IntraspeciŽc and nomenclature of theDanube crested , Triturus dobrogicus

SpartakN. Litvinchuk 1,2,LeoJ. Borkin 1

1 ZoologicalInstitute, Russian Academy ofSciences, 199034St. Petersburg, Russia 2 Instituteof Cytology, Russian Academy ofSciences, 194064St. Petersburg, Russia

Abstract. ThePannonian and Delta populationsof Triturusdobrogicus are well separated morpholog- icallyfrom each other.We recognizetwo of theDanube crested newt.The nominotypic subspecies inhabitsthe . Triturusdobrogicus macrosomus (Boulenger,1908), comb. and stat. nov., occurs in theremaining part of the species’ range.

Introduction

TheDanube crested newt has been described as a varietyof the crested newt, Triton cristatus var. dobrogicus Kiritzescu,1903, from lakesin the environsof Sulina, and theDanube River Delta in the northern part of Dobrogeain Romania (Kiritzescu, 1903). However,the name was placedin the synonymy of thenominotypic subspecies of Triturus cristatus (“Tritoncristatus typus”— Schreiber,1912: 110; “ Molgecristata cristata ” — Nikolsky,1918: 202). W olterstorff(1923) created a newsubspecies of the crested newt underthe name “ Tritoncristatus danubialis ”.He consideredthe “ forma dobrogica” as a localDobrogean form includedin hisnew subspecies. Nevertheless, Mertens and Mü ller (1928,1940) regardedboth forms astwo different subspecies, Trituruscristatus dobrogicus and T.c.danubialis .Accordingto these authors, the former subspeciesoccurs in the DanubeDelta, and the latter is distributed over the remaining part of thespecies’ range. Someother authors (e.g., C alinescu,Æ 1931; Fuhn, 1953) did not support this opinion. They believedthat both taxa occurred in the Danube Delta. Indeed, Fuhn (1953) recognized threevarieties of thesubspecies T.c.danubialis inRomania, namely: T.c.danubialis sensu stricto, T.c.danubialis var. dobrogicus and new T.c.danubialis var. intermedia. The two former varietieswere sympatricin the Delta, with a preponderanceof theŽ rst one.Based onthe geographic deŽ nition of subspecies and the general similarity of danubialis and c KoninklijkeBrill NV ,Leiden,2000 Amphibia-Reptilia21: 419-430 ° 420 SpartakN. Litvinchuk,Leo J. Borkin dobrogicus,someauthors synonymized them as Trituruscristatus dobrogicus (Fuhn,1960; Mertensand W ermuth,1960; Fuhn and Freytag, 1961), a solutionthat became widely accepted. Theapplication of biochemicaland cytogenetic methods to the systematics of crested newtsover the last decades has resulted in the elevation of thesubspecies to full species rank(Bucci-Innocenti et al.,1983; Macgregor et al., 1990), and this status was evidenced bydataobtained from studieson thecontact zones of thesetaxa (W allisand Arntzen, 1989; Litvinchuket al., 1994, 1997, 1999). The new concept of the T.cristatus superspecies stimulatedre-evaluation of the taxonomic structure of the “ new”species. Recently, two subspecieswere recognizedwithin T. carnifex and T.karelinii ,respectively(Kalezi c´ et al., 1997;Litvinchuk et al.,1999). The range of T.dobrogicus consistsof two parts, restricted to the Pannonian and Dobrogeanlowlands, respectively (Arntzen et al., 1997; Ž g.1).Based on thesmall range, relativerarity and rapid loss of habitat,Arntzen et al.(1997) considered the conservation status of T.dobrogicus asvulnerable. The species is protected by the Berne Convention (theAnnex II) andis included to the 1996 IUCN RedList of ThreatenedAnimals under thecategory “ datadeŽ cient” (Oliveira et al.,1997). Thegoal of thepresent paper is to re-evaluatethe taxonomic composition of T. dobrogi- cus andits nomenclature.

Materialsand methods

Westudiedthe Pannonian T.dobrogicus from threeY ugoslavianlocalities, and from eleven localitiesin the Ukrainian Transcarpathians ( ZakarpatskayaProvince). The Dobrogean D sampleswere studiedfrom fourUkrainian localities (Odessa Province) in theDanube Delta (Žg. 1). W ealsostudied a Moldaviansample, and a specimenwith uncertain origin (the holotype of Molgemacrosoma ). For morphometricanalysis, we used163 adult specimens of T.dobrogicus (tables 1 and2). The following characteristics were measuredwith dial callipers (to the nearest 0.1mm) for eachindividual: TL istotallength; L isbody length, from thesnout to thefront edgeof thecloacal split; Lcd is taillength, from theanterioredge of thecloacal split to the tailtip; LiE is distancebetween fore andhind extremities; Pa andPp are fore- andhindlimb length,respectively; Lc is head length, from thesnout to the border of themouth angle; Ltcis thehead width between the mouth angles. WI isthe “ WolterstorffIndex” , givenas ratiobetween Pa andLiE (in percents). Principal component analysis was performedfor bothsexes on thestandardized residuals of theregression of naturallogarithm of sizedata withSYST AT5.0(Wilkinson, 1989). Discriminant analysis was performedseparately for malesand females on thenatural logarithm of sizedata with ST ATISTICA 6.0.Lcd was removedin these analyses because of incompletenessof data. Taxonomy of Triturusdobrogicus 421

Figure 1. Distributionof Triturusdobrogicus ,withlocalities studied (1 — UkrainianTranscarpathians: Batevo, Chop,Chornyi Potok, D’ yakovo,Dobroselje, Drisina, Gat’ , MalyeGeevtsy, Minai, Mukachevo, Shalanka; 2 — Yugoslavia:; 3 —Yugoslavia:Radoevo; 4 —Yugoslavia:Obetska Bara; 5—Moldavia:Kagul; 6 — Odessa Province:Reni; 7 —Odessa Province:Izmail; 8—Odessa Province:Vilkovo, Leski).

Anatomicalpreparations (dissection, “ dry”skeleton, alizarin treatments) and larval externalsegmentation were usedfor examinationof thenumber of rib-bearingvertebrae (atlasexcluded). Additional information about color and belly spots distribution was studiedon alive in the Ž eldor in captivity, and on preserved specimens (only patterning).The total sample consisted of 320adults, 21 juvenileand 58 larvalspecimens. In1994-1997, we crossednewts representing various members of the T.cristatus superspecies;within T.dobrogicus ,animalsfrom variousparts of the range were also involved.All crosses were carriedout at room temperature (20-25 ±C)inaquaria 60-100 l volume.Eggs deposited on vegetation were collecteddaily and transferred to shallow bowlswhere development continued. Larvae were keptin identicalconditions and reared in aquaria(20-30 l volume),with density between 10 and40 larvaeper aquarium, depending onthestage of developmentand the size of animals(exclude larval cannibalism). Embryo andlarval survival rates were determinedas the number of hatchingembryos (beginning activefeeding) and metamorphosed juveniles, respectively, in relation to the number of fertilizedeggs.

Results

Theresults of the principal component analysis demonstrated separation of the Danube Deltaand Pannonian populations of T.dobrogicus .Inthe two-dimensional plot (Ž g. 2)we 422 SpartakN. Litvinchuk,Leo J. Borkin s e * u h 0 5 ; . . r 0 0 0 5 9 e 1 0 3 5 0 0 7 4 0 8 0 T . 1 ...... 2 ...... u ) 5 n 0 t . e e 6 3 2 7 . 0 0 6 5 3 1 2 1 3 . 1 0 6 9 9 5 i 9 o n g g 2 r 1 6 6 3 2 2 8 1 7 1 1 8 7 5 2 2 8 1 6 1 4 z o n ------n ------T s i a t a 5 0 5 2 3 0 8 8 1 3 0 5 0 0 2 0 7 5 7 5 i t f ...... c D a R R a o a i 3 1 1 9 5 6 5 7 7 2 5 2 2 9 2 3 5 7 9 2 a t t t l n v 9 5 4 2 1 1 4 1 9 5 4 2 1 1 3 1 l s ) ( n e e e e 7 o l d a D D c 2 t p l d e e e r e m b D b a h 0 6 5 0 2 2 6 5 2 0 a 4 D t u ...... 5 1 9 1 2 7 9 9 1 u d s s ...... n s n 8 4 4 3 2 2 0 0 6 1 4 n n e ( o 8 7 7 4 2 2 0 0 4 1 a a t a 1 a b t 4 t l u D s e D e s s n i , a a D o n 4 1 1 9 2 6 0 9 6 7 4 l ...... a s a n 9 t i D e c 5 1 4 3 0 1 7 8 9 4 . 5 5 0 2 0 9 3 7 4 l e o a ...... b t 1 6 5 3 2 2 5 1 e 0 s e d 4 6 8 7 8 6 9 5 4 u l 1 2 M . e D 6 5 3 1 1 4 1 a n t s M 1 a u c e u e q b c D l i e l u ) g d o n 2 e a o c n ) i r a r a a 1 v D s b D 8 0 0 0 1 0 5 2 1 8 0 0 5 3 a ...... e ...... o n 2 , l 2 5 d – – n e D 5 0 5 7 6 8 7 2 s a 4 6 7 8 . . 0 3 l d 1 ( p i g 6 4 1 1 3 1 n 6 3 1 1 6 8 5 1 o s n s v ------n a a – – m i ( u i a i a 0 9 6 0 9 2 4 0 a M r ...... v h d 2 s t R l u a 3 5 8 7 5 8 7 3 s t a o 2 d 6 3 1 1 4 1 i n l p r ; a r o M i ) T 0 6 0 0 7 2 a ...... , h 1 4 f c e t m M 8 9 0 0 . . 9 4 n s o a g k 7 4 2 2 7 9 4 1 a n i p n ------– – s a r 0 a n 2 0 1 0 8 3 3 9 e r a ...... l 2 o R c 7 1 4 0 1 7 3 6 6 7 5 7 7 0 T p a n ...... s n i 7 0 6 3 1 1 3 1 e n 8 3 3 5 . . 7 4 ) n m a v ) m g a 5 6 4 2 2 6 9 5 1 a a h a 2 o n ------t r – – l s P r 1 a s 6 9 0 0 3 4 3 6 f T D ...... e a e o 9 2 6 5 6 0 9 2 t r R , 3 1 5 8 5 7 3 1 ...... h l s – – d D g n o t 3 5 3 1 1 4 1 6 5 1 1 0 1 4 1 e e ( u f n n m o D Y i ( t o ( e l a s m s i a b e a 2 5 4 9 4 5 0 9 n u ...... l v 7 9 8 6 3 4 8 1 u t u s – – a x ...... a a 4 3 2 1 0 0 5 0 q a n e e – – l 9 2 7 8 6 8 1 2 t e a s m s 6 4 1 1 4 1 s i M o e e D n r f r g e c a d u n d 7 8 7 0 9 2 6 1 . n m a ...... Y n 1 i T a e – – 2 7 8 1 5 8 9 3 s 0 a c . 6 3 1 2 4 1 h n 9 5 0 3 0 0 4 e n M ...... t 2 a a 6 3 a i p e i 3 4 7 0 9 1 . . 0 4 i s m g h 7 5 5 1 2 7 9 5 1 w 1 v t f o ------n - 2 a a s e a o s 0 0 5 0 0 0 6 8 6 5 d ...... p o n l n r R r ) r 5 4 5 1 5 6 5 7 6 1 o o a e 7 8 0 0 3 5 2 a c 7 i ) 5 5 3 1 1 3 1 2 z ...... b c 4 a h 1 e 9 1 M 0 3 2 4 . 0 4 5 t s t g 1 m , c m 7 4 2 2 7 1 6 1 a n n ------D u – – a n a p t e a 0 8 7 0 7 2 6 2 r r a n D ...... i n n g a T R l 0 6 9 3 2 7 5 5 3 8 ( 8 4 5 7 5 7 3 1 e n o c ...... n s o 4 2 1 1 4 1 s ( h c o 6 5 0 4 1 1 0 0 3 0 2 t m n n s M e o a d n n r h f r c a f a t o i T P L n 9 4 6 7 4 6 4 4 2 8 d h d ...... e t a e e n d 0 8 7 8 4 8 0 1 p 5 6 6 0 7 8 6 8 3 2 a e ...... h n s – – o n 2 6 5 4 1 1 4 1 t i y p 6 4 1 1 0 0 5 1 t a y r M 1 f e o a m l o L b c a o s s x T s h n e e n l n s 0 4 0 5 2 7 3 3 4 6 a 3 8 0 0 5 1 a e r ...... s ...... a a r 4 9 i e e e h 9 1 5 5 9 0 6 8 4 3 n . . 3 2 0 0 1 3 T t t h m g e 9 6 4 3 1 2 5 1 8 5 2 2 7 9 4 1 t c M n ------1 n – – a a n m i s a i 9 5 2 0 0 0 9 2 r i p ...... n a r s c R s 5 7 5 0 5 7 4 1 r a a h r e i ) 6 4 1 1 3 1 e c c e p t h t 1 s 8 ) s t r c . c 1 n 5 1 7 5 0 7 6 a 9 f a o a 0 ...... a r f 0 8 p r 1 e o r . . 2 4 4 7 2 6 6 4 r a D 9 4 7 1 5 7 2 6 a g * 1 7 5 4 2 2 7 9 5 1 r ...... T a s – – h h n ------D e c 5 3 1 1 0 0 2 0 ; n c c a 4 2 0 4 0 1 3 0 0 8 n ( s b 1 ...... n a c n c R ( 5 7 i i 5 5 0 1 4 8 3 1 i e m a r l r n 1 5 5 3 1 1 4 1 r t u t 1 i b 1 n e e s n T a 3 2 0 6 6 7 9 9 . a a ...... r t n s e m e – – m t k 4 6 8 8 6 8 8 1 a o e i h o n 7 4 1 1 3 1 t U M h h h h e t t , p c p e 9 9 2 8 2 9 5 6 6 8 a d r r ...... r s n h p c i e o o 2 4 2 3 2 1 0 0 3 0 t r p L s a e m m n c i n p n d f f 5 9 7 2 7 2 0 5 1 5 i s o y ...... n o i o d t n 2 4 0 n 6 7 7 2 4 1 9 2 t a 3 6 7 1 6 6 7 2 2 n e a ...... o a a t n n 8 9 8 6 2 2 1 4 1 e l r 6 8 3 7 4 1 8 9 1 0 e L c n o o v o 1 T 1 6 5 3 1 2 5 1 i i i e g T t t l i g M 1 H l a a s s i i o s r e r r i . c e a a d s t s L V V c r u r e / e a . c . e e e l c i t r t t s 2 1 p c c a g L a a e e h o m l L l L r r * * r c — o / / a * * a a b b i d d b I I E c c E c c r t S 2 h h L c i c t t L c i c t t a p a p a a o ; a o T r C T L L L P P L L W L 1 f T d C T L L L P P L L W L 1 Taxonomy of Triturusdobrogicus 423

Figure 2. Plotfor males (darkŽ gures)and females (openŽ gures)of T.dobrogicus fromBatevo (squares), Radoevo(triangles) and Vilkovo (circles) inthe space ofŽ rst andsecond principal component axes (PCA).

showthe results of this analysis for Batevo,Radoevo and V ilkovopopulations, as most distantfrom thecontact zones with other members of T.cristatus superspecies.The Ž rst axishas high loadings of all variables, but with contrasting sign (LiE versus the others). Itexplains 55% of totalvariance. The second axis, explaining 22% of the total variance, hashigh loadings of the strongly sexually dimorphic characters — Pa,Pp, Lc and Ltc. Theresults of thediscriminant analysis showed that most specimens from thePannonian (85.4%for malesand 94.3% for females)and Danube Delta (77.6% for malesand 96.3% for females)populations are well differentiated from eachother. The Moldavian sample andthe specimen from anuncertainlocality (the holotype of Molgemacrosoma ) are close tothePannonian populations. Triturusdobrogicus from theDanube Delta was usuallycharacterized by 16rib-bearing vertebrae(table 3). On other hand, we notedthat the Pannonian populations of the speciespredominantly have both 16 and17 rib-bearingvertebrae. Moreover, samples with prevalenceof 16 rib-bearingvertebrae were collectedonly close to thecontact zones with 424 SpartakN. Litvinchuk,Leo J. Borkin othermembers of T.cristatus superspecieswhere extensive genetic occurs (see Litvinchuk,1998; Litvinchuk et al.,1999). Morepolished skin, more obvious costal grooves on thebody sides and usually bright orangeor redbelly with small sparse rounded black spots also differentiated the Danube Deltaadult animals from thePannonian ones. In half of thesample from Vilkovoand in 10-20%of the Pannonian samples, the black spots merge on the belly, forming a long dorso-ventralblack stripe with uneven edges. Small larvae of T.dobrogicus hadlight (with blackspots) or darkcoloration. All large larvae (more than50 mm) oftheDanube Delta becamedeep black, with the exception of gilltips and a stripein themiddle of belly.The Pannonianlarge larvae usually have a somewhatlighter coloration. Wefoundthat the survival rate of the offspring obtained in crosses between the Pannonianand Danube Delta proved to bequitelow at embryostage (table 4). For

Table 3. Thenumber of rib-bearing vertebrae inthe holotype of Molgemacrosoma andin the Pannonian, Moldavianand Danube Delta samples of T.dobrogicus .

Locality,country n Mean s Numberof rib-bearingvertebrae 15 16 17 18

Holotype 1 16 1 Transcarpathians 1 , Ukraine 33 16.7 0.5 11 22 Transcarpathians 2 , Ukraine 57 16.4 0.6 2 29 25 1 Ostrovo,Y ugoslavia 2 17.0 2 Radoevo,Y ugoslavia 8 16.9 0.4 1 7 ObetskaBara, Yugoslavia 7 16.0 0.6 1 5 1 Kagul,Moldavia 3 17.0 3 Reni,Ukraine 1 16 1 Vilkovo,Ukraine 30 16.2 0.6 3 19 8

1;2 —see designationsin the table 1.

Table 4. Embryoand larval survival rates (inpercents) incrosses between members ofthe Trituruscristatus superspecies.

Crosses Localities Date ofegg-laying Number of Embryo Larval fertilitedeggs survival survival dob dobVilkovo(f1) Vilkovo(m1) 12.01.-17.01.1994 94 28.7 23.4 £ £ dob dobVilkovo(f2) Vilkovo(m1) 19.01.-25.01.1994 24 33.3 33.3 £ £ mac dobMinai(f1) Vilkovo(m1) 5.01.-13.02.1994 156 1.9 1.9 £ £ dob mac Vilkovo(f3,4) ObetskaBara(m1) 19.09.-23.10.1997 279 13.3 6.8 £ £ kar karAdzhi-Su(f1) Adzhi-Su(m1) 14.06.-14.07.1995 239 42.7 21.3 £ £ kar car Kutuzovka(f1) DonjaLokanj(m1) 10.09.-15.09.1997 59 20.3 10.2 £ £ car car DonjaLokanj(f1) DonjaLokanj(m1) 11.10.-5.11.1997 102 29.4 15.7 £ £ cri cri Taitsy(eggs collected in the Ž eld)27.04.-22.05.1999 1223 28.4 – £

“dob” is T.d. dobrogicus (Odessa Province),“ mac”is T.d. macrosomus (UkrainianTranscarpathians and Yugoslavia),“ kar”is T.k. karelinii (theCrimea), “car”is T.carnifex macedonicus (Yugoslavia),“ cri”is T.cristatus (St.Petersburg Province), “ f”is female, “m”is male. Taxonomy of Triturusdobrogicus 425 instance,the lowest survival (1.9%) was recordedin the cross of a male T.dobrogicus from Vilkovo(Odessa Province) with a female T.dobrogicus from Minai(Ukrainian Transcarpathians).The embryos from thecrosses between a malefrom ObetskaBara ()and two females from Vilkovoshowed low survival rates, too (13.3%). In contrast, ourobservations on thecontrol Vilkovo sample demonstrated that the Vilkovo x Vilkovo crossesprovided higher embryo survival rates (28.7-33.3%). It should be mentioned thatŽ ftypercent of mortality might be explained by lethal homozygosity on the Ž rst chromosome(Macgregor et al.,1990). Interestingly, even the survival rate of theF 1 hybrids obtainedfrom thecross of afemale T. carnifex and a male T.karelinii provedto be higher (20.3%)than that from thePannonianand DanubeDelta samples of T.dobrogicus (table 4).

Discussion

Thedifferences in the morphological features and crossing experiments support the idea thatthe newts from theDanube Delta and Pannonian Lowland should be considered as differenttaxa at thesubspeciŽ c orspeciŽc level.However, low values of geneticaldistances (Nei’s D 0.00-0.09)between the samples of T.dobrogicus from OdessaProvince and D UkrainianTranscarpathians based on allozymedata do notindicate their deep divergence (Litvinchuket al., 1994; Mezhzherin et al., 1997; Litvinchuk, 1998). W efailedto Ž nd anysigniŽ cant differences in the size (DNA owcytometry) of these samples (Litvinchuket al.,1999). Based on ourstudywe haveno evidence of sympatryof twoforms of T.dobrogicus intheDanube Delta. Therefore, taking into accountobvious contradictions betweenvarious data sets, we inclineto recognizetwo different subspecies of T.dobrogicus ratherthan two different species. Unlikethe Danube Delta subspecies, the nomenclatural status of the Pannonian sub- speciesis unclear. According to Mertens and W ermuth(1960), the following synonyms haveto beconsidered: Tritoncristatus danubialis Wolterstorff,1923; Tritoncristatus danu- bialis forma werneri Wolterstorff,1923; Tritoncristatus danubialis forma smederewana Karaman,1948, and Trituruscristatus danubialis var. intermedia Fuhn,1953. The last threeinfrasubspeciŽ c namesare unavailable — see1.3.4. and 45.5. of the International Codeof Zoological Nomenclature (Anonymous, 1999). Moreover, an allocation of the name Trituruscristatus danubialis var. intermedia Fuhn,1953 based on animalsfrom the Bucharestand Jassy regions should be re-evaluated. Examination of Fuhn’ s description andillustrations (Fuhn, 1953; Ž gs.3 and8) allowedus toassignthe variety intermedia to T.cristatus ratherthan to T.dobrogicus . Tritoncristatus danubialis hasbeen described from theenvirons of ,, Pecs(including Fü nfkirchen), Brasov (formerly Kronstadt), Bucharest, and the Dobrogea (Wolterstorff,1923); the typespecimens (except Dobrogean ones) were keptat theMagde- burgMuseum, Germany (W olterstorff,1925). Moreover, W olterstorff(1923) recognized thesamples from Kronstadtand Bucharest as a transitionalfrom T.c.danubialis to the 426 SpartakN. Litvinchuk,Leo J. Borkin typicalsubspecies, and the V iennaand Danube Delta samples as the forma werneri and forma dobrogica within T.c.danubialis ,respectively(W olterstorff,1923). Later, Mertens andMü ller (1928) restricted the type territory of T.c.danubialis toBudapest.Therefore, the name danubialismaybe appliedas validto the Pannonian newts. However,we founda seniorname belonging to T.dobrogicus .InJuly 1908, George Boulengerdescribed a newspecies, Molgemacrosoma .He hadone female presented by CaptainFlower to the British Museum. Flower reportedly obtained the from Ismail BeyChakir in Cairo in 1903, but because the Bey bought it from adealerin Vienna, thenewt may be of Europeanorigin. Indeed, no occur in Egypt. According toBoulenger (1908: 33), the new species “ isquite distinct from anyEuropean newt, beingmore nearly related to Molgecrocata (Neurerguscrocatus , Cope, Molgestrauchi , Stdr.)from AsiaMinor” . Later,Thorn (1969: 223) applied the name to the subspecies T.cristatuscarnifex . Thedetailed description and excellent drawing (Boulenger, 1908: plate IV) aswellas ourexamination of theholotype of Molgemacrosoma allowus tore-identifythe newt as T.dobrogicus .Indeed,palatine teeth arranged in twoparallel series, four Ž ngersand Ž ve toes,head shape, laterally compressed tail, and absence of parotoid glands evidence that thespecimen belongs to salamandridsand the Triturus,whilewarty skin and large bodysize indicate the T.cristatus superspecies.The body proportions (table 2), feebly wartyskin, presence of costalgrooves and lack of whitedots on thesides of thehead and trunkallow us toassign it to T.dobrogicus . Thelarge size and black belly seemed to be somewhat strange peculiarities of Molge macrosoma.However,our observationson T.dobrogicus inthelabshowed that in captivity femalescan reach quite a largesize (L upto91 mm), andthat, sometimes, black spots on thebellycan extend to cover much of thebelly.Only the tipsof Žngersand toes,cloaca and loweredge of thetail remain orange or yellow;such a colorationpattern (“ yellow”) isin accordwith the original description of Molgemacrosoma .Moreover,many other authors (e.g.,Dü rigen, 1897; Schreiber, 1912; Gislé n andKauri, 1959; Arntzen and T eunis,1993) notedsome crested newts with black bellies, too. Thetype locality of Molgemacrosoma isunknown, but, probably, it would be the environsof Viennawhere the species is still quite common (Tiedemann, 1990; Thonke et al.,1994). Boulenger’ s nameis olderthan Tritoncristatus danubialis Wolterstorff,1923, andyoungerthan Tritoncristatus var. dobrogicusKiritzescu,1903. Hence, as aseniorvalid name,it shouldbe appliedto thePannonian subspecies of T.dobrogicus ,andwe propose anewcombination and a newstatus, Triturusdobrogicus macrosomus (Boulenger,1908), comb.et stat. nov. Taxonomy of Triturusdobrogicus 427

Taxonomicaccount Triturusdobrogicus dobrogicus (Kiritzescu,1903) Tritoncristatus var. dobrogicus Kiritzescu,1903: 262. — Terra typicarestricta (Mertens andMü ller, 1928): lakes in the environs of Sulina,Danube River Delta, Romania. Lectotype:male, Museum “ Gr.Antipa”in Bucharest (MGAB), No 8(Fuhnand Freytag, 1961). Paralectotypes:4 malesand 4 females,Museum “ Gr. Antipa”in Bucharest(MGAB), No 8 (Fuhnand Freytag, 1961).

Figure 3. Reportedvalues of the“ WolterstorffIndex” (mean s andrange) for males (a) andfemales (b)of C T.dobrogicus .W—Wolterstorff,1923 (Austria, Hungary and Romania); H —Herre, 1932(Austria); F — Fachbach,1974 (Austria); Z1 —Zavadilet al.,1994 (Austria); Z2 —Zavadil,1996 (Czech Republic);L —Lác, 1957(Slovakia); FF1 and FF2 — Fuhnand Freytag, 1961 (Hungary and Romania with types of T.d. dobrogicus , respectively);A —Arntzenand Wallis, 1994; K —Kalezic´ et al.,1990 and 1997 (the former Yugoslavia);P — presentpaper (1 — theholotype of Molgemacrosoma ;2—Yugoslavia;3 —UkrainianTranscarpathians, beyond thecontact zone (Batevo, Chop); 4 —UkrainianTranscarpathians, near thecontact zone; 5 —Moldavia;6 — DanubeDelta, Ukraine).Sample sizes (ifknown) are presentedat thetop of the bars. 428 SpartakN. Litvinchuk,Leo J. Borkin

Diagnosis: T.d.dobrogicus differs from T.d.macrosomus bybodyproportions. The ratio Ltc/Lisusually higher than 13.8% for malesand 13.3% for females(see tables1 and2), andthe “ WolterstorffIndex” is usually more than 54.0% for males(see table1; Žg.3). As arule,the nominotypic subspecies has 16 rib-bearingvertebrae and brighter (almost red) colorationof thebelly. Range: T.d.dobrogicus isendemicto theDanube Delta.

Triturusdobrogicus macrosomus (Boulenger,1908),comb. et stat.nov . Molgemacrosoma Boulenger,1908: 32. — Terra typica:unknown; probably, the environs ofVienna,Austria. Tritoncristatus danubialis Wolterstorff,1923: 120. — Terra typicarestricta (Mertens and Müller, 1928): Budapest, Hungary. Tritoncristatus danubialis forma werneri Wolterstorff,1923: 121 (nomen illegitimum). — Lang-Engersdorf,the environs of Vienna,Austria. Molgecristata danubialis forma smederevana Karaman,1948: 52; tables 1-7 (nomen illegitimum).— ,Serbia, Y ugoslavia. Holotype:female, British Museum (BM), No 1946.9.6-29. Diagnosis:Differs from T.d.dobrogicus bythe ratio Ltc/ L,whichis usually less than 13.8%for malesand 13.3% for females(see tables 1 and2), and by the“ Wolterstorff Index”, whichis usually lower than 54.0% for males(see table1; Žg.3).As arule,the subspecieshas 17 rib-bearingvertebrae and orange or yellowcoloration of thebelly. Range: T.d.macrosomus inhabitsthe Pannonian and V iennabasins, as wellas anisolated areaalong the Lower Danube,probably, eastward to Reni(Ž g. 1).

Acknowledgements. We thankG. Duki c´ andM. Kalezi c´ ()for samples fromY ugoslavia,and to A.A. Fedorchenko(Kiev) for arranging a work-permitfor the Ukrainian Danube Delta Reserve “Dunaiskieplavni” . We are gratefulto I.G. Danilov (St. Petersburg) for assisting with anatomical preparations.B.T. Clarke (London), C.McCarthy(London), V .F.Orlova (Moscow) and the late N.N.Szczerbak (Kiev)kindly offered the museum collectionsavailable for study. Our special thanksto J.W. Arntzen and two anonymous reviewers whoread an earlier draftof the paper and made manyhelpful comments. TheSt. Petersburg Association of Scientists and Scholarsprovided facilities forpreparation of themanuscript. The research was fundedby grants of the Russian Foundationof Basic Research (96-15-97880)and INTAS (97-11909).

References

Anonymous(1999): International Code of ZoologicalNomenclature. Fourth edition. London, International Trust forZoological Nomenclature. Arntzen,J.W., Bugter, R.J.F., Cog alniceanu,Ï D., Wallis,G.P .(1997):The distribution and conservation status of theDanube crested newt, Triturusdobrogicus .Amphibia-Reptilia 18: 133-142. Arntzen,J.W .,Teunis,S.F.M. (1993): A sixyear studyon the population dynamics of the crested newt( Triturus cristatus)followingthe colonization of anewlycreated pond.Herpet. J. 3: 99-110. Arntzen,J.W., Wallis, G.P .(1994):The “ WolterstorffIndex” and its value to the taxonomy of the Crested Newt superspecies.Abh. Ber. Naturk.,Magdeburg 17, Sonderband: 57-66. Taxonomy of Triturusdobrogicus 429

Boulenger,G.A. (1908): Description of anew newt.Ann. Mag. Natur. Hist., ser. 8, 2: 32-33. Bucci-Innocenti,S., Ragghianti,M., Mancino, G. (1983): Investigations of karyology and hybrids in Triturus boscai and T. vittatus,witha reinterpretationof the species groupwithin Triturus (Caudata:). Copeia 1983: 662-672. Calinescu,Ï R.I. (1931): Contribu¸ tiunisistematice si¸ zoogeograŽ ce la studiulamphibiilor si¸ reptilelor din Romania. Acad. Romana.Mem., Bucure¸ sti,Sec¸ t.Sti.,¸ ser. 3, 7: 119-291. Dürigen, B. (1897):Deutschlands Amphibien und Reptilien. Magdeburg, Creuz’ sche Verlagsbuchhandlung. Gislén, T., Kauri, H. (1959):Zoogeography of the Swedish and reptiles with notes on their growth andecology. Acta Vertebratica, Stockholm 1: 195-397. Herre, W.(1933): V ergleichendeUntersuchungen an den Unterarten des Trituruscristatus Laur.Zeitschrift Anat. Entwicklungsgeschichte 99: 1-62. Fachbach,G. (1974):Das Serum-Eiweissbildder Trituruscristatus Unterarten.Z. zool.Syst. Evolut.-forsch. 12: 22-30. Fuhn,I.E. (1953): Contribu¸ tiila cunoa¸sterea solomázdrelor-de-ap aÏ (tritonilor)din R.P .R.II.Studiul subspeciilor si¸ varieta¸Æ tilor de Trituruscristatus Laur. Bul. Sti.,¸ sec¸ t.sti.¸ biol., agronom., geol. geogr. 5: 625-640. Fuhn,I.E. (1960): Amphibia. Fauna Republicii Populare Romî ne, vol. 14, fasc. 1,Bucure¸sti,Acad. Rep.Popul. Romîne. Fuhn,I.E., Freytag, G.E. (1961): Taxonomische und ö kologischeStudien ü ber Trituruscristatus inRumä nien. Zool. Anz. 166: 159-173. Kalezic,´ M.L.,D uki c,´ G.,Mesaroš, G.,Crnobrnja-Isailovi c,´ J.(1997):The crested newt( Trituruscristatus superspecies) inex-Y ugoslavia:morphological structuring and distribution patterns. Univ. Thought, Nat. Sci., Priština 4: 39-46. Kalezi´c, M.L.,D uki´ c, G.,Stamencovi´ c, S.,Crnobrnja-Isailovi´c, J.(1990):Morphometrics of the crested newt (Trituruscristatus complex)from Y ugoslavia:relevance fortaxonomy. Arhiv Biol. Nauka, Belgrade 42: 17- 37. Kiritzescu, C.(1903):Contributions à la faunedes Batraciens deRoumanie.Bull. Soc. Sci., Bucuresci 12: 243- 265. Lác, J.(1957):Prí spevok k poznaniugeograŽ cký ch rá s mlokavel’ ké ho ( Trituruscristatus Laur.)na Slovenskua poznámky k ichekoló gii. Bioló gia, 12: 724-745. Litvinchuk,S.N. (1998): Systematics anddistribution of newts ofthe Trituruscristatus complex(Salamandridae) inRussia andadjacent countries.Unpublished Ph.D. thesis, St. Petersburg, Zoological Institute, Russian Academy ofSciences (inRussian). Litvinchuk,S.N., Borkin, L.J., D uki c,´ G.,Kalezi c,´ M.L.,Khalturin, M.D., Rosanov, J.M. (1999): Taxonomic status of Trituruskarelinii ontheBalkans, with some comments aboutother crested newttaxa. Russ. J. Herpet. 6: 153-163. Litvinchuk,S.N., Rosanov, J.M., Borkin, L.J. (1997): A contactzone between thenewts Trituruscristatus and Triturusdobrogicus inthe Ukrainian Transcarpathians: distribution and genome size variation.In: HerpetologiaBonnensis, p. 229-235. Bö hme, W .,Bischoff,W .,Ziegler,T., Eds, Bonn, Societas Europaea Herpetologica. Litvinchuk,S., Sokolova, T.M., Borkin, L.J. (1994): Biochemical differentiationof the crested newt( Triturus cristatus group)in the territory of theformer USSR.Abh. Ber. Naturk.,Magdeburg 17, Sonderband: 67-74. Macgregor,H.C., Sessions, S.K., Arntzen, J.W .(1990):An integrative analysis of phylogenetic relationships amongnewts ofthe genus Triturus (familySalamandridae), using comparative biochemistry, cytogenetics andreproductive interactions. J. Evol.Biol. 3: 329-373. Mezhzherin,S.V .,Morozov-Leonov,S.J., Piontkovska, O.A. (1997): [Allozyme variation in species endemic tothe eastern Carpathians].In: Biodiversity in the Carpathian Biosphere Reserve, p.352-367. Didukh,A.A., Ed., Kiev, Naukova Dumka (in Ukrainian). Mertens,R., Mü ller, L. (1928):Liste derAmphibien und Reptilien Europas. Abh. Senck. Nat. Ges. 41: 1-62. Mertens,R., Mü ller, L. (1940):Die Amphibienund Reptilien Europas. Abh. Senck. Nat. Ges. 451: 1-56. Mertens,R., Wermuth, H. (1960):Die Amphibienund Reptilien Europas. Frankfurt am Main,W. Kramer. Nikolsky,A.M. (1918): Amphibia. In the series: Faunaof Russia andadjacent countries.Petrograd, Russian Acad. Sci.(in Russian). 430 SpartakN. Litvinchuk,Leo J. Borkin

Oliveira,M.E., Daszkiewicz, P.,Gauvrit,B. (1997):Summary of the conservation statuses andlevels ofthreats inEurope. In: Atlas ofAmphibians and Reptiles in , p. 408-412. Gasc, J.-P.,Cabela, A.,Crnobrnja- Isailovic,´ J.,Dolmen, D., Grossenbacher, K., Haffner, P.,Lescure, J.,Martens, H., MartinezRica, J.P.,Maurin, H.,Oliveira, M.E., SoŽ anidou, T.S., V eith,M., Zuiderwijk,A., Eds,Paris, Societas EuropaeaHerpetologica. MuseumNational d’ HistoireNaturelle. Schreiber,E. (1912):Herpetologia Europaea. Jena, GustavFischer. Thonke,A., Jehle, R., Hödl, W. (1994): Structure, dynamics and phenology of a populationof the Danube warty newt (Triturusdobrogicus )onthe Danube Island near Vienna(a preliminaryreport). Abh. Ber. Naturk., Magdeburg17, Sonderband: 127-133. Thorn,R. (1969“ 1968”): Les Salamandres d’Europe,d’ Asie et d’Afriquedu Nord. Paris, Paul Lechevalier. Tiedemann,F., Hrsg. (1990): Lurche und Kriechtiere Wiens.Wien, J &VEditionWien. Wallis,G.P .,Arntzen,J.W .(1989):Mitochondrial-DNA variation in the crested newtsuperspecies: limited cytoplasmicgene  owamong species. Evolution 43: 88-104. Wilkinson,L. (1989):SYSTA T:The System of Statistics. Computer program. Evanston, Illinois: SYST ATInc. Wolterstorff,W .(1923):Uebersicht derUnterarten und Formen des Tritoncristatus Laur.Blä tt. Aquar.- Terrarienk. 34: 120-126. Wolterstorff,W. (1925): Katalog der Amphibien-Sammlung im Museumfü r Naturund Heimatkunde. Abh. Ber. Mus.Nat. Heimat., Magdeburg 4: 155-234. Zavadil,V .(1996):Dopl nujíÏ cí pozná mka k cláÏ nku ColekÏ dunajský Triturusdobrogicus (Kiritzescu, 1903)nový m druhemobratlovce CeskéÏ republiky. Ochrana P Ïrírody 51: 6-8. Zavadil,V .,Piálek, J., Klepsch, L. (1994):Extension of the known range of Triturusdobrogicus :electrophoretic andmorphological evidence for presence inthe Czech Republic.Amphibia-Reptilia 15: 329-335.

Received: January8, 1999.Accepted: April4, 2000.