Acta Herpetologica 13(1): 13-19, 2018 ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah DOI: 10.13128/Acta_Herpetol-21171 The first demographic data and body size of the southern banded newt, Ommatotriton vittatus (Caudata: Salamandridae) Abdullah Altunişik Recep Tayyip Erdoğan University, Faculty of Arts and Sciences, Biology Department, Rize, Turkey. Email: abdullah.altunisik@erdogan. edu.tr Submitted on: 2017, 26th August; revised on: 2017, 28th October; accepted on: 2017, 14th November Editor: Fabio M. Guarino Abstract. In this study, the age structure and some life history traits were revealed for the first time in a population of Ommatotriton vittatus, living in Tarsus (mid-south part of Turkey), at the western limit of the species’ range. Maxi- mum longevity was recorded to be eight years in both females and males and age at maturity was estimated as three years for males and four years for females. Although mean age did not differ significantly between sexes, males had significantly larger snout-vent length (SVL) than females. Age and body size were positively correlated with each other for both females and males. Since the populations of the Southern Banded Newt in Turkey are in decline, the present study that provides preliminary data on life history traits of this newt could be helpful for future biological studies. Keywords. Amphibia, age, demography, skeletochronology, life history, newt. INTRODUCTION The study of growth in amphibians is important because it directly influences age at sexual maturity, timing of reproductive events, fecundity, and longevity (Hemelaar, 1988; Smirina, 1994; Miaud and Merilä, 2000; Navas, 2006). Therefore, knowing age structure of a pop- ulation is essential for obtaining information about the life history of the species (Altunışık and Özdemir, 2015). Although there are several methods (e.g., body size correlation, mark and recapture and skeletochronology) to estimate the age and growth of amphibians and reptiles, skeletochronology which is based on counting the lines of arrested growth (LAGs) visible in histological sections, has been the most preferred method by researchers in recent years (Carranza et al., 2014; Bionda et al. 2015; Sinsch et al., 2015; Altunışık et al., 2016; Bülbül et al., 2016), as it yields reliable results. As previously described in many amphibian species, periosteal bone growth depends on ecological factors, e.g., changing favorable and unfavorable periods of bone growth. Unlike active growth seasons, in the winter season the periosteal bone growth is reduced due to the influence of the cold climate and a different growth rate occurs, which forms the LAGs. The differences in body size between males and females of the same species is a key feature that leads to significant biological insights (Denoël et al., 2009) and several hypotheses have been put forward to explain these variations. It is well documented that the basic factors that elicit sexual dimorphism are generally sex- ual selection, natural selection and fecundity selection (Andersson, 1994; Fairbairn, 1997; Pincheira-Donoso et al., 2008; Liao et al., 2015; Altunışık, 2017). In addition, sexual differences in age and growth rate associated with life history hypothesis may also cause the emergence of sexual size dimorphism (SSD), which is a well-known population property (Young, 2005; Cadeddu et al., 2012; Liao et al., 2013). A member of the cosmopolitan family Salamandri- dae, Ommatotriton genus has three species, Ommatotri- 14 Abdullah Altunışık ton nesterovi, Ommatotriton ophryticus (Northern Band- ed Newt) and Ommatotriton vittatus (Southern Banded Newt; van Riemsdijk et al., 2017). Here we focus on the Southern Banded Newt, which is a medium-sized newt of 90–110 mm in total length. During the breeding period it inhabits shallow, still or slowly flowing waters with a lot of vegetation, otherwise found in wooded areas or open places with loose rocks or stones not far from water. O. vittatus ranges from mid-south of Turkey, through western Syrian Arab Republic, Lebanon and northwest- ern Jordan, south to Israel (Olgun et al., 2009; IUCN, 2017; Fig. 1). Mid-south of Turkey represents the western limit of the species’ range. There are two subspecies of O. vittatus, namely O. v. vittatus and O. v. ciliensis. While the distribution area of the first subspecies is Turkey, Syria and Israel, the distribution area of the second one is with- in the borders of Turkey (Bogaerts et al., 2013). So far, demographic investigations have been con- fined to Northern banded newt (Kuzmin, 1999; Tarkh- nishvili and Gokhelasvili, 1999; Kutrup et al., 2005; Özcan and Üzüm, 2015) of Ommatotriton genus and no data are available for the Southern Banded Newt. The aim of this study is to determine age structure (longevity, age at sexual maturity, survivorship, adult life expectancy), mean body size and SSD of a population of O. vittatus in the Southeastern Turkey, at the western limit of the spe- cies’ range, using skeletochronology. MATERIAL AND METHODS Study area and species The study site (25 m a.s.l.) is located in Tarsus, Mersin (36°54’N, 34°53’E) in the mid-south part of Turkey and has a climate that is Mediterranean and moderately continental. According to climatic data obtained from the meteorological sta- tion situated near the study site (Tarsus Meteorological Station) for the years 1950-2016, the average temperature during summer was 27.16°C, and 10.9°C in the winter (www.mgm.gov.tr). A total of 40 adult O. vittatus individuals (23 males and 17 females) were caught by hand or using a dip net in the day- time during the breeding season in 2017. After anesthesia in MS-222, their snout-vent length (SVL) was measured using a digital vernier caliper (500-706-11, Mitutoyo, Tokyo, Japan) to the nearest 0.01 mm precision and the fourth toe of the left foot was clipped. In order to use toe samples in skeletochronological analysis, firstly they were fixed in 10% formalin and later pre- served in 70% ethanol. After sampling, all newts were released back to their habitats. The sexes of the individuals were deter- mined by externally visible secondary sexual characters (promi- nent cloaca and dorsal crest in males) (Bogaerts et al., 2013). Skeletochronological analysis We applied standard skeletochronology procedure that was described by Castanet and Smirina (1990) for laboratory proto- cols to assess the age structure. Firstly, we washed the phalan- ges kept in 70% ethanol in tap water then decalcified in nitric acid (5%) from 30 to 90 min depending on the bone structure. Using a freezing microtome (Shandon Cryostat, Germany), preferentially second phalange was cross-sectioned at 12-16 μm and stained with Harris hematoxylin for 10 min. The selected cross-sections from the narrow part of the phalanx diaphysis were placed in glycerin for scanning under a light microscope (Olympus BX51) at 200x and 400x magnifications and their photographs (1-10 for each cross-section) were taken by a cam- era (Pixera), which is connected to this microscope. The num- ber of LAGs was assessed independently by two observers. The distance between two adjacent LAGs is a good indicator of indi- vidual growth in a given year (Kleinenberg and Smirina 1969). Therefore, in this study age at sexual maturity was determined using the distance between LAGs. Where we observed an obvi- ous decrease in spacing between two subsequent LAGs, we took it to mark the age when sexual maturity was achieved (Ryser, 1988; Özdemir et al., 2012). Data analysis The normality of data was analyzed by Shapiro-Wilk test and the homogeneity of variances was analyzed by Levene test. After confirming that all data were normally distributed (P > 0.05), independent sample t-test was used to test if females and males differ in mean SVL or in mean age. SSD was quanti- fied with Lovich and Gibbons (1992) sexual dimorphism index Fig. 1. Distribution map of O.vittatus (IUCN, 2017). 15Age structure of Ommatotriton vittatus SDI = (size of larger sex / size of smaller sex) ± 1 (+1 if males are larger or -1 if females are larger), and arbitrarily defined as positive when females are larger than males and negative in the converse case. Survivorship rate for adult individuals was calculated from the age structure according to following formula (Robson and Chapman, 1961): S= T/(R+T–1), where S is the finite annual sur- vivorship rate estimate, T = N1 + 2N2 + 3N3 + 4N4 + …. + nNn, R = ΣNi, and Ni is the number of individuals in the age group i. Adult life expectancy (ESP), the expected life span of the salaman- ders that have reached sexual maturity, was computed according to the formula of Seber (1973): ESP = [0.5+1/(1-S)] where S rep- resents the survivorship rate. Pearson’s correlation coefficient was computed to measure the degree of relationship between SVL and age and in order to determine the correlation equation between the two variables; regression analysis was also performed. The growth patterns were estimated according to the von Bertalanffy growth model, which has been used earlier in several studies on amphibians (Ryser 1988; Miaud et al., 2000; Guarino et al., 2011). The general form of the von Bertalanffy growth equation used was Lt = L∞.(1 – e-k(t-t0)), where Lt is length at age t, L∞ is a parameter representing asymptotic maximum size, e is the base of the natural logarithm, k is a growth coefficient, and t0 is the age at metamor- phosis, which is the starting point of the growth curve in the pre- sent study. Because data on the size at metamorphosis of the stud- ied population were lacking, we used size at metamorphosis (Lt0 = 21.5 mm), provided by Tarkhnishvili and Gokhelasvili (1999). The parameters L∞ and k were estimated using Microsoft Excel pro- gram. Data analysis was conducted using SPSS 17 statistical soft- ware package (IBM SPSS Statistics for Windows). RESULTS LAGs recorded in phalange bones were clearly visible in all cross-sections, as it can be seen in Fig. 2. According to the results of age reading obtained by direct observa- tion in the microscope and photographs on the computer, the maximum lifespan was 8 years in both females and males. Females of O. vittatus were on average 5.82 ± 0.30 years old (range: 4-8) and males had an average age of 5.65 ± 0.29 years (range: 3-8; Table 1). No significant difference was detected between the sexes with regard to mean age (t = 0.407, df = 38, P > 0.05) and age distribution (Kolmogorov-Smirnov test, P > 0.05). As it can be seen from age distribution of the population (Fig. 3), the age group with the highest fre- quency is six (14 individuals). Skeletochronological pat- tern showed that males and females attained sexual matu- rity at age of 3 and 4, respectively. Since age structure did not differ between the sexes, S and ESP calculations were combined and S was found to be 0.79 and ESP was 5.26 years. SDI was -0.055, indicating a male biased size dimor- phism. Independent sample t-test (t= -2.327, df= 38, P <0.05) showed that males had significantly larger SVL than that of females. For both sexes, the maximum SVL recorded was higher than the estimated asymptotic SVL (SVLasym: males, 48.74 mm; females, 47.28 mm). The growth coefficient was higher in males than in females (k: males, 1.80; females, 1.49). Fig. 2. Cross-section of a phalanx of an individual at the age of 7. White arrows show LAGs. m.c. = marrow cavity. Table 1. Descriptive statistics for the studied population. Sex N Age (years) SVL (mm) ESP (years) S Mean ± SE Range Mean ± SE Range Females 17 5.82 ± 0.30 4-8 45.60 ± 0.79 40.38-52.30 5.26 0.79 Males 23 5.65 ± 0.29 3-8 48.12 ± 0.72 41.72- 54.15 16 Abdullah Altunışık Body size was positively correlated with age in both females (Pearson’s correlation r = 0.589, P < 0.05; Fig. 4A) and males (r = 0.648, P < 0.01; Fig. 4B). DISCUSSION The first demographic data on an Ommatotriton vitta- tus population was provided in this study. The results sug- gest that Southern Banded Newt has a lower lifespan than the Northern Banded Newt. We found that maximum age could be reached at 8 years in both males and females in Tarsus population. In congeneric O. ophryticus sensu lato, maximum lifespan was recorded as 21 years for Georgian populations (Kuzmin, 1999; Tarkhnishvili and Gokhelas- vili, 1999) and 10 and 16 years for the low and high alti- tude Turkish populations, respectively (Kutrup et al., 2005; Özcan and Üzüm, 2015). In the present study, no signifi- cant difference was detected between females and males with regard to mean age. A similar result was recorded for Bahçesultan and Erbaa populations of O. ophryticus sensu lato (Özcan and Üzüm, 2015). Delayed maturity and longer lifespan are expected phenomenons for high-elevation populations, because they are exposed to colder temperatures and shorter growth seasons in comparison with low-elevation popula- tions. In line with this trend, the studied low-altitude pop- ulation of O. vittatus had lower lifespan and age at matu- rity than that of high-altitude populations of O. ophryticus sensu lato, formerly referred to as Triturus vittatus ophryti- cus, (Kuzmin, 1999; Tarkhnishvili and Gokhelasvili, 1999; Kutrup et al., 2005; Özcan and Üzüm, 2015). It is expected that amphibians with indetermined growth should enhance not only their age at maturity but also division of resources between reproduction and growth over the years following sexual maturation (Kozlowski and Uchmanski, 1987). Generally, amphib- ians reach sexual maturity when they reaches a certain size, therefore sexual maturity may vary between differ- ent species and populations (Gibbons and McCarthy, 1984; Hemelaar, 1986; Ryser, 1988). The age at maturity, which is determined to be 3-4 years, is compatible with the two populations of O. ophryticus (Kutrup et al., 2005). However, in the Tosya and Bahçesultan populations of O. nesterovi and Erbaa populations of O. ophryticus located at higher altitudes, the age at maturity was determined as 4-6 years (Özcan and Üzüm, 2015). Kuzmin (1999) stated that the sexual maturity was attained at 3-5 years in O. ophryticus for Caucasus populations. Similarly, Tarkh- nishvili and Gokhelasvili (1999) established the earliest age at maturation in Georgian O. ophryticus populations from near Tbilisi as 4 years for females and 3 years for males (Table 2). Sexual size dimorphism is common among amphib- ians. More specifically, about 90% of the anurans and 61% of the urodeles indicate a female-biased SSD (Shine, 1979; Kupfer, 2007; Reinhard et al., 2015). Interestingly, our results show that males of O.vittatus did not conform to this tendency but indicate male-biased dimorphism. Similar results were reported by Kutrup et al. (2005), Çiçek and Ayaz (2011), Başkale et al. (2013), Özcan and Üzüm (2015) and Cvijanović et al. (2017) for dif- Fig. 4. Relationship between age and body size (SVL). Females (A) and males (B). Fig. 3. Age distribution of O. vittatus from Tarsus population. 17Age structure of Ommatotriton vittatus ferent populations of O. ophryticus sensu lato. Although rarely found in the literature, males of some urodele spe- cies (e.g., Lissotriton vulgaris, Halliday and Verrell, 1988; Mertensiella caucasica, Üzüm, 2009; Lyciasalamandra atifi, rewieved in Reinhard et al., 2015) are larger than females in terms of mean body size. Most of the variation in size dimorphism has been considered to result from differences in age structure between sexes (Monnet and Cherry, 2002; Cadeddu et al., 2012; Liao et al., 2013). The annual survival rate of this newt differs from the other congeneric species (O. nesterovi), because it was calculated as 0.35 for a long-term mark-recapture and skeletochronological study (Başkale et al., 2013). The fact that Southern banded newts have a higher annual sur- vival rate than Northern Banded newt may lead to dif- ferences in the potential reproductive lifespan between these two newts. It is widely assumed that age and body size are positively correlated in amphibians (Halliday and Verrell, 1988). Although there is no positive correlation for some species, in some cases this correlation may vary depending on sex. Similar to O.nesterovi’s Tosya, Bahçe- sultan (Özcan and Üzüm, 2015) and Sakarya (Başkale et al., 2013) and O. ophryticus’s Erbaa (Özcan and Üzüm, 2015) populations, in the studied O. vittatus population, body size of both males and females showed positive sig- nificant correlations with age. Contrarily, no correlation was reported between body size and age in both popu- lations of Northern banded newt (Kutrup et al., 2005). This discrepancy among the congeneric species may be explained by the decrease of growth rates after sexual maturity is achieved, as well as by the different interan- nual reproductive effort of individuals, resulting in differ- ent individual growth curves (Kutrup et al., 2005). In conclusion, as mentioned before the relatively low longevity and age at maturity observed in Southern banded newt could result from local ecological condi- tions (e.g., altitude, temperature, length of the activity period). It is also necessary to investigate more popula- tions exposed to various conditions in order to evaluate more precisely the interpopulational variation of these life-history characteristics in O. vittatus. ACKNOWLEDGEMENTS I am grateful to M. Emin Altunışık for his assis- tance in collecting animals from the field. This study was supported by Recep Tayyip Erdoğan University BAP under grant number 2016.53007.102.03.01. All experi- ments were performed in accordance with the Turkish law (number of permission to capture: 72784983-488.04- 45006) and with permission for animal experiments of the local ethics committee of the Recep Tayyip Erdoğan University (approval reference number: 2015/71). I kindly Table 2. Demographic life history traits in Ommatotriton species throughout its geographical range. Species Location Altitude (a.s.l.,m) Sex Mean age (years) Age at maturity (years) Longevity (years) Mean SVL (mm) References O. ophryticus Different parts of Georgia - Male - 3-5 8-21 - Kuzmin (1999) Female - 3-5 8-21 - O. ophryticus Akhaldaba, Georgia 750 Male 5.53 3 8 - Tarkhnishvili and Gokhelashvili (1999)Female 6.97 4 12 - O. ophryticus Gürbulak, Trabzon, Turkey 300 Male 4 2-3 10 61.7 Kutrup et al. (2005) Female 4 2-3 10 53.3 O. ophryticus Hıdırnebi, Trabzon, Turkey 1300 Male 8.2 4 16 59.6 Kutrup et al. (2005) Female 9.2 4 16 51.0 O. nesterovi* Sakarya, Turkey 31 Male 4.92 3 8 90.79 Başkale et al. (2013) Female 4.90 2 6 86.06 O. nesterovi* Bahçesultan, Bilecik, Turkey 1015 Male 7.4 6 10 72.39 Özcan and Üzüm, 2015 Female 7.4 6 10 63.40 O. nesterovi* Tosya, Kastamonu, Turkey 1202 Male 6.4 4 9 64.07 Özcan and Üzüm, 2015 Female 7.06 5 9 55.57 O. ophryticus Erbaa, Tokat, Turkey 1239 Male 8.2 6 12 71.07 Özcan and Üzüm, 2015 Female 8.4 5 11 60.36 O. vittatus Tarsus, Mersin, Turkey 25 Male 5.65 3 8 48.12 < Female 5.82 4 8 45.60 SVL, snout-venth length; *referred to as O.ophryticus 18 Abdullah Altunışık thank three anonymous reviewers for constructive sug- gestions on improving the manuscript. REFERENCES Altunışık, A. (2017): Sexual size and shape dimorphism in the Near Eastern fire salamander, Salamandra infraimmaculata (Caudata: Salamandridae). Anim. Biol. 67: 29-40. Altunışık, A., Kalayci, T.E., Uysal, İ., Tosunoğlu, M., Özdemir, N. (2016): Age, adult survival rate, and adult life expectancy of a Podarcis tauricus population (Reptilia: Lacertidae) from Saros Bay, Turkey. Russ. J. 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Acta Herpetologica Vol. 13, n. 1 - June 2018 Firenze University Press Species diversity and distribution of lizards in Montenegro Katarina Ljubisavljević1,2,*, Ljiljana Tomović3, Aleksandar Urošević1, Slađana Gvozdenović2, Vuk Iković2, Vernes Zagora2, and Nenad Labus4 The first demographic data and body size of the southern banded newt, Ommatotriton vittatus (Caudata: Salamandridae) Abdullah Altunişik Diet and helminth parasites of freshwater turtles Mesoclemmys tuberculata, Phrynops geoffroanus (Pleurodira: Chelidae) and Kinosternon scorpioides (Criptodyra: Kinosternidae) in a semiarid region, Northeast of Brazil Antonio Marcos Alves Pereira*, Samuel Vieira Brito, João Antonio de Araujo Filho, Adonias Aphoena Martins Teixeira, Diêgo Alves Teles, Daniel Oliveira Santana, Vandeberg Ferreira Lima, Waltécio de Oliveira Almeida Electric circuit theory applied to alien invasions: a connectivity model predicting the Balkan frog expansion in Northern Italy Mattia Falaschi1,2,*, Marco Mangiacotti3, Roberto Sacchi3, Stefano Scali2, Edoardo Razzetti4 First report of Bufo bufo (Linnaeus, 1758) from Sardinia (Italy) Ilaria Maria Cossu1, Salvatore Frau1, Massimo Delfino2,3, Alice Chiodi4, Claudia Corti1,5*, Adriana Bellati4 Natural history and conservation of the Nurse Frog of the Serranía del Perijá Allobates ignotus (Dendrobatoidea: Aromobatidae) in northeastern Colombia Hernán Darío Granda-Rodríguez1,2, Andrés Camilo Montes-Correa3,*, Juan David Jiménez-Bolaño3, Marvin Anganoy-Criollo4,5 Exploring body injuries in the horseshoe whip snake, Hemorrhois hippocrepis Juan M Pleguezuelos*, Esmeralda Alaminos, Mónica Feriche How effectively do European skinks thermoregulate? Evidence from Chalcides ocellatus, a common but overlooked Mediterranean lizard Grigoris Kapsalas, Aris Deimezis-Tsikoutas, Thanos Georgakopoulos, Ismini Gkourtsouli-Antoniadou, Kallirroi Papadaki, Katerina Vassaki, Panayiotis Pafilis Thermal tolerance for two cohorts of a native and an invasive freshwater turtle species Jun Geng, Wei Dang, Qiong Wu, Hong-Liang Lu* Diet of a restocked population of the European pond turtle Emys orbicularis in NW Italy Dario Ottonello1, Fabrizio Oneto1,2, Monica Vignone2, Anita Rizzo2, Sebastiano Salvidio2,* Helminths of the lizard Colobosauroides cearensis (Squamata, Gymnophthalmidae) in an area of Caatinga, Northeastern Brazil Aldenir F. da Silva Neta*, Robson W. Ávila