abbreviations:pfc, groove premaxillary-frontal for contact;plp, posterolateral process. Siale bar: z mm. thesquamosal is relativelylong and directed posterodorsally; sectionbears a rostrumdirected anteroventrally. On the lar its medialside contacts the crista parotica of the prootic. The eralsurface of the squamosalthere are two malncrista: one anteriorramus is of thesame length or somewhat shorterthan is inclinedposterolaterally and runs along the lateralside the posteriorramus and articulateswith posterolateral the from the posteriorprocess to the distal end of the ventral processof thefrontal, forming a completefronio-squamosal process;the other starts from thedorsolateral margin of the arch,On the dorsomedialside of the anteriorramus, there anteriorramus and meets the lateral crest at theposterolateral is a smallbut distinctprocess, which, at leastin largerspe_ cornerof thesquamosal. At thatpoint a prominentknob_like cimens(e.g. MMP.2005.37l/1,43611) couldbe in contacr structureis produced(Fig. 3D, E). with the lateralprocess of the parietal.In MMp.2005.436/l .-lentary. Although all specimensare fragmentary, the ventralramus is brokenoff distally,but both posterior availablematerial includes specimens with eithei the an_ andanterior rami arewell developed and thedistal anterior terioror the posteriorsection of boneallowing a complete NEwsALAMANDRID FRoM Mtoole MtocEHe or HulclnY 47

MAtraszdl6sz. A, B,MMP.zoo 5.?7812lrontalt C, MMP'zoo5'gzr/r squamosah Ff$re 3 corpothotritonmatroensisgen. et sp. nov. from rightfirst ceratobranchiah H,l' MMP'zoo5'4o8/r D,E, MMp.2oo5.3Zrlz squamosah F, MMP.2oo5.389h orbitosphenoid; G, MMP.zoo5.4o3 E,F, l, lateratviews; D, posterior view; H, anterior view' Scale quadrate;l, tUmp,zooS.+olr/rrib; A, ventral view; B, G, dorsat views; G, l, bar: r mm,

completelyat thelevel of theseventh or eighth reconstruction.The labial surface is broadly convex and row andcloses position.The symphysisis rathersmall. smooth,with a shallow groove along the labial side of the tooth The prearticularis elongateand baseof the dentalparapet' In MMP.2005.437I1,represent- i'rearticular+angular. mediolaterally and is providedwith a inga distinctlylarger specimen (Fig. 5F), the labial surface is slightly compressed coronoidprocess and a low lateralflange bor- scilpturedwith longitudinalstriations, grooves and foramina rrtutiutty high Meckel's groovelingually' The dorsalmargin of andihereis awell-defined keel along the lowerdental margin' dering tire proc€ssbears an imprint indicatingthat it was The anterior(i.e. toothed)section ofthe dentaryis relatively the c6ronoid overlappedby the dorsalprocess of the dentaryposterior ra- mus (seeabove). The surfaceof the Meckeliancartilage is orientatedposterodorsallY. Hyobianchial skeleton.The only referablebony ele- ment is MMP.2005'403, a right first ceratobranchial (Fig. 3G). This is curvedmediodorsally, the headis slightly OititeO ana has an oval shapeand the stem is thickened distallyand slightly flattened dorsoventrally. The distal plate 'presumably contacts the epibranchial. Atlas-.The availablematerial consists of atlantalcentra from individualsof varioussizes (Fig. lJ, K). Centrumlength approximatelyequals centrum width. The subcentralarea is an'o*-sttapeOund .on.uu" ventrally,displaying several pits borderedby bony ridges.The articularsurface of the paired atlantealcondyles is faintly convexand elliptical in shape' The odontoid processis relatively wide and displaystwo well separatedand shallowly convexventrolateral articular surfaceiof oval shape.The atlantealcotyle is circular' The foramina for the exit of the first spinal nervesare situated 4a M. VENczEL

l,ilft liri,lr,ii, 1,:r

(MNCN19275);B' cynops pyrrhogastel (MNcN Flgure4 Premaxillaeof Recentsalamandrids in posteriorviews' A, Ttiturusmormorufus (MNCN zi1zz\C,Tarichagronulosa (MNCN u83o); D, Ommatottitonviffotus (MNCN 4o462);E, Lissotriton helvetrcus t9265); (MNCN Abbreviations:dp, t, paromesotritonhongkongensis (MNcN zlssz); G,calotriton asper (MNCN t6tiz)tH, Lissotritonboscai r8r87). dorsalprocess; lh, lonino horizontalis; pd, pors dentoris, Scale bar = z mm'

posteriorto the ventrolateralmargins of the atlantealcon- areais completelydelimited by a sharpdorsolateral crest, lam- iyles. In one atlas(MMP.2005.409) the odontoidprocess is which runs from the posterolateralborder of the neural rither smalland asymmetriclacking its left lobe and the left ina to the diapophysis,The postzygapophysealcrest extends articularsurface on its ventrolateralside, respectively. anterodorsally,merging into the dorsolateralcrest. On the Trunk vertebrae.MMP 200538A1 (Fig' lM-P) rep- right lateralside, the dorsolateral crest is comparativelylower without resentsan anterior trunk vertebrain which both prezyga- in height.In anteriorview, the condyle is circularand pophysealareas and the right transverseprocesses (= rib a notochordalfossa. The neuralcanal is pear-shaped,while edge bearers)are broken olT. In lateralview, the centrumis opis- the neural spineis extremelyhigh with a thin leading ventralview, the thocoelousand moderatelyelongated. The condyle is re- anda somewhatthickened dorsal surface. In con- latively small, rounded and separatedfrom the vertebral centrumis somewhatconstricted medially betweenthe extendbetween centrumby a distinctconstriction. The transverseprocesses dyle and the cotyle. Prominentbony ridges andthe vent- arewidely spacedand, except for their distalend, are con- theventrolateral sides of thecondyle and cotyle resulting nectedwith a bony lamina. The rib-articulatingsurface of ral surfaceof the parapophyses,respectively. The diamond-shapedsubcentral area is deeplyconcave and pen- etratedby severalforamina of different size' In dorsalview, the neuralarch is relatively short.The dorsalsurface of the neural spine is relatively thin with a gradualposterior en- largementbearing pit and ridge sculpture.In posteriorview, thecotyle is deeplyconcave and circular.MMP. 2005.382/2 closelyresembles MMP. 2005.382/l(Fig. lQ) but hasmore HUNGARY NEW SALAMANDRIDFROM MIOOIT MIOCEHE OF 49

Ilium. The iliac shatl is extremelylong, bearsa line longitudinalstriation on its lateralsurface and is roughlythe sarie thicknessalong its length with somewaisting in the preacetabulararea (Fig. 68, E). In dorsal or ventral view itt" iliu. shaftis curvedmedially, the acetabulumis oval and

to the femur. Ischium. Available specimens (MMP'2005'402/l' 4O212)come from relatively small individualsand display C a very simplemorphology. The ischiadicplate is slightly elongited with a convexmedial margin; the ischiadicpro- cessis orientatedposterolaterally and thereare two small foraminapiercing the ischiadic plate near its base. Femur.Inanterolateral or posteromedialview, this bone hasa faintlysigmoidal shape. In larger(i.e. older) individu- als the epiphy.sesare completelyossified (Fig' 6C)' The

Flgure5 DentariesofCorpothottiton matroensis gen' et sp' nov' fromM5traszdlds z.A, MMP.zoos.393lr dentary; B'D' MMP.2oo5.393/2dentary; €,MMP.zoo5.393l3 dentary; E'F' MMP.2oo5.437lrdentary. A,C, D, E, lingual views; B, F, labial views' Scalebar=rmm.

ovoid articularsurfaces. At leastsome were provided with

afticular surfacerounded and connectedto the moderately prominentventral humeral crest (Fig. 6A)' The surfaceof ihe humeralcrest, especially in largerindividuals, is covered with lateralstriae and pierced by severalsmall foramina' The dorsalhumeral crest is a thin and rather short bony lamella of roughlytriangular shape, its heightusually diminishing matroensisgen. et sp' nov'from well aSov-ethe livel of the ventral humeralcrest' The distal Flgure6 Corpathotriton z.A, D, MMP.zoos.398/r, 398/z humeri; B' E' sectionof thehumeral shaft is slightlywidened and provided M6traszdlds leftitium; C, F, MMP'zoo5.4o5lr femur' A-C, lateraI with a rathershallow ventral cubital fossa(Fig' 6D)' The MMP.zoo5.4or/r IFF,ventral views. Scale bar = 1mm. entepicondyleis slightlylarger than the ectepicondyle' views, 5o M. VeHczrl headof the boneis roundedand slightly thickenedwhile this has a completefronto-squamosal arch with secondary the ventralmargin is concaveand delimits a relativelyshal- sculptureon the frontals,parietals and squamosals. From the low ventraldepression. The trochanterpossesses a spur-like figureof Rodek(I996a: text-fig.5) theonly discemableele- processconnected to a well-definedcristatrochanterica run- mentsof thehead region are the epibranchials. Finally, com- ning alongthe anteroventralmargin of the femur (Fig. 6C' paringthe orientation of theappendicular skeleton and tail in part F). The distal part of the shaft is moderatelywidened and theabove hgures, it is clearthat Estesand RoEek figured dorsoventrallyflattened with a shallowdistal embayment. and counterpartof the samespecimen and cameto different MMP.2005.4I l/1-3 representwell-preserved femurs, each conclusions.In a differentarticle, Rodek (1996b:196) stated with a tuberosityjust below the trochanter.Because there is thatBrachycormus is closely relatedto Chelotritonand that no fractureline present,the femoralshaft was presumably both havea fronto-squamosalarch. Thus, it is not clearyet deformeddue to pathologicalproliferation of bonytissue. if the materialreferred to Brachycormasby Estesand Rodek representtwo different genera(one closeto theTylototriton REMARKS.The frontalin Carpathotitonis like thoseseen groupand the other close to Triturus)or differentontogenetic in somemembers of Lissotriton (e.g. L. boscai andL. heL- stages(including pre- andpostmetamorphic individuals) of veticus')and Ommatotriton(O. ophryticus and O. vittatus) thesame genus. with a completefronto-squamosal arch, but its posterolateral The fused premaxillaeassigned to Carpathotritonare processis stouterand tendsto widen distally(Fig. 3A, B)' unique in having an undevelopedpalatine process. The In othersalamandrids the frontal bears a secondarypustular premaxillaeare paired in Archaeotriton,Chelotriton, Chio- sculpture,asinrybbtiton andrelated genera (e'g. Chelotri' glossa,Echinotriton, Mertensiella,Salamandra, Salaman- ton,Echinotriton: Nobel 1928;Westphal 1978, 1980; Estes drina, Pleurodelesand Tylototriton, but they are fused in l98l: Bailon1989, l99l; Haller-Probst1998), or a pit and the remainingsalamandrid genera. In mosttaxa with fused ridge sculptureas in Cynops,Pachytriton, Paramesotriton premaxillaethe distal sectionof thedorsal processes end free andSalnmandrina (Chan et al. 2001; pers. obs') (Fig. 2E' with their stem laterally compressed.In contrast,when the I). The sculptureis somewhatattenuated in Pleurodeles, dorsalprocesses are fused distally, as exemplifiedby Calo' Taricha(Fig. 2A, F) and in membersof the Trituru,tgroup triton, Paramesotriton,some Ussolriton and Triturus, they (exceptT. marmoratuswhich has an extensivedorsal sculp- are dorsoventrallyflattened (Fi-e. 4F-H). Ln Carpathotriton ture:Fig. 2D) and,usually, is reducedto the orbitalmargin thedorsal process is reminiscentof newtshaving distally free in Calotriton,Euproctus, Lissotriton, Mesotriton, Ommato- and divergentpremaxillary dorsal processes, as observedin triton (Fig. 28, C, G) andCarpathotriton (Fig. I B' D). The Taricha,Triturus and in some speciesof Cynopsand Lis- frontal is devoidof sculpturein membersof theSalamandra sotriton.However, in Carpathotritonthe imprint left on the group.The dorsalsurface of the frontal is flat or slightly anteromedialside of the frontal by the premaxillarydorsal concavein the Tylototriton Eroupand in Cynops,Pachytri- processsuggests that the distal section of thelatter was dor- ton,Paramesotriton and Pleurodeles.In the remaining gen- soventrallyflattened. era and Carpathotritonthe dorsal surfaceof the frontal is The parietal in Carpathotriton has a taperinglateral more or less convex. In the membersof one Tylototriton process,which probably contactedthe squamosalover the group the sculpturecovers the fronto-squamosalarch (e'g. otoccipital,as in severalother salamandridgenera (e.g' seeRodek 2005: text-flg. 28, C) and the supraorbitalcrest Salamandra,Cynops, Calotriton, Neurergus, Parame sotri- is extremelyprominent. Comparison with Brachycormusis ton, Pleurcdelesand membersof the Tylototriton group: equivocalbecause of conflictingpublished inlerpretations. Duellman& Trueb 1986;Haller-Probst & Schleich1994; The monotypicB. noachicuswas seenby Estes(1981) as a Chan et al. 2001). In the Tylototriton group, the parietal 'Group memberof II' salamandersand included with Pleur- dorsalsurface bears a secondarypustular sculpture, while in odeles,Tylototriton and relatedextinct genera(e'g' Chelo- someoriental genera(e.g. Paramesotriton)a pit and ridge triton andPalaeopleurodeles). In the absenceof a holotype sculptureis present. Estes(1981) designated specimen SUB 1307as the neo- The orbitosphenoidin Carpathotriton, comparedto typeof Brachycormusnoachicus and gave the following dia- othernewts, has a rathersimple morphology, with a relatively gnosis:'skull longerthan broad;fronto-squamosal arches smalloptic nerve foramen, a posteriorlysituated occulomotor well developed;skull roof broad,sculptured with inegular nerveforamen and no posteromedialextension. In all newts, dermalbone that on the squamosalsbecomes pustular; ver- including Carpathotrito,n,the optic nerveforamina are situ- tebraewith relativelyhigh neuralspines capped with pus- atedwithin theorbitosphenoid border (Fig. 7). In themem- tular dermalbone.. .'. Rodek(1996o), reviewing the type bersof Lissorri tun, M es otr ito n, Ommat o t ri ton, Tritur us and, materialfrom Orsberg,Germany, observed that the neotype to a lesserdegree, in Cynops,Paramesotriton and Taricha, designatedby Estes(SUB 1307: PIUB Ro 4429),is in fact the posteriorsection of the orbitosphenoidis producedinto theholotype figured by Goldluss (1831: pl. 13,figs 6' 7)'The a distinct posteromedialextension facing posterodorsally. In 'Brachycor- diagnosisgiven by Rodek(19964:481) stated: theSalamandra group the occulomotor nerve foramen is situ- nus is closelyrelated to Triturusbut differsfrom it in that ( 1) atedclose to the posteriorborder or is sometimesmarginal it is neotenous,with ossifiedhyobranchial skeleton, specific- (pers.obs.). ally the copulaanterior, ceratobranchial l, ceratohyaland The squamosalin theSalamandra group is rathersimple epibranchials,and with extemalgills apparentlypresent, and with a relativelylong ventralramus, but without well de- (2) the skeletonshows incomplete ontogenic development, velopedanterior and posteriorrami. The rami in Mesotriton with frontotemporalarch absent. . .'. Moreover,both authors andTriturus are only slightly expanded,while in Lissotriton alsoreferred to BMNH 30268,which representsa complete theposterior ramus is alwayslonger than the anterior ramus. salamanderskeleton in dorsalview (Estes1981: 79; Roiek In Pleurodelesthere is a long anteriorramus and a relatively 1996a483).Accordin-e to thefigure of Estes( 198l: tig.20A) reducedposterior one. In Cynops,Paramesoffiton, Taricha NewsaumnNDRlD FRoM Mtooue MtoceHe or HuNennv ,t

E*3

C' rids.A, corpothotritonmatroensis gen. et sp.nov' (MMP'zoo5'l8g/t); B' Ffgure7 orbitosphenoidsof fossiland Recent salamand (MNCN granuloso(MNCN il83o); F' Triturusmarmoratus Lissotritonhelveticus(MNCN r9z6s); D, Cynopspynhogosfer 48zz);E,Toricho z3lsz); ''ommatotritonv'ttafus (MNCN 40462); (MNCNrgzzs); G, Lissotritonboscai(MNCN r8r8z); H, Poramesotritonhongkongensis(MNCN pme,posteromedial extension' onf,optic nerve foramen; ocnf, occulomotor nerve foramen; l, pleurodeleswaltl (MNcN rg64g). Abbreviations: A,G-1, lateralviews; B, dorsal view. Scale bar = z mm'

dra salamandrathe prearticular + angularwas not fusedwith

Chio glos sa,M ertensiella, Salamandra, P le urode le s andTrit' spineis moderatelyhigh in the membersof "itifn"neuralLissotriton,Mesotriton, Ommatotriton,Calotriton and 54- Iamandrina,butis very high in Cynops,Pachytrtton' Para' mesotritonand Archaeotriton (Fig. 10)' In Carpathotitun and Archaeotriton the humeral length/distalwidth and the femoral lengtlr/distalwidth ra- tios'are distinctly higher than in other salamandrids'while 52 M. VEtczEt-

motmorutus(MNCN tgz75);8, illesotriton olpestis (MIC Ff$re E Squamosalsof Recentsatamandrids in right lateral view. A, Triturus pleurodeles helveficus(MNCN tgz65);E, Taricha gronulosa (MNCN rr83o); F, Lissottitonboscoi 44So);C, woltl (MNCN196z18); D, Lissotriton ramus;pr, posterior ramus; vr' ventralramus' scale (MNCNrgrgz); G, Cynopspyrrhogaster (MNCN 23823). Abbreviations: ar, anterior b?r= 2 ml|l.

(see Appendix and Table l). Characterscores the relativelength of the ilium is similarly higher'In both drid taxa mainly on first hand examinationof disso- Carpathotritonand Archaeoffiton these three attributes may wcre based skeletonsor on alcian blue and aliz- relaieto their adaptationto an aquaticmode of life' ciated salamandrid arin red colouredspecimens, but some data were obtained from the literature (Duellman & Trueb 1986',Zhao & Hu 1988;Haller-Probst & Schleich1994; Roiek 19964; ETlcRELATIo NsH IPS B6hme 1998;Haller Probst1998; Rage & Hossini2000; PHYloeen Chanet al.200l). 'all OFCARPATHOTRITON A hypothetical zero' ancestorand Salamandra sa- 'true' lamandra,a memberof the salamander'sgroup' were designatedas outgroups,and the most parsimonioustrees werJfound usingthe branch-and-boundalgorithm in PAUP* (Swofford 2001j. Polarity decision for characterstates is basedon outgroupcomparison (Bryant 2001)' All charac- ters were unorderedand equally weightedand both the ac- celeratedtransformation (ACCTRAN - favouringreversals over convergencesif both interpretationsare equally prob- - able) and dilayed transformation(DEUIRAN favouring convergencesover reversals if both interpretationsare NEwsalnneNDRtD FRoM Mtoole MtoceHe or HuncRnv 53

'all ingroup Tabler Datamatrix of 35characters for two outgroup taxa (Solonondro solamondro and an zero'hypothetical ancestor) and 13 taxa,

Characters ooooo ooool 11111 77772 22222 22223 33333 o/o Terminaltaxa 72j45 6t89o 12345 6789o 12345 6789o 72345 missing Hypotheticalancestor ooooo ooooo ooooo ooooo ooooo ooooo ooooo o Solomandrosolamandra oo110 00000 ooooo ooo10 o1001 00000 ooooo o Pleurodeleswaltl ooool 10000 oo100 10100 10020 00101 10101 0 Tylototriton oooo2 11100 10110 00100 10010 00111 10111 0 Echinotriton oooo2 11100 10111 00100 10010 00111 10111 0 Salamandrino ooool 10011 oo110 00111 10021 10000 ooooo o Calotritonaspel 11111 10101 o2110 10111 11110 11111 71127 0 Ommatotriton 11011 10111 o2110 10110 10120 11111 7 7 7 27 0 Lissot ti ton he lv et i cu s 11111 10111 o2010 10110 70720 11111 77727 0 Mesotiton 11111 00011 o2000 10110 10120 11111 77727 0 Triturus 11111 00011 o2000 10110 10120 11111 77127 0 Carpothottiton gen. nov. 10111 1010? oo111 ttt?? ????? ??t?? ????7 46 ?. Archaeotriton ooool 10011 o?1oo ?tt?? ??7?t ?????.? ? ? ? 51 Cynopspyrrhogoster 10001 10100 o1110 10100 10120 11111 77727 0 Poromesotiton 77t72 11100 01110 10110 10120 11111 t? tzt 3 o, t, 2,r€present character states; ?, unknown. The final column lists the percentage of missingrecords.

equally probable)optimisations were applied in mapping tentionindex:0.81: Fig. ll). The monophylyof newtsis the distdbutionofcharacter states on trees'The branch-and- well supported(bootstrap:937o, de'cay index:4 steps)by boundsearch was used also for the bootstrapand decayana- six derivedcharacter states [5(l), 6(l), l3(l), 18(1),2l(l), lyses.Bootstrap values resulted from 2000 replicates,while 24(l)l indepndent of optimisation(Fig. 12).Carpathotriton the decayindices (steps)of Bremer (1994) were obtained forms an unnamedclade along with Cynops,Paramesotri- by a searchfor trees successivelylonger than the shortest ton, Calotriton, Ommatotrito n, Lissotriton,M esotriton and tree until all branchescollapsed and the consensusof all Triturus.This clade is supportedby six apomorphiesusing cladogramsappeared as unresolved. both theACCTRAN andthe DELTRAN optimisations[l(1)' The branch-and-boundsearch yielded a singleshortest 23(l), 26(l), 27(l), 32(l), 34(2)I andtwo morecharacters by tree (tree length:62 steps,consistency index:0'63, re- eitherACCTRAN [ 12( I )] or DELTRAN optimisationI I 6( I )]

FlgulegRightdentariesofRecentsalamandridsinlingualview.A,Poramesotritonhongkongensis(MNCNzllsz);B,Lissotritonhelveticus (MNCN16rlz); l, Triturus (MNCN19265); C, Ommatottitonviftatus (MNCN qoq6z\ D, Tarichagronuloso (MNCN fl83o); E,Colottiton asper (MNCN Lissotritonboscoi (MNCN t8r8Z). mormoratus(MNCNr9zZ5); G, Cynopspyrrhogaster (MNCN nSzz); H, Pleurodeleswotfl rg6+8),l, Abbreviations:dp, dental parapet; mg, Meckelian troove; pr, posterior ramus. Scale bar: z mm. 54 M. VENczEL

(MNCNr9z7l:J;c'F' D,Tarichagranulosa (MNCN u83o); B, E,Triturus mormoratus Ffgurero Trunkvertebraeot Recentsalamandrids. A, (MNCN Lissotritonhelveticus (MNCN 19265); K' !' parunesotriton zlssz); G,n, cynopspyriogaster z38zz);l't' hongkongensis(MNCN views;D-F' H' diapopnysl;pa, parapophysis; ns, neural spine. A-c, G, l, K,dorsal I' ommatotritonyitfotus(MNCN 4o452). Abbreviations: da, lateralviews.Scale bar = z mm.

50Voof the included characters\n Archaeotritoncould be

semblageor remain unresolved.A more extensrvemor- phofogftatdata set (includingcranill,. axial.and appendic- ult ."h*u"r".s) is, therefore, needed to obtain a robust solution. NewselnmaNDRlD FRoM Mtoot-E MtocrHe oF HUNGARY 55

PRmeoEnvlRoNMENTAL AND PALAEOGEOGRAPHICALIMPLICATIONS

from M6trasz6lSsI the frequencyof Palaeobatrachusvar- ied between 28.4-37'7Vo, while that of Rana esculentais between65.1-74.37o.In Mdtrasz6l6s 2, thefrequency of Pa- Iaeobatrachusranges-betweenbetween 8.8-16.37o, while that of Raza esculentaranges 52.7-55.7Vo.The frequencyof otheranurans was much lower. Latonia (2'5-9'3Vo), Disco'

Flgureu Shortesttree based on 35 informativecharacters scored forr4 salamandridterminal taxa and a hypothetical'alIzero' ancestor' Indicesof supportfor clades are given to theteft of eachnode (bootstrapvalue (o/o)) for zooorepticates, while the decay index (steps)is givento theright of eachnode'

Salamandnsalamandn 3(l) 4(t) t90) 22(l)25(1) z0(l)26(l) 14(r)le(r) Salamandrina 5(l) 9(r)r0(r) re(I) zo(I)2s(l Archacotrinn 6(l) r3(t) Pleurodeleswaltl E(l)l6(r) lE(l) Tylototiton verrucosus 2r(l) 28(l)30(l) 24\1' 34(t) 3r(l)33(l) 8(t) 5(2)?(l) 1l(l) t6(0)24(r) Echinotriton 35(l) r4(l) Cynopspwhogaster 29(r) r(t) t2(t) 34(t) l6(l)23(t) Carpathottina ,2(0)r5(l) l7(l) 25(1\ 21(t) Paramcsotnton 32(r)34(2) 4(l) C,alotritonasryr t9(t) 20(1>22(r) 24(r) Ommatotnton

Lissotiton helveticus

Mesotriton

Tituns cristatus

optimisations'characters in italics of derivedcharacter states mapped using ACCTRAN and DELTRAN Fiture12 shortesttree with distribution in boldare in Romantype are supported by DELTRANoptimisation only; characters aresupported by AccrRAN optimisation only; characters supportedbY both oPtimisations' 56 M. VENczEL

studied.Dr BorjaSanchiz imparted much un- Desmanellasp.probably also lived in thesunounding area of fossillocalities dataand helped with producingthe micrographs' freshwaterlaies (G6l et al. 1999b).Based on the faunallist published micrographswere processedby L. Tormo at the publishedby Hir & K6kay (2004)the rodentfauna was rather fne Snfr4 MuseoNacional de CienciasNaturales, Madrid. Dr J' E' iin".t" (Eurotagusfontannesi, Spermrtphilinus bredai, M us- kindlyfacilitated the study ofthe recentcollections cardinus aff. sansaniensis,Eliomys truci' Bransatoglyssp" Gonz6lez at the Museo Nacional de CienciasNaturales, Eomyopsoppligeri, Keramidomysmohleri, Megacricetodon in his care grateful to two anonymousreviewers and to minor,' Democricetodon mutilus, D. cf .freisingensis, Criceto' Madrid, I am B. Smith for providing very helpful comments don sp.,Eumyarion medius, Anomalomys gaudryi) but' with Dr Andrew the eiception of Megacricetodonminor they are very infre-

RerenEucrs Anderson, J' 1871. Description of a new genus of newts from western of the Ztological Societyof lnndon l87l: 423- of planktonicalgae (Spirogyrascorbicu- Yunan.Proceedings the abundance 425 braunii) and the cormophyte Myrica lata, Botryococcus Arntzen, W & Olgun, K.2000' Taxonomyofthe bandednewt, Trllurus 2002)' J. (Szuromin6-Korecz& Nagy-Bodor virtctrs: morphologicaland allozyme data Amphibia-Reptilia2l: 155- an accu- Sedimentationat M6trasz6l6s I startedwith 168 Bailon, S' 1989. Les amphibienset les reptiles du Pliocdnede Balaruc II (H6rault, F rdnce\.Pa I ae ov e rt eb rata 19: 1-28. - t99 I . Amphibienset reptilesdu Pliocine et du Quaternairede France et d'Espagne: mise en place et evolution des faunes PhD Thesis: Universityof ParisVII, Biologieet Sciencesde la Nature;499 pp' Bocage,J. V. B, 1864.Note sur un nouveaubatracien du Ponugalet sur une grenouillenouvelle de I'Afrique occidentale'Revue et Magasinde Zoologie Pure et Appliquie 16:248-254. fauna(licshwater molluscs, Iishes, newts' remainsol local Bell, T. I 839. A History of British Reptiles.Van Voorst,London, 142 pp' Bolkay, St. J' 1928. Die Schlidel der Salamandrinenmit besonderer Riicksicht auf ihre systematischeBedeutung T*itschrift AnatomieEn' nvicklungsge s chichte E6: 259-3 19. Biihme, M. 1998.Archaeotriton basalticus (v Meyer, 1859) (Urodela' Salamandridae)aus dem Unteroligoziin von Hammerunterwiesenthal (FreistaatSachen). Abhandlungen des StaatlichenMuseums fiir Min- eralogie und Geologie zu Dresden 43144:265-562' - 2003. The MioceneClimatic Optimum: evidencefrom ectothermic vertebratesof Central Eut ope Palaeog'eography Palaeoc limatology PalaeoecologY195: 389-401 - & Ilg A' 20O3' FosFARba.re.Available at: www wahre-staerkecom/' - & Riissfer, R. 2002. Fund eines zweiten Skelettes von Archaeotri' ton basalticus(Urodela, Salamandridae)aus dem Unter-OliSoziinen Maar von Hammerunterwiesenthal(Erzgebirge)' Verdfentlichungen Museumftir Naturkunde Chemnitz25: 63-68' Brtmer, K. 1994. Branch support and tree stability' Cladistics 10: 295- analysisshows that Carpathotriton is closelyrelated to some 304. polzuity and the rooting of cladograms nsl atic f orms(Cy nop s, Pachy t riton, Parame s o t ri ton)' Based Bryant, H' N. 2001. Character Pp. 319-338 in WagnerG. P. (ed.) The Character Conceptin Evolu' on mitochondrial-DNAsequence analysis, the earliest di- tionary Biology. Academic Press,New York' R' Caccone, A., Millnkovitch, M. C., Sbordoni, V' & Powell' J' 1994.Molecular biogeography:using the Corsica-Sardiniamicro- plate disjunction to calibrate mitochondrial rDNA evolutionaryrates in mountain newls (Euproctus)' Joumal of Evolutionary Biology 7: 227-245. Carranza, S' & Amat, F. 2005. Taxonomy,biogeography and evolution of Euproctus(Amphibia: Salamandridae),with the resunectionof the gewi Calotriton and the description of a new endemic speciesfrom ih" Ib".iun Peninsula.Zoological Journal of the Linnean Society145: 555-582. of the Chan, L. M., Zamudio, K. R. & Wake, D' B' 200l Relationships salamandrid generaParamesotiton, Pachytriton' and Cynops based on mitochondrial DNA sequences.Copeia 2001: 997-1009' Duellman' W. E. & Tfueb, L' 1986 Biotogv of Amphibians' McGraw- Hill'NewYork'670pp AcTTowIEDGEMENTS Dumbril,A. M. C' 18O6'Zoologie analy'ique' ou mithodenaturelle de des animaru,rerulue plus Jbcile d I'aidede tableaux I am deeply indebtedto Dr JiinosH(r who made the material tlttssiJication synoptiques.Paris, 344 pp' availaute for study and provided relevant information on the NewsnlemnNDRtD FRoM Mtoole MtocEHE or Hurucenv 57

-2O4 In Hungarianwith English Estes, R. l9El. Gymnophiona, Caudata. Hendbuch der Paldoherpe- M egye i M rtzeumok Ev kbnyv e 22 17| [ tologie Part2, GustavFischer Verlag, Stuttgart, I l5 pp. abstractl. -, -, 2001. El6zetes -, Hecht, M. K. & Hoffstetter, R. 196T Paleoceneamphibians from Venczel, M., G6l, E. & Kessler, E. leldhelykomplex Cemay,France. American MuseumNovitates 2295" 1-25. besz6mol6a felsdt6rk6nyi'Giiddr-kert' n. 6sl6nytani 25: 4l- - & Holfstetter, R. 1976.Les uroddlesdu Miocdne de Ia Grive-St-Alban fjravizsg6lati4r6l.Folia Historico Naturalia Musei Matraensis (fsdre,France). I ulletin Musium National Histoire Naturelle (Sciences 64. [n Hungarianwith Englishabstract]. Pliocenelvanovce local- de Ia Terre) 398: 297 -343 Hodrova, M. I 984 Salamandridaeof the Upper - Fftzinger, L, J. F. J. 1826.Neue Classifit'atittnder Reptilien nath ihren ity (Czechoslovakia).Acta Universitatis Carolinae Geological9U: Natiirlichen Venuandtschaften. N ebst einer Verwandtsc hafts -TafeI und 331-352. einem Verzeichnisseder Reptilien'Sammlungdes k k. zoologischen Kordos, L. 1981. A hasznosi felso-mioc6n gerinceslel6hely kora Museumszu tlien. J. G. Huebner,Vienna, 66 pp. eml6szon6ci6 alapjdn. Magyar Allami Fdldtani Intdzet ielentise with English G6l, E,, Hfr, J., Kesster, E., K6kay, J., M6szdms, L. & Venczel' M. az t979 ivr6'1, pp. 459463. [tn Hungarian 1999a. A M6traszo'l6sl. Lel6hety kdz6ps6 miocdn dsmaradvdnyai abstractl (Cricetidae' (Riivid szinopszis).Ndgrdd Megyei Milzeumok Evkdnyve23: 4148. - 1986. A hasznosi6s szentendreifelso-mioc6n hiircstigiik 6s r6tegtani vizsgillata. Magyar Atlami [In Hungarianwith English abstract]. Mammalia) rendszertani -, - & - 1999b. Kiiz6psd-miocdn 6smaradvdnyok,a Fbldtani Intizet jelentise az I 979. ivrol, pp. 523-553' [In Hungarian M6traszdl6s,R6k6czi-kdpolna alatti ritbev6g6sb6l.l. A M6traszcil6sI with English abstractJ lel&rely Folia Historico Naturalia Musei Matraensls 23: 33-78. [ln Larsen, A., Weisrock, D. W. & Kozak, K. H.2003' Phylogeneticsys- Hungarianwith English abstract.l. tematicsof salamanders(Amphibia: Urodela), a review.Pp. 3l-108 in -, -, -, -, - & - 2000. Ktiz6ps6-mioc6n dsmaradviinyok a D. M Sever.(ed.) ReproductiveBiology and Phylogenyof Urodela- M6traszdlds,Rrik6czi-kiipolna alatti ritbevdSdsb6l.II. A Mritrasz6lds2 SciencePublishers, Enfield (New Hampshire)& Plymouth lefolrely. Folia Historico Naturalia Musei Matraensis A:39-75. lln Laurenti, J. N. 1768. Specimen Medicum exhibens Synopsin Rep' Hungarianwith Englishabstractl. tilium emendatamcum experimentis circa venena et antidota reP- Godlnoi M., Broln, F. de, Buffetaut,li', Rage, J. C. & Russel,D' 1978 tilium austriacorum. Joan, Thomae Nob. de Trattnern, Vienna, Dormaal: une des plus anciennesfaunes 6ocdnes d'Europe Comptes 214 PP Rendusde I'Acadimie des Sciences,Paris,D 287: 1273-1276 Lltr'lnchuk, S, N,, Zulderwflk, A., Borkln, L. J. & Rosanov,J. M. 2005. Goldfuss. G. A. 1820 Hantlbuch der Zoologie Vol 2' JohannLeonard Taxonomic status of Triturus vittatus (Amphibia: Salamandridae)in vertebrae sizeand allyzome data. Schrag,Niimberg, 696 PP westernTurkey: trunk count, Senome -323. - I 83l. Beitrlgezur Kenntnissverschiedener Reptilien der Vorwelt. Nava Amphib ia-R e p til ia 26: 305 Acta Physico- M edica Academia Caesareae k opoldino- Carolinae Milner, A. R. 2000. Mesozoic and Tertiary Caudataand Albanerpeton- (eds) Akodemieder Naturforscher 15: 6l-128 tidae. Pp. 1412-1444in H. Heatwole & R. L. Carrol Amphibian The Evolutionary History of Am- Gray, J. E. l85O Catalogueof the Specimensrf Amphibia in the Collec- Biology. Volume4, Palaeontology, tion ofthe BrirtshMuseum Part II. Batrachia,Gradientia, e/c.Trustees phibians. Surrey Beatty & Sons,Chipping Norton 43-275 in RamosM. A. of the British Museum, London, 72 PP. Montori, A. & Herrcro, P. 2004. Caudata. Pp. - 1858.Proposal to separatethe familiy Salamandridae,Gray' into two et al. (eds\ Faunalberica, Volume24,Amphibia, Lissamphibia.Museo families, accordingto the form of the skull' Proceedingsof the Zoolo- Nacionalde CienciasNaturales, CSIC, Madrid. gical Socieryof Inndon lE58: 136-144. Nav6s, L, 1922. Algunos fosiles de libros (Teruel). Boletin Sociedad Haeckel, E. 1866.Generale Morphologie der Organismen Vol. 2, Verlag Ibeica Ciencias Naturales2l: 52-61. offossil and recentsalamanders (Am' von Georg Reimer, Berlin, 462 PP Naylor, B, 1978.The systematics verlebral column and Halfer-Pmbst, M, 1998 Studieson Tylototriton verrucosus(Amphibia: phibia: Caudata),with special referenceto the of Alberta, Urodela). Contributions to the osteology of rybbtilon veruucosus trunk musculature.Unpublished PhD Thesis:Univenity Anderson,l87l and T. shaniingNussbaum et al., 1995(Amphibia' Edmonton;857 pp Franceschi, D. de, Caudata: Salamandridae) Verdffentlichungen aus dem Fuhlrcu- Nel, A., Pliieg, G, de, Dejax, J., Dutheil, D., Aug6' M.' Museum4:35-56. Gheerbrant, E,, Godinot, M., Herret, S., Menler, J' J., Hua, S., Haller-Probst, M. & Schleich, H. H' 1994 Vergleichendeosteologische Bignot, G., Cavagletto, C., Duffaud, S., Gaudant, J., C.' Beuchet' R Untersuchungenan einigenUrodelen Eurasiens. (Amphibia: Urodela, Jossang,A., Broin, F. L. de, Pozzi, J. P', Palcheler, J. ir plantes, Salamandridae.Proteidae) Courier ForschungsInstitute Senckenberg & Rage, J. C' 1999. Un gisementspamacien exceptionnel (Oise, 173:23-77. arthropodeset vert6br6s(Eocbne basal, MP 7): I* Quesnoy Paris,Sciences Herre, W. 1941.Puluenpleuroclelet huuJlinov. gen. nov spec',ein fossiler France).Comptes Rendus de I'Acaddmiedes Sciences, Schwanzlurchaus dem Miozlin von SiiddeutschlandTnologischer An' de la terre et desplanets 329:65-72. Amphibia of zoogeographicimport- zeiger 134: l-17 Nobel, G. K. 1928.TWo new fossil Novitates303: - 1950. Schwanzlurcheaus dem Paleociinvon Walbeck. Taologischer ancefrom the Miocene of Europe.Ameican Museum AnzeigerSupplement 145: 286-301 l-13. of the salamandrid - 1955. Die fauna der mioziinen Spaltenftillungvon Neudorf a d Nussbaum, R. A. & Brodie, E. D. 1982.Partitioning (Amphibia: with descriptionof March (eSR.), Amphibia (Urodela) Sitzungberichte Osteneichische genusTylototiton Anderson Caudata) Al

- 199fu. The salamanderBrachycormus noachicus from the Oligocene Cranial characters of Europe, and the role of neoteny in the evolution of salamanders Palaeonto lo gy 39: 47'1495. l. Premaxillae:paired (0), fused(l) (Wake&6zeti 1969 - 1996b.Skull of the neorcnicsalamandrid amphibian Triturus alpestris ill;Tirus & Larson1995 [l];Wang & Evans2006 [t]). and abbreviateddevelopment in the Tertiary Salamandidae.Joumal 2. Premaxillaralary processes:free (0), at least50Vo of of Morphology 230: 187-197. theirlength fused (l). - palaeontographia 2005. Late Miocene Amphibia from Rudabdnya. 3. Premaxillary-frontalcontact: absent (0), present(l) I talic a 90 (20O4): | | -29 (Wang& Evans2006 [2, modified]). Sanchlz, B. 1998. Vertebratesfrom the Early Miocene lignite depos_ 4. Nasalconract: presenr (0), absent(l) (Wake& its of the opencastmine Oberdorf (Western Syrian Basin, Ausrria): 6zeti 1969 Titus& Larson1995 2. Amphibia. Annalen Naturhistorisches Museum Wien 99A: 13_ [4]; t5l; Wang& Evans2006 29. 15,moditiedJ). - 5. Frontal & Mlynarski,M. 1979.Remarks on the fossil anuransfrom the polish sculpture:absent (0), weak sculpture present ( I ), Neogene.Acta zoologica CracoviensiaA: 153-174 strongsculpture present (2). Scopoll, J. A,1777. Intoductio ad historiam naturalem sistemsgenera 6. Fronte-squamosalarch:. absent (0), at leastpartially de_ lapidum,plantarum, et animalium hactenusdetecta, caracteribus es_ veloped(1) (Wake & Ozeti t969 t2l; Titus & Larson sentialibus donata, in tribus divisa, subinde ad leges naturae Gerle, 199512,modifiedl). Praga, 506 pp. 7. Parietalsculpture: absenr (0), present (l). Swofford,D. L.2001. (Version PAUP- 4.0b10for 32-bitMicrosoft Win_ 8. Parietal:longer than frontal dows) or equalsit in length(0), SinauerAssociates, Incorporated Sunderland, Massachusetts. shorter(l). Szuromin6-Korecz, A. & Nag5r-Bodor, E. 2OOZ.Edesvizi mioc6n 9. Parietalsquamosal contact: absent (0), present ( osuacod6k 6s Sporomorpha a mdtraszdlcisiR6k6czi-k6polna alatti I ) (Wang & Evans2Cfr6r37D. ftbevdgdsb6l.IMiocene freswater Ostracodaand Sporomorphafrom the road-cut close to the Rdk6czi Chapel ar Mdtraszrildsl.pp. 30, in 10. Maxillarylength: reaches or nearlyreaches the quadrate (0), abstractvolume of 5. Magyar dslinytani V6ndorgyu16s,p6szt6. [In extendsbeyond the eye, or falls shortofthe posterior Hungarianl. marginof rheeye (l) (Wake& 6zeti 1969t3l; Titus& fitus, T. A. & Larson, A, 1995. A molecular phylogenetic perspecrtve Larson 1995 [3, modifiedl; Wang & Evans2006 [3, on the evolutionaryradiation of the salamanderfamily Salamandridae. modifiedl). SystematicBiology 44: I 25-l 5 | . I l. Maxillary-pterygoidjoint: absenr(0), present (l) (Wake Velth, M. & Steinfartz,S. 2004. When non-monophyly resulrsin raxo_ & Ozeti1969 [a0]; Titus& Larson1995 nomic consequences- the caseof Mertensiel/cwithin the t4l). Salaman_ 12. Orbitosphenoidpostero-dorsal extension: (0), dridae (Amphibia: Urodela) Salamandra 40: 67 -80. absent postero-dorsalextension small (l), postero-dorsal Venczef,M,2004. Middle Mioceneanurans from the CarpathianBasin. ex_ PalaeontographicaA. 27l: | 5 | -lj 4. tensionlarge (2). 13. Von Meyer, H. 1860.Salamandrinen aus der Braunkohleam Rheinund Squamosalfrontal process: absent or reducedto a small in Bdhmen.Palaeontographica 7: 47-7j. process(0), long (l). Wake, D, B, & dzeti, N. 1969. Evolutionary relationshipsin the family 14. Squamosalposterior process: absent or reducedto a Salamandridae.Copeia 1969: 124-13j . smallprocess (0), long (l). Wang, Y. & Evans, S. E.2006 A new short-bodiedsalamander from I 5. Quadrateanterolateral process: absent (0), present (l the Upper palaeontologica ). Jurassic/LowerCretaceous of China. Acta 16. Parasphenoidposterior ventral crest: absent(0), pre_ Polonica5l:. 127-13O. sent(l). Welsrock, D, W., Papenfuss, T. Macey, J,, J. R., Lltvinchuk, S. N.. 17. Dentarytooth row: long (0), tooth Polymenl, R., Ugurtas, l'IJ.,Zhao, E,n row abouthalf the Jowkar, H, & Larson, A. lengthof wholedentary (1). 2006.A moleculiuassessmenr of phylogeneticrelationships or shorter and lin- 18. Prearticular+angular: eageaccumulation rates within the family Sa.lamandridae(Amphibia, not fused with articular (0), (l). Caudata).Molecular Phylogeneticsand Evolution 4l: 368-3g3. fused Westphaf, F. 1978. Tylototriton (Amphibia, Urodela) aus dem Ober- 19. Operculum:ossified or mineralised(0), composedof mioz?invon Ohningen.Neues Jahrbuch Geologie pakiontologie Mat- unmineralisedcarrilage (l) (Wake& Ozeti 1969 t5l; thausen1978:49 l-50 l. Titus& Larson1995 [6]). - f980. Chelotriton robustus n. sp, ein Salamandrideaus dem Eoziin Gene ra I anatomic der Grube Messel bei Darmstadt Senckenbergianalethaea 60: 415 al charac te rs 487. 20. Lungreduction: well developedlungs (0), lungs weakly Wolterstorff, W. 1925.Katalog der AmphibienSammlung im Museum developedor absenr(l) (Wake & Ozeti 1969 tgl; fi.ir Natur- und Heimatkunde zu Magdeburg Ersrer Teil: Apoda, Titus& Larson1995 t8l). Caudata.Abhandlungen und Berichte ausdem Museumfiir Naturkunde 21. Skin texture: smooth (0), rough or keratinised(l) und Vorgeschichte,Magdeburg 4: 155-310. (Wake& Ozeti 1969[9]; Titus & Larson 1995 Zhao,E, & Hu, Q. 1988.The classificationand evolutionof theChinese tgl). 22. Epidermis:presence of an epidermallining SalamandridaePp. 12-26in E Zhao,Q Hu, y Jiang& y yang(eds) in the an- Studieson Chinesesalamanlers SSAR, Ithaca,Ny terior half of the femalecloacal chamber (0), absence Zittef, K. A. v. 1890.Handbuch der Palaeont

26. Radii: two pairs ofradii present(0), single pair ofradii 32. The rectus abdominis profundus and the rectus ab- pr€sent(l) (Wake & 6z'eti 1969 [15], Titus & Lanon dominissuperficialis: the musclesare difierentiated and from eachother lee5 [26]). sepamte(0), the musclesare not distinct 27. lntenadial cartilage:absent (0), present(l) (Wake & ( I ) (Wake& O zeti 1969[30] ; Titus & Lanon I 995 ['t0]). Oz,etl1969 [ l6]; Titus & Larson 1995[27, modified]). 33. The geniohyoideusand genioglossusmuscles: they are 28. First ceralobranchial: cartilaginous (0), bony (l) unconnected(0), or attachedby denseconnective tissue (Wake& Ozeti 1969[l8]; Titus & Larson 19951291). at the anteriorends of the ceratohyals(l) (Wake& Ozeti 1969132);Titus & Larson 1995142,modifiedl)' Hyobranchialmus c ulat ure 34. Rectuscervicis profundus: insertion of slipsof therectus 29. Rectuscervicis profundus: inserted via a singlehead (0)' cervicisprofundus may be absentfrom thefirst basibran- with severalinsertions (1) (Wake & Ozeti 1969 [23]: chialand radii (0),present on theposteriorpartofthe first Titus & Iarson 1995[34]). basibranchial(l), or presenton both the first basibran- 30. Hebosteoypsiloideusmuscle: more differentiated(0), chial and radii (2) (Wake & 6z,eti 1969 [34]; Titus & lessdiffeientiated (l) (Wake&6znti 1969t25l; Titus & Larson 1995[44]). Larson1995 [36]). Axial skeleton 31. Inter-ossquadratamuscle fibres: fall entirely short of the raphe(0), few fibrcs extendto the medial rapheor' 35. Atlantal centrumroughly of samelength as postatlan- the muscleis well developed(l) (Wake & 6zeti 1969 tal centra(0), much shorterthan postatlantalcentra (l) [26]; Titus & Larson 1995[37 , modified]). (Wang& Evans2006 [62, modified]).