North-Western Journal of Zoology Vol. 5, No. 1, 2009, pp.74-84 P-ISSN: 1584-9074, E-ISSN: 1843-5629 Article No.: 051109

A skeletochronological study of age, growth and longevity in a population of the Caucasian , Mertensiella caucasica (Waga 1876) (Caudata: ) from Turkey

Nazan ÜZÜM

Adnan Menderes University, Faculty of Arts and Sciences, Department of Biology, 09010 Aydın, Turkey, E-mail: [email protected]

Abstract. Age structure, size, and growth patterns of the local population of the Caucasian Salamander, Mertensiella caucasica (Waga 1876) were determined by using skeletochronology. Lines of arrested growth (LAGs) recorded in phalanges were used to estimate the age of juveniles and adults. Results show that juveniles were from 1 to 3 years old. The average age was significantly higher in males (7.3 ± 1.28 yrs) than in females (6.0 ± 1.15 yrs). The maximum observed longevity was 10 years in males and 9 years in females. Age at maturity in this population was 4-5 years in both sexes. The average SVL (snout-vent length) did not differ between sexes (males: 67.5 mm and females: 63.8 mm). The average weight of males was 4.7 g, and significantly higher than the weight of females (3.7 g). The growth curves for M. caucasica population were well described with a von Bertalanffy growth model. K (0.348±0.026 for males and 0.129±0.009 for females) and SVLmax (72.03±1.02 mm for males and 102.00±3.80 mm for females) were significantly differed between males and females in this population.

Key words: skeletochronology, age structure, growth, longevity, Mertensiella caucasica, Turkey.

Introduction part of the range belongs to the watershed, while the eastern part drains to Mertensiella caucasica (Waga 1876), the the Caspian Sea. M. caucasica is an ovipa- Caucasian Salamander, is one of at least six rous species whose breeding depends upon known representatives of the salamander water (Schultschik 1994, Tarkhnishvili subfamily Salamandrinae, which includes 1994). Its presence in North East Anatolia is the genera Chioglossa, Mertensiella, Lycia- known from Vilayets Giresun, Rize, salamandra and Salamandra, with a distri- Trabzon, Artvin and Gümüşhane (Atatür & bution over mountainous regions of Europe Budak 1982, Baran & Atatür 1998, Franzen and the Near East (Nöllert & Nöllert 1992, 1999). Griffiths 1996, Frost et al. 2006). It is a rare The male of Mertensiella is characterized species with uneven spatial distribution. by the presence of a conspicuous tubercle The range of M. caucasica is restricted to projecting from the skin of the dorsal Turkey and . The larger western surface of the tail base. The sister taxon of

©NwjZ, Oradea, Romania, 2009 North-West J Zool, 5, 2009 www.herp-or.uv.ro/nwjz Oradea, Romania Age structure in Mertensiella caucasica 75

M. caucasica is not the Anatolian Lycia- sex of each individual was determined according to salamandra Veith & Steinfartz 2004, but the external secondary sexual characters: Male has a fleshy protuberance at the base of the tail, whereas Iberian Chioglossa lusitanica Bocage 1864 female has a prominent cloaca but without the (Titus & Larson 1995, Veith et al. 1998, Frost protuberance. Juvenile, is differentiated from adult et al. 2006). M. caucasica is listed in the one in having smaller body length and lacking both IUCN Red List of Threatened as the protuberance and the prominent cloaca (Başoğlu Vulnerable (Hilton-Taylor 2000). Popula- et al. 1996, Baran & Atatür 1998). were measured from the tip of the snout to the posterior tions of this species in Georgia display margin of the vent (SVL) by using a dial caliper with significant fluctuations, and the population an accuracy of 0.02 mm. Body mass was also in Turkey has probably declined signi- weighted to the nearest 0.01 g. Sexual size ficantly over the past decade. In Turkey, dimorphism was estimated with the Lovich & only around 12% of suitable forest habitat Gibbons (1992) sexual dimorphism index (SDI): remains within the species range (Özhatay ⎛ sexlargerofsize ⎞ SDI = ⎜ ⎟ m 1 et al. 2003). ⎝ size smallerof sex ⎠ The aim of this study was to determine life history characteristics (e.g. size and age +1 if males are larger or -1 if females are larger, and arbitrarily defined as positive when females are at maturity, longevity, growth) of a popu- larger than males and negative in the converse case. lation of the rare, narrow ranged Caucasian salamander, M. caucasica by using skeleto- chronological analysis. Post metamorphic Skeletochronological analysis growth, minimum age at sexual maturity The longest finger of the forelimb (i.e. second or and life span were estimated, providing third) was cut and preserved in 70% ethanol. Age crucial information to the better under- was determined by using skeletochronological standing of the species population dyna- analysis previously described (e. g. Castanet & mics as well as ecological and conservation Smirina 1990, Olgun et al. 2005): The phalanges stored in 70% ethanol solution were dissected; the requirements. largest bone of the phalange was washed in running water for about 24 hours, decalcified in 5% nitric acid for 2 hours and then washed again in running water Material and Methods overnight. The mid-diaphyseal region of each phalanx was sectioned at 15-18µm using a freezing Study area, climate and sampling microtome. The resulting sections were stained in Ehrlich’s hematoxylin. The sections were mounted The population studied was located in Kümbet on slides and observed with a light microscope. Bone Plateau in Dereli/Giresun (40°33′ N, 38°26 ′ E) at an sections from each individual were photographed (at altitude of 1575 m a.s.l. (Figure 1). The vegetation the same magnification) using a camera lucida, was typical Black Sea region woodland with mainly allowing for simultaneous comparison and coniferous (Abies nordmanniana and Picea orientalis) facilitating the analysis of the bone growth pattern. mixed forests. The active period of M. caucasica starts Two different persons who have similar experience in May and ends in October, a period when with the technique counted the annular rings of temperature allows breeding and feeding activities growth in each section (Olgun et al. 2001, Olgun et (N. Üzüm, personal comment). al. 2005). The salamanders used in the present study (19 LAGs appeared as colored stripes of various males, 13 females and 3 juveniles) were all caught in widths. Counting the number of LAGs in cross the daytime on the 09 September 2007. The method sections of the phalanges revealed the number of of capture was by hand, with individuals taken from arrested growth periods experienced by each under stones in the shallow water of the stream. The individual. Due to the salamanders were caught at

North-West J Zool, 5, 2009 76 Üzüm, N. the beginning of autumn, the periphery of the cross linear regression of juveniles: body size = -0.38 x age sections was not regarded as the outer LAG. + 42.83 (r=-0.58, P=0.06047, n=3). The von Bertalanffy growth model was fitted to the average growth curve using the least square procedure. The para- Data analysis meters SVLmax, K and their asymptotic confidence intervals (CIs) were also estimated by non-linear All age-classes and all body measurements (SVL and least-square regression. weight) were normally distributed (Kolmogorov- Smirnow D test, all P > 0.05), thus allowing comparisons using parametric tests (t test). All tests were processed with STATISTICA 6.0 (Statsoft Inc., Results USA) and Excel (Microsoft), and interpreted at α= 0.05. Line of arrested growth and age estimation Growth was estimated using the von Bertalanffy (1938) model, taking into account only the juvenile Bone growth pattern at the phalangeal and adult stages of the life cycle (Arntzen 2000, Olgun et al. 2001) as: midshaft diaphyses were similar to those described previously for Salamandra atra ( −− ttK met ) t SVLSVL max (SVLmax −−= met )eSVL and S. lanza (von Fachbach 1988, Miaud et al. 2001) and M. luschani (Olgun et al. 2001). where SVLt= average body size at age t, SVLmax= At the diaphysis level, some cartilage may average maximum body size, SVLmet= average body size at metamorphosis (tmet), t= number of growing remain in the marrow cavity in adults. The season experienced (age), tmet= proportion of the marrow cavity also can be enlarged by growing season until metamorphosis (age at endosteal resorption which creates an metamorphosis), and K= growth coefficient (shape of erosion of the periosteal bone on the edge of the growth curve). The corresponding tmet intercept on the time axis was fixed at zero. Because we did the marrow cavity. Comparison of the osteal not collect juveniles at metamorphosis, size at meta- cavity diameters in juveniles and adults morphosis (SVLmet) was estimated from the age/size showed that resorption sometimes locally

Figure 1. Map showing the locality of studied samples of Mertensiella caucasica.

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destroys the first and the second lines of Body size, weight and growth arrested growth (LAGs) and sometimes completely destroys the first LAG as seen in According to the results of the t-test (t= Figure 2A and Figure 2B, respectively. 1.77, df= 30, p=0.085), SVL of males was Moreover, according to Tarkhnishvili & not statistically different than that of Gokhelashvili (1994) who give the pre- females. The sexual dimorphism index was liminary data of the age structure of M. found to be -0.06. The difference in weight caucasica, just before metamorphosis larvae between the sex was significant (t = 2.62, df have at least one year ring. Sometimes indi- = 30, p = 0.013), males were larger and viduals could be distinguished as having a heavier than females. Table 2 summarizes second or third ring and they concluded the relationships of the variables. The SVL that those rings corresponded to 2 to 4 years of the Caucasian salamander was poorly of life. Thus, it can be considered the age of correlated with age in both sexes, while each salamander to be at least 3 years more there was a strong positive correlation in than the number of visible year rings. The both sexes with SVL and weight. error might be ±1 year. The von Bertalanffy (1938) growth Adult males and females caught during model correctly fitted the age/body-length their activity season possessed external relationship of the sampled individuals secondary sexual characters and were (Figure 4). R=0.897 and percentage of considered mature. Juveniles did not exhibit variance explained (VE) was 80.4% in males such characters. Descriptive statistics on and R=0.798 and VE=63.7% in females. age, body size and weight are given in Table Males and females significantly differ in 1. growth parameters: the average maximum Thirty-five bones (representing 3 juve- length (SVLmax) was 72.03±1.02 mm for niles, 19 males, and 13 females) were scored males and 102.00±3.80 mm for females. The and all of them (100%) allowed for an growth coefficient K was 0.348±0.026 for estimation of the individual’s age. Juveniles males and 0.129±0.009 for females. were from 1 to 3 years old. The minimum age for adult salamanders was found to be 5 and 4 years for males and females respecti- Discussion vely. The maximum observed longevity was 10 years in males while it was 9 years for Skeletochronology is widely used to females. The average ages of the males and estimate age in and reptiles females were calculated as 7.26±1.28 and (Castanet, 2002). Although the winter 6.00±1.15 years, respectively (Table 1). The growth marks can be weakly expressed in difference in terms of age between the sex warm, temperate climate regions (e.g. in R. was found statistically significant (t = 2.84, perezi from southwest Spain [Esteban et al. df = 30, p = 0.007). The youngest female was 1996]), the LAGs in the Caucasian sala- 4 years old, the youngest male was 5 years mander were clearly stained. This has been old, and the oldest juvenile was 3 years old. observed previously in species exposed to I thus estimated age at maturity to be 4-5 harsh environmental conditions of high years old (Figure 3). altitude or latitude (Esteban & Sanchiz 2000,

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Figure 2. Cross sections through phalanges of male and female Mertensiella caucasica. (A) Male 69.66 mm SVL. Six LAGs were observed in the periosteal bone. The innermost and the second LAGs were almost completely destroyed by endosteal resorption. This individual was 6 years old. (B) Female 68.52 mm SVL. Five LAGs were observed in the periosteal bone. The first LAG was completely and the second LAG was partially destroyed by the endosteal resorption. This individual was 6 years old.

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Table 1. Descriptive statistics of age, SVL and weight in Kümbet specimens.

Age (years) SVL (mm) N Mean ± SD Range N Mean ± SD Range Juveniles 3 2.00 ± 1.00 1-3 3 42.07 ± 0.65 41.46-42.76 Males 19 7.26 ± 1.28 5-10 19 67.53 ± 2.38 63.10-73.36 Females 13 6.00 ± 1.15 4-9 13 63.77 ± 8.83 51.28-76.60

Weight (g) N Mean ± SD Range Juveniles 3 1.11 ± 0.19 0.92-1.31 Males 19 4.72 ± 0.71 3.51-5.73 Females 13 3.66 ± 1.53 2.12-7.25

Lai et al. 2005, Leclair & Laurin 1996, Leclair mental influences in the frog Rana sylvatica et al. 2000, Miaud et al. 1999, Miaud et al. (Berven 1982) and the salamander 2000). It was determined that only one LAG Desmognathus ocoee (Bernardo 1994). In the was formed during each hibernating period latter study, the experimental design and they could easily be identified in the revealed that growth and maturation specimens examined in this paper. evolved independently. In newts, age and Endosteal resorption was observed in all size variation would result from phenotypic cross sections of adult M. caucasica. Several plasticity (Miaud et al. 2000). authors suggested that resorption may be linked to environmental conditions (Smirina 1972), with e.g. less resorption for popu- Table 2. Correlations among variables recorded in the Caucasian salamander Mertensiella caucasica lations living in high altitudes than for from Kümbet Plateau, Giresun. Rs = Spearman lowland populations (Esteban et al. 1996, correlation coefficient, N = sample size, p = 1999) or the opposite (Caetano & Castanet significance. 1993). Age at maturity was estimated at 4-5 years in Kümbet specimens while N Rs p Tarkhnishvili & Gokhelashvili (1994) Males reported that after metamorphosis, at least 9 Age - SVL 19 0.17 0.50 years pass (9 to 13 years) before maturation. Age - Weight 19 0.21 0.40 Since these two populations live in different SVL - Weight 19 0.74 0.00 habitats and are exposed to different Females ecological and biological factors, they show some variations in their life history traits. Age - SVL 13 0.59 0.03 Differences in age between different en- Age - Weight 13 0.54 0.05 vironments were also found to result from SVL - Weight 13 0.92 0.00 the combination of genetic and environ-

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9 Male 8 Female 7

6

5

4

3

Number ofindividuals Number 2

1

0 45678910 Age (ye ars)

Figure 3. Age distribution of adult Mertensiella caucasica.

In this study, age between the sexes was rainfall, food supply, predation, compe- found to be statistically significant (t = 2.84, tition, etc.) and ultimate (genetic) factors. df = 30, p = 0.007), indicating that adult These factors can result in differentiation males were on average older than females. among local populations (Morrison & Hero Lima et al. (2000) also found a significant 2003, Sorci et al. 1996). More interpopu- difference in age structure between the lational and interspecific studies are needed sexes in Chioglossa lusitanica, but indicating on the proximate effect of various environ- that adult males were on average younger ments on salamander life-history traits to than females. The maximum life span of test hypotheses on local adaptation versus Kümbet specimens was estimated as 10 phenotypic variation. years in males while it was 9 years in The SVL of individuals in the Kümbet females. These results are similar to the population was similar to the length range result found for its sister species, C. of salamanders reported by Tarkhnishvili & lusitanica where longevity was estimated at Gokhelashvili (1994). SVL of adult males 10 years. However, Tarkhnishvili & varied between 63.10-73.36 mm and 51.28- Gokhelashvili (1994) reported that adult M. 76.60 mm in females (these values de- caucasica can live for ten years or more after termined as 61-73 mm in males and 70-78 maturation as long as 26 years. I suggest mm in females for Georgian population). that the plasticity of life history trajectories Sexual size dimorphism is observed in of M. caucasica depend upon different many amphibians, and females are often environments. Sources of variation in life larger than males in many salamanders and history traits for organisms include newts (e.g. Diaz-Paniagua et al. 1999, Miaud proximate (environmental: temperature, et al. 2000, Olgun et al. 2001, Olgun et al.

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2005). Contrary to this, males have larger having better access to females than smaller average SVL than females in our indi- males. Sexual selection for body size is one viduals as in Triturus vulgaris (Halliday & such factor where increased male body size Verrell 1988) and Ommatotriton ophryticus is an important determinant of male mating (Kutrup et al. 2005). Males larger than success; males should not be expected to females are not common, but the reason forego growth in order to breed earlier than could be of sexual selection, i.e larger males females (Monnet & Cherry 2002).

Figure 4. Growth curves in males (A) and females (B) in the Caucasian Salamander (Mertensiella caucasica). Growth curves were fitted to von Bertalanffy’s growth equation.

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The age of adult animals is significantly ledge use of Maptool program for analysis and graphics correlated with their size (r = 0.50, p<0.05), in this paper. Maptool is a product of Seaturtle.org (information is available at www.seaturtle.org) although Tarkhnishvili & Gokhelashvili (1994) found the contrary result for their studied population. The growth curve of References Caucasian salamander is relatively flat in females (small K value) while the Arntzen, J. W. (2000): A growth curve for the newt asymptotic size is larger (Figure 4). In C. Triturus cristatus. Journal of Herpetology 34: 227- 232. lusitanica, it was also found that K was Atatür, M. K., Budak A. (1982): The present status of higher for males than for females while Mertensiella caucusica in Northeastern Anatolia. SVLmax was higher for females than for Amphibia-Reptilia 4: 295-301. males (Lima et al. 2000). Information on Baran, İ., Atatür, M. K. (1998): Turkish herpetofauna (amphibians and reptiles). Republic of Turkish such asymptotic size-K relationships is Ministry of Environment, Ankara. needed in other urodelan species. The Başoğlu, M., Özeti, N., Yılmaz, İ. (1996): Türkiye opposite relation was found in R. temporaria Amfibileri (The Amphibians of Turkey). Ege Üniversitesi Fen Fakültesi Kitaplar Serisi, No: 151. (K increase when asymptotic sizes decrease: İzmir. Miaud & Guyétant 1998) and C. lusitanica Bernardo, J. (1994): Experimental analysis of allocation in (Lima et al 2000). Such a negative two divergent, natural salamander populations. relationship was also found in several American Naturalist 143: 14-38. Berrigan, D., Charnov, E. L. (1994): Reaction norms for species of fishes (Berrigan & Charnov 1994). age and size at maturity in response to temperature: This study has great importance to give a puzzle for life-historians. Oikos 70:474-478. new information on age, growth and Berven, K. A. (1982): The genetic basis of altitudinal variation in the wood frog Rana sylvatica. I. An longevity of M. caucasica from different experimental analysis of life history traits. Evolution habitats. As it seen from this study, age 36:962-983. structure, growth and longevity of this Caetano, M. H., Castanet, J. (1993): Variability and population showed differences from microevolutionary patterns in Triturus marmoratus from Portugal: age, size, longevity and individual another population studied by growth. Amphibia-Reptilia 14: 117-129. Tarkhnishvili & Gokhelashvili (1994). The Castanet, J. (2002): Amphibiens et Reptiles non aviens: interpopulational differences in life history un matériel de choix en squelettochronologie. Bulletin de la Société Herpétologique de France 103: traits suggest that environmental conditions 21-40. could play an important role on the Castanet, J., Smirina, E. M. (1990): Introduction to the populations’ structures. More studies are skeletochronological method in amphibians and thus needed on the proximate effect of reptiles. Annales des Sciences Naturelles, Zoologie 11:191-196. various environments on this newt life his- Diaz-Paniagua, C., Mateo, J. A. (1999): Geographic tory trait and detailed studies are required variation in body size and life-history traits in for consistent conservation measures. Bosca’s Newt (Triturus boscai). Herpetological Journal 9:21-27.

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