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the sore- and hi nil limbs. Vii. Shape of the tali cnd - l. the tip tapers oss gradually to a sine poliat. 2. the tip ta pers oss gradu atly to a blunt poliat. 3. willi a distinct shori si lament. d. xv illi a distinctmoderately long si lament. 5. with a distinct long si lament. VIII. Toe naps on the hi nil limbs- l. oh-
as the leuci sor signi sicanCC. individua is received a score on cach qualitative character consisting os a discrete value sorthe colour and or sit ape trait. These data were statisti catly analyZed using a corresponde iace analysis sol lowing the algorii hin os CREENACRE sl 984). The input data sor this analysis xv c re the two-waycontingen cy tables os the observed freque iacies os character states in the analyZod populations using the PROC CORRESP procedure. The ouli ut os such an analysis was the coordinates os the row populations) and column states os charactei s) corresponde iace axes superimpo sed on the scalter diagram. The proximi ly bet cen populations is interpreteil to evaluate not only the relations hipsa mong each set os variabies, but also ille relations among the row and column Vari abies. The interpretation is based on potnis soli nil in approximate ly the fame direction sim in the origin und in approximate ly the sanae region of the space desined by corresponderace axes. Distances bet cenpoinis do not have a Strat gliis orward interpretation in multiple corresponde iace analysis CREENACRI & HASTI E l98J). so the obtained resulis allowed only evaluation os the relations hips among populations according to the scaturos os the corresponde iace axes estimated by the positions os the columia variabies on the scat ter diagram. Si Ze disserences bet con se males and males xvcre assessed using the single classi sic alio nanalysis os variance sor uia halanceii data applying the PROC CL M procedure that reve aled signis i cant dissere iaces bet ween the sexus sor most os the analyZed characteristi cs. To determine theamount os dissere iaces bet con sexus and among populations in ternas os ex te mal morphology. a multi variate analysis os variance MA NOVA) was u scd. Ali test criteria si catelling 's trace. Wilk'stam da and Pillais trace) showed a significant variation lictween hoth sex es anil populations respective ly. The data sor the sexus were analyZed separate ly in sui ther analyses.
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Canonicat variate analysis hicli maximi Zed the variation hel vecia groups. was used locharacteri Ze the degrue os divergeiace among the populations JAM lis & MCCULLOCH l9903. The canonicat variates were calculaled CANDI SC) and the centroi is os cach sample population. wilhout previ ousty es Priori designatisag populations to particular taxon. xvere plotted on the sit si lxvo ca
Correspondeiace analysis indicates Id dimensions in the correspondeiacet able . hut mosi os them are triviai and minor or os negligibie influence on the mutuat corresponde iace bet ween rows and columns. The c2 os ille fit st three singularva lues depicis about 40 in os total χ' 39.2 in and 42.4ta in males and se males. re speciive ly). e soti nil a great deal os variation in qualitative character istic s. e speciallyin males villa considerable interpopulation overtas' even among dissereni sub- species. Ne Veriheless. population Samples are grouped in I recogniZable manner. xv illi more or less obvious taxonomic mea ning Fig. 2). Populations si ona solitheria Hei Zego vina and Monte negro, named a tonitiXinii sorna see Discussion). vere main ly discern able si om ille re maining populations on the basis of the dominant phenotype characteri Zed by the sol lowing seatures: spois on the ventral patet os ille tali sin sit uated on the tipper tali sin margin
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pale stanti background with reticulate pigmentatiori dorsal crest stat ting fui therbackward and the tali ending with a distinc t. main ly long si lament. Two populations frona the former Yugos lav Republic os Macedonia T. V. graec HS) were Veryclo se to this groti p. In dividuals of the se populations were characteriZed as fol-lows: the lail eniis with a distinc t. shori filament. the ventral part of the tali sin is without pigmentation spois on the ventral part of the tali sin are sit uated on theupper and lower paris with a pale mi dille pelvic line or the spois are fuse l. se- males are with a whi te stripe bet ween the back eyebali margin and the corner of
Most of the populations frona Sio venia and Croatia. which indisputat, ly be- long to T. V. meridic HialiS according to their geographical position were main lyin the tipper test quadrant be ing mostly concentraled around the Zero value of thesii St two corresponden Ce axes. These populations were mostly characteri Zed by the following traiis: a pale throat with su sed dark biotches or with distincti ve dari spois, a low and smooth dorsal crest stari ing frona the back part of the head. i hetati tis' gradu atly tapers os f to a sine poliat, moderate ly devel Oped dorso- laterat foliis on the body. pale stan k background willi distinctive spois; a head with ad ark stripe clown the mihi die. Most of the populations geographi catly be longing to the nominate sub- species were siluated in the tipper right quadrant. These smooth newts typicat lyhail up to three distinct dark spois on the ventral part of the tali sin. pale flant background with unclear spois or without illem the naiddie dark stripe not obvio his on the top of the head intensive colour on the central belly Zone wilh weal marmoration, a deniculated crest under 2 nam; ille tali lip ta pering oss gradu atly toa blunt poliat. the hi nil limbs without si aps; dorso- laterat solds absent or weal lycle vel ope d. Some of the Pannoni an populations Nos 50. β l. 53. 54) moved to theles t. to ward T. v. meridionialiS and the populations frona intergradation Zones. In
species involved. iii med a distinct group sit uated in the Lipper test quadrant. These populations were characteri Zed by tali colour with a pale backg round Un Clearly sp Otted, pale throat without spois, pale central belly Zone with weal marmoration. marginal belly Zone with weali spois, deniculated crest. weli above
Corresponde iace analysis of sthe emales silowed previ ous ly known indistinct taxonomic qualitative characters. Also; ille se males of most populations 3Jout os βο) are morphologicalty very similar. i. e. with almost the sanae characters states concentraled mostly in the lest quadrant, close to the Zero value Fig. 3). The position os other populations presenis no more than a stight Variation distribu ted hi nequat ly between populations but wilhout any taxonomic mean ing.
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Tu hic l. Correlations os the botri cen population varia hi lily os analysed traiis illi the sit si three canoni C:
Fig. 3. Plois os populations sor semales on the sit si and the second corresponde iace axes DIM). Fornumbers of the localities seu the Appendix
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Analysis of the body si Ze snout-vent tength) os intraspecies grotis' differen-ces revealed that males frona the former Yugos lav Republic of Macedonia T. P. grciccus) were statisticatly significantly smaller than ita e sanae sex of the other sub species studi ed. including males frona the intergradation Zones Schesse 'smultiple t-test). Also, males frona areas of subspecific hybridi Zation were significantly smaller than tomiasinii males. Concerning femules, T. V. meridiori iliX were significantly larger than the Other grOUPS. The fit st two canonicat variates depicted accounted for 56 in and β9 in of the totai dispersion in males and females, respective ly. The patieria of character correlations bet ween population Variabili ty and canonicat Variates was sonaewhat different bet ween the sexes Table l). The fit si canonicat variates appeared bipo-lar in both sexes: the distance bet ween nostrils In) and eyebali tength so) were opposed with the distance he tween fore and hind limbs D). In addition. in se males the variabies In and O were contrasted willi tho se of the other measti redi ratis. Correlations os most meas ured traiis with interpopulation variabili ty on thesecond canonicat variates were hi gh and significant in both sexes. This suggeststhat discrimination bet ween populations based on the seconii canonicat Variates may be caused by o Veruli si Ze. Both in females and males only the majori ty of the tom&SiHii form and T. V. gria ocuS 9 of l3. and l l of l3. respective ly) are groupeis together with the hig-hest value on both axes Figs 4 and β). These populations were discriminate d
a. Plois os populations centroids sor males on the sit si and the seconis canonicat axes CAN).For numbers of the localities see the Appendix
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Fig. S. Plois os populations centroids sor scinales on the sit si and the seconii canonicat axes CAN). Eor numbers of the localities sco the Appendix
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grcio MX populations s Mount Ko us . No. 40) is welt with in the tomasinit grotis' os populations s Fig. 2). Certa in ly our data are not conclusive in establis hing tomaXinii as a separate taxonomic unit sub species). Mucii more extensi Ve T. V. g GECHS population sampling is nee led to draw reli able conclusions. However the resulis of Our stud y do support the opinion that the 'griaecuXV problem is not clo sed . Hecata now adit additional proos in favour of this vlew. BOI KAY sl 928) stated that the si uti traiis of individuals of the tomesXinii forna clearly differ frona those of T. V. griae HS individuals frona Corsu is lanii. Also smooth newt populations rangingi rom the River Neretva to Montenegro Caia be distinctly recogniZed frona populations in the former Yugos lav Republic os Macedonia when comparing the phen O- types of the two gene loci sKALEZIC l 984).we suppose that besides the former Yugos lav Republic os Macedonia an d
aled belween T. V. moridionialiX and nominate subspecies. This Zone covers then Orthwest contact be tween these subspecies s Fig. l) as demonstrated earlier byelectrophoretic data KALEZl C l 984). The intergradation Zone of western gr&ECHSV and T. V. VulgiariS subspecies is mostly locali Zed in the Oromediterranean part of the Dinaric Alps while the rest is in the Mediterranean and sub
Mediterranean Zone. The intergradation Zone Os T. V. meridioricillS and T. V. VHl-
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ween T. V. ALGECHS And T. V. VulgiariX in Serbia has nol been confirmed probab lydue to the lack os relevant population sample S. As clai med earlier by SCHMIDTLER L SCHMlDTLER l 983) we confirmed that T. P. XChrotheri cloes not constitute an in dependent taxonomic unit. lndividua is frona the type locali ty os uechroiberi Ravni Kotari. No. id) dici not disserfrona the nominate smooth newis Fig. 2). interest ingly it seems that T. V. Vulgari, frona Ravni Kotari is separated si ona ille rest of the smooth newt range by a narrow Zone with the alpine newt strituruX est i Stri, KALEZl C et est l990). Assar a s we know it is the only casu os fragmentation of the Bali an smooth ne trange without morphological and genetical disserentiation occurring at the sanae time. It seems that smo ait, ne vi populations frona Ravni Kotari and ille Dalma tian Zagora sNos i l9. respective ly) have recent ly been in conlact with the resios T. P. VulgiariX through the large Dinaric karsi stellis; as weli as a long the valle ys of the Sava River tributaries of the area in question. These were the old palh-ways os possit, te amphibi an population communications in the Neogene . in ternasos the distribution os water hodies and laniis during the M iocene KRSTIC l096). Jur state ment scenas to he supported by the sact thal morphological iratis os the Glamoc karsi si eid specimens bolong to the nominotype No. 24. Figs i and 2). Unsortunate ly. only one male in breed ing dres sing was a vallabie frona the locali tyin question. lis morphological traiis were welt with in the T. V. VulgiariS spacebo unded by the fit st two corresponding axes Fig. 2). so it seenas reason able tostippo se ii belongs to the nominale sub species. Tlais population is probab ly a relici of the a bove- mentione d connection pallis. SCllVIDTLER and SCHMIDTLER l 983) suggested iliat the smooth ne vi populations in the Sava River valle y disser morphological ly si om ille rest os thenominotypical populations. e found that populations frona the easteria Pannonian area Vojuod ina) including one ac ros s the Danube No. 50) were groti pedtogether on the fame pote os T. V. VulgiariS morphological variation pallern Fig. 2). Neveriheles s. il would be premature to consider easteria Pannoni an popula tions as an in dependent ly evolving Unil.