Contributions to Zoology, 68 (3) 181-203 (1999)

SPB Academic Publishing bv. The Hague

Geographic variation and taxonomy of crested ( cristatus superspecies): morphological and mitochondrial DNA data

² J.W. Arntzen¹ & Graham+P. Wallis

1 Department of Zoology and Anthropology, Faculty of Sciences, University of Porto, CECA/1CETA/UP,

2 Campus Agrdrio de Vairao, 4480 Vila do Conde, Portugal; Department of Zoology and Centrefor Gene

Research, University ofOtago, P.O. Box 56, Dunedin, New Zealand

crested Keywords: asymmetric hybridisation, biogeography, , geographic variation, morphology,

Triturus mtDNA, taxonomy, cristatus

Abstract Introduction

Within the in the the newt genus Triturus, large-bodied species Studies of geographic variation within and among

T. show cristatus (crested newt) superspecies an unusual degree closely related taxa have provided crucial infor-

ofvariation in relative trunk length as a result ofamong-taxon mation on the nature of species and the process of variation in interlimb vertebral count. Here we examine the speciation (Mayr, 1963). Studies of spatial mor- systematic value of this feature as assessed by both exterior of phological heterogeneity are interest from the measurement (Wolterstorff Index) and direct radiographic count of view of adaptation since differences in ofrib-bearing vertebrae, with particular reference to a number point

ofconfoundingfactors (sex differences,hybridisation, geographic morphology are likely to be functional or at least

mtDNA variation, allometry, preservation effects). Using our have consequences on fitness and ecology. A proper

which with haplotype data, are largely concordant geographic analysis of morphological variation requires inde-

distributionof species, wefind that direct count ofthe rib-bearing pendent assessment of the phylogenetic relation- vertebrae performs more reliably (14% misclassification) than ships of morphotypes (e.g. Losos, 1990). A central external measurement (31% misclassification) as a species question to the systematist is: does geographically identifier. We therefore recommend this feature as a taxonomic

differentiation reflect local tool, although (like external measurement) it breaks down near bounded, morphological

for the observed which would be hybrid zones. To account biogeographicalpattern adaptation (in case convergence

- scenario is and phenotype genotype discrepancies, a presented expected to be common), or is it a function of phylo- that combines movement ofthe contact zonebetween taxa the which geny (in case morphotypes should be mono- with This asymmetric hybridisation. scenario applies to spe- phyletic) (Harvey and Pagel, 1991; Bernatchez, cies interactions in eastern Yugoslavia and western France. 1995)?

Adaptation and phylogeny are not, of course,

mutually exclusive explanations of character dis-

tributions. Contents Adaptive morphological change can

in occur and be proliferated by cladogenesis the

absence of reversal. To resolve this, the first Introduction 181 step

Material and methods 182 is to determine whether different character sets

Results 183 define the same taxa. Concordance oftaxon bound-

Discussion 193 aries resolved different by means, e.g., morphol- Performance of WI and RBV for species diagnosis 193 ogy and genetics, is strong evidence for their real- Adaptation 194 ity (Hillis, 1987; Avise and Ball, 1990). Indeed, Taxonomy and phylogeny 194 often becomes Distribution and biogeography 195 morphological divergence only

Acknowledgements 197 apparent after the taxa have been resolved geneti-

References 197 cally (e.g., Arntzen and Garcia-Paris, 1995;

Appendix 199 McDowall and Wallis, 1996). Here we look at

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geographic variation in vertebral count in the Tri- of the five phenotypes was associated with one or turus cristatus superspecies (or Artenkreis sensu more characteristic mitochondrial genotypes (Wallis

and in WI Rensch) and re-evaluate this character as a tool Arntzen, 1989). Significant differences for species identification, with particular reference were observed between groupscarrying these geno-

be tool for to the several hybrid zones present in the group types, and therefore WI can used as a

Limb interlimb dis- and our genetic analyses of these zones (Wallis species diagnosis. length and and Arntzen, 1989; Arntzen and Wallis, 1991; JW tance are sexually dimorphic characters, making it

Arntzen, in prep.). necessary to calibrate WI for males and females

Wolterstorff revised technical mtDNA data In his classic paper, (1923) separately. For reasons, were the taxonomy of the crested newt, Triturus cristatus available for (almost exclusively) femalenewts only

index. WI for using a morphological He distinguished four (Wallis, 1987). However, we measured males taxa at the subspecies level [carnifex (Laurenti, from the same populations and because (with a few

1768), cristatus (Laurenti, 1768), dobrogicus (Kiri- exceptions) they belong to the same species, WI tzescu, 1903) and karelinii (Strauch, 1870)] and can be calibrated for males as well. The number of provided compelling phenotypic descriptions for rib-bearing vertebrae (RBV) varies in parallel with each of these. He provided a crucial discriminator the inverse of WI (Arntzen and Wallis, 1994; for taxon identification, which has become known Crnobrnja-Isailovic et ah, 1997). Similar patterns as the Wolterstorff Index (WI). WI is the ratio have been documented for lizards (Greer, 1987; between forelimb length (PaL) and interiimb dis- Griffith, 1990; Caputo, Lanza and Palmieri, 1995),

* WI = PaL 100 tance (LiE) and is defined as / fish (Lindsey, 1975) and other salamanders (Jock-

LiE. We have reviewed the available data on WI ush, 1997). and have discussed some of the practical and theo- Here we present additional data to calibrate WI

and and document the existence retical pitfalls associated with its use (Arntzen and RBV of geographic

Wallis, 1994). WI succinctly summarises the mor- variation across and within species. The results

differentiation to much the confirm the limited value of WI for phological among taxa diagnostic

RBV same extent as more sophisticated methods of purposes. The character on the other hand is multivariate morphometric analysis used in crested shown to be diagnostic and without much intraspe-

cific variation. it to newts and other urodeles (Kalezic et ah, 1990; geographic Moreover, appears

Arntzen and Wallis, 1994; Sket and Arntzen, 1994; be largely immune to observer bias, developmen-

Arntzen and Sket, 1997; Cvetkovic, Kalezic and tal or sexual variation, or preservation artefacts.

Dzukic, 1997). From a statistical standpoint, indi- Applying the character at the population level pro-

undesirable duces coherent of variation ces may have certain properties (Atchley a pattern geographic

that et ah, 1976), but the WI has become a popular tool coincides with that of the species, although

external it for species identification of crested newts in the (like measurements) breaks down near field (for example Grillitsch et ah, 1983). How- hybrid zones. To account for the observed pheno- ever, to avoid circular reasoning, the calibration of type - genotype discrepancies and biogeographi-

WI requires independent criteria, i.e., an identifi- cal patterns in eastern Yugoslavia and western

is that combines the cation without reference to body shape. France, a scenario presented

In and its the crested newt superspecies sister movement of the contact zone between taxa with taxon T. marmoratus (Latreille, 1800), WI increases asymmetric hybridisation. in the order: dobrogicus — cristatus — carnifex —— karelinii —marmoratus, describing a morphologi- cal series from slender and short-legged to stout Material and methods and The also character- long-legged. taxa possess istic colouration patterns of that are described on Forelimb length and interlimb distance weremeasured with plastic

Plates I-II (for approximate species distributions callipers (0.1 mm precision) on preserved or live material, the representing 142 populations across total range. X-ray see Fig. I). We previously gathered mtDNA RFLP photographs were taken on preserved or sedated specimens data from female newts and concluded that each (representing 116 populations) with an ‘Elinax 90/20’ and an

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Fig. I. Distribution of Triturus marmoratus and five taxa in the T. cristatus superspecies (after Arntzen, 1995). The crossed hatching

covering Yugoslavia and adjacent regions refers to T. carnifex macedonicus, a taxon recognition resurrected from Karaman (1922).

of0.7 seconds 4mA D10DW exposure at 38kV, on Agfa-Gevaert type (Table I). For reference, we include published and X-ray film. Additional data on WI (31 populations) RBV data from studies dealing with three or more taxa

(38 populations) were obtained from the literature(Vallee, 1959; that were identified phenotypically (Table I). Statis- Kalezic et ah, 1990; Crnobrnja-Isailovic et ah, 1997). Analysis tically significant differences were found between ofvariance (ANOVA) and othertests followed Sokal and Rohlf

all combinations for WI one-tailed t- (1981) and Siegel and Castellan (1988). Logistic equations were species (four

determinedwith SPSS 1990), tests were P < (SPSS, Non-parametric tests between consecutive groups for each sex: performed with StatView (StatView, 1988), Association of 0.001 in all cases, except for males T. cristatus - intraspecific morphological variation and geographical distance T. carnifex and males T. carnifex - T. karelinii was tested for with the Mantel-test (NTSYS 1.80, Rohlf, 1993). with P < 0.05 and females T. karelinii - T.

marmoratus with P < 0.01) and for RBV (four

G-tests Median between five consecutive groups Results with P < 0.001 in all cases). Males consistently

have an average WI of 8-10 points greater than We estimates ofthe Wolterstorff Index present (Fig. females of the same species. A weighted regres- and the number of vertebrae in 2) rib-bearing newts the sion was used to determine best fitting logistic from characterised mtDNA populations by geno- in curve separating species that are adjacent mor-

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Table I. Number of rib bearing vertebrae observed in crested dobrogicus from all other species. On the basis of and marbled Triturus. Mean ± SD; size in newts, genus sample WI, three males (4.2%) and three females (2.9%) brackets. The populations used in the present study were would have been classified as belonging to a spe- idententified to the species onphenotype and mitochondrial DNA cies with (T. cristatus) pheno- (Wallis & Arntzen, 1989). non-corresponding

type. Much higher levels of misclassification were Herre, Lanza et al., present obtained over all species (31%), with no signifi- 19321932 1991 study cant difference in misclassification between the

sexes > G-test of T.T. dobrogicus 17(3) 17.617.6 ±0.51 (14) 17.5 ±0.58 (47) (P 0.05; independence). Using RBV, good separation is obtained between: T. T. cristatus 16(3) 16.016.0 ±0.15 (44) 16.0 ±±0.410.41 (122) dobrogicus and T. cristatus (6.6% misclassified),

T. cristatus and T. carnifex (6.8%), T. carnifex and T. carnifex 15(3) 15.1 ±0.30(148)±0.30 (148) 14.9 ± 0.33 (26) T. karelinii (16.4%) and T. karelinii and T.

T. karelinii 14(1) 14.1 ±0,30±0.30 (40) 14.2 ±±0.440.44 (41) marmoratus (6.7%). Overall misclassification is

13.7%.

T. marmoratus 13,113.1 ±0,34 (49) Application of the RBV cut-off point criteria to

inferred population means (Table II) provides spe-

cies distributions that are largely internally con- The WI for each crested phometric space. typical sistent, in line with prior knowledge, and hence newt species is as follows: males: T. dobrogicus < reflect documented distributions with some excep- 54.0, T. cristatus 54.0-63.69, T. carnifex 63.7-67.09, tions as follows. Populations 1 and 2 are inferred T. karelinii >67.1; females; T. dobrogicus < 46.2,

T. karelinii’ and T. marmoratus’, but are within T. cristatus 46.2-53.89, T. carnifex 53.9-59.19, T. the known geographic and phenotypic of karelinii > 59.2. For RBV the modal values ob- ranges

T. marmoratus and T. karelinii, respectively (Fig. served are: 13 in T. marmoratus, 14 in T. karelinii,

3). Similarly, populations 3-5 are obviously 15 in T. carnifex, 16 in T. cristatus and 17 or 18 in

’ wrongly classified (T. dobrogicus within the T. T. dobrogicus. Limited intraspecific variation is

cristatus and T. karelinii ’ within the observed with 86% of the investigated newts show- range range

of T. carnifex carnifex). Sites 6-8 are known ing modal values. Ranges for the identification of

- T. and T. T. > syntopic T. cristatus marmoratus populations are set as follows: dobrogicus 16.5,

cristatus - T. dobrogicus populations with bimo- T. cristatus 15.5-16.49, T. carnifex 14.5-15.49, T.

dal character state distributions karelinii 13.5-14.49 and T. marmoratus < 13.5. (Vallee, 1959;

Wallis and Arntzen and Median Fisher’s exact tests demonstrated the ab- Arntzen, 1989; Wallis,

for this 1991; Arntzen et al., Individuals with mixed sence of significant sexual dimorphism 1997).

in T. - T. at site 9 character, except T. dobrogicus where females dohrogicus carnifex phenotype

and 10 had RBV scores for T. cristatus{ i.e., tendto have a higher count than males (modal RBV typical

in between the values for T. - of 18 versus 17, P < 0.05). A trend was observed typical dohrogicus

for increasing overlap of character states between T. carnifex) and hence remain unclassified in Fig.

species with increasing WI and decreasing RBV. 3. Populations 11 and 12 were in the field identi-

WI alone is usually sufficient to distinguish T. fied as T. carnifex, showed RBV counts typical

Fig. 2. Reported values ofthe WolterstorffIndex for seven studies that involved three or more taxa in the crested newt superspecies.

= W = = and Fachbach Wolterstorff (1923): range, interrupted lines refer to extreme values; H Herre (1932): mean ± SD range; F

= al. tables 1 3 allocated (1974): range; K Kalezic and Stevanovic (1980) and Kalezic et (1990: to on page 34-35, with populations to

species following Kalezic et al, (1997) but excluding the population from Tresnja which Wallis and Arntzen (1989) identified as a

= mixed T. - T. SD and L Gentile and Torricelli genetically dohrogicus kareliniipopulation: average weighted mean± range; Lanza,

and A = Arntzen and Wallis ± SD and PI and P2 = ± SD and (1991): mean range; (1994): mean range; present paper: mean range.

in identified the distributionsas in 3. Solid bars refer to Populations figuring ‘P2’ were aposteriori, following species Fig. populations

classified on the basis mitochondrial DNA genotype. Sample sizes (if known) are presented at the top of the bars. Horizontal bars

indicate optimal interspecies boundaries with respect to W1 (details see text). Data on T. marmoratus are given for comparison.

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Fig. 3ABC. Newt populations for which mean NVR (number of rib bearing vertebrae) and mtDNA haplotype was determined. Solid lines the of = Triturus cri = T. dob = T. connect records describing edge species ranges: car carnifex carnifex, cristatus, dobrogicus,

= = Numbered in the text. No kar T. karelinii, mac T. carnifex macedonicus. populations are discussed big-bodied newts are known

for the areaapproximately coinciding with Bosnia-Hercegovina. The area marked with a *?’ in northwestern Bulgaria - adjacent parts of Yugoslavia is not devoid of crested newts (see Kalezic et ah, 1997), but the taxon to which they belong is undetermined. The

line C describes with mtDNA characteristic for T. and interrupted heavy on map an area newt populations possessing karelinii (Wallis

Amtzen, 1989); studied populations shown by triangular symbols. Arrows refer to hypothesized dispersal events discussed in the text.

Note that the River connects the Pannonian and Dobrogean Plains through the Iron Gate. For the comprehensive distribution of T. dohrogicus see Arntzen et ah, 1997.

for T. cristatus, an mtDNA haplotype typical for lively low RBV scores (populations 30-33, Fig.

< T. dohrogicus and also remain unclassified. Indi- 3c; mean RBV 15.9) are all situated at the fringe

of the close viduals with mixed T. carnifex - T. karelinii phe- T. cristatus range, to T. karelinii which

notype (site 13) were classified as T. carnifex on typically has low RBV counts. Among 16 T. do- the basis of RBV. Similarly, a newt with mixed T. brogicus populations with N < 5, those with rela-

< - RBV 17. cristatus T. karelinii (site 14) phenotype was tively low counts (mean 5, see Table

classified T. karelinii. Newts from sites 15-20 with situ- as II) are, one exception (population Alap)

are phenotypically T. karelinii, but have RBV scores ated along the southern edge of the T. dohrogicus

higher than average for that species. Sites 6-19 range, whereas those with higher counts (mean RBV

> are all located at the fringe of species distributions 17.5) are, with one exception (population Jamena) and close another where situated from the of the to species, one might expect away edge species range.

interbreeding if the various taxa of crested newts Application of the W1 criterion to the identifica-

were not genetically isolated. No T. dohrogicus or tion ofindividuals and populations produces mixed

T. cristatus samples (except for those mentioned results. For T. dohrogicus, the distribution pattern

above) had RBV scores that did not correspond with obtained with W1 identifications neatly fits the

the 2 documented whereas for the other phenotype. However, among larger (N distribution,

T. Table 5) samples, four cristatus populations with rela- species the results are erratic (compare II

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Table II. Sample site coordinates, sample size (N), mean Wolterstorffindex (WI), mean number ofrib-bearing vertebrae (RBV) and mitochondrial DNA haplotypes observed in Triturus cristatus superspecies and in T. marmoratus. Numbers in brackets refer to sites discussed in the and marked in 3, the text Figure Italicized values are outside range typical for the taxon (no range was identified for

WI in T. marmoratus) or atypical for the taxon (mtDNA haplotypes). Identifications were made by phenotype.

coordinates males females

Phenotype (see Plates I-III)I-II1) latitude longitude N WI N WIWl N RBV N Haplotype ftf.

Triturus carnifex carnifex

Acerra, Napoli, Italy $ 40 50 14 15 3 58.8 2 14.50

Belovar Moravce, Croatia $ (21)(21) 4545 51 16 10 1 74.6 1 15.00

Benevento, Italy (5) 4141 08 14 46 3 14.3314.33

Bobinaco, nr. I’Aquila, Italy $ 4242 21 13 24 3 71.7 3 62.262.2 8 15.13

Etzmandorf, nr. Eggenburg, $(11)$(11 ) 4848 39 15 45 1 61.2 2 16.00

Farma, Torniella, Italy $ 43 08 11 10 3 73.0 1 61.5 66 115.005.00

Firenze, Italy $ 43 47 11 15 1 65.9 5 52.952.9 66 15.00 3 CARS

1 CAR 6

I1 CAR 7

Fuscaldo, Italy $ 39 25 16 02 5 70.1 6 61.6 II11 14.91

Haidlhof, nr. Baden, Austria $ (38) 4747 58 16 10 2 60.1 3 50.750.7 5 15,20 3 DOB II

* Ig (Podstrmec), Slovenia 45 48 14 3434 2626 65.8 19 55.2 20 14,95

Istarske Toplice, Croatia * 45 21 13 54 20 15.0515.05

Kleinmeiseldorf, Austria $$ (12) 4848 4040 15 45 2 62.9 9 54.6 10 16.00 9 DOB II11

Kramplje, nr. Nova Vas, Slovenia $ 4545 44 14 30 2 72.9 6 58.7 88 14.88 6 CAR 8

Lackcnbach, Austria $$ 47 36 16 27 5 14.6014,60

* Licki Osik, Croatia (23) * 4444 36 15 25 10 63.463.4 9 56.3 19 14.95

Locarno, Switzerland $$ 46 10 08 48 1 64.6 4 63.6 66 15.00 3 CAR 9

Pisa, Italy $ 4343 43 10 24 4 56,1 77 15.00 4 CAR 5

Plitvice, Croatia (22) 4444 52 15 37

Radenci (Turjanci), Slovenia * 46 42 16 0404 1111 65.2 13 53.9 18 15.06

* Stanjel (Goce), Slovenia 4545 4949 13 55 13 63.1 18 56.2 20 14.95

Salakovci, Croatia * 4545 03 14 05 20 63.0 15 52.452.4 17 15.06

Sinac, Croatia $ 4444 49 15 22 I1 65.0 2 15.00

* Velika Vala, Slovenia = 4545 40 a= 14 00 20 14.95

** Zumberak (Budinjak), Croatia 45 47 15 30 14 66.1 2020 56.0 1818 15,06

Taxon sample size and weighted mean 113 65.3 141 56.1 234 15.03

Triturus carnifex macedonicus

Ano Kaliniki, Greece $ 4040 52 21 26 33 63.1 8 53.6 5 15.00 8 CAR? 10

** F.R. = 4242 25 18 30 20 15.35 BjcloSi,BjeloSi, Yugoslavia (27) =

Dimitrovgrad, F. R. Yugoslavia (29) 4343 00 22(4722 \ 47 Divcibare, F.R. Yugoslavia $ 4444 06 19 5656 5 71.7 6 64.3 II11 14.91

* Dobrsko Selo, F.R. Yugoslavia 4242 23 19 05 1155 72.3 1188 61.1 19 115.115.11

Donja Cadjavica, Gornja Cadjavica,

Bosnia-HcrccgovinaBosnia-Hercegovina $ 4444 48 19 02 2 60.5 1 15.00

Dubrava Tuzla, Bosnia- Donja cq. Tuzla,

* Hercegovina (24) 4444 28 18 42 10 68.2 17 57.8 19 15.00

* Donji Stoj, F.R. Yugoslavia 4141 54 19 20 15 68.5 13 54.4 18 15.28

* Galicica (Djafa), Macedonia * 4141 04 20 5252 30 63.6 32 55,3 20 14.95

Gornja Cadjavica, Bosnia-Hercegovina $ 4444 45 19 05 4 80.9 5 62.6 1010 15,27

* 42 21 18 58 15.06 Gornji Ceklin, F.R. Yugoslavia = = 18 16

Grcak, F. R. Yugoslavia $ (35) 43 2828 20 57 2 68.1 6 58.5 8 14.6314.63 4 KAR? 12

Karan, F.R. Yugoslavia $ 43 54 19 52 2 73.8 4 64.6 66 15.00

Karbinci, F.R. Yugoslavia $ 41 46 22 14 1 64.6 1 15.00

* Lesnovo, Macedonia 42 0101 22 14 11 75.2 13 67.1 20 14.90

Livadia, Greece 4141 17 23 01 1 64.7

Lucane, F. R. Yugoslavia $ (36)(36) 42 25 21 43 11 61.5 15 14,67 12 KAR? 12

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Table II. Continued.

coordinates males females

Phenotype (see Plates I-I1I)I-III) latitudelatitude longitude N WI N WIWl N RBV N Haplotype 0£

BrdoBrdo * 42 21 Novo cq. Grmija, F.R. Yugoslavia 42 36 21 26 20 64.9 20 58.2 26 14.96

* Petrovec, Macedonia 41 56 21 37 13 62.0 9 55.6 19 15,0015.00

* Prilep, Macedonia * 41 19 21 31 16 60.1 II 53.2 18 14.89

Probistip, Macedonia $ 41 59 22 10 2 80.5 3 62.8 4 14.50

Radosice, F.R, Yugoslavia * 43 17 20 42 11 62.8 25 58.3 17 14.94

* Rtanj, F.R. Yugoslavia 43 46 21 56 16 56.7 19 51.5 17 14.94

Sarajevo, Bosnia-FlercegovinaBosnia-FIercegovina (25) 43 52 18 26

Stanisinci, F.R.F.R. Yugoslavia $ 43 31 20 53 2 66.9 2 65.1 4 15.00

Tavna Monastire, Bosnia-Hercegovina $ 44 36 19 04 4 69.3 5 65.0 9 14,78

Valjevo (Bukovac), F.R. Yugoslavia * 44 11 19 59 14 63.4 13 58.1 20 14.70

Visegrad, Bosnia-Hercegovina $ 43 47 19 20 4 74.1 11 64.8 15 14.73

* Vlasina, F.R. Yugoslavia 42 42 22 23 17 64.9 17 59.2 32 14,78

Vranje (Sv. Ilija), F.R. Yugoslavia * 42 37 21 52 21 67.8 11 59.8 1818 14.94

Zelengora Mountain, Bosnia-Hercegovina

((26)26 ) 43 21 18 32

.

Zupa (Rataje) cq. Rataje,

* F.R. Yugoslavia * 43 29 21 08 33 67.5 24 55.3 1818 14.9414,94

* Zupa, F.R.F.R. Yugoslavia 43 29 21 08 •• 21 14.71

TaxonTaxon sample size and weighted mean 271 66.1 306.. 58.2 427 14.93

Triturus crislaluscristatus

Ambleteuse, France $ 50 46 01 37 1 71.0 1 53.8 2 16.00

Biel, Switzerland $ 47 09 07 16 4 55.3 1 51.3 5 16.2016.20

Bogdana, Romania 47 02 23 01 9 62.1 10 51.5

Bor, F. R. Yugoslavia $ (37) 44 02 22 08 2 60.5 2 45.4 5 16.10 2 KAR? 12

Bumbe§ti-, Romania 45 10 23 23 11 66.9 10 55.4

Canterbury, UK $$ 51 17 01 05 2 65.1 2 51.2 2 16.00

Cimpeni, Romania $ 46 23 23 05 3 63.5 7 51,3 9 16.0016.00 5 CR1 1

Craciune§ti, Romania 47 58 23 58 11 63.8 11 56.4

Cra§na, Romania 45 35 26 13 11 58.7 10 48.3

Doetinchem,Netherlands $ (4) 5151 58 06 1717 I1 61.0 1 17.0017.00

Drighiu, Romania 4747 11 22 40 7 59.8 4 50.950.9

Galole§ti, Romania 45 34 27 00 6 66.8 14 55.8

Halma§d, Romania 4147 06 22 21 8 60.0 5 48.6

Hoensbroek, Netherlands $$ 50 55 05 55 1 65.0 2 52.5 2 16.00

Houthem, Netherlands $ 50 52 05 46 1 62.0 1 16.00

Hurezani, Romania 44 50 23 09 6 63.8 4 55.7

Huta-Certeze, Romania 47 55 23 29 7 64.1 71 54.1

Jabukovac, F.R. Yugoslavia $ 44 20 22 24 2 57.0 3 46.646.6 7 16.00 2 CRI 2

Jitia, Romania 45 35 26 44 10 66.4 71 54.8

Klokocevac, F.R. Yugoslavia $ (30) 44 20 21 12 2 60.6 3 46,646,6 5 15.80 3 KAR? 12

Lanckorona, nr. Wadowice, Poland $ 4949 54 19 29 1 65.0 6 56.8 8 16.00

Limanowa, Poland $ 49 43 20 25 10 51.5 9 16.00 10 CRI 1

Maidstone, UK 51 17 00 32 2 CRI 1

Marghita, Romania 4747 21 22 20 2 56.7

Mayenne, France (6) 48 18 - 00 37 86 16.03

Mayenne, France $$ (6) 48 18 - 00 37 31 60.7 52 55.0 115 16.00 40 CRI 1

Milanovac, F.R. Yugoslavia $ (31) 44 11 21 36 2 55.8 5 55.3 10 15.9015.90

* Mogielnica, Poland 51 42 20 41 19 15.95

** Negotin, F.R. Yugoslavia 44 14 22' 33 13 57,757.7 16 48.8 1919 16.00

Ottenstein, nr. Zwettl, Austria $ 48 28 14 17 11 52.9 2 15.50 1 CRI 1

- Oxford, UK 51 46 - 01 1515 1 CRI 1

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Table II. Continued.

coordinates males females

Phenotype (see Plates I-I1I)l-III) latitude longitude N WI N WI N RBV N Haplotype 0(4

Peterborough, UK $$ (3) 52 3535 - 00 1515 19 58.0 17 55.0 1 17.00 1 CRI 1

Picaturile, Romania 44 29 23 52 9 63.2 6 52.2

§ebi§, Romania $ (7) 46 22 22 06 2 62.7 1 51.951,9 3 16.00 I1 CRI 1

Sinaia, Romania $ 45 20 25 33 10 48.3 II11 16.00 6 CRI 1

St L6, France 49 07 - 01 05 1 64.6 I1 CRI 1

St. Michielsgestel, Netherlands $ 51 38 05 21 1 16,00

Stubik, F.R. Yugoslavia * (32) 44 1818 22 22 14 56.4 15 50,4 16 15.88

Stubik, F.R. Yugoslavia $ (33) 44 1616 22 22 1 58.158,1 4 50.4 5 15,8015.80 4 CRI 2

Ta§nad, Romania 47 28 22 36 1 55.8

Tirgoviste, Romania $ 44 56 25 27 19 63.4 14 52.3 9 16,0016.00 3 CRI I1

Turt, Romania 47 59 23 12 I1 60,3

Videle, Romania $$ 44 16 25 31 3 54.6 7 48.7 10 16.00 3 CRI 3

I1 CRI 4

Vipere§ti, Romania 45 14 26 28 9 59.9 3 50.3

Virfuri, Romania $ (8) 46 17 22 28 19 63.3 12 52.5 10 16.00 3 CRI 1

Winterswijk, Netherlands $ 51 58 06 44 12 16.08

size and Taxon sample weighted mean 248 61.4 286 52.7 385 16.04

Triturus dobrogicus

(Pannonian)

Alap, Hungary $ 46 48 18 41 2 43,4 6 17.33 I DOB 11

Albertirsa, Hungary $ 47 14 19 36 I1 44.1 1 18.00

Andrid, Romania 47 31 22 21 11 41.7 3 37.7

Babe§ti, Romania 47 58 23 06 I1 60.5 I1 44.0

Beograd, F.R. Yugoslavia $ 44 50 20 30 6 41.9 2 17.50 6 DOB 1111

Cegenydanyad, Hungary 47 56 22 30 1I 50.2

Debrc, F. R. Yugoslavia $ (34) 44 36 19 52 16 51.1 10 43.2 29 17.38 8 DOB 11

KAR? !212

Drbsing, Austria $ 48 33 16 55 4 18.00

Dugo Selo, Croatia $ 45 49 16 15 1 36.8 1 18.00

Ecka, F.R. Yugoslavia $ 45 18 20 27 7 49,6 5 42.0 12 17.73 2 DOB 11

Gherteni§,Ghcrtenis, Romania 45 23 21 45 1 59.0 2 48.0

Glusci, F.R. Yugoslavia $ 44 53 19 32 5 45.4 6 42.9 7 17.00 I1 DOB 11

* Ivanovo, F.R. Yugoslavia 44 44 20 43 31 46.5 20 38,9 14 17.43

Jamena, F.R. Yugoslavia $ 44 54 19 02 4 50.0 4 42.7 99 17.56 I Kormend, Hungary $$ 47 01 16 36 1 40.3 1 18.00

* Novi Knezevac, F.R. Yugoslavia 46 02 20 07 10 51.2 14 40.0 20 17.75

Obedska Bara cq. Obrez,

* F.R. Yugoslavia 44 44 19 59 14 48.8 17 41.5 17 17,24

Ocsod, Hungary $ 46 56 20 23 3 48.7 4 39.9 9 17.89 I1 DOB 11

Opovo, F.R. Yugoslavia $ 45 02 20 26 1 48.4 7 40.7 13 17.54

Orle, Croatia * 45 41 16 14 16 52.9 15 42.3 17 17.18

Pi§colt, Romania 47 35 22 18 1 44.5

Podgorac, Croatia $ 45 29 18 12 1 35.1 2 17.50 1 DOB 11

§ebi§, Romania $ (7) 46 22 22 06 I1 50.8 2 37.5 3 17.67 2 DOB 11

Senta, F.R, Yugoslavia $ 45 55 20 06 4 47.8 6 46.446.4 13 17.62 1 DOB 11

Slavonski Brod, Croatia * 45 II11 18 03 14 53.7 10 42.5 20 17.00

Svetozar Miletic, F.R, Yugoslavia * 45 52 19 18 10 46.2 16 41.4 17 17.71

Szolnok, Hungary $ 47 10 20 10 I1 47.8 1 17.00

Tadten, Austria $ 47 46 17 00 3 17.33

Virfuri, Romania $ (8) 46 17 22 28 1 37.9 I1 18,0018.00 1 DOB 11

Vrsani, Bosnia-Hercegovina $ 44 50 19 01 8 41.6 15 17.07 5 DOB 11

Zupanja, Croatia $ 45 05 18 42 ' 6 39.5 7 17.0017,00 4 DOB 11

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Table II. Continued.

coordinates males females

Phenotype (see Plates I-III) latitude longitude N WI N WI N RBV N HaplotypeFlaplotype 0

(Dobrogean)

Amara§ti de Jos, Romania 43 59 24 06 2 56.5 11 42.0

Ariciu, Romania 45 22 27 31 5 50.2 2 45.545,5

Hogioaia, Romania # 45 11 27 57 11II 45.0 1515 39.6

Kladovo, FR Yugoslavia $ 44 36 22 33 2 48.7 2 17.00

Obrejita, Romania 45 2929 27 08 1212 56,556.5 9 44.1

Prahova Forest, N of Bucure§ti, Romania 44 4949 25 52 4 55,355.3

Svistov, Bulgaria $ 43 37 25 21 2 42.4 2 18.00 1 DOB 11

Tartal, Romania 44 10 24 08 4 58.3 5 5/.51.88

Zimnicea, Romania $ 43 39 2525 21 2 37.6 2 17.00 11 DOB 1111

Taxon sample size and weighted mean 182 50.1 205 41.6 250 17.49

Triturus karelinii

(Europe)

Arandjelovac, F.R. Yugoslavia $ 44 19 20 3535 6 68.5 33 62.6 9 14.00 3 KAR?ICAR? 1212

Bansko, Macedonia $ (19) 4141 23 22 46 11 61.3 1 15.00

* Berovo * cq. Smojmirovo, Macedonia

(18) 41 44 22 5050 9 •70.4-70.4 13 62.0 19 14.53

Bigla, Macedonia $$ 4141 56 22 40 2 71.0 3.67,13. 67.1 5 14.00

Dafnohori, Greece $ (20) 40 57 22 48 22 61.6 11 62.5 3 15.00

Djurinci, F.R. Yugoslavia $ (16) 44 30 20 38 11 72.1 11 56.056.0 2 15.00

Gornja Sabanta, F.R. Yugoslavia $ (2) 43 54 21 00 1 65.3 I1 13.00

Grivac, F.R. Yugoslavia $ 43 58 20 40 3 72.6 6 63.6 9 13.89

Guberevac, F.R. Yugoslavia $ 4343 49 20 4646 2 63.7 4 62.462.4 5 14.00 33 KAR? 1212

11 KAR? 1414

Istanbul, Turkey $ 41 02 28 57 6 68.6 3 60.5 9 14.20 22 KAR 17

Karlovo, Bulgaria $ 42 38 24 49 3 75.0 1 57.157.1 5 14,0014.00 11 KAR? 1212

Kentriko, Greece $ 41 10 22 54 55 74.374.3 8 62.8 14 14.2914.29

* Kozuf (Visoka Cuka), Macedonia 41 17 2222 20 16 67.767.7 22 55.7 19 14.26

Levski, Bulgaria $ 43 25 2525 08 4 67.967.9 55 61.2 10 14.2014.20 5 KAR? 12

Mitrasinci, Macedonia $ 41 45 22 46 2 72.6 10 60.9 1414 14.29

Rakovski, Bulgaria $ 42 16 2424 58 4 59.5 4 14.25 3 KAR? 12

Resaskaica Pecina (Resavica),

F.R. Yugoslavia $ (17) 44 0303 21 36 11 65.165.1 2 63.6 3 14.6714.67

Sevlievo, Bulgaria $$ 43 28 2525 13 33 65.665.6 3 14.33

Sisevac,Sisevac, F.R. Yugoslavia $ 43 56 21 37 3 71.5 1 63,0 3 14.25

* Tresnja, F.R. Yugoslavia * (15) 44 36 2020 35 13 59.559.5 56 50.1 20 14.35

Tresnja, F. R, Yugoslavia $ (15) 44 3636 2020 35 99 50.5 9 14.6714.67 3 DOB II11

33 KAR? 12

33 KAR? 13

(Asia)

Adapazari, Turkey $ 40 45 30 23 3 73.3 3 59.4 16 14.06 1 KAR? 15

1 KAR? 16

1 KAR 17

Bartin, Turkey $ 41 37 32 20 4 78.0 6 64.4 10 14.10

Gokgekent, Turkey # 4040 42 36 37 15 67.2 20 62.7

Karacabey, Turkey $ 4040 15 28 18 1 68.5 3 14.00

Resadiye, Turkey # 40 24 37 19 50 64.3 33 59,259.2

Serafiye, Turkey # 40 09 3f37' 46 50 68.5 29 64.0

Tokat,Tokat, Turkey # .40 20 36 3535 42 70.3 35 63.3

Taxon sample size and weighted mean 245 67.8 281 58,9 196 14.43

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Table II. Continued.

coordinates males females

Phenotype (see Plates I-I1I)I-III) latitude longitude N WI N WI N RBV N FlaplotypeHaplotype 0

Triturus marmoratus marmoratus

Barcelona, Spain $ (1) 41 25 0202 1010 1 79.0 2 14.00

Confolcns,Confolens, FranceFrance $$ 46 01 0000 40 1 74.8 I 13.00 1 MAR 18

El Berrueco, valley of Rascafria, Spain $ 40 5454 - 0303 53 55 79.379.3 55 73.7 8 13.38

Mayenne, FranceFrance (6) 4848 18 - 00 37 64 13.30

Mayenne, FranceFrance $ (6) 48 18 - 0000 3737 15 75,075,0 4040 68.9 83 13.12 1919 MAR 18

1 MAR 20

Rochechouart, France $ 45 49 0000 5050 4 66.4 8 13.38 33 MAR 18

11 MAR 19

Salamanca, Spain $ 40 58 - 0505 40 2 61.5 4 13.25

- Santillana del Mar, Spain 43 24 - 0404 06 3 65.3

Valongo, Portugal $ 41 II11 - 08 3030 22 74.574.5 3 13.00

Vilar Formoso, Portugal $$ 40 37 - 0606 5050 11 71.071.0 22 13,0013.00

Triturus m. pygmaeus

- 04 23 11 73.0 11 13.00 Archidona - Loja, Spain $ 37 06 04 23 11 73.0 11 13,00

Cadiz, Spain 36 32 - 06 18 3 13.33

FloyoHoyo de Manzanares - Torrelodones,

- Spain 40 35 - 03 56 3 67.2

- 12 13.00 10 PYG 21 Puerto de Galiz, nr. Ubrique, Spain $ 36 41 05 27 12 65.5

Rio Alberite, Spain $ 36 24 - 05 39 22 13.09

Venta del Charco, Spain $ 38 12 - 04 16 10 62.2 13 13.08 10 PYG 22

Villalba, Spain $ 40 38 - 04 00 5 71.9 66 68.1 15 13.06

24 75.9 55 68.8 175 13.22 Taxon sample sizesize and weighted mean 24 75.9 55 68.8 175 13.22

mixed

Donja Cadjavica, Bosnia-Hercegovina $$

(10) 44 48 19 02 3 67.3 4 54.1 11 15.64

Lukovo, F. R. Yugoslavia $ (14) 43 48 21 51 11 71.8 1 14.00

France =4848 18 -00- 00 37 64 14.73 Mayenne, t (6) = = 37 14.73

= = Mayenne, France $ (6) =4848 18 -00- 00 37 20 66.9 36 59.6 84 14.44 10 CR1CRI 1

Vedropolje, Croatia $ (9) 45 19 1616 36 11 62.6 2 16.50

Vitanovac, F. R. Yugoslavia $ (13) 43 43 20 48 3 74.2 2 60.560.5 4 15.0015,00

undetermined I1

Sibenik - = = 1^16 - Split, Croatia (28) =4343 30=30 00

i see Wallis & Arntzen (1989) and Arntzen & Wallis (1991)

$ voucher material deposited at the Institute for Systematics and Population Biology (Zoological Museum), University ofAmsterdam

II Vallee(1959)

* Kalezic et al. (1990) and Crnobrnja-Isailovic et al. (1997)

# data courtesy of D. Cogalniceanu and K. Olgun

with Fig. 1). On the basis of the reconstructed spe- 50% in T. carnifex and 39% in T. karelinii. No cies distributions (Fig. 3) the frequency of incor- significant differences in classification scores are

identification of individual observed between the for karelinii rect crested newts ap- sexes, except T.

the WI in plying can be estimated post hoc manner for which males are more often misclassified than

11 % in as T. dobrogicus, 42% in T. cristatus, > females (G-test of independence, P < 0.001). Clas-

Downloaded from Brill.com10/08/2021 06:56:32PM via free access Contributions to Zoology, 68 (3) - 1999 193

sification success is affected of and classification made not significantly by cacy WI compromise

in T. < Mann- The WI sample size, except carnifex {P 0.05, solely on this basis. purports to capture

Whitney U-test, one-tailed). information useful in taxonomy but in fact con-

Mantel tests indicate a marginally significant founds the variables limb length, vertebral num- association between geographic distance and WI ber, and possibly vertebral length.

for males and females T. cristatus and male T. Using mtDNA haplotype (Wallis and Arntzen,

~ while the association dobrogicus {P 0.05), of 1989) we showed that WI makes a good approxi-

with RBV geographical distance is significant (P mation to species classification, but can only be

< 0.05 for T. cristatus and P < 0.01 for T. dobro- used with confidence for discriminating adult T. gicus). A series of ANOVA’s with ‘sex’ nested under dobrogicus from the other species. In contrast, the

in WI ‘group’ confirmed the difference between numberof rib-bearing vertebrae (RBV) as assessed

the sexes {P < 0.001) and between the following a by radiography eliminates all but one of these prob- priori identified groups (cf. Fig. 1): T. m. lems. Because RBV is a direct discrete meristic marmoratus versus T. m. pygmaeus (Wolterstorff, count, stable through the lifetime of the individual,

1905) {P < 0.01), Pannonian versus Dobrogean T. and with limited intraspecific geographic differen-

T. in dobrogicus (P < 0.01) and carnifex carnifex versus tiation its use conjunction with a diagnostic genetic

T. carnifex macedonicus (Karaman, 1922) (P < character leaves only the issue of hybridisation to

caused 0.05), whereas no significant difference was ob- be addressed. Variation by hybridisation near served for Asian versus European T. karelinii. A regions of parapatry is difficult to disentangle from significant difference was also found between T. intraspecific geographic variation on the basis of carnifex carnifex and T. carnifex macedonicus in morphological data alone. Under both scenarios,

RBV (Median (7-test, P < 0.05) but not between T. variation will be most clearly expressed when m. mannoratus and T. > from of the in. pygmaeus (P 0.05). samples remote parts geographic range

The limited sampling of Dobrogean (versus are compared. However, the observation that char-

Pannonian) T. dobrogicus and Asian T. karelinii acter state changes are consistently in the direction

(versus European T. karelinii) precluded their test- to that of the neighbouring species supports the

ing. hypothesis of hybridisation, rather than that of in-

trinsic geographic variation. For example, the sig-

nificantly different values in T. carnifex Macedonians

Discussion (high WI, low RBV compared to T. carnifex

carnifex) may well be a result of introgression from

Performance ofWI and RBVfor species diagnosis T. karelinii, whose mtDNA prevails in some T. carnifex Macedonians populations (Wallis and

In that the meristic our previous work concerning the capacity of Arntzen, 1989). Note, however,

in not the Wolterstoff Index to discriminate females of count hybrids is necessarily intermediate to

that the taxa, we identified several potential problems of the parental species, as documented for

salmonid fishes Allendorf and (Arntzen and Wallis, 1994): 1) statistical represen- (Leary, Knudsen,

crested tation (the need for a mean value), 2) non-biologi- 1985). If hybridisation betweentaxa of newts is it cal variation (preservation and measurement dif- a common phenomenon, should be possible to

ferences), 3) sexual variation (WI is higher formales find genetic markers for the species covarying with

than for females), allometric variation(WI decreases interspecific morphological variation. The observed

with breakdown of the of RBV in size), 4) geographic variation (nearby animals diagnostic power ar-

the of may tend to be more similar within species), 5) eas where taxa meet may reflect true nature

characters in contact hybridisation (hybrids between two species can have a zone.

It is that RBV influenced intermediate values typical of a third species), and possible is early on in

circular is often 6) reasoning (the need for an independent development. Indeed, vertebral count highly

character set to determine the significance of WI). labile in fish and salamanders (e.g. McDowall, 1970;

All of these factors to some extent reduce the effi- Jockush, 1997), with cooler conditions generally

Downloaded from Brill.com10/08/2021 06:56:32PM via free access 194 J.W. Arntzen & G. P. Wallis - Geographic variation in crested newt superspecies

slowing development and increasing several mer- ter, T. carnifex four months, and T. cristatus five

istic counts (Barlow, 1961 and references in Jock- months (Bouton, 1986; Griffiths and Mylotte, 1987;

ush, 1997). In Triturus vulgaris the average RBV Andreone and Giacoma, 1989), while the aquatic

count increases with the temperature at which the phase of T. dobrogicus usually lasts six months

embryos are raised (Orska and Iraiolek, 1962), while (Karaman, 1948; Jehle et ah, 1997). No data are

for salamander T. other species more complicated en- available on the phenology of karelinii. This

is short vironmental effects were found (Lindsey, 1966; species predicted to have a aquatic phase,

intermediate that of and Peabody and Brodie, 1975). To address the ques- of length to T. carnifex T.

tion to what extent the variation in RBV is geneti- marmoratus. In terms of performance, the sinu-

cally determined and to gain insight into the rela- soidal swimming ability shouldbe best in the lowest

tionship between embryonic development and adult WI / highest RBV count taxon, he., T. dobrogicus

morphology requires experimental work. Triturus and relatively poor in T. marmoratus. Indeed, the

dobrogicus might be the best species to work with ecological niche of T. dobrogicus is different from

because it is naturally polymorphic for RBV. that of the other big-bodied species. It may co-

exist with fish in oxbows, river margins and other

non-temporary water bodies (Arntzen et ah, 1997).

Adaptation The observed ventral aposematic coloration pat-

T. tern in the cristatus superspecies versus the dorsal

the colouration of T. marmoratus Elongation of the body and a reduction in length aposematic (particu-

of limbs in vertebrates generally indicates a more larly evident in the entirely terrestrial juveniles) further piscine locomotion by sinusoidal body undulation. provides support to our interpretations. Predictions about such in the Although this can be associated with some terres- performance, as gath-

in trial habitats (as in sand-swimming skinks), it is ering of food or predator avoidance behavior

more usually an adaptation to a more aquatic mode the aquatic versus terrestrial habitat could be tested

of life. Reduction of trunk size and the develop- experimentally.

ment of robust legs is more unequivocally associ-

ated with a terrestrial mode of life where the body

requires more support (Young, 1950;Lande, 1978). Taxonomy and phytogeny

The wide variation in body shape in the leg-less

of the Gymnophiona, ranging from stout to thread-like, The depth differences among taxa, and the

with behavior relative of the led may also be associated locomotory sharpness contact zones us to

and ecological adaptation (Renous and Case, 1989). follow earlier suggestions to raise the taxa to full

In the salamander extensive status and genus Batrachoseps, species (Bucci-lnnocenti, Ragghianti

geographic variation in RBV, possibly related to Mancino, 1983) as have others (Frost, 1985). The

fossoriality, has been observed, most ofwhich was ' available data, unfortunately, do not support a single

shown to be genetically determined (Jockush, 1997). phylogenetic hypothesis for the four taxa comprising

female T. cristatus RBV Selection on fecundity (correlated with inter- the superspecies. is primitively

limb length) and male sexual performance (stat- 14 in the genus Triturus (B. Lanza et ah, in prep.),

also ure at display correlated with leg length) may rendering RBV of 13 an autapomorphic character

for - play a role (Arntzen and Wallis, 1994). state T. marmoratus and RBV of 15 18 a

Morphology predicts the aquatic period of T. synapomorphic character state series for T. carnifex

marmoratus to be short, that of T. carnifex to be - T. cristatus - T. dobrogicus. This character alone

intermediate and that of T. cristatus -and T. dobro- would suggest that T. karelinii represents the old- gicus in particular - to be long. While of course est extant crested newt lineage, followed by T. carni-

the and This phenology of breeding may be different from fex, T. cristatus T. dobrogicus. evolutionary

to this corrobo- classification is somewhat year year, prediction appears to be supported ambiguously

rated by field data. Triturus marmoratus spends by the phenetic analysis of protein electrophoretic

annually approximately three months in the wa- data (Crnobrnja, Kalezic and Dzukic, 1989) but

Downloaded from Brill.com10/08/2021 06:56:32PM via free access Contributions to Zoology, 68 (3) - 1999 195

1922. contradicted by another such study (Litvinchuk et nii var. macedonica Karaman, Considering ah, 1994). The phylogenetic analysis of molecu- the morphological and genetic differentiation be-

different the this lar data (mtDNA RFLP’s) suggests a tween forms, we propose raising taxon to phylogeny. Looking at the most-parsimonious the subspecies level and, supported by phyloge- mtDNA tree (Wallis and Arntzen, 1989: Fig. 4), a netic arguments, classifying it as belonging to T.

karelinii tree that optimizes RBV character-statechange [tree: carnifex (not T. as suggested by Karaman, character structure (MAR,PYG: 13)/ancestor: 14/ 1922). Therewith, Triturus carnifex var. albanicus

1959 T. (KAR?,KAR: 14(CAR?,CAR: 15(CRI: 16,DOB: 17,18)))] Dely, is a junior synonym of carnifex involves moving only the ‘DOB’ branch (with ter- macedonicus (Karaman, 1922). Following our cor- minal taxon number 11). [CAR? and KAR? refer respondence with co-author J. Crnobrnja-Isailovic to deeply differentiated haplotype lineages within (J. W. Arntzen, in letter, 1996) this taxonomic solu-

T. carnifex and T. karelinii. We now recognize the tion is accepted by Kalezic et al. (1997), although

T. the referred in feminine first of these as belonging to carnifex mace- taxon is incorrectly to

donicus (see below) while the other will be sub- gender.

T. ject to taxonomic description at the subspecific level The two more massive newt species, karelinii

show (S. Litvinchuk et al., in. prep.)]. If DOB were placed and T. carnifex, much greater restriction site with the T. cristatus (CRI) haplotypes, increased variation than the other two species (Wallis and

also have RBV becomes a derived character interior to the Arntzen, 1989). They slightly larger

basal. mitochondrial and for tree, with the more massive built newts genomes a greater tendency

has 70 insertions in the control These Although this haplotype tree steps as op- region (Wallis, 1987). posed to 67 in the published maximum parsimony factors suggest that the two northern species may tree (Wallis and Arntzen, 1989: Fig. 4), there is have been subjected to long-term small popula-

above 50% for of the tion size during glaciations (Wallis and Arntzen, no bootstrap support any

and it is conceivable that the crested newt species-level structure. That is to say, 1989), evolutionary

the relationship DOB(KAR(CAR,CRI)) is only change in vertebral count is related to this popula-

defined by three synapomorphies in total and the tion genetic feature. tree could more conservatively be depicted as a

four-way polychotomy at this level. This incom- plete resolution is appreciated by Wallis and Arntzen Distribution and biogeography

(1989: 99) and emphasised by further analysis (Faith

and Cranston, 1991; Faith, 1992). However, strong Crested newts appear to be absent from the larg-

support is obtained from the ‘CAR’ and ‘CAR?’ est part of Bosnia-Hercegovina (see for example

mtDNA haplotypes for the sister taxon status of Schmidtler and Schmidtler, 1983; Kalezic, Dzukic

Italian and central Balkan crested newts. These and Tvrtkovic, 1990; Kalezic et al., 1997). The

the groups of populations are also united by the synapo- southeasternmost localities of T. carnifex to

character state RBV = 15. We therefore northeast of the in the distri- morphic perceived gap species

consider the crested newts from the central Balkan bution are sites 21-23 [Belovar Moravce (Table

The of this to belong to T. carnifex. range species II); Plitvice (Fejervary-Langh, 1943) and Licki Osik is disjunct (see below). Newts from both parts of (Kalezic et al, 1990)]. Further to the southeast T.

the range are phenotypically distinct: T. carnifex carnifex is found at sites 24-27 [Donja Dubrava

from the western (Italian and Slovenian) part of (Kalezic et al., 1990), Sarajevo (Bolkay, 1929; com-

the ill-defined black G. Moun- range typically have few, large, municated by Dzukic), the Zelengora dots on the bellies, whereas T. carnifex from the tain (Bolkay, 1928) and Dobrsko Selo (Kalezic and

of the R. and recorded local- eastern part range (F. Yugoslavia Dzukic, 1990)]. The easternmost

Greece) have ventral coloration patterns with many ity is site 29 at Dimitrovgrad (Radovanovic, 1964).

sharp-edged spots, as Freytag (1988) observed, not Crested newts of unknown taxonomic affinity were

unlike that of T. cristatus (Plates I-II). Crested newts recorded at the Dalmatian coast [site 28, situated

of the described kareli- eastern group were as Molge in between Sebenico (= Sibenik) and Spalato (=

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also mentioned Buresh On distri- Split) (Werner, 1897, by a gross geographic scale, phenotype

and Zonkov, 1941)], but with no clearly indepen- butions, and mtDNA haplotype distributions are

dent confirmation for over a century we doubt the concordant. However, in northern Yugoslavia the

of this mtDNA validity record. Dzukic (1993) considers ‘KAR?’ haplotype is more widespread than

the distribution of T. be the T. karelinii distribution would carnifex not to interrupted phenotype sug-

but continuous, following a strip of land to the south gest (Fig. 3) (Wallis and Arntzen, 1989). The

of the river, without, however, presenting data ‘KAR?’ mtDNA haplotype is locally found in T.

this supporting view. The area where crested newts dobrogicus, T. cristatus, and T. carnifex macedoni-

coincides with the ofthe karst The are absent core area cus populations. reverse situation, with a for-

(Sket, 1994), where most natural water bodies are eign haplotype in T. karelinii, has been observed

ephemeral and do often not support the larva! de- once (the ‘DOB’ haplotype in population 15).

of with larval with either velopment species a prolonged phase, Populations foreign haplotypes possess

such as crested newts. The small-bodied newts such two haplotypes - the original plus an alien, such as

as T. alpestris and T. vulgaris in contrast are wide- at site 15 and 34 in T. karelinii and T. dobrogicus),

spread and locally abundant. They may reproduce or just the alien haplotype [‘KAR?’ in T. cristatus

successfully in shallow and temporary ponds such (site 30 and 37, N = 5) and in T. carnifex mace-

a wheel ruts (Winkler and Brauns, 1990) and the donicus (site 35 and 36, N= 16; Wallis and Arntzen,

for the small newt T. is To account for these observations dispersal rate vulgaris es- 1989)]. we sug-

timated to be higher than that for the big newt T. gest the following scenario. In former times T.

cristatus (Stensjo, 1998). While most contempo- karelinii was more widespread than at present, with

rary newt ponds are man-made and rarely desic- a range approximately coinciding to the present

cate (i.e., watering holes for cattle), the puddles day distribution of the ‘KAR?’ haplotype. By dis-

formed fallen and southwards and by trees springs may originally persing northwards, respectively have been the habitat for the small T. and T. macedonicus typical breeding cristatus carnifex super-

T. which the bodied species. seded karelinii, in process range of

The distribution of the four crested newt spe- T. karelinii south ofBelgrade became isolated from cies in F. R. Yugoslavia is complex (Fig. 3c). Tri- the main stock (see the arrows in Fig. 3c). The tiums dobrogicus is found all over the Pannonian genetic interactions between T. karelinii at one side

and T. and T. and Dobrogean Plains. Both parts of the range are cristatus carnifex macedonicus at probably connected by the Danube where flowing the other where such that the formation of FI hy- through the Iron Gate (Arntzen et al., 1997). Triturus brids was asymmetric, with hybrid offspring and

has wide the cristatus a European range, is widespread subsequent backcrosses possessing the (mater- over Romania and reaches southwards over the Iron nally inherited) T. karelinii mtDNA. This scenario

Gate into Yugoslavia. Triturus carnifex macedoni- is surprisingly similar to the one we described for

is of the — cus widespread over most Yugoslavia, T. cristatus T. marmoratus interactions in west-

Former Yugoslavian Republic of Macedonia, Al- ern France (Arntzen and Wallis, 1991). In France, bania, and northern Greece. Triturus karelinii is T. cristatus supersedes T. marmoratus, forming T. found south and southeast of enclaves in the immediately . marmoratus process. Hybridisation

The available evidence suggests that the local dis- between the species is strongly asymmetric, with

tribution is - in a small pocket an enclave, geo- FI adults derived from matings of T. cristatus graphically isolated from the main T. karelinii mothers and T. marmoratus fathers significantly distribution in Bulgaria, Thrace, and Turkey (Fig. outnumbering the reverse combination. The facts

3c). However, a link between the parts, along a responsible for this phenomenon are largely un- narrow in northeastern in known but involve the strip Yugoslavia (as may genetic incompatibil-

1995 and in Kalezic et 1997: of the nuclear and mtDNA Arntzen, al, fig. 6), ity genomes (J. W.

Arntzen cannot be excluded. The further surveying ofeastern et ah, in prep.). By comparing past and

Yugoslavia and northwestern Bulgaria is required present distributions, the rate at which T. cristatus to settle this issue. takes over from T. marmoratus has been estimated

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km The be Arntzen JW, Bugter RJF, Cogalniccanu D, Wallis GP. 1997. as averaging one a year. process may The distribution and conservation status of the Danube triggered, or accelerated, by the removal of crested newt, Triturus dobrogicus.Amphibia-Reptilia 18: 133- hedgerows, modifying a landscape with terrestrial 142. features favourable to T. marmoratus, the most Arntzen JW, Garda-Pan's M. 1995. Morphological and the into terrestrial among big-bodied newt species, allozyme studies ofmidwife toads (genus Alytes), including one favourable to the more aquatic T. cristatus. the description of two new taxa from Spain. Contrib. Zool.

65: 5-34. The habitat preferences of the various crested newt Arntzen JW, Sket B. 1997. Morphometric analysis of black species in eastern Europe, with the exception of T. and white European cave salamanders, Proteus anguinus. J. are understood and it is un- dobrogicus, poorly Zool. 241: 699-707. clear which affect their dis- six the ecological parameters Arntzen JW,Tennis SFM. 1993. A year study on popu- tribution or change in distribution. Another area lation dynamics ofthe crested news (Triturus cristatus) fol-

the colonization of a created pond. of complexity is that around , where T. lowing newly Herpetol. 1 3: 99-110. carnifex, T. cristatus and T. dobrogicus meet (Fig. Arntzen Wallis GP. 1991. Restricted flow in JW, gene a mov- 3b). At sites 12 and 36 newts were found with T. ing hybrid zone of the newts Triturus cristatus and T. and the mtDNA carnifex phenotype haplotype typi- marmoratus in western France. Evolution 45: 805-826.

cal for T. dobrogicus, matching a similar observa- Arntzen JW, Wallis GP. 1994. The ‘WolterstorffIndex’ and its

value to the of the Crested Newt superspecies. Abh. tion at Tasovice in the Czech Republic (48°49’ N, taxonomy Ber. Naturk. Vorgesch., Magdeburg 17: 57-66. 16°09’ E; J. Pialek, V. Zavadil and J. W. Arntzen, Gaskins Andersen D. 1976. Statistical Atchley WR, CT, prop- unpubl.). erties of ratios. I. Empirical results. Syst. Zool. 25: 137-148.

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Appendix

Plate I. Sexual display by male Triturus dobrogicus. Symptomatic ofthe species are the bulging black and white throatand high head

the of Photo M. crest, giving appearance increased size. by Sparreboom.

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Plates II. III. Ventral surface offive taxa ofcrested newts in the Triturus cristatus superspecies.

d, f, T. T. Plate III: 1, Triturus carnifex carnifex; n, T. carnifex Plate II: b, c, e. Triturus dobrogicus; a, cristatus; j, o, p, q-w, of the - T. karelinii. and T. karelinii. carnifex carnifex; k, T. carnifex macedonicus; g i, macedonicus, m, Geographical origin individuals is: 1 and Benevento, Farma, t, Geographical origin of the individuals is: T. dobrogicus from p, Italy; o, Italy; n, v,

of Yugoslavia; r, Tavna Tadten, Austria; T. cristatus from Mayenne, France; j, Farma, w, Divcibare, Federal Republic q,

Federal Karan, Federal of Italy; k, Visegrad, Bosnia-Hercegovina; g, Djurinci, Monastire, Bosnia-Hercegovina; s, Republic and Istanbul, Republic of Yugoslavia; h, Istanbul, Turkey, and i, Karacabey, Yugoslavia; u, Visegrad, Bosnia-Hercegovina, m,

Turkey. Turkey.

can be that males h, n - and females (the others) by For geographical coordinates see Table 2. Note (a, e, g, j, r, u, v) distinguished and In males shape, colour and size of the cloaca, by the colour of the underside of the tail and by digit and limb length body shape. flattened. males the ventral side of the tail is the cloaca is large, black and two-lobed and in the female it is small, yellow and In

females the tail underside is most or all ofits yellow only directly behind the cloaca or it is entirely black; in yellow along lenght.

not shorter in males than in females. Note that some toes are size (such Digits and limbs are longer and inter-limb distance is original

but from L.A. van der Laan. as one ofthe inner toes in newt ‘a’) regenerating marking by toe-clipping. Photography by

Description of phenotypes

tail blase, sides, black throat with white Triturus dobrogicus. Lean build, short legs, narrow heavily white-stippled large angular and fuse in ventral surface with roundish black Notes: Ventral tend to line-up as (b). spots, deep orange many sharp, spots. spots

than in females. A function of the throat coloration is shown by Plate III. Throat spots are larger in males possible conspicuous

sides, throat a muddied mix of Triturus cristatus. Lean-mediumbuild, medium sized legs, narrow tail base, heavily white-stippled

black and yellow with fine white stippling, ventral surface yellow-orange with irregular black spots. Notes: The black spots grow and

the with & Teunis, and individual is to be younger than the others. spot pattern becomes denser age (Arntzen 1993) (a) likely throat colour Triturus carnifex carnifex. Medium build, large legs, tail base medium wide, little or no white stippling on sides,

ill-defined and to black Notes: Males variablewith white stipples. Ventral surface yellow with few large, roundish, muddy-gray spots.

than females. The newts T and ‘o’ sides of the tend to have darker throats with more and larger white stipples represent opposite

range.

sides throat Triturus carnifex macedonicus. Medium to heavy build, large legs, tail base medium wide, densely white-stippled, a

medium Ventral surface to with a dense muddiedmix of black and yellow with many, sized white stipples. yellow orange-yellow variable and individuals resemble T. pattern of small, irregular spots. Notes: The coloration characteristics are particularly may

dobrogicus (n), T. cristatus (s), T. carnifex carnifex (t) or T. karelinii (v).

Triturus karelinii. Heavy build, large legs, wide tail base, heavily white-stippled sides, ventral surface yellow-orange with many

in and continuous with throat where tend to be small to medium-sized black spots, extending on to the tail (especially females) spots

between T. different mtDNA and angular. Notes: no obvious coloration characters distinguish karelinii possessing haplotypes (KAR

KAR?, see Wallis and Arntzen, 1989).

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Plate II Downloaded from Brill.com10/08/2021 06:56:32PM via free access Downloaded from Brill.com10/08/2021 06:56:32PM via free access Contributions to Zoology, 68 (3) - 1999 203

Plate III Downloaded from Brill.com10/08/2021 06:56:32PM via free access