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While it is generally assumed that only the Northern Crested Newt (Triturus cristatus) occurs in Germany, there are early reports of Crested Newts with features typical of the Italian Crested Newt (T. carnifex) from the Berchtesgadener Land (Bavaria). We study a panel of eight nuclear DNA SNP markers for 26 individual Crested Newts to test if T. carnifex alleles occur in this region. All but two of the studied individuals contain alleles diagnostic for T. carnifex, with the largest percentage of T. carnifex alleles observed in a single individual being 37.5%. The sampled individuals show morphological characteristics typically associated with T. carnifex. We conclude that the natural hybrid zone between the Northern Crested Newt and the Italian Crested Newt reaches further west than previously realized and extends into the extreme southeast of Germany.

[No abstract available], We would like to thank S. Guldemond for the photo of a grass snake used in Figure 2. Frank Glaw and Jörn Köhler made valuable comments on an earlier manuscript version. The present study was conducted in the Senckenberg Dresden Molecular Laboratory (SGN-SNSD-Mol-Lab).

Triturus anatolicus sp. nov. Type material. Holotype. RMNH. RENA. 48232, an adult male (Fig. 3) from G��lk��y, near Kalecik, Turkey (coordinates: 40.077 ��N, 33.341 ��E, elevation: 1230 meter a.s.l.; Fig. 4), collected 6 April 2014 and donated to Naturalis Biodiversity Center by Kurtuluş Olgun from the Department of Biology at Adnan Menderes University, Aydın, Turkey. Paratypes. Six males (RMNH. RENA. 48233-48238), six females (RMNH. RENA. 48239-48245), and a juvenile (RMNH. RENA. 48246), with collection details identical to the holotype. For imagery, see Fig. 3 and online Appendix 1 and 2. Diagnosis. The new species is assigned to the genus Triturus because it possesses the characteristics of crested newts (the T. cristatus superspecies), in particular the combination of a denticulated crest (in males in breeding costume), a dark brown dorsal coloration and an orange ventral side covered with black spots. Based on mtDNA sequence data, crested newts from the type locality are firmly placed in the genus Triturus, within the distinct ���central T. karelinii sensu lato ��� mtDNA lineage (Wielstra et al., 2010). Although identification based on mtDNA is possible across the majority of the range of the new species, mtDNA derived from T. ivanbureschi has introgressed at the western part of the range (Wielstra et al., submitted; Fig. 2). The pattern of asymmetric introgression is mirrored by certain individual nuclear DNA markers and we consider this to represent a genomic footprint of hybrid zone movement (Wielstra et al., submitted; Fig. 2). As yet only multilocus nuclear DNA sequence data distinguish T. anatolicus sp. nov. unambiguously from T. ivanbureschi ��� and from T. karelinii, with which both T. anatolicus sp. nov. and T. ivanbureschi were until recently considered conspecific (Wielstra et al., submitted). No morphological features are currently known to distinguish the three species comprising T. karelinii sensu lato. How to separate T. anatolicus sp. nov., T. ivanbureschi and T. karelinii from the other crested newt species is discussed in Wielstra et al. (2013 b). Description of holotype. A relatively robust crested newt, with a broad head (wider than neck) and well developed limbs. Four fingers and five toes. Toes fringed but interdigital webbing absent. Relative length of fingers left 1 Description of paratypes. The morphology of the paratypes resembles that of the holotype in general, but the pattern of black spots on the ventral side is highly variable among individuals (online Appendix 1). Compared to males (such as the holotype), adult females have non-swollen cloacae, lack the denticulated crest and possess a less pronounced tail fin. All paratypes have an NRBV count of 13 except one, which has an NRBV count of 12 (Table 1; online Appendix 2). In Table 1 the same measurements as taken for the holotype are provided for the paratypes. Distribution. The distribution encompasses Asiatic Turkey south of the Black Sea, reaching up to c. 200 kilometers inland (usually less), but not into inner Anatolia. To the west the new species reaches the Bosphorus at the northern side of the Marmara Sea. On the southern side of the Marmara Sea it meets T. ivanbureschi sensu stricto, east of Lake Ulubat and west of the city of Bursa. The two species form a hybrid zone here (Wielstra et al., submitted). To the east the new species reaches the town of Yomra, just east of the city of Trabzon. The nearest known Triturus localities further east are from the extreme NE of Turkey, over 150 km away, and probably concern T. karelinii sensu stricto (Wielstra et al., 2013 a). An outline of the distribution of the new species is provided in Fig. 1. A database of distribution records is provided in Wielstra et al. (2014 b). Etymology. The specific epithet reflects the distribution of the new Triturus species. Triturus anatolicus sp. nov. is endemic to Anatolia. It is the only Triturus species to which this applies. It should be noted that the range of T. ivanbureschi sensu stricto covers western Anatolia (Wielstra et al., 2013 a; Wielstra et al., submitted) and the range of T. karelinii sensu stricto probably protrudes into northeastern Anatolia (Wielstra et al., 2010). Proposed vernacular name. We propose to use the vernacular name Anatolian Crested Newt for T. anatolicus sp. nov. This name highlights its status as an Anatolian endemic. We suggest to use the vernacular name Balkan Crested Newt for T. ivanbureschi sensu stricto (rather than Balkan-Anatolian Crested Newt previously applied to T. ivanbureschi sensu lato). Although T. ivanbureschi sensu stricto also partially occurs in Anatolia, the main part of its range is in the Balkan Peninsula., Published as part of Wielstra, B. & Arntzen, J. W., 2016, Description of a new species of crested newt, previously subsumed in Triturus ivanbureschi (Amphibia: Caudata: Salamandridae), pp. 73-80 in Zootaxa 4109 (1) on pages 74-79, DOI: 10.11646/zootaxa.4109.1.6, http://zenodo.org/record/260655, {"references":["Wielstra, B., Litvinchuk, S. N., Naumov, B., Tzankov, N. & Arntzen, J. W. (2013 b) A revised taxonomy of crested newts in the Triturus karelinii group (Amphibia: Caudata: Salamandridae), with the description of a new species. Zootaxa, 3682, 441 - 453. http: // dx. doi. org / 10.11646 / zootaxa. 3682.3.5","Wielstra, B., Baird, A. B. & Arntzen, J. W. (2013 a) A multimarker phylogeography of crested newts (Triturus cristatus superspecies) reveals cryptic species. Molecular Phylogenetics and Evolution, 67, 167 - 175. http: // dx. doi. org / 10.1016 / j. ympev. 2013.01.009","Wielstra, B., Sillero, N., Voros, J. & Arntzen, J. W. (2014 b) The distribution of the crested and marbled newt species (Amphibia: Salamandridae: Triturus) - an addition to the New Atlas of Amphibians and Reptiles of Europe. Amphibia- Reptilia, 35, 376 - 381. http: // dx. doi. org / 10.1163 / 15685381 - 00002960"]}

Triturus ivanbureschi sp. nov. Arntzen & Wielstra Type material. Holotype. RMNH. RENA. 47200, an adult male from Ostar Kamak, Bulgaria (coordinates: 41.878 ��N, 25.853 ��E, elevation: 240 meter), collected 8 May 2012 by J.W. Arntzen, N. Tzankov and B. Wielstra (Fig. 4). Paratypes. Three males (RMNH. RENA. 47201-47203) and 15 females (RMNH. RENA. 47204-47218), with collection details identical to the holotype. For imagery, see online Appendix 5. Diagnosis. The species is assigned to the genus Triturus because it possesses the characteristics of crested newts (the T. cristatus superspecies), in particular the combination of a denticulated crest, a dark brown dorsal coloration and orange ventral side covered with black spots. Based on mitochondrial DNA sequence data, crested newts from the type locality are firmly placed in the genus Triturus, within the western T. karelinii s. l. mitochondrial DNA lineage (Wielstra et al., 2013 b). As yet only molecular genetic data are available to distinguish T. ivanbureschi sp. nov. from its sister species T. karelinii. More details are provided in the section ���A review of phylogenetic studies with Triturus karelinii s. l. ���. How to distinguish T. ivanbureschi sp. nov. + T. karelinii from the other crested newt species is discussed in the section ���The name T. arntzeni is a junior synonym of T. macedonicus ���. Description of holotype. A relatively robust crested newt, with a broad head (wider than neck) and well developed limbs. Four fingers and five toes, slightly fringed but interdigital webbing absent. Relative length of fingers 1 Description of paratypes. The morphology of the paratypes resembles that of the holotype in general, but the pattern of black spots on the ventral side is highly variable among individuals (online Appendix 5). All paratypes have an NRBV count of 13, except RMNH. RENA. 47205 (and probably RMNH. RENA. 47202), which has an NRBV count of 14 (Table 2; online Appendix 6). Compared to males (such as the holotype), adult females have non-swollen cloacae, lack the denticulated crest and possess a less pronounced tail fin. In Table 2 the measurements taken for the holotype are provided for the paratypes. Distribution. The European distribution encompasses the south-eastern Balkan Peninsula, covering most of Bulgaria, the eastern parts of Greece, Macedonia and Serbia, as well as European Turkey. An isolated distribution pocket is found in Serbia that is disconnected from the main range by other intervening Triturus species (T. cristatus in the north and T. macedonicus in the south; Arntzen & Wallis, 1999; Wielstra & Arntzen, 2012). The Asian distribution is restricted to Asiatic Turkey, along the coast of the Aegean Sea, the Marmara Sea and the Black Sea, reaching up to c. 300 kilometers inland (usually less) but not inner Anatolia. An outline of the distribution is provided in Fig. 1 (T. ivanbureschi sp. nov. encompasses the western and central gene pools of T. karelinii s.l., which meet in northwestern Asiatic Turkey). An incomplete listing of locality data is provided in Wielstra et al. (2012). Etymology. We dedicate the newly recognized species of crested newt to the memory of Dr. Ivan Buresch (1885-1980), scientific director of the Institute of Zoology, Sofia (see pp. 234-235 in Adler, 2012). Dr. Buresch, together with his assistant Jordan Zonkov, laid the foundation for herpetology in Bulgaria and indeed the Balkan region with the publication of the four part monograph, ��� Untersuchungen ��ber die Verbreitung der Reptilien und Amphibien in Bulgarien und auf der Balkanhalbinsel��� (Buresch & Zonkow, 1933, 1934, 1941, 1942). The work is published in the Bulgarian language with extensive summaries in German. The compound name ivanbureschi was chosen to avoid confusion with ��� Triturus cristatus karelinii forma bureschi ��� (Wolterstorff, 1925) (nomen illegitimum, Litvinchuk et al., 1999; Mertens & Wermuth, 1960). Proposed vernacular name. Balkan-Anatolian Crested Newt or Buresch���s Crested Newt., Published as part of Wielstra, B., Litvinchuk, S. N., Naumov, B., Tzankov, N. & Arntzen, J. W., 2013, A revised taxonomy of crested newts in the Triturus karelinii group (Amphibia: Caudata: Salamandridae), with the description of a new species, pp. 441-453 in Zootaxa 3682 (3) on pages 448-450, DOI: 10.11646/zootaxa.3682.3.5, http://zenodo.org/record/215933, {"references":["Wielstra, B., Crnobrnja-Isailovic, J., Litvinchuk, S. N., Reijnen, B. T., Skidmore, A. K., Sotiropoulis, K., et al. (2013 b) Tracing glacial refugia of Triturus newts based on mitochondrial DNA phylogeography and species distribution modeling. Frontiers in Zoology, 10, 13. http: // dx. doi. org / 10.1186 / 1742 - 9994 - 10 - 13","Arntzen, J. W. & Wallis, G. P. (1999) Geographic variation and taxonomy of crested newts (Triturus cristatus superspecies): Morphological and mitochondrial data. Contributions to Zoology, 68, 181 - 203.","Adler, K. (2012) Contributions to the history of herpetology. Vo l. 3. Society for the study of amphibians and reptiles, Vancouver, Canada, 564 pp.","Buresch, I. & Zonkow, J. (1933) Untersuchungen uber die Verbreitung der Reptilien und Amphibien in Bulgarien und auf der Balkan halbinsel. I. Teil: Schildkrotten (Testudinata) und Eidechsen (Sauria). Bulletin des Institutions royales d'Histoire Naturelle, Sofia, 6, 150 - 207.","Buresch, I. & Zonkow, J. (1934) Untersuchungen uber die Verbreitung der Reptilien und Amphibien in Bulgarien und auf der Balkan halbinsel. II Feil: Schlangen (Serpentes). Bulletin des Institutions royales d'Histoire Naturelle, Sofia, 7, 106 - 188.","Buresch, I. & Zonkow, J. (1941) Untersuchungen uber die Verbreitung der Reptilien und Amphibien in Bulgarien und auf der Balkanhalbinsel III Teil: Schwanzlurche (Amphibia, Caudata). Bulletin des Institutions royales d'Histoire Naturelle, Sofia, 14, 171 - 237.","Buresch, I. & Zonkow, J. (1942) Untersuchungen uber die Verbreitung der Reptilien und Amphibien in Bulgarien und auf der Balkanhalbinsel IV Teil: Froschlurche (Amphibia, Salienta). Bulletin des Institutions royales d'Histoire Naturelle, Sofia, 15, 68 - 154.","Wolterstorff, W. (1925) Katalog der Amphibien-Sammlung im Museum fur Natur- und Heimatkunde. Abhandlungen und Berichte aus dem Museum fur Natur- und Heimatkunde zu Magdeburg, 4, 155 - 234.","Litvinchuk, S. N., Borkin, L. J., Dzukic, G., Kalezic, M. L., Khalturin, M. D. & Rosanov, J. M. (1999) Taxonomic status of Triturus karelinii on the Balkans, with some comments about other crested newt taxa. Russian Journal of Herpetology, 6, 153 - 163.","Mertens, R. & Wermuth, H. (1960) Die Amphibien und Reptilien Europas. Verlag Waldemar Kramer, Frankfurt am Main, 252 pp."]}

The name T. arntzeni is a junior synonym of T. macedonicus Litvinchuk et al. (1999) described the subspecies Triturus karelinii arntzeni based on differences in genome size, protein variation and morphological characteristics. Subsequently, Espregueira Themudo et al. (2009) elevated this subspecies to species level, i.e. Triturus arntzeni Litvinchuk, Borkin, Džukiċ and Kaleziċ, 1999 (in Litvinchuk et al., 1999). However, concern has been expressed that the type locality of T. arntzeni ���Vrtovaċ, near Pirot in eastern Serbia (Fig. 1)���in fact represents the species T. macedonicus and not T. karelinii s. l. (Arntzen & Wielstra, 2010; Stoyanov et al., 2011). We review the species identity of crested newts from Vrtovaċ below. Because the bulk of evidence points towards T. macedonicus being the crested newt species occurring at Vrtovaċ, we identify the name T. arntzeni as a junior synonym of T. macedonicus and conclude that it does not represent a taxon newly distinguished in T. karelinii s. l. Geography. The population in Litvinchuk et al. (1999) representing T. arntzeni comprises seven different localities (14 a-g; see Fig. 1), all of which are situated at, or close to, various crested newt contact zones. Note that locality 14 c concerns the actual type locality of T. arntzeni. For the latest overview of Triturus distribution, including a database of locality data, see Wielstra et al. (2013 b). The species composition of the seven T. arntzeni populations we evaluate as follows: Locality 14 a, Trbusaċ, Serbia is T. dobrogicus (Wielstra et al., 2013 b). Species identification is based upon the phenotype of material deposited at the Institute of Biological Research, ���Sini��a Stankoviċ���, University of Belgrade (JWA, BW). Locality 14 b, Tre��nja, Serbia represents the western T. karelinii s. l. nuclear gene pool with T. dobrogicus mitochondrial DNA and some limited presence of, or genetic admixture with, T. dobrogicus (Arntzen et al., submitted; Wallis & Arntzen, 1989). Locality 14 c, Vrtovaċ, Serbia, is positioned inside the T. macedonicus range (Wielstra & Arntzen, 2012). Neighboring crested newt species are, in order of increasing distance, T. cristatus, T. dobrogicus and western T. karelinii s. l. nuclear gene pool (Fig. 1). Locality 14 d, Vlasi, Serbia, has western T. karelinii s. l. nuclear gene pool and T. macedonicus in syntopy (Wielstra & Arntzen, 2012). Species identification is based on examination of animals observed in the field (JWA, BW). Locality 14 e, Vlasina, Serbia has T. macedonicus (Wielstra & Arntzen, 2012). Species identification is based upon the phenotype of material deposited at the Institute of Biological Research, ���Sini��a Stankoviċ���, University of Belgrade and of animals observed in the field (JWA, BW). The number of rib-bearing vertebrae (details below) reported in this population (n= 7 with NRBV = 13, n= 25 with NRBV = 14) is typical for T. macedonicus (Crnobrnja-Isailoviċ et al., 1997). Locality 14 f, Stracin, Macedonia has the western T. karelinii s. l. nuclear gene pool and T. macedonicus in syntopy (Wielstra & Arntzen, 2012). Species identification is based upon the phenotype of material deposited at the Institute of Biological Research, ���Sini��a Stankoviċ���, University of Belgrade (JWA, BW). Note that a population of T. macedonicus is located c. 4 kilometers to the west (Rugince, Macedonia in Wielstra & Arntzen, 2012). Locality 14g, Berovo, Macedonia is western T. karelinii s. l. nuclear gene pool. Species identification was confirmed based upon the phenotype of material deposited at the Institute of Biological Research, ���Sini��a Stankoviċ���, University of Belgrade and of animals observed in the field (JWA, BW) and conforms to species identification by Kaleziċ and Hedgecock (1980). Another confirmed T. karelinii s. l. nuclear gene pool locality is located 9 kilometers to the northwest (Mitra��inci, Macedonia in Wielstra et al., 2013 a). Throat and belly pattern. In the field crested newts are best identified by morphotype (see introduction) and by their coloration pattern (Arntzen, 2003; Arntzen & Wallis, 1999) as follows: Triturus karelinii s. l.: Stocky build. Heavily white-stippled sides. Ventral surface yellow-orange with many small to medium-sized, frequently angular black spots and continuous on the throat. Triturus carnifex: Medium stocky build. Little or no white stippling on sides. Throat color variable with white stipples. Ventral surface yellow with few large, roundish, ill-defined and muddy-gray to black spots. Triturus macedonicus: Medium stocky build. Sides densely white-stippled. Throat dark black or a muddied mix of black and yellow with many, medium sized white stipples. Ventral surface yellow to orange-yellow with a dense pattern of small, irregular spots. Triturus cristatus: Slender build. Heavily white-stippled sides. Throat a muddied mix of black and yellow with fine white stippling. Ventral surface yellow-orange with irregular black spots. Triturus dobrogicus: Very slender build. Heavily white-stippled sides. Black throat with large angular white spots (especially in males), ventral surface deep orange with many sharp, roundish black spots. For a picture overview of throat and belly patterns of the different crested newt species, see Arntzen and Wallis (1999). Pictures of the throat and belly pattern of the holotype and paratypes of T. arntzeni (online Appendix 1), preserved material stored at the Institute of Biological Research, ���Sini��a Stankoviċ���, University of Belgrade, and live newts from the type locality as observed by several of us (JWA, BW) most resemble T. macedonicus and not T. karelinii s. l. In particular we point to the absence of the distinctive T. karelinii s. l. throat pattern, with small angular dark spots surrounded by yellow, similar to the pattern on the belly. These spots are also absent in the holotype of T. arntzeni. To ascribe the Vrtovaċ newts to T. macedonicus by coloration pattern alone is not straightforward: as noted by Freytag (1988), T. macedonicus overlaps considerably with T. cristatus in patterning and coloration. Number of rib-bearing pre-sacral vertebrae. The Triturus morphotypes were initially distinguished with the help of the ���Wolterstorff index���, a measure of overall shape, defined as ���forelimb length divided by inter-limb distance��� (Wolterstorff, 1923). Arntzen and Wallis (1994) found the number of rib-bearing pre-sacral vertebrae (NRBV) in Triturus to represent a taxonomically discriminating character superior to the Wolterstorff-Index (the two are strongly negatively correlated across species). Accordingly, the Triturus morphotypes are characterized by a modal number of rib-bearing pre-sacral vertebrae (NRBV) (Arntzen & Wallis, 1999). Modal NRBV count increases from 12 in T. marmoratus ��� T. pygmaeus, via 13 in T. karelinii s. l., 14 in T. carnifex ��� T. macedonicus and 15 in T. cristatus, to 16 / 17 in T. dobrogicus (Arntzen, 2003; Arntzen & Wallis, 1999). A disadvantage of NRBV (similarly applying to the Wolterstorff index) is that, although informative at the population level, it is not unambiguous at the individual level, due to intraspecific variation. Moreover, intermediate scores due to interspecific hybridization may point to a third species not involved (Arntzen & Wallis, 1994). We determined NRBV for 63 individuals from the T. arntzeni type locality (the holotype and 12 paratypes, deposited at the Zoological Institute, Russian Academy of Sciences, St. Petersburg; 46 specimens deposited at the Institute of Biological Research, ���Sini��a Stankoviċ���, University of Belgrade and four specimens deposited at Naturalis Biodiversity Center, Leiden, see online Appendix 2). Three specimens with different NRBV (13 versus 14) on either sides of the body were ignored. Eleven individuals (18.3 %) have an NRBV count of 13, NRBV is 14 in n= 44 (73.3 %) and NRBV is 15 in n= 5 (8.3 %). The distribution of modal versus non-modal NRBV scores in the Vrtovaċ population is not significantly different from that of T. carnifex / T. macedonicus (G-test of independence, d. f. = 1, P> 0.05; reference data in Arntzen, 2003). The NRBV count of 15 shown by the holotype of T. arntzeni is indicative since this value fits T. macedonicus better than it does T. karelinii s. l. Genome size. The size of the genome may be a source of relevant taxonomic information (Green & Sessions, 2007; Kron et al., 2007). Litvinchuk et al. (1999) observed a difference in genome size between T. arntzeni and T. karelinii s. l., with non-overlapping confidence intervals. We here interpret the genome size of T. arntzeni from Vrtovaċ (n= 13) not as ���different from other T. karelinii��� (as in Litvinchuk et al., 1999), but as ���not different from T. macedonicus��� (Fig. 2; constituent data in online Appendix 3). It is important to note that these genome sizes also differ from those of T. cristatus and T. dobrogicus. In contrast to Litvinchuk et al. (1999) we do not invoke convergent evolution or introgressive hybridization to explain the data but conclude that newts from Vrtovaċ actually are T. macedonicus. The genome size of the holotype of T. arntzeni in particular falls within the range of T. macedonicus but outside of the range of T. karelinii s. l. The fourteenth specimen labeled as T. arntzeni in Litvinchuk et al. (1999) is from Tresnja (population 14 b, see above) and has a genome size that is intermediate of T. karelinii s. l. and T. dobrogicus; it is excluded from Fig. 2. Allozyme data. Litvinchuk et al. (1999) presented data for nine allozyme loci and analyzed these at the population level. We here re-analyze the data at the level of the individual and in a phylogenetic framework for 42 individuals with a complete dataset and 48 individuals with data missing at one locus (online Appendix 4). Unfortunately the dataset for the holotype of T. arntzeni is highly incomplete, but several paratypes can be included in the analysis. We first determine the number of distinctive gene pools that is best supported by the data probabilistically using BAPS v. 5.3 (Corander et al., 2008). The recognized groups were clustered in a dendrogram on the basis of Rogers��� genetic distance with BIOSYS- 1 (Swofford & Selander, 1981). All ten included T. arntzeni individuals and three T. cristatus are in one group. Other BAPS groups are composed as follows: all T. karelinii s. l. individuals with representatives of all three T. karelinii s. l. nuclear gene pools, the n= 3 included T. macedonicus individuals, two T. cristatus groups and four T. dobrogicus groups and, finally, a mixed group with one T. cristatus and one T. dobrogicus (online Appendix 4). In the UPGMA dendrogram (Fig. 3) the BAPS group that contains all T. arntzeni clusters with T. cristatus and it does not cluster with either T. karelinii s. l. or T. macedonicus. A distance-Wagner tree yields the same result (data not shown). We further note that the two loci considered to distinguish T. arntzeni from T. karelinii s. l. by Litvinchuk et al. (1999) are actually represented by the same alleles in T. cristatus and T. macedonicus. However, the reference sample size is small, in particular for unquestionable T. macedonicus (n= 3) and western T. karelinii s. l. nuclear gene pool Nuclear DNA sequence data. Wielstra et al. (2013 a) provided a framework to cluster Triturus newts to species based on three nuclear DNA markers developed by Espregueira Themudo et al. (2009). Unfortunately, our attempts to obtain nuclear DNA sequence data from the type material (the holotype and 12 paratypes) of T. arntzeni were not successful, presumably because of degraded DNA. However, Wielstra et al. (2013 a) presented sequence data for another five individuals from Vrtovaċ (voucher ID 2533 - 37 stored at Naturalis Biodiversity Center, Leiden). These individuals were all assigned to T. macedonicus (see Appendix S 1 in Wielstra et al., 2013 a). Mitochondrial DNA. As noted above we did not manage to obtain usable DNA extract and were not able to sequence mitochondrial DNA for the type material. However, we note that mitochondrial DNA could not be used to identify these newts anyway. This is because T. macedonicus in and around Serbia contain asymmetrically introgressed western T. karelinii s. l. mitochondrial DNA (the area for which this phenomenon has been documented is detailed in Wielstra & Arntzen, 2012). Similarly, the nearest Serbian T. cristatus populations contain asymmetrically introgressed western T. karelinii s. l. mitochondrial DNA (Wielstra & Arntzen, 2012; Wielstra et al., 2013 b). So, from the perspective of mitochondrial DNA, T. macedonicus and T. cristatus are indistinguishable in the Vrtovaċ region. In conclusion, our analyses confirm that newts from Vrtovaċ, the type locality of T. arntzeni, including the holotype of T. arntzeni, are dissimilar to T. karelinii s. l. and resemble T. macedonicus (NRBV, genome size, nuclear DNA sequences) or T. cristatus (allozyme data). The throat and belly pattern suggest they are T. macedonicus, but we acknowledge that the distinction with T. cristatus is difficult. On balance, we consider the population from Vrtovaċ to represent T. macedonicus (with perhaps some influences of T. cristatus). Consequently, we consider the name T. arntzeni to be a junior synonym of T. macedonicus and we conclude that the name T. arntzeni should not be used for the central/western T. karelinii s. l. species., Published as part of Wielstra, B., Litvinchuk, S. N., Naumov, B., Tzankov, N. & Arntzen, J. W., 2013, A revised taxonomy of crested newts in the Triturus karelinii group (Amphibia: Caudata: Salamandridae), with the description of a new species, pp. 441-453 in Zootaxa 3682 (3) on pages 443-448, DOI: 10.11646/zootaxa.3682.3.5, http://zenodo.org/record/215933, {"references":["Litvinchuk, S. N., Borkin, L. J., Dzukic, G., Kalezic, M. L., Khalturin, M. D. & Rosanov, J. M. (1999) Taxonomic status of Triturus karelinii on the Balkans, with some comments about other crested newt taxa. Russian Journal of Herpetology, 6, 153 - 163.","Espregueira Themudo, G., Wielstra, B. & Arntzen, J. W. (2009) Multiple nuclear and mitochondrial genes resolve the branching order of a rapid radiation of crested newts (Triturus, Salamandridae). Molecular Phylogenetics and Evolution, 52, 321 - 328. http: // dx. doi. org / 10.1016 / j. ympev. 2009.03.024","Stoyanov, A., Tzankov, N. & Naumov, B. (2011) Die Amphiben und Reptilien Bulgariens. Chimaira, Frankfurt am Main, 582 pp.","Wielstra, B., Crnobrnja-Isailovic, J., Litvinchuk, S. N., Reijnen, B. T., Skidmore, A. K., Sotiropoulis, K., et al. (2013 b) Tracing glacial refugia of Triturus newts based on mitochondrial DNA phylogeography and species distribution modeling. Frontiers in Zoology, 10, 13. http: // dx. doi. org / 10.1186 / 1742 - 9994 - 10 - 13","Wallis, G. P. & Arntzen, J. W. (1989) Mitochondrial-DNA variation in the crested newt superspecies: Limited cytoplasmic gene flow among species. Evolution, 43, 88 - 104. http: // dx. doi. org / 10.2307 / 2409166","Crnobrnja-Isailovic, J., Dzukic, G., Krstic, N. & Kalezic, M. L. (1997) Evolutionary and paleogeographical effects on the distribution of the Triturus cristatus superspecies in the central Balkans. Amphibia-Reptilia, 18, 321 - 332. http: // dx. doi. org / 10.1163 / 156853897 x 00378","Kalezic, M. L. & Hedgecock, D. (1980) Genetic variation and differentiation of three common European newts (Triturus) in Yugoslavia. British Journal of Herpetology, 6, 49 - 57.","Wielstra, B., Baird, A. B. & Arntzen, J. W. (2013 a) A multimarker phylogeography of crested newts (Triturus cristatus superspecies) reveals cryptic species. Molecular Phylogenetics and Evolution, 67, 167 - 175. http: // dx. doi. org / 10.1016 / j. ympev. 2013.01.009","Arntzen, J. W. (2003) Triturus cristatus Superspecies - Kammolch-Artenkreis (Triturus cristatus (Laurenti, 1768) - Nordlicher Kammolch, Triturus carnifex (Laurenti, 1768) - Italienischer Kammolch, Triturus dobrogicus (Kiritzescu, 1903) - Donau- Kammolch, Triturus karelinii (Strauch, 1870) - Sudlicher Kammolch). In: Grossenbacher, K. & Thiesmeier, B. (Eds.), Handbuch der Reptilien und Amphibien Europas. Schwanzlurche IIA. Aula-Verlag, Wiebelsheim, pp. 421 - 514.","Arntzen, J. W. & Wallis, G. P. (1999) Geographic variation and taxonomy of crested newts (Triturus cristatus superspecies): Morphological and mitochondrial data. Contributions to Zoology, 68, 181 - 203.","Freytag, G. E. (1988) Erinnerungen an Willy Wolterstorff. Wie kompliziert ist die Rassengliederung des Kammolches (Triturus cristatus [LAURENTI, 1768])? (Amphibia, Caudata, Salamandridae). Ein blick in die Geschichte der Salamanderkunde. Zoologische Abhandlungen fur Tierkunde Dresden, 44, 1 - 10.","Wolterstorff, W. (1923) Ubersicht den Unterarten und Formen des Triton cristatus Laur. Blatter fur Aquarien-und Terrarienkunde, 34, 120 - 126.","Arntzen, J. W. & Wallis, G. P. (1994) The ' Wolterstorff Index'and its value to the taxonomy of the Crested Newt superspecies. Abhandlungen und Berichte fur Naturkunde, 17, 57 - 66.","Green, D. M. & Sessions, S. K. (2007) Chapter 6 - Karyology and cytogenetics. In: Heatwole, H. (Ed), Amphibian Biology Volume 7. Surrey Beatty & Sons, Chipping Norton, NSW, Australia, pp. 2757 - 2842.","Kron, P., Suda, J. & Husband, B. C. (2007) Applications of flow cytometry to evolutionary and population biology. Annual Review of Ecology Evolution and Systematics, 38, 847 - 876. http: // dx. doi. org / 10.1146 / annurev. ecolsys. 38.091206.095504","Corander, J., Marttinen, P., Siren, J. & Tang, J. (2008) Enhanced Bayesian modelling in BAPS software for learning genetic structures of populations. BMC Bioinformatics, 9, 539. http: // dx. doi. org / 10.1186 / 1471 - 2105 - 9 - 539","Swofford, D. L. & Selander, R. B. (1981) BIOSYS- 1: A FORTRAN program for the comprehensive analysis of electrophoretic data in population genetics and systematics. Journal of Heredity, 72, 281 - 283."]}

Systematics of T. karelinii s. l. The type locality of T. karelinii is ���the northeast of the Iranian region along the southern shore of the Caspian Sea��� (Strauch, 1870). Litvinchuk and Borkin (2009) restricted the type locality to the coast of the Gulf of Gorgan, Iran, based on the report of the expedition during which the type material was collected (Karelin, 1883). The type locality is within the range of the eastern T. karelinii s. l. species (Fig. 1). Therefore, the name T. karelinii sensu stricto should be applied to the eastern T. karelinii s. l. species. As a vernacular name we suggest Caucasian Crested Newt or Karelin���s Crested Newt. A considerable number of localities (n = 235) are known from the former Soviet Union (reviewed in Litvinchuk & Borkin, 2009) but distribution data for Iran are sparse (n = 13 in Wielstra et al., 2012). Its presence in the extreme north-east of Turkey is suspected but awaits confirmation (Wielstra et al., 2013 a). Considering that no previously proposed name is applicable to the central/western T. karelinii s. l. species, we provide a new name, for which the authors are J. W. Arntzen and B. Wielstra, Published as part of Wielstra, B., Litvinchuk, S. N., Naumov, B., Tzankov, N. & Arntzen, J. W., 2013, A revised taxonomy of crested newts in the Triturus karelinii group (Amphibia: Caudata: Salamandridae), with the description of a new species, pp. 441-453 in Zootaxa 3682 (3) on page 448, DOI: 10.11646/zootaxa.3682.3.5, http://zenodo.org/record/215933, {"references":["Strauch, A. (1870) Revision der Salamandriden-Gattungen nebst Beschreibung einiger neuen oder weniger bekannten Arten dieser Familie. Me moires de l' Acade mie i mpe riale des sciences de St. - Pe tersbourg Serie 7, 16, 17 - 426.","Litvinchuk, S. N. & Borkin, L. J. (2009) Evolution, systematics and distribution of crested newts (Triturus cristatus complex) in Russia and adjacent countries. Evropeisky Dom Press, St. Petersburg, 590 pp.","Karelin, G. S. (1883) Trip of G. S. Karelin on Caspian Sea. In: Zapiski Imperatorskogo Russkogo Geograficheskogo Obschestva po Obschey Geografii (Otdeleniyam Geografii Matematicheskoy i Fizicheskoy), Oнikс, St. Petersburg, pp. 1 - 497.","Wielstra, B., Baird, A. B. & Arntzen, J. W. (2013 a) A multimarker phylogeography of crested newts (Triturus cristatus superspecies) reveals cryptic species. Molecular Phylogenetics and Evolution, 67, 167 - 175. http: // dx. doi. org / 10.1016 / j. ympev. 2013.01.009"]}

We characterized the adult body form of the crested newt (Triturus cristatus superspecies) and explored its evolution. From seven morphometric traits, we determined that body size, interlimb distance and head width define the body form. None of the morphometric traits showed a phylogenetic signal. Three body-shape morphotypes (Triturus dobrogicus+T. cristatus, Triturus carnifex+Triturus macedonicus and Triturus karelinii+Triturus arntzeni) and three body-size morphotypes (T. dobrogicus, T. cristatus and all other crested newts) could be recognized. The ancestral phenotype (a large body with a short trunk and a wide head) characterized T. karelinii and T. arntzeni. Triturus carnifex and T. macedonicus had a somewhat different phenotype (large body and wide head, accompanied by mild body elongation). The most derived phenotype included body size reduction and more pronounced body elongation in T. cristatus and, especially, in T. dobrogicus. Body elongation occurred by trunk lengthening but not head and tail lengthening. Additionally, contrary to other tetrapods, evolutionary axis elongation in crested newts was followed by a decrease in body size. We advocate the hypothesis that ecology drives the evolution of body form in crested newts. © 2011 Blackwell Verlag GmbH.

Data files accompanying "Phylogenomics of the adaptive radiation ofTriturusnewts supports gradual ecological niche expansion towards an incrementally aquatic lifestyle"

Data files accompanying "Phylogenomics of the adaptive radiation of Triturus newts supports gradual ecological niche expansion towards an incrementally aquatic lifestyle"