Volume 1, Number 11 December 1990 ISSN 0268-0130 THE HERPETOLOGICAL JOURNAL
Published by Indexed in THE BRITISH HERPETOLOGICAL SOCIETY Current Contents HERPETOLOGICAL .JOURNAL. Vol. I. pp. -181--192 (1990) 48 1
REVIEW: A REVIEW OF THE SMOOTH NEWT (TRITURUS VULGARIS) SUBSPECIES, INCLUDING AN IDENTIFICATION KEY
C. J. R..\.'\ WO RTllY
/)ep11rt111c111 of liiolog1·. The Open l 111ireni1r .. \/i/1011 f.:1Tlll''" ,\/A" 7 li.-1.·f. 1:"11g!ru11/.
(·ll"Cl'f'/er/ /8.8.89)
ABSTRACT
A taxonomic rev1s1on of the Smooth N ewt . Tri111rns l'lllgaris leads to the recognition or seven subspecies: T. 11• l'lllgaris. T. 1•. 111cridio11alis. T. r. grncc11s. T. 1·. /ant::i. T. 1•. a11111c/c11sis. T. 1'. kos.1·11·igi and T. 1·. sc/1111id1/cmm111. T. 1•. horcalis and T. 1•. tataicnsi.1 arc considered tobe synonvms of T. r. rnlgaris. A n identification kev and upd;1tcd distri but ion map is prnvicled These subspecies a re rccogn is eel bv char;1cters which reprcsen t non-independent p;1ttcrns of r;1cial 1·;1riation. prolx1hh· produced as a dirL'Ct consequence of :1llnp;1tric divergence in isolated ghcial rcfugia. Based nn the biological -;peciL'S concept there can be no _justification in raising these t;1.xonnmic units tn species r;1nk.
INTRODUCTION lateral folds and pigmentation ol" the male (f'ig. 1 ). However all these features only fully develop during The recognition of subspecies has ahv;1ys been the breeding season when newts arc aq uatic. Unless considered to be of importance to taxonomists fully developed thev are of little taxonomic value. This working on European Salamanclricls. Many species proble m is discussed by Steinitz ( 1965) in relation to show clear racial differences between geographical T. vi11a111s t<1xonomy. He found that only two of the areas, particularly within the genus Trit11rns (see five described subspecies were in fact valid and blamed Thorn. 1968). At present there arc over 30 described earlier workers for using small samples and animals subspecies of European ne wts. although this will which were clearly out of full breeding condition. certainly be reduced fo llowing a fu ll revision of the group. The Smooth newt Tri111ms l'lllgaris 1s exceptional among the Trit11rns species in that the donohtua.lfold degree of secondary sexual character variation seen between subspecies is far greater than that found in the other 11 species. This has resulted in the description of many T. vu/garis subspecies and. in the past. some tail fibmrnt considerable taxonomic confusion. The aims of this paper are to review and revise the subspecific taxonomy of the Smooth newt, Tri111rns pig mental.ion v11/gari1, and discuss the values and applications of the subspecies concept. This taxonomic study represents part uf a more detailed comparative investigation of the subspecific evolution of courtship behaviour and seconda ry sexual characters in the Smooth Newt Fig. Some scxuallv dimorphic characters of ( Raxworthy, 1989 and in pre p . ). Triturus rn/garis.
A HISTORICAL SUMMARY Secondary sexual characters such as the crest. toe flaps and tail filament may all regress over a matter of The original descriptions of the T. vulgaris subspecies clays if animals are kept under stressful conditions or were based on museum material which was considered starved. Also, these characters are not fully developed to show significant variation between different at the very start and end of the aquatic breeding period geographical populations. Usually material was (Verrell, Halliday and Griffiths, 1986; Griffiths and available to museum taxonomists in only very small Mylotte, 1989). samples, and this has sometimes resulted in subspecies T. v11/garis was originally described by Linnaeus in having been described based on insignificant variation. 1758 which he placed in the genus Laccrta with the Male secondary sexual characters have been used European lizards. His fo rm became the fi rst nominal extensively to recognise racial variation, indeed most subspecies upon the discovery more than one hundred racial variation is restricted to these characters in and twenty years later of the next subspecies, T. vu!garis. These secondary sexual characters include T. v. meridiona/is which was described by Boulenger the crest, tail tip, toe flaps, body cross section, clorso- (1882). At around the turn of the century many more 482 C. .J. RAXWORTHY
SUBSPECIES ORIGIN/\L NAME 1\1 D SYNONYMS
Trir11rus l'lllgari1. · l'lllgaris l.accrra l'lllgaris (Linnaeus 1758) Triton 1•11/garis subsp. lrf1ica forma Ka111111crcri (Woltcrstorff 1907) Tri1011 l'lllgaris forma Schreihffi (Woltcrstorff 1914) Tri 111rus 1'11/garis horca/is Ka u ri 1959 Trifllru1. · rn/garis tataimsis Dclv 1967
Tritums rn/garis 111cridio11a/is Molgc 1•11/garis suhsp. 111cridio11a/is (Boulen!!cr I 882) .\fo/gc \'11/garis subsp. kaf1clrma (Mchclv 1905) Mo/gc \'11/garis Bo11/c11gcri (Dunn 1918)
Trirur11s \'11/garis graccus Tri1011 \'11/garis suhsp. gracca (Woltcrstorff 1905) Triton \'11/garis subsp. gracca form a 1·orcrre11si.1· ( \Voltc-r'1or!T 1908) Tritm1 1•11/gari.1 subsp. gracca forma Tonl!lsinii (vVtJltcrstorll 1908)
1-fvhricl T r. rnlgaris x T I'. gmcrn.1 Tri1011 l'lllgaris subsp. dal111!111tirn (Kolomb;1tm ic 1907) Tri1011 l'lllgaris suhsp. i111cm1edia (Kolomh;1to1·ic: 1907)
Tri111rus 1•u/gari.1· /a111::i Tri111rus ru/gari1. subsp. npica forma f_11111::i (Woltcrstor!T 1914)
Tri111rus rnlgaris !llllfl!'lcnsis Tri111rus rn/garis (/lllf'C/c11.1is ruhn 1951 Tritums rnlgaris rnlgaris (;1111pclcnsis-form) Ful111 and frc·1·tag 1952
Tri111rus rn/garis kos.1·11·igi Trifllru1. · l'lllgari.1 ko.l.l'll'igi rrcytag 1955
Tri111rus l'u/garis sclu11id1/craru111 Triturus 1·11/garis sc/1111id1/eri Raxll'orthv 1988
TABLE I: Original subspecific names and post-1882 svnonvms of T 1·11/garis. subspecies were described. The most prolific taxonomist fo rms by making a detailed survey of the Adriatic of this time was Wolterstorff who described in all a region. They considered T. v. schreiberi to be a total of five fo rms. although only two are still synonym of T.v. vu!garis, which they found established recognised. The post- 1882 synonyms are given on in the type locality area. T. v. da/maticus and Table I for all recognised subspecies. All earlier T.l'. inll'rmedia are considered to be forms collected in synonyms refer to the nominate form and are given by the intergradation zone between T. v. vulgaris and Mertens and Wermuth (1960). A brief summary of T.v. graecus. while T. v. romasinii is a synonym of recent taxonomic changes since then is given below: T.v. graecus (Schmidtler and Schmidtler 1983). Mertens and Wermuth ( 1960) included T.it a!iC11s as The latest subspecies to be described is a subspecies of T. vulgaris, despite the detailed and well T.l'. schmidtll'rorum collected from Turkey (Rax illustrated description given by Peracca (1898a,b) who worthy. 1988). It had been suspected earlier to deserve rightly considered this to be a good species. Mertens subspecific status by both Eiselt ( 1966) and Schmidtler and Wermuth's taxonomic view was followed by and Schmidtler (1967), although this view was not Steward ( 1969) in his review of the European Urodeles. supported by Ozeti (1964). The subspecific name of However Mancino ( 1961) provided strong evidence this subspecies has been changed from Trirurus vulgaris that T.ita! irns was a good species based upon his schmid1 frri to Triturus vu!garis schmidtlerorum. because hybridisation experiments between T. ita!iC11s and this subspecies was named after two people: J. J. T.vu!garis. This was supported by Thorn( 1968) who Schmidtler and J. F. Schmidtler and therefore the gave T.italicus species rank in his review of Tri/llrus original subspecific name was grammatically incorrect and this has been followed by all more recent studies. (Raxworthy. 1989). Thorn ( 1968) recognised all the presently subspecies A population of T. vu/garis discovered in the Sava with the exception of T. v. schmidt/crorum (which had Valley. north Yugoslavia, resembles the nominate not been decribed then) and also T.v. tataiensis (which form but may deserve subspecific status (Schmidtler had only just been described by Dely in 1967). He also and Schmidtler 1983) following further studies. There included three other subspecies: T. v. schreiberi. is also a possibility that specimens from the Danube T. v. dalmaticus and T. v. borea!is. basin may also represent another new subspecies The situation regarding the Dalmatian coast (Schmidtler, pers. corn.). T. vu/garis subspecies remained confusing with five subspecies having been described from this region of PROPOSED REVISION Yugoslavia. This is especially evident when referring to the descriptions given by Steward ( 1969) for Trirurus vulgaris borealis Kauri 1959 T.11• dalmaticus,T v. graeC11s and T.v. schreihcri. The T.\'. horea!is was recognised by its low crest, smaller diffe rences he gives (based on the original descriptions) body spots, general marked 'longitudinal ridges' on are strikingly unconvincing. Schmidtler and Schmidtler the back and in some specimens a thread like ( 1983) finally resolved the taxonomic status of these appendage (Gilsen and Kauri 1959). The description TR/TURUS VULGARIS SUBSPECIES 483 was based on 11 males caught in NorthernSwede n. I Guillaume (1984) have found the electrophoretic have examined six specimens of T. v. borea/is kindly studies were unable to separate the metamorphic form loaned to me by J. Elmberg which were collected from from the paedomorphic fo rm .. Stokarret (Skelleftea), Alnbn (Medelpad), Vindel gransele (Lykselle) and Strigstjarn(S abra). There is no evidence of true dorso-lateral folds or a tail fi lament THE TRITURUS VULGARIS SUBSPECIES (as defined by Raxworthy. 1988) and body pigmentation does not diffe r significantly from the nominate form. A total of seven subspecies are recognised in this The crest however is slightly lower (mean = l .4mm, study. These are: n = 6) than that seen in good breeding condition males Tri111rus v11!garis v11/garis (Linnaeus 1758) from England (mean= 2.5mm, n = 19), although the Tri111rus v11/garis meridionalis (Boulenger 1882) small sample size does not allow a firm conclusion to Triturus v11/garies graecus (Wolterstorff 1905) be made on this. Kauri found that this northernrace is (Wolterstorff 1914) subject to a poor diet and has a shorter period of Tri111rus v11/garis /antzi activity than populations further South. Crest Tri111rus v11/garis a111pe/c1 1is Fuhn 1951 development is known to be dependent upon food Tri111rus v11/garis kos.1·11·igi Freytag 1955 availability (Halliday, 1977) and therefore the slightly Tri111rus v11/garis sch111idtleron1111 Raxwort hy 1988 lower crest seen in T.v. borea/is might be due to local A brief description of the seven recognised environmental factors. Translocation experiments subspecies is presented below. Because rully developed would readily support or reject this hypothesis. male secondary sexual chara cters are used to However based on the material examined I conclude distinguish subspecies. only these features arc that these specimens belong to the nominate considered here. Body size is not a diagnostic fe ature of subspecies. subspecies (Raxworthy. 1988). Because of the well known differences in body size between populations of Triturus vu!garis tataiensis Dely 1967 newts (Bell. 1966. Tucic and Kalezic. 1984. Clifford. Dely ( 1967) described a paedomorphic subspecies 1986) body size is considered to be largely innuenced T.v. tataiensis from Tata. Hungary. The adults had by local environmental fa ctors. external gills and the males had poorly developed secondary sexual characters. This paedomorphic Tri 111rus \' 11/garis v11/garis (Linnaeus 1758) Common Smooth newt condition. which is seen in T. v11/gari.� is most suitably be described as facultative partial neoteny (Raxworthy, The dorsal crest is high along the body (> l .Omm at 1989, in prep.). From the description of T. v. tataiensis mid-body) and deeply notched by rounded denti it is clear that it is separated from the nominate form by culations. There is no tail filament, the tail usually paedomorphic features only. Paedomorphic specimens tapers to a blunt end. There are no dorsal-lateral folds. examined during this study from Smilic. Yugoslavia. The body cross section is rounded. The toe naps are show similar features. although they belong to the moderately or poorly developed. Fig. 2 and Fig. 8. nominate subspecies. The paedomorphic and metamorphic condition Triturus v11/garis meridiona!is (Boulenger 1882) represent the two states of a polymorphism. The Southern Smooth newt phenotypic expression of the paedomorphic condition The dorsal crest is low and smooth edged (
Fig. 2 T. 1·. vu/garis. Oxford. England. (T. R. Hallidav).
Fig. 3 T 1'. 111eridio11a/is. Torino. Italy. (C. .I. Raxworthy). TRITUIWS V u;ARIS SUBSPECIES 485
Fig. 4 T. I'. gra1·rns. Vil11si. Yugoslavia. (C. .I. Raxwnrthy).
Fig. 5 T. I'. !ant::i. Tbilisi. U.S.S.R. (B. Ardabyevski). 486 C. J. RAXWORTHY
Fig. 6 T v. ampe/cnsis. Valea Dosului, Rumania. (D. Cogalniccanu).
-..
Fig. 7 T v. kos.migi. Adapazari. Turkey. (C. J. Raxworthy). nun Rl s I l 'I (; I/US Sl IBSl'l:Cll'S
Fig. X T. 1·. 1·chl/lidrl<'rnrt1111. Karacahcv. Turkcv. (C. .1. R:IX\\' Fig. 9 Top: T. 1·. rn/garis. Milton Kevnes. England. Bottom: T. 1•. sc/1111id1/eran1111. Karacabcy. Turkey. (C. .I. Raxworthy). 488 C. J. RAXWORTHY Triturus vu/garis ampelensis (Fuhn 1951) hnishvili (pers. comm.). A very similar distribution Apuseni Smooth newt map is given by Macgregor, Sessions and Arntzen (in The dorsal crest is of medium height (>l.Omm at prep.). mid-body) and weakly denticulated by rounded serrations. The tail gradually tapers to a fi ne thread A IDENTIFICATION KEY (but there is no tail fi lament). There are no dorso lateral folds. The body cross section is slightly square This key will only identify aquatic males which are in dorsally. The toe naps are large and well developed. full reproductive condition. It is preferable that Fig. 6. material examined should not have been kept in captivity for more than 48 hours prior to examination Triturus vulgaris kossirigi (Freytag 1955) since important diagnostic characters such as toe naps Kosswig's Smooth newt and crest denticulations may start to regress quickly The dorsal crest is smooth edged and very low on the under some conditions. The features used in this key body (<1 .0mm at mid-body) but high at the basal area can all be examined readily on live unrestrained of the tail. The tail ends in a long tail filament. There aquatic animals, but the key works equally well with are strongly developed dorso-lateral folds and the museum material provided that the secondary sexual body cross section is square-shaped dorsally. The toe characters are fu lly developed. This can be determined naps are broad and very well developed. Fig. 7. by examination of the crest and toe naps. In all subspecies, one of these characters is always well Triturus vu!garis schmidtlerorum (Raxworthy 1988) developed for animals in fu ll breeding condition. Schmidtler's Smooth newt Occasionally animals may have suffered natural The dorsal crest is of medium height (>2.0mm at damage to the tail tip. Under these circumstances it is mid-body) and denticulated by almost spine shaped best to avoid using the tail tip features in the key. serrations. The tail gradually tapers to an elongated point (but no tail filament). There are no dorsal-lateral The crest is obviously denticulated along the body. folds but the body is slightly square-shaped dorsally. Fig. I l(a,b). 2 The tow naps are only very weakly developed. Fig. 8 The crest is approximately smooth edged along its and Fig. 9. entire length. Fig. I l(c). 4 Fig. 10 shows the geographical distribution of each subspecies based on the distribution information given 2 The tail tip gradually tapers to a fi ne point. by Schmidtler and Schmidtler ( 1967, 1983), Kalezic frequently with a very fi ne thread like end. (1983), Sparreboom and Arntzen (1987) and Tark- Fig. 11(d). The body cross sect ion is slightly squares � L:...:...:l vulgaris � meridionalis � ompclcnsi.r E3graccus 1�: j � j� (Ischmidc/uorum ITIIillkosswigi �la1111i . -·� . · . . � . Fig. 10 The subspecies distribution of T. 1•11/garis. TRITURUS l'ULGAR!S SUBSPECIES 489 ---- poorly developed. Fig. 11(111). Usually a few large � . AAAA ,. spots on the bellv (>I. 5111111 dia.). T. 1•. ko\'.\'ll'igi Cc> �;�. /�{ : ... _ _ ji (a) ,.'.=>': _;::;, {i: ./{�. . (b) 5��r _J!.:: /��: 6 The crest is always smooth edged. There arc weak dorso-lateral folds on the body. T. "· 111eridio11a/is (d)======(c) � (f)� The crest may be weakly denticulated. There arc no dorso-lateral folds on the body although this is slightly square shaped dorsally. T. v. a111pele11sis DISCUSSION PHYLOGENETIC ASPECTS The two most closely re la tcd species to T. v11lgaris are T. hefrcric11s and T.1110111m1d o11i. The three species fo rm their own monophyletic group (Rafinski and Arntzen. 1987. Arntzen and Sparrcboom. 1987, Macgregor et al. in prep.). Hybridisation studies (Macgregor er al. in prep.). cytological evidence (Ragghianti, Bucci-Innocent and Mancino, 1978) and � � electrophoretic evidence (Rafinski and Arntzen, 1987) (l) (m) all indicate that T.vul garis is most closely related to T. 111onrando11i. Sympatric populations arc known in Central Europe and natural hybrids have been Fig. 11 reported which may be common in some populat ions (Hofmann, 1908, Geyer, 1953, Fuhn, Sov;1 and Dumitrescu. 1975, Pecio and Rafinski, 1985). shaped dorsally, Fig. I I . The crest denticulations (j) Within T. vulgaris two main groups can be may be pointed. Fig. 1 l(a), or rounded, Fig. I l(b). 3 recognised (Raxworthy. 1989). The 'graecus' group is comprised of T. v. graecus and T.v. ko.1·.1wigi. The The tail tip usually tapers gradually to a blunt point, 'vulgaris' group is comprised of T. v. vulgaris, T.v. lantzi Fig. I l(e). The body cross section is rounded in the and T. v. schmidrlerornm. T. v. rneridionalis and dorso-lateral region, Fig. I l(i). The crest denti T. v. ampelensis represent intermediate forms, with culations are rounded, Fig. 1 l(b). T. v. vu!garis stronger affinities to the 'graecus' group fo r T.v. meridionalis, and stronger 'vulgaris' affinities for 3 The toe flaps are not obvious, Fig. I I (g). The crest T.v. ampelensis. Ou tgroup comparison suggests that denticulations arc almost pointed and extend onto the 'graecus' group represents the ancestral state, while the tail past the cloaca. Fig. I !(a). the 'vulgaris' group subspecies show more derived T. v. schmidtlerorum characters (Raxworthy, 1989, in prep.). Subspeciation The toe flaps can be seen easily, Fig. l l(h). The of T. vulgaris appears to have occurred in the denticulations may be rounded or pointed and Pleistocene. when populations during glacial pe r iods extend onto the tail past the cloaca. T. v. lantzi became fragmented and isolated into small refuges The toe flaps are large and very obvious, Fig. I l(h). areas. It is suggested that i n allopat ry these forms The crest denticulations are rounded Fig. I l(b) and evolved independently, and that subspccific fusion was usually do not extend on to the tail past the cloaca. limited to secondary contact zones only (Raxworthy, T. v. ampelensis 1989, in prep.). 4 The tail tip ends in a long tail filament which is THE SUBSPECIES CONCEPT devoid of tail fin and shows a distinct transition Taxonomists have always considered it useful and from the rest of the tail, Fig. 11(f). There are practical to use subspecific names to draw attention to obvious well developed dorso-lateral folds on the interesting regional variation (Amadon and Short, body, Fig. I l(k). 5 1976), and in the case of T. vulgaris, this species The tail tip tapers gradually to a fine point, showing exhibits clear racial differences which can be no distinct transition from the rest of the tail, conveniently labelled using the subspecies nomen Fig. I l(d). Dorso-lateral folds may be present, clature. However the subspecies concept has been Fig. 11(k) or absent, Fig. 11(j). 6 frequently criticised, with for example Wilson and Brown ( 1953) arguing that the subspecies is so 5 No large dark spots typically touch the lower tail fin arbitrary a concept that it should be abandoned. margin and the pale pelvic line is usually very Regarding the situation seen in the T.vul garis obvious, Fig. I !(I). Only small spots on the belly subspecies, all subspecies are defined, based on the ( logical characters do not show independent patternsof Huntley and Birks ( 1983) shows close correlations with distribution and therefore this leads to the recognition the centres of current subspecies distributions of of clearly defined (and non arbitrary) subspecific taxa. T. vu!garis subspecies (Raxworthv. 1989). It is By use of such characters subspecies boundaries therefore possible to propose the appropriate refugia become real biological concepts represented where most subspecies were confined during at least by intergradation zones which can be readily the last glacial period. identified (Fig. 12). The T. alpcs1ris subspecies also The biogeographical congruence of subspecific appear to show non-independent geographical phylogenies, the congruence of different taxonomic variation for both morphological and electrophoretic data sets and the correlation of subspecies distributions characters. Arano ( 1988) has fo und that most of the with glacial refugia provide strong evidence that these T. a/peslris subspecies (described using morphological subspecies represent valid biological entities. Therefore criteria) are supported by electrophoretic patterns of I conclude that these subspecies are not arbitrary variability and the Yugoslavian T. vu/garis subspecies concepts. but rather represent real non-independent have also been supported by electrophoretic evidence patterns of geographical variation. (Kalezic, 1984). SUBSPECIES OR SPECIES') Is it appropriate to consider raising the taxonomic status of the T. l'lllgaris subspecies to the species lcveJ'l a) clinal variation Biological species arc defined by Mayr ( 1942) as 'groups of actually or potentially interbreeding natural populations which are reproductively isolated from other such groups'. chararacter state Natural hybridisation is usually rare between Tri11ir11s species (the exception being between T.1110111a ndoni and T. l'lllgaris. and T. cris1a111s and T. 111armora111s). When hybridisation 1s induced artificially in the laboratory. post-zygotic isolation mechanisms always result in unfit hybrids. e.g. between T. 111ar111ora111s and T. cris1a111s (Lantz. 1947). T 1•11(�aris and T hcfre1irns (Scali and Mancino. 1968) and the distance transect T. cris1a111s superspecies (Callan and Spurway. 1951. Wallis and Arntzen. 1989). In all these cases post zygotic isolating mechanisms prevent genetic fusion and are therefore contribute to the reproductive subspecies A isolation of each species. Gever ( 1953) however has reported viable hybrid T. v11/garis-1110111andoni in the b) subspecificvariation first and second generation. Although wild T. vulgaris x 111011/andoni hybrids are well known (see earlier). recent studies have found mutually exclusive parapatric distribution patterns between these two species chararacterst:1te (Rafinski pers. comm .. in Arntzen and Sparreboom submitted). Evidence for ecological and ethological pre-mating isolation mechanisms have been fo und in subspecies B some species (Halliday. 1977. Arntzen. 1986) which would prevent gene now between species. Only a very limited amount of work has been distance transect undertaken concerning hybridisation experiments Fig. 12 The diffe rence between cl inal variation and between the T. v11/garis subspecies. Lantz (1947) noted subspecific variation. that intra-specific crosses between various subspecies of T F11/garis were obtained without difficulty. yielding The subspecific phylogenies of T.vul gari.� and offspring which are normal 111 every respect. T.al pesrris show a surprising level of biogeographical Observations made during this study found both congruence. based on completely diffe rent types of reciprocal mixed pairs of T. v. 1'11/garis and data: morphological and courtship behaviour for T. 1•. 111cridiona!is showed successful courtship with T. vulgaris (Raxworthy. 1989) and electrophoretic sperm transfer. evidence for T. a/pesrris (Arano and Arntzen. 1987. T. 1111/garis subspecific intergradation zones have Arano. 1988). Both phylogenies have ancestral Balkan now been recognised and described throughout much subspecies and a more derived lineage giving rise to the of Europe. Intergradation zones have been found in: Italien and widespread Central European subspecies Yugoslavia: T. 11• vulgaris x T. v. meridionalis and (Arntzen and Raxworthy. in prep.). The T. vu!garis T v. F11/garis x T. F. graecus (Schmidtler and Schmidtler, subspecies appear to have arisen as a direct 1983). Rumania: T. v. vu!garis x T. v. ampe!ensis consequence of climatic changes in the Pleistocene. Cogalniceanu pers. comm.) and in Turkey T.v. vulgaris The glacial refugia identified by tree pollen analysis by and T. "· kossirigi (Tabrizi, 1980). TR ITURUS VU/. C.-IRIS SUBSPECIES 49 1 The size of some of these intcrgradation zones (up to Sal:unandriadal') du st1cl de la Ynugoslavie. Rull. Soc. 160km in length along the Dalmatian coast) and the Z1111/. Fr. 109 (4). 377-.\X9. reported norm a I fi tness of hybrids do not provide any 13reuil. M. and Thunt . M. (191n). E t lw -ecolngv of neolenic evidence of reproductive isolation betwccn t hcsc a l pine m: \\'t (Tri111rus alpr'l·tris 11101111·n1•gri1111.1· subspecies. There is obvious gene now occurring at R ad ovanm · i c . 1951) in la k e 13ukumir (Montcnegr<1. y ugosl :l \'i:l ) : Exa mi11:1t ion or lakc Clllll lllU ll i t ies k:ll lll'l'S these secondary contact zone� . although subspe�ific rn i 1 n l ' fusion mav be very slow if migration is low and there is and propo:;aI of an ecologica I det.: 1 1 a t ion for ne t<:I \ . (;/as. l? !-puhl. 7.arnda 7.agst. l'riodc l'rirod11jackog some degree of subspccific assort ativc mating .\lu:efa. Ti111grnd11. 16. 85-%. (Raxworthy. 1989). Clearly. there can be no question Callan. H. (i. and Spur\\'av. H. ( 195 I). .'\ stud\' or meiosis in of raising these taxonomic units to species rank based intermediate hvbritb or the 11.:w1 Tri 111rus cris111111s. on the biological species concept. ./11urn11/of l,cnctics. 50. 2.15-249. Cliffo rd . T. ( 1986). Not.:s on the 1nnrpl10metrics and spot pattern:;or female Smooth newts (Tri111rus l'lllgaris) a t :1 ACKNOWLEDGEMENTS cna :;t :il sit<: in l.i ncolnshir.:. //apctologirn/./. I. 9.\-96. Del\'. 0. G. ( 19(17). N.:unc· /\11g:1bc11 1ur Ke11n111is de:; l should like to thank all the fo llowing colleagues Nen1.:11ischrn T.:ich1110khe:; ( 1·ri111ms l'lllgaris 1 .. ). who have contributed information and advice. In .·l cta :1111/. hung. IJ (.\-4 ). 25.1-270. part icular I should like to mention Bcgc'inaAr ano. Pim Dunn. E. R. ( 1918). The colkct ion or /\ m ph i h i :1 C:1udata Arntzen. Franco Andrconc. Nick Arnold.John Baker. of the Museum of Comparatiw Zonlngv . !lull. tl f11s. Terry Burke. Barry Clarke. Dan Cogalniceanu . .Johan co1111i. Zoo!. !Ian-. 62 (9). 445-4 71. Elmberg. Cristina Giacorna. Richard Griffi ths. Tim Eisclt. .J . ( 1966). Ergehhnisse rnolllgischer sammelreiscn in Halliday. Lottie Hosie. Milos Kalczic. Herbert d<:r Tii rkei: /\mphibia Ca ud at:1 . .'ln11/. na111rh. J\ fus. Macgregor. Giorgio Malacarne. Giorgio Mancino. ll'f('//. 69. 427-445. Phil McGowan . .Jan Rafinski . .J ulie Roberts . .Joseph Fre v t a g . G. E. ( 1955). Ein neucr Teich mo lch aus clcrTiirkcl. Schmidtler. Max Sparrcboom. Da vid Tarkhnishvili. 7.oo/. /111:. 15.t, 195-200. Paul Verrell and Robert Vignali. This studv was l-ul111. I. E. ( 1951 ) . Conributiuni la sistemat i ca salamaz supervised by Tim Hallidav and fu nded by the Science d rc lnr din republica populara Ro m a nana I. Studiul and Engineering Research Council. I am grateful to catorva popul a t ii de Tr iturus l'lllgaris. L. /]11/ctin sti. Pim Arntzen and .Joseph Schmidtler for their !lead. Rep11h. />Of'. ron1. Sa. geo/. geog. hio/. st. teh. ag. (3). 501-5 12. comments on an earlier version of this work. SN't. St. /Jio/. ugron. gco/. geog. 3 Ful111. I. E.. Sova. C. a nd Dumitrcscu. M. (1975). Unc popula tion hybridogcnc Triturus 1·. 1•u/garis x Boul. du lac Crauclc (Mts. N em i ra REfERFNCES T. n10111a11do11i . Depart . Bacau). Stud Co111. Mus. S1i111. Na111r. Rar. 8, 225-236. Amadon. D. and Short. L. L. ( 1976). Treatment of subspecies Gabrion . .J (1977). approaching the species 'talus. S.1 '. 1·1. 7.nnl. 2 5. (2). .. Scntein. P. and Gabrion. C. Les 161-167. po pul a t i o ns n eo tcni q ue de Tri111ru" hc/l'('tic11s des ca u sses et du bas- l angucd oc I. Repartition et Arano. B. ( 1988 ) . A.1pcctns fi/11ge11eticos 1/c·I gc11cro Tri11rru" ch:tractcriqucs. Terre Vie 3 1 . 489-505. COii especial co11sidcrocin11 11 /11 crnlucin11 de/ co1111>/cjn ( 195:1). O Triturus a/pestris. Ph D Thes i s. Madrid. Dcpartamcntn Gevcr. H. ber Hastarde 1. wischcn Karpatcnmolch de Zoologia. Musco Nacinnal de Cicncias Ri ologica s . ( Triturus mnntandoni) und Tei chmolc h (Triturus vulgaris 1'11/garis) und ihre F:- Na ch k omm en. Mill. Naturk. Arano. B. and Arntzen . .J . W. ( 1987). Genetic diffe rential ion Vorgcsch. l\fus. t-:ult.-Gcsch. Magde/J. 3, 185-195. in the Alpine newt Tr iturus alpestril'. In !'rocecdi11gs n( ( 1959). the fo urth ()rdi11an· (ic11cra/ .\lcc1i11g n( the Societas Gislcn. T.and Kauri. H. Zoogeographv of the Swedish ro Fuopaea !lcrpcto/ogirn. 21-24. \ a ll Gc ld� r . . J . .J .. amphibians and rept iles with notes on their g wth and (3). 1%-397. St rij bosch. H. and Bcrgers . P . .J. M. (Fl1'). Nijmegen. ecol ogv. Acta Vert. 1 F:.1ctiltv of Sciences. N ij mcgen. Griffiths. R. /\. and Mvlotte. V . .J . ( 1989). Observations on Arntzen. .J . W. ( 1986). N ot c·s 't1r la conistence d 'c.:s pece the development of secondarv sexual characters of male sv mpatri que de tritons du genre· Triturus. Rull. Soc. newts Tr illlrus l'lllgaris and Tri111ru.1· he/c1•e1icus. .!. !lcrp. 476-480. f-!crp. Fra11ce. 37. 1-8. 22. Arntzen . .J . W. a n d Spa rrcbon111. M. ( 19X7). The use ofbill Hallidav. T. R. (19 77). Thc courtship ol' Europcan newts. An ch emica l and behavioural data rn r t he phvlngcnv of evolutionary perspective. In Th e Rcproductil'e bin/og.1· Old World n C\l't s. genus Tri1u1·us. In l'roceedi11gs of the o( 11111phihia11s. 185-232. Taylor. D. H. and Guttman fo urth ()u/i11an· l,c11cra/ Mccti11g nfthc Socictas Fuopaca (Eds.). New York. Plenum Press. 1-frrpctn!ogica. 25-28. van Gelder. . 1 . .J .. Stri.i bosch . H. Hofmann. 0. ( 1908). Ober Tritnn Mo111ando11i Big. in Galizien. and Bergers. P . . 1. M. (Fds.). N i j megc n. Facultv of Wschr.. 1 quar. - 11. Terraric11k. 1908, 65-66. N Sciences. ij m cgcn. Huntley. B. and Birks. H . .J . B. (1983). An Atlas o(pasl and Bell. G. A. C. ( 1%6). The size of a series nf Lei cestershi re present pollc11 maps fo r Furope: n- /.IO()() rears ago. newts. Brit. .!. 1-frrp. 3. 279-284. Cambridge. Cambridge University Press. Boulenger. G. A. ( 1882). Catalnguc ofthe Ratrachia l,radirntia Kalezic. M. L. ( 1983). Geograph i ca l aspects of genetic s. Caudata rmd R111rachia Apoda in the collect inn of the variabilitv in the Smooth newt (Triturus l'lllgaris). Brirish \111sc11111. Second Edition . London. British l,enetika 15 (I). 93- 103. Museum. Kalezic. M. L. (1984). Evolutionary divergences in the Breuil. M. and Guillaume. C. P. (1984). Etude electro Smooth newt Triturus vu!garis (Urodela. Sala phoretique de quclques population de Tritons alpcstres mandridae): electro phoretic evidence. Amphibia neoteniq ues (Tri111rus alpcstris. A m ph i b i a . Caudata. Rcpti!ia 5, 221 -230. 492 C .J. RAXWORTHY Kolombatovic. G. (1907). Contribuzioni alla fa una dei Rocek. Z. (197.:1). Biometrical i1l\'estigations nf central vertebrati dclla Dalmazia. (i/a.rn. h1T. 110rndos/. nmsl. European populations of the alpine newt Trir11ru.1· 19. 1-24. alpnrri.rn/f!C\'lris (Laurenti. 1768). (Amphibia: Urodcla\. Lantz. L. A. ( 194 7). H vbrids between Tri1111·11s cri.1H1111s Laur. Ac1a U11i1·. Cami. Rio!. 1972. 295-.'1 7.1 . and Tri111ru.1· 11wr111ora111.1· L.atr. Proc. Zoo!. Soc. /.rmd. Scali. V. and Mancino. G. ( 1968). Observazioni sulla 117. 247-258. spermatogenesi di ibridi poliploidi Tri111rus hc!i·C1ic111 Linnaeus. C. ( 1758) . S1·st{'ll/a 11a111ral' f'CI' r{'g11a 1ria 11l11111·a{' I 0. 1·11/garis fe male x T. rnlg ari.1· male. . \femoric Alli. Soc. Stockholm. lose. Sci. 1w1. 75. 8-18. Mancino. G. (1961 ). Anomalie meiotiche nella spcrmato Schmid tier. .I . .I. and Schmidtler. .1 . F. ( 1967). U ber die Verbreitung der Molchgatllln!.! Trir11ru.1· in Kleinasic·n. genesi dell' ibrido Tri111rus i/{//icus fe male x Tritums - - rnlgaris male. Roll. Zoo!. 28. 69 1-70 1. Sa/a111rmdra 3. 15- 36. Mayr. E. ( 1942 ). S1·s1c111a1ics and !he origin o(sf!ccics. Ne\\. Schmidtler. .I . .I. and Sehmidtlcr. . I. !-. ( 1983). Verbreitung. York. Columbia Universitv Press. Okologie und innerartliche Gliederu ng \' O il Trililrus Mayr. E. ( 19 63). Animal Sf!CCi{'s and crnli11io11. Cambrid!.!e. rnlgaris in den adriastischcn Kiistengcbieten. Spil'i11na - Mass.: Belknapp Press. 6 (.\). 229-249. Sparreboom. M. and Arntzen. P. ( 1987). O ber die Amphibien Mehely. L. ( 1905). Die herpetologischen Verhiiltnisse des Mecsek-Gebirgcs und der Kapcla. Ann. M11s. Nat. !lung. in der Umgebung von Adap;1zari . Turkei. l/c1pc1o(a11na 9 (50). 27-34. 3, 256-273. Steinitz. H. ( 196'.i). (.Jcnvns). geographic Mertens. R. and Wermuth. H. ( 1960). Die Amphibicn und Trirurus ri11a111s distribution and taxonomic subdivision. /sra{'/ ./. 700/. Reptilien Euro pas 3. Senck{'///Jcrg. hiol. .• 8. 1-264. I .:I. 234-240. Ozeti. N. ( 1964). rstudics Oil the morphologv. taxonomic Steward . J. W. position. seasonal activity and t hcrmotaxis beha vinu r of ( 1969). Th e tai/{'d amf!hihir//1.1· o( F11mf!C. Newton Abbot . David ;111d Charles. Tri111rus rnlgari.1 (Linnaeus) in the Aege;1 11 region. I Sci. R{'f'. Fae. Sci.. Fgc Un i1·. No. 15 . 1-49. rin Turkish Tabrizi. F. H. (1980). ron some new material of Trir11ru.1· with English summarv] rn/gari1· (l..) from the ;1rea between Abant (Bolu) and Bospl10rusin NW Anatolia] Eg{' U1 1i1·. Fen. ra k. Ocr. (8). Pecio. A. and Rafinski . .I. ( 1985). Scxu:i l behaviour of the 115- 140. rin Turkish with English summarv]. Montandon's newt Tri111ru.1· 1110111m1drmi (Bou\cngcr) .:I. � f (Caudata. Salarnandridae). /1111{!hihia-R{'{!filia 6. 11 :2:2. Thorn. R. (1968). Les sa!a111a11drcs d'Furopc. l 'Asic et dA(riq11{' du :\lord. Paris. Lechavalier. Peracca. M. G. ( J 898a ). Descrizioninc di una nuov a specie di Tri tone Italiano. Roll. :\fusci. Zoo!. .�na1. COll/f'. R. Un ir. Tucic. N. and Kalczic. M. L. ( 1984 ). Morphological variation To rino. 13 (:117). 1- 6. within and among populations of the Smooth newt . Tr ililru.1· rnlg11ris. Rio si11c111a1ika I 0 ( 1). .:15-58. Peracca. M. G. (1898b). Note on an Italian newt. .\fol<..:c i/(//ica. Proc. Zoo!. Soc. /.011d ( 18 98). 482-487. Verrell. P. A .. Hallidav. T. R. and Griffi ths. M. (1986). The annual re productive cvclc of the South newt (Tri111ru.1· Radovanovic. M. ( 195 1 a). Vodo::c111cii r;mi::al'ci 11asc ::cml. ic. rnlgaris) in England . ./. 700/. /_(lfllf. 210, 101-1 19. Srf!sko hio/o.1ko dms11·0. Beograd ..Nanc na Kmjig;1 . Wallis. G. P. and Arntzen . .I. \\I . ( 1989). Mitochondrial DNA Radovano\·ic. M. (195 lb). A new race of Alpine newt fro m variation in the Crested newt superspecies: limited Yugoslavia. Rrit . ./. /-IC!p. I (5). 93-97. cvtoplasmic gencllmv among species. Fl'o/11rio11 .:13 ( 1 ). Rafinski. J. and Arntzen . .I . W. ( 1987). Biochemical aspects 88- 104. of the Old World newts. genus Tritums. lsozvme- d;1ta. - Wilson. 0. E. and Brown. W. L. ( 1953 ). The subspecies Herf!cto!ogica 43 (4). 446-457. concept and its taxonomic implic;1tions. S!. '.1/. Zoo!. 2. Ragghianti. M .. Bucci-Innocent. S. and Mancino. G. (19 78). 97-111. Karvologv of the Carpathian newt Tritums 111on1andoni Wolsterstorff. W. ( 1905). 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