Turkish Journal of Zoology Turk J Zool (2015) 39: 328-334 http://journals.tubitak.gov.tr/zoology/ © TÜBİTAK Research Article doi:10.3906/zoo-1403-60

A new subspecies of Lyciasalamandra flavimembris (Urodela: ) from Muğla, southwestern Turkey

Nazan ÜZÜM*, Aziz AVCI, Emin BOZKURT, Kurtuluş OLGUN Department of Biology, Faculty of Science and Arts, Adnan Menderes University, Aydın, Turkey

Received: 28.03.2014 Accepted: 04.08.2014 Published Online: 27.02.2015 Printed: 27.03.2015

Abstract: A new Lycian subspecies, Lyciasalamandra flavimembris ilgazi subsp. nov., is described from southwestern Anatolia, Turkey. It differs from nominate subspecies by its characteristic color pattern, especially the ground color of the dorsum, tail, upper eyelids, and limbs. Its distribution is limited to the type locality (Kötekli, Muğla Province, Turkey).

Key words: Lyciasalamandra flavimembris ilgazi subsp. nov., Urodela, Turkey, new subspecies

1. Introduction across these species (Sever et al., 1997). Therefore, Lycian of the genus Lyciasalamandra were tubercular structures are accepted to have originated first described by Steindachner (1891) from Dodurga independently in the 2 species of Mertensiella (Weisrock (Muğla, Turkey) as Molge luschani. Wolterstorff (1925) et al., 2001). Through recent molecular studies (Weisrock then classified the Lycian salamanders together with the et al., 2001; Veith and Steinfartz, 2004), the of Caucasian salamanders in the genus Mertensiella. Nine Lycian salamanders has been reorganized and the genus subspecies of M. luschani have been described, accepting Lyciasalamandra has been established. According to the Dodurga specimens as the nominate race (Pieper, 1963; detailed morphological (Öz et al., 2004) and molecular Başoğlu, 1967; Başoğlu and Atatür, 1974, 1975; Başoğlu studies (Veith and Steinfartz, 2004; Weisrock et al., 2006; and Baran, 1976; Baran and Atatür, 1980; Franzen and Veith et al., 2008), this genus has been accepted to be Klewen, 1987; Başoğlu et al., 1994; Mutz and Steinfartz, represented by 7 allopatric species [L. antalyana (Başoğlu 1995). and Baran, 1967), L. atifi (Başoğlu, 1967), L. billae (Franzen After a comparative morphological study with 2 and Klewen, 1987), L. fazilae (Başoğlu and Atatür, 1974), species of Mertensiella, Mertens (1942) concluded that L. flavimembris (Mutz and Steinfartz, 1995), L. helverseni M. luschani should be considered more primitive than M. (Pieper, 1963), and L. luschani (Steindachner, 1891)] in the caucasica. However, Özeti (1967) investigated the external Mediterranean basin, and their phylogenetic relationships anatomy and osteology of M. luschani, M. caucasica, and have been examined in detail. According to Weisrock et al. salamandra, and concluded that the hedonic (2001) and Veith et al. (2008), Lyciasalamandra subspecies gland, which is not present in Salamandra, was the shared originated from vicariant speciation as a result of almost character between M. luschani and M. caucasica. For this simultaneous divergence from each other during the Late reason, she indicated that M. luschani and M. caucasica Miocene, approximately 6 to 8 million years ago, when were assigned to the subgenus Mertensiella within the the uplifting of Anatolia occurred in response to the genus Salamandra. However, phylogenetic and molecular northward movement of the Arabian plate. Six of these studies (Titus and Larson, 1995; Weisrock et al., 2001; Veith 7 species were endemic to Anatolia and distributed from et al., 2008) did not support the monophyly of the genus an area between Marmaris (Muğla) and Alanya (Antalya) Mertensiella, requiring independent origins of the dorsal along the Mediterranean coast of Turkey and on some tail tubercle (hedonic gland) observed in males of the 2 adjacent islands (Veith et al., 2001; Öz et al., 2004; Veith species. In addition, histological studies in morphologies and Steinfartz, 2004; Franzen et al., 2008; Baran et al., of the tubercle glands in the 2 species of Mertensiella 2012). Finally, in addition to the previously reported taxa, suggested that these tubercles were not homologous 3 new species (L. irfani, L. arikani, and L. yehudahi) were * Correspondence: [email protected] 328 ÜZÜM et al. / Turk J Zool described from Muğla and Antalya through morphological Material compared and serological studies (Göçmen et al., 2011; Göçmen and L. flavimembris (N = 7). 1 ♂, 1 ♀, 2 juv., Çiçekli village Akman, 2012). With these studies, the number of species - Ula/Muğla, 27.02.2010, leg. Kurtuluş Olgun, Aziz Avcı; 1 belonging to the genus Lyciasalamandra has been raised ♂, 1 ♀, 1 juv., Çiçekli village - Ula/Muğla, 27.02.2010, leg. to 9 in Anatolia. In all Lyciasalamandra species, only L. Kurtuluş Olgun, Aziz Avcı, Emin Bozkurt. luschani is represented by 3 subspecies. The following morphometric measurements and During several field trips carried out between February ratios of our samples were determined according to the 2010 and January 2014 in the vicinity of Muğla, we found published literature (Baran and Atatür, 1986; Mutz and 8 specimens of Lycian salamanders in Kötekli/Muğla, Steinfartz, 1995; Öz et al., 2004; Göçmen et al., 2011; which were easily recognized by distinct morphological Göçmen and Akman, 2012): total body length (TBL), tip characters from the known species near this area. New of snout to tip of tail; rostrum–anus length (RA), tip of specimens differ from L. flavimembris, which has a snout to posterior end of the cloaca opening; length of more southern distribution range, in terms of coloration trunk (LT), length from gular fold to the anterior edge of and pattern characteristics and some morphometric cloaca opening; tail length (TL), length from the posterior measurements. end of the cloaca opening to the tip of tail; head and body In the present paper, we describe the Kötekli/Muğla length (HBL), length from snout to the anterior end of population as a new subspecies of L. flavimembris, the cloaca opening; nostril–eye distance (NED); distance between nostrils (DBN); eye diameter (ED); head length accepting that the populations living in the area between (HL), distance from the snout to the gular fold; head width Çiçekli and Marmaris represent the nominate subspecies (HW); parotid length (PL); parotid width (PW); fore limb (L. f. flavimembris). length (FLL); hind limb length (HLL); distance between fore and hind limbs length (DFHL). We also used ratios of 2. Materials and methods HW/HL, TL/TBL, PW/PL, and NED/HL in our study. All The specimens used in this study were obtained from morphometric measurements were taken using a digital Kötekli/Muğla and the village of Çiçekli in Ula/Muğla. caliper (Mitutoyo) with accuracy of 0.01 mm. They were collected under stones, especially during the Statistics of the morphometric values and ratios of day, on different dates in 2010 and 2014. We took photos of the new subspecies and other specimens were conducted the specimens at the places of capture. Color and pattern with STATISTICA 7.0. We used a t-test for comparing the characteristics were recorded while specimens were metric characters and took the index values of PERCRA still alive. We injected specimens with 96% ethanol after (percent of rostrum–anus length; [each metric character / they were etherized and put them in glass jars with 70% RA] × 100) according to Werner (1971). We evaluated all ethanol. They were deposited in the Zoology Laboratory statistical analyses using the statistical significance level of of the Department of Biology at the Faculty of Science and P ≤ 0.05. Arts, Adnan Menderes University. Material examined 3. Results Holotype: 1 ♂ Kötekli, Muğla, 23.01.2014, leg. Nazan Diagnosis. Adults of Lyciasalamandra flavimembris and Üzüm, Aziz Avcı, Emin Bozkurt, Ömer Barış Üzüm. our samples resemble each other in terms of having small Paratypes: (N = 7). 1 ♀, 2 juv., Kötekli, Muğla, yellowish spots on the dorsum. In addition, color and 07.03.2011, leg. Serdar Özcan, Yaşar Ayyıldız; 2 ♀♀, 2 pattern characteristics of juveniles are relatively similar to juv., Kötekli, Muğla, 17.01.2014, leg. Kurtuluş Olgun, Aziz each other as well. Thus, we concluded that our samples Avcı, Emin Bozkurt. belonged to L. flavimembris (Figure 1).

Figure 1. General view of juvenile: (a) L. f. flavimembris (Çiçekli - Ula/Muğla) and (b) L. f. ilgazi subsp. nov. (Kötekli/Muğla).

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Although there are some similarities between our lighter than the dorsum (Figure 3). Subdigitals are purple. samples and L. flavimembris, our new population is easily The upper side of tail is lighter brownish. The ventral side distinguished from all known populations of this species is flesh-colored without any yellow spots. in terms of coloration and pattern characteristics (Figure L. f. ilgazi subsp. nov. has a significantly higher PW/ 2). The ground color of the dorsum in both sexes and PL ratio and PW in the PERCRA index than the nominate juveniles is purplish dark brown, with small yellowish subspecies (P ≤ 0.05). spots. These spots are disordered and few in number. The Description of the holotype. There are no differences flanks are also purplish dark brown with yellowish flecks. in body form between L. f. ilgazi subsp. nov. and other Upper eyelids are mainly the same color, but darker than Lycian salamander taxa. TBL and TL are 147.11 and 68.88 the dorsum. The ground color of the parotids is also the mm, respectively. Head length is longer than head width same color as the dorsum. The color of fore and hind (HW/HL: 0.76). The posterior of parotids is bigger than limbs of our samples is purplish dark brown, but relatively the anterior, and their lengths are longer than their widths

Figure 2. General view of adult: (a) L. f. flavimembris (Çiçekli - Ula/Muğla) and (b) L. f. ilgazi subsp. nov. (Kötekli/Muğla).

Figure 3. Differential coloration ofL. f. flavimembris and L. f. ilgazi subsp. nov.: (a) head coloration of L. f. flavimembris; (b) Head coloration of L. f. ilgazi subsp. nov.; (c) hind limb coloration of L. f. flavimembris; (d) hind limb coloration of L. f. ilgazi subsp. nov.

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(PW/PL: 0.35). The ground color of the dorsum is purplish The altitude of specimens is 650 m a.s.l. (GPS dark brown. There are small yellowish spots on the dorsum, coordinates: 37°09′42.0″N, 28°22′50.7″E). Three parotids, head, and extremities. The ground color of the specimens were found on 7 March 2011 when the weather parotids is also purplish dark brown with small black was rainy, at around 2300 hours. Four specimens were spots. Upper eyelids are darker than the head. Extremities collected on 17 January 2014 when the weather was sunny, and tail have the same coloration but are lighter than the from 1600 to 1700 hours. We gathered 1 specimen on 23 dorsum. The upper side of the tail is lighter brownish with January 2014 when the weather was cloudy, between 1045 small dark spots. The ventral side of body is flesh-colored, and 1300 hours. During the field studies, we also observed including extremities, tail, and cloacal region. Subdigitals Mediodactylus kotschyi (Steindachner, 1870) in Kötekli. are purple. The gular fold is easily distinguished (Figure 4). The habitat is close to the city center and the Kötekli The measurements and ratios of the holotypes are as Campus of Muğla Sıtkı Koçman University. For this reason, follows: RA, 78.23 mm; LT, 59.43 mm; HL, 15.35 mm; HW, there is much heavy construction activity near the habitat 11.72 mm; NED, 3.42 mm; DBN, 4.90 mm; ED, 5.08 mm; of L. f. ilgazi subsp. nov. These activities constitute a great PL, 8.51 mm; PW, 2.97 mm; FLL, 22.52 mm; HLL, 26.27 threat to this unique population of the new subspecies. mm; DFHL, 42.86 mm; TL/TBL, 0.47; NED/HL, 0.22. Hence, strict conservation precautions are very important Paratypes and variations. All of the specimens for the future of the population. examined from Kötekli/Muğla on the material list without Distribution. Lyciasalamandra flavimembris ilgazi a holotype are accepted as paratypes (N = 7). Variations observed in some measurements for adults and juveniles subsp. nov. is at present only known from the type locality are given in the Table. (Figure 7). There is no sexual dimorphism within the population Etymology. The new subspecies is named for the (P > 0.05) in either raw data or PERCRA values. The surname of the Turkish herpetologist Prof Dr Çetin Ilgaz, coloration of paratypes would largely apply to the holotype. who has made valuable contributions to the knowledge of There are no differences between female and male adults Turkish herpetofauna. (Figure 5). Juveniles are lighter than adults, and they have more spots on the dorsum. The coloration of the tails of 4. Discussion juveniles is flesh-colored. Parotids of juveniles are the L. flavimembris was first described by Mutz and Steinfartz same color as the dorsum, but there are large yellowish (1995) from approximately 6 km north of Marmaris as maculations. Ventral side of all specimens is the same. a subspecies of Mertensiella luschani. It was found to be Color and pattern characteristics of juveniles are relatively different from other sister taxa of M. luschani in terms similar in the nominate and new subspecies. of the coloration of the dorsum, parotids, and eyelids Habitat and range. All of the specimens were collected (Mutz and Steinfartz, 1995; Veith et al., 2001; Öz et al., from under mossy stones. The area is very humid and 2004). Through recent molecular studies (Veith and rocky and is covered with Pinus brutia. Other vegetation Steinfartz, 2004), all known taxa of M. luschani were elements are Juniperus oxycedrus, Salvia fruticosa, Quercus transferred to another genus called Lyciasalamandra, and coccifera, Phillyrea latifolia, and Cistus sp., a typical the flavimembris subspecies were assigned a species rank Mediterranean maquis. The habitat is shown in Figure 6. with 6 other subspecies of M. luschani. Known localities of this species are 6 km north of Marmaris (type locality) and the village of Çiçekli (Mutz and Steinfartz, 1995; Öz et al., 2004; Franzen et al., 2008; Baran et al., 2012). In this study, we describe a new subspecies belonging to L. flavimembris, which has been known as a monotypic species up to now. By this newly described subspecies, the known range of the species has been extended by about 13 km of air distance to the north. Lyciasalamandra flavimembris ilgazi subsp. nov. is the most northern taxon of Lyciasalamandra identified to date. When we look at the current distributions of L. flavimembris and the newly described subspecies, we can assume that Figure 4. L. f. ilgazi subsp. nov.: general view of holotype (adult rotation could have divided a main ancestral population, male). Photo: K Olgun. and isolation would have led to the speciation of these

331 ÜZÜM et al. / Turk J Zool SD 0.49 11.18 0.54 2.28 1.67 5.95 1.33 0.80 2.65 6.62 1.55 4.98 2.25 1.13 3.49 4.56 0.03 0.42 0.00 0.19 0.04 0.47 0.01 0.13 1.05 0.17 1.38 0.97 1.79 0.94 1.47 1.24 1.24 Range 1.23–2.20 60.44–82.45 3.47–4.54 169.81–173.88 12.37–15.64 33.69–45.47 32.26–34.92 93,79–95.21 12.88–17.76 35.42–48.48 33.82–36.63 23.82–33.68 18.42–22.80 67.25–69.47 47.03–53.75 25.02–33.97 0.75–0.81 69.81–70.64 0.41–0.41 2.14–2.48 0.25–0.34 5.12–6.04 0.21–0.22 3.38–3.61 7.45–9.54 2.85–3.18 6.29–8.98 9.87–11.79 24.32–27.87 7.90–9.55 19.70–22.30 4.82–7.00 12.04–14.44 Mean 1.69 70.29 4.03 171.25 13.82 39.09 33.59 94.71 14.73 41.31 35.60 28.34 20.91 68.49 50.93 28.98 0.79 70.71 0.41 2.27 0.30 5.54 0.21 3.46 8.49 3.03 7.46 10.75 26.21 8.46 20.61 5.57 13.43 Juveniles N 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 3 SD 0.38 13.37 0.48 3.82 2.64 6.52 1.86 1.09 1.35 6.17 2.02 4.47 4.34 0.36 3.02 7.31 0.04 3.82 0.01 0.50 0.05 0.43 0.02 0.65 0.65 0.81 1.01 2.11 1.20 1.12 0.37 0.89 0.45 Range 2.22–3.15 115.23–147.88 2.93–3.99 174.67–183.07 17.96–24.00 61.52–77.06 27.23–31.63 93.26–95.44 22.32–25.58 65.97–80.78 29.80–34.23 48.11–58.70 35.76–45.61 72.67–73.44 50.09–56.46 49.26–67.10 0.76–0.84 74.67–83.07 0.43–0.45 2.88–3.94 0.26–0.37 3.85–4.88 0.19-0.21 4.61–6.07 6.08–7.51 3.52–5.26 4.64–7.04 13.41–18.36 20.33–22.73 11.32–14.01 16.48–17.34 7.17–9.21 10.36–11.40 Mean 2.70 132.49 3.64 178.02 21.72 70.30 29.16 94.50 24.09 74.34 32.49 54.34 39.31 73.09 52.85 58.14 0.79 78.02 0.44 3.21 0.33 4.32 0.20 5.12 6.89 4.48 6.04 15.96 21.41 12.63 16.98 8.15 10.96 Adults Lyciasalamandra flavimembris flavimembris flavimembris Lyciasalamandra N 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 samples. 1 = Values in raw data; 2 = values in PERCRA; N = number of samples; SD SD samples; of N = number in PERCRA; 2 = values data; in raw 1 = Values samples. SD 0.46 13.81 0.54 4.97 2.66 6.60 1.56 1.24 3.22 7.12 1.96 4.64 5.59 2.02 4.10 6.81 0.05 4.97 0.02 0.39 0.03 0.75 0.03 0.40 0.49 0.38 0.53 2.12 0.85 1.05 1.06 1.21 0.77 flavimembris Range 2.02–3.00 70.04–102.07 4.32–5.46 164.92–174.09 12.55–18.37 40.95–55.45 27.71–31.33 93.56–96.42 13.23–20.92 42.47–58.63 31.15–35.68 30.38–40.32 19.00–31.42 67.36–72.12 44.74–53.59 27.57–43.44 0.71–0.82 64.92–74.09 0.39–0.43 1.71–2.52 0.36–0.42 4.03–5.46 0.16–0.22 3.48–4.42 7.54–8.69 3.68–4.62 7.88–9.14 10.32–15.13 23.98–25.81 8.46–10.76 18.31–20.36 5.38–7.94 11.93–13.54 Mean 2.35 82.22 4.85 169.72 14.45 45.75 29.79 94.67 16.50 48.35 33.96 33.59 23.18 69.56 47.50 33.87 0.77 69.72 0.41 2.29 0.38 4.76 0.19 3.88 8.07 4.13 8.61 12.04 24.84 9.26 19.23 6.14 12.64 Juveniles N 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 SD 0.32 6.21 0.47 3.92 1.00 2.08 1.61 0.89 1.35 2.77 1.56 2.50 1.12 1.52 2.16 4.03 0.02 3.92 0.01 0.36 0.03 0.39 0.03 0.29 0.56 0.36 0.29 1.25 1.37 1.05 1.16 0.70 0.75 Range 2.97–3.71 139.22–152.88 3.80–4.76 178.74–188.05 22.52–24.68 72.43–77.50 28.79–32.40 93.74–95.67 24.43–27.60 76.18–82.68 31.37–35.18 55.09–60.96 40.41–42.86 72.32–75.97 50.15–54.79 61.33–70.20 0.76–0.80 78.74–88.05 0.44–0.47 2.88–3.68 0.33–0.39 3.78–4.73 0.17–0.22 4.90–5.43 5.93–7.13 5.08–5.86 6.49–7.10 15.35–18.32 19.62–22.76 11.72–14.26 14.98–17.51 8.51–10.22 10.88–12.55 Mean 3.40 145.06 4.33 184.21 23.65 74.81 30.06 95.02 26.27 78.75 33.38 58.36 41.83 74.11 53.16 66.31 0.78 84.21 0.46 3.39 0.36 4.30 0.20 5.15 6.56 5.41 6.87 16.90 21.45 13.16 16.71 9.44 11.98 N 4 subsp. nov. subsp. ilgazi flavimembris Lyciasalamandra Adults 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 1 1 2 2 1 1 2 2 1 1 2 1 1 2 2 1 1 2 1 1 1 2 1 1 2 1 2 1 2 1 2 1 2 L. f. and nov. subsp. ilgazi L. f. of ratios (in mm) and characters Mensural = standard deviation. Abbreviations of characters are given in Section given 2. are characters of Abbreviations deviation. = standard Table. PW TBL FLL HBL HLL RA LT DFHL TL HW/HL TL/TBL NED PW/PL NED/HL DBN ED HL HW PL

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Figure 5. L. f. ilgazi subsp. nov.: general view of paratype (adult Figure 6. Habitat of L. f. ilgazi subsp. nov.: Kötekli, Muğla female). Photo: K Olgun. Province, Turkey.

Figure 7. Distribution of L. f. ilgazi subsp. nov. (star) and L. f. flavimembris (triangle): 1) 6 km north of Marmaris; 2) Çiçekli village - Ula/Muğla; 3) Kötekli/Muğla. closely related subspecies. The similarities in mensural southern piedmont of the Madran Baba Mountains. Thus, characters, especially in color and pattern characteristics, we can conclude that Lycian salamanders are possibly undoubtedly support our assumption as well. However, to distributed not only in the western but also in the eastern obtain a better understanding of the relationship between part of the Batı Menteşe and Madran Baba Mountains. these 2 subspecies, detailed molecular studies are required. In addition, Mutz and Steinfartz (1995) suggested that Acknowledgments the possible distribution of Lycian salamanders could We are most grateful to Dr Ömer Barış Üzüm, Serdar include the western part of the Batı Menteşe Mountains. Özcan, and Yaşar Ayyıldız for their help during the In this study, we found the new population at the fieldwork, and to Dr Özkan Eren for description of the southeastern-most side of this mountain chain, near the vegetation.

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