Odabas_Buelbuel_etal_Lacerta_trilineata_Age_Growth_TR:HERPETOZOA.qxd 12.02.2019 14:56 Seite 1

HerPeTOzOa 31 (3/4): 183 - 193 183 Wien, 28. Februar 2019

age structure and growth in a Turkish population of the Balkan Green , trilineata BedriaGa , 1886 (: Sauria: )

altersstruktur und Wachstum in einer türkischen Population der riesensmaragdeidechse Lacerta trilineata BedriaGa , 1886 (Squamata: Sauria: Lacertidae)

YaSeMiN OdaBaş & U FUk BüLBüL & a Li İHSaN erOğLU & H aLiMe kOç & M UaMMer kUrNaz & B iLaL kUTrUP

kUrzFaSSUNG

die Studie bringt informationen zum alter und Wachstum von Lacerta trilineata BedriaGa , 1886 in der Population von Sergen, Westtürkei (420 m ü. M.). das alter bei erreichen der Geschlechtsreife, geschlechtsbe - dingte Größenunterschiede (SSd) und Wachstumsraten werden angegeben. dazu wurden Querschnitte von zehenknochen mit skeletochronologischen Methoden untersucht. in der Stichprobe aus erwachsenen individuen (14 Männchen und 12 Weibchen) betrug das alter für beide Geschlechter zusammen 6-13 (  = 8,77) Jahre, wobei der Unterschied zwischen Männchen mit 7-13 (  = 9,07) Jahren und Weibchen mit 6-10 (  = 8,42) Jahren nicht signifikant war. die Geschlechtsreife erreichen beide Geschlechter im dritten Lebensjahr. Weder bei Männchen noch Weibchen zeigte sich eine signifikante korrelation zwischen kopf- rumpflänge (SVL) und alter. die Männchen der Sergen Stichprobe waren kaum größer als die Weibchen (SSd = 0.014). der Wachstumskoeffizient (k) war bei Weibchen kleiner als bei Männchen (k ± konfidenzintervall, Männchen: 0.96 ± 0.22; Weibchen: 0.75 ± 0.41). die Wachstumsraten der Geschlechter unterschieden sich nicht voneinander. aBSTraCT

This study provides information about age and growth of Lacerta trilineata BedriaGa , 1886, in the popu - lation from Sergen, West , at an altitude of 420 m a.s.l. age at maturity, sexual size difference (SSd) and growth rate of the sample are presented. Cross-sections of phalangeal bones were examined based on the skeletochronology method. in the adult sample (14 males and 12 females), the age ranged from 6-13 (  = 8.77) years for both sexes collectively, the val - ues 7-13 (  = 9.07) years in males and 6-10 (  = 8.42) years in females not differing significantly from each other. Sexual maturity was attained in the third year of life in both sexes. There was no significant correlation between the ’ body size (SVL) and age for both males and females. a slightly male-biased sexual size dimorphism (SSd = 0.014, not significant) was observed. The growth coefficient (k) was lower in females than males (k ± con - fidence interval, males: 0.96 ± 0.22; females: 0.75 ± 0.41). There was no difference in the growth rate between sexes. keY WOrdS reptilia: Squamata: Sauria: Lacertidae; Lacerta trilineata ; ecology, life history, skeletochronology, LaGs, lines of arrested growth, von Bertalanffy growth curve, age, growth, longevity, body size, Turkey

iNTrOdUCTiON

The Balkan Green Lizard, Lacerta tri - western Turkey in the east (mostly according lineata BedriaGa , 1886, is the largest spe - to UeTz et al. 1995-2018). it is classified as cies of the . it has a wide distribution LC (Least Concern) in the iUCN red List area from the coastal regions of , (BöHMe et al. 2009) due to its wide distribu - , , Monte - tion, presumed large population and the negro and southwards across unlikeliness to decline fast enough to quali - , , Macedonia and to fy for listing in a more threatened cate gory. Odabas_Buelbuel_etal_Lacerta_trilineata_Age_Growth_TR:HERPETOZOA.qxd 12.02.2019 14:56 Seite 2

184 Y. O daBaş & U. B üLBüL & a. İ. e rOğLU & H. k Oç & M. k UrNaz & B. k UTrUP

age determination studies are carried ronmental conditions affect the lizards’ life out by researchers to reveal the life history history traits such as age at maturity, traits of species ( eSTeBaN et al . 2004; longevity, reproductive phenology, survival rOiTBerG & S MiriNa 2006; G UariNO et al. rates and growth rates ( adOLPH & P OrTer 2010). Skeletochronology is the most 1996). appropriate and reliable method for deter - üSTeL (2010) reported on longevity mining the age of lizards. Many studies on and the relationship between age and SVL lizards in Turkey used skeletochronological in the L. trilineata population from the methods ( arakeLYaN et al . 2013; aLTUNişik Gelibolu and Biga Peninsulas, Province of et al . 2013; TOk et al . 2013; GüL et al. 2014; çanakkale, Turkey, and the paper by üzüM et al . 2014; YakiN & T Ok 2015; kaLaYCi et al. (2018) dealt with life-history kaNaT & T Ok 2015; G üL et al . 2015; üzüM traits such as average and maximum age, et al . 2015). This method is based on count - age upon arrival at maturity and various ing of the lines of arrested growth (LaGs) morphometric parameters relative to age, that appear in long bone sections of individ - comparing Turkish lowland (edirne, 17 m uals with strongly reduced activity during a.s.l.) and high altitude (Bolu, 1,250 m the hibernation, estivation or starvation a.s.l.) populations. The aim of this study is periods ( HeMeLaar 1981; C aSTaNeT & to add life-history information on NW SMiriNa 1990; S MiriNa 1994). during their Turkish Lacerta trilineata based on skele - periods of surface activity, climate and envi - tochronological methods.

MaTeriaLS aNd MeTHOdS

a total of 26 adult specimens (14 males are larger than females and −1 if the males and 12 females) were caught from a opposite is true. The result is arbitrarily population in Sergen, Province of kırkla- defined as positive if the females are larger reli, european Turkey, during the breeding and negative if the males are larger (L OViCH season (capture permission no. 72784983- & G iBBONS 1992). 488.04-42844 issued by the Turkish Minis- From each lizard, the longest (4th) toe try of Forest and Water affairs). The was clipped at the second phalange and pre - Sergen population (41°42’31” N, 27°42’ served in 10 % formalin solution for subse - 27” e) is located at an altitude of 427 m quent histological analyses. after toe-clip - a.s.l. The lizards were sexed by examina - ping, the lizards were released back into tion of the secondary sexual characters their original habitats. The specimens were (presence of light blue color at the ventral treated in accordance with the guidelines of side of the head in adult males) and by the ethics committee of the karadeniz sounding for the presence or absence of Technical University (kTü.53488718- hemipenis pockets. 417/2016/38). at the collecting site, the lizard sur - after ablation of the skin, the toes face-active period lasts from early March to were put in 5 % nitric acid solution for 2.5 November. The lizards were captured on hours to decalcify the bone tissue. after the 12-16 august 2017 between 11.00 a.m. and toe samples had passed a tissue processing 4.00 p.m. The average temperature at the system (Leica ® Tissue processor), they were time of sampling was 29 °C according to embedded in paraffin with a tissue embed - climate data provided by 1st edirne Mete - ding device (Thermo ®). deparaffinized orology regional directorate. cross-sections (10 μm, rotary microtome) of Snout-vent length (SVL) was meas - the phalanges were stained with hema - ured to the nearest 0.01 mm using a digital toxylin using the procedure described by caliper; sexual size dimorphism was quanti - BüLBüL et al. (2016), then mounted on fied by the Sexual dimorphism index (Sdi) microscope slides, closed using entellan ® as described by the formula: Sdi = (mean and observed under a light microscope. SVL of the larger sex / mean SVL of the Skeletochronology is widely applied smaller sex) ± 1. The value +1 is used if to gain information about the life histories Odabas_Buelbuel_etal_Lacerta_trilineata_Age_Growth_TR:HERPETOZOA.qxd 12.02.2019 14:56 Seite 3

Life-history traits of Lacerta trilineata BedriaGa , 1886, from Sergen, Turkey 185

of individuals with the aim to describe age Sample kolmogorov-Smirnov Test, P > structures, growth rates and maturity ages 0.05) within the sample, the parametric within the studied population ( CaSTaNeT et independent Samples T-Test was used for al. 1993; üzüM & O LGUN 2009). There is comparison of means and Spearman’s rank agreement among researchers that this Correlation Test ( P < 0.01) to analyze cor - method produces reliable results when relations. all statistic tests were processed clipped phalangeal bones are analyzed, so with SPSS 21.0 for Windows and the level that even endangered populations can be of significance set at 0.05. studied since the individuals are not de- Using the von Bertalanffy’s model, tached from their habitat (C aSTaNeT & growth curves were formed as described in SMiriNa 1990; GUariNO et al. 2010). Study - the literature (J aMeS 1991; W aPSTra et al. ing cross-sections of long bones, the age of 2001; rOiTBerG & S MiriNa 2006; G UariNO individuals is estimated counting the lines et al . 2010). The authors used the general of arrested growth (LaGs) that are formed form of the von Bertalanffy equation, L t = -k (t-t ) during the hibernation period. Summer heat L∞ (1 – e 0 ), where L t is SVL at the can lead to estivation which results in an age t, L ∞ is the asymptotic maximum accessory LaG preceding the following length, e is the base of the natural loga - hibernation-caused LaG, both appearing as rithm, k is a growth coefficient, and t 0 is double line ( CaBezaS -C arTeS et al. 2015; the age at hatching. due to the unavail - CaSTaNeT et al. 1993; YakiN & T Ok 2015). ability of data on the hatching size in the To minimize the subjective error margin, studied population L t = 27.0 mm was three of the authors (a. İ. eroğlu, Y. Odabaş used, which is the mea0n value provided by H. koç) independently counted the LaGs JaMeS & S HiNe (1988). The parameters L ∞ on the cross-sections and their results were (asymptotic maximum SVL) and k, and compared and harmonized. double lines their asymptotic confidence intervals (Ci), were counted as single lines. arrival at sex - were estimated using a non-linear regres - ual maturity was assumed where any obvi - sion procedure. Growth rates (r) were cal - ous decrease in space between two subse - culated as r = k (L ∞ – L t). Growth curves quent LaGs was observed ( rYSer 1988; were considered to be significantly differ - YiLMaz et al . 2005; özdeMir et al . 2012). ent if their 95 % confidence intervals did Since age classes and SVL measure - not overlap ( JaMeS 1991; W aPSTra et al . ments were normally distributed (One- 2001).

reSULTS

a growth zone and thin hema - (i.e., after their third hibernation) for both toxylinophilic outermost line corresponding sexes in all 26 specimens. to a winter line of arrested growth were The means of the adult SVL, age and present in the phalangeal cross sections of growth rate values were, 109.37 ± 5.27 mm; all 26 individuals (Fig. 1). in several cases, 8.77 ± 1.6 yrs; 0.07± 0.1 mm/yr for all indi - the first (innermost) LaG was not com - viduals of L. trilineata (110.12 ± 6.09 mm; pletely decomposed by endosteal resorption 9.07 ± 1.6 yrs; 0.037 ± 0.05 mm/yr in the or the resorption zone did not even reach the male and 108.51 ± 4.21 mm; 8.42 ± 1.37 first LaG. in all cross-sections, the exten - yrs; 0.32 ± 0.34 mm/yr in the female speci - sion of the resorption zone was clearly mens) (Table 1). restricted to endosteal bone and never a dif - age ranged from 7-13 years in males ficulty for age determination. double lines and 6-10 years in females. The mean age of and endosteal resorption were observed in the specimens was not significantly differ - 17 (65.4 %) and 4 (15.4 %) individuals, ent between the sexes (independent Sam- respectively. The oldest females were 10, ples T-Test; t = 1.020, df = 24, P = 0.318). the oldest males 13 years old (Fig. 2). The intersexual differences in body size (SVL) age upon arrival at maturity was three years were slightly male-biased (SSd = 0.014). Odabas_Buelbuel_etal_Lacerta_trilineata_Age_Growth_TR:HERPETOZOA.qxd 12.02.2019 14:56 Seite 4

186 Y. O daBaş & U. B üLBüL & a. İ. e rOğLU & H. k Oç & M. k UrNaz & B. k UTrUP

Fig. 1: Cross section (10 μm) of a toe bone of a six-year-old female (117.23 mm SVL) of Lacerta trilineata BedriaGa , 1886 from the Sergen population in Turkey. The age was derived from the presence of six lines of arrested growth (marked with six > symbols in the upper portion of the picture) surrounding the resorption line (rL). MC – marrow cavity; eB – endosteal bone; rL – resorption line; dL – double line; P – periphery. abb. 1: Querschnitt (10 μm) eines zehenknochens eines sechs Jahre alten Weibchens (117.23 mm kopf-rumpflänge) von Lacerta trilineata BedriaGa , 1886 aus der Population von Sergen (Türkei). das alter wurde aus der anzahl von sechs Linien verlangsamten Wachstums (markiert durch sechs > Symbole im rechten oberen Bildbereich) außerhalb der resorptionslinie (rL) abgeleitet. MC – Markhöhle; eB – endostaler knochen; rL – resorptionslinie, dL – doppelline, P – Peripherie.

The mean SVL was not significantly differ - 27.0 mm was slightly shorter than the max - ent between sexes ( t = 0.771, df = 24, P = imum SVL recorded (SVL asym ± Ci, males: 0.448). The correlation between SVL and 108.40 ± 25,68 mm; females: 109.21 ± age was weak for both males (Spearman’s 44.21 mm). The growth coefficient was rank correlation: r = 0.318, P = 0.268) and higher in males than females (k ± Ci, males: females ( r = -0.675, P = 0.016). The growth 0.96 ± 0.22; females: 0.75 ± 0.41). There curves estimated by von Bertalanffy’s equa - was no significant difference in growth rate tion are shown in Fig. 3. For both sexes, the between the sexes (independent Samples T- estimated asymptotic SVL based on L = Test; t = -1.85, df = 8, P = 0.100). t0 Odabas_Buelbuel_etal_Lacerta_trilineata_Age_Growth_TR:HERPETOZOA.qxd 12.02.2019 14:56 Seite 5

Life-history traits of Lacerta trilineata BedriaGa , 1886, from Sergen, Turkey 187

N

age (yrs) / alter (a)

Fig. 2: Frequency distribution of the age in 14 male (black) and 12 female (white) Lacerta trilineata BedriaGa , 1886, from the Sergen population (Turkey). N – Number of individuals. abb. 2: Häufigkeitsverteilung des alters bei 14 Männchen (schwarz) und 12 Weibchen (weiß) von Lacerta trilineata BedriaGa , 1886 in der Population von Sergen (Türkei). N – anzahl der individuen. ) m m (

L r k

/

) m m (

L V S

age (yrs) / alter (a)

Fig. 3: Von Bertalanffy growth curves for females (open square, dashed F-line) and males (solid square, solid M-line) of Lacerta trilineata BedriaGa , 1886. For the mean SVL of the lizards at hatching 27.0 mm was taken according to JaMeS & S HiNe (1988). abb. 3: Von Bertalanffy Wachstumskurven für Weibchen (offenes Quadrat, gestrichelte F-Linie) und Männchen (gefülltes Quadrat, durchgezogene M-Linie) von Lacerta trilineata BedriaGa , 1886. die mittlere kopf-rumpf- länge (krL) der eidechsen beim Schlupf wurde nach JaMeS & S HiNe (1988) mit 27,0 mm angenommen. Odabas_Buelbuel_etal_Lacerta_trilineata_Age_Growth_TR:HERPETOZOA.qxd 12.02.2019 14:56 Seite 6

188 Y. O daBaş & U. B üLBüL & a. İ. e rOğLU & H. k Oç & M. k UrNaz & B. k UTrUP . s a ) t u

n diSCUSSiON a a e e h 6 n c i 8 l b i 9 8 i r 8 1 e t .

estimating age and growth parameters

, 0 4 8 W a a -

t 5 2

i of L. trilineata in individuals from Sergen 3 G r 7 r 1 3 e . . 0 e a b i 0 0 c 0 (NW Turkey), the authors found the oldest r a e 0 s . r d L e 0 e h age observed (females 10, males 13 years) l f a B J o a

to exceed the maximum values reported by a 0 r t n m 1 e a o

t e i üSTeL (2010; four and five years) and ka - e l . t d 0 ) n F 3 7 a n a i 2 s l

6

1 l 7 LaYCi u r et al. (2018; seven and five years). / 3 . i - u

2

. a + r 8 6 p 9 e t e

0

o

, Significant differences in longevity g y s a

p 8 t

a e

0 r ,

l and age upon arrival at sexual maturity are n 1 e 7

e

a c , 8 d g a

6 species-specific and influenced by environ - 8 r n

. 6 M L e ; 7 a

7 9

n S 7

n mental factors. GUariNO et al. (2010) found 1 7 9 L

e 3

6 o 1 3 e , h 3 . V - 2 v h 8 c life history characteristics such as maximum 0 9 5 t

S ] . , n

0 3 a 1 f . 7 n / 1 age, age at sexual maturity, longevity and o 7 ä ,

m 9 6 ]

M r :

growth rates to vary widely between the m i s y

, / e e l b

r subpopulations of one and the same species. m

a [

e m r m e STeL

in the study of ü (2010), the L. triline - t e h , f a a 1

r r J ; 9 s

[ ata specimens came from nineteen different

s

8 r 3 . 4 8 m e a 1 t 0 5 2 u populations (nine from Gallipoli Peninsula

e - t a 5 r 1 3 d y r s . . 3

n h 0 0 and 10 from Biga Peninsula) and the num - 4 3 h u c t 0

1

. a

. 1

w ber of specimens per population (1 – 4) was 0 s d 1 W o

e n

, r t r a r

9 low. Moreover, the number of juveniles was g

e

e

r , 1 l h s e 8 1 w a c t

higher than the number of adult specimens

l e i l , , u l 8 e l 0 2 7 r 9 t n a 9

( t 2

a 1 4

h in his study, which may be an explanation ,

i n / 3 . - 1

ä . 8 a n 8 j 6 m

e

0

M e , for shorter longevities than found in the

g s e d d 7 s s

n n a : e F a a s l u Sergen population. Small sample size and d

e k ] ] l r 3 s s a a e r the large proportion of juveniles may be the r 2 [ l .

e y 7 t m h r 7 0 [ l ( 1 e e

reason for the young maximum age of five 1 2 t f 1 L s e a l 1 7

2 d 5 e - g r . r 1 V a 1 aLaYCi s years found by k et al. (2018) in the 8 a e

a 8 2 s S

] . 4 d , d a 0 .

] l 1 l n

m population from edirne (17 m a.s.l.), which 1 1 c h a

m 0 t m a e

1 .

S is about 100 km from Sergen. Short , m z g

)

a L , – n e

i a f longevity was also reported for other lacer - L V e d e o S

V S [ r UariNO m

tids. For example, G et al. (2010)

S N 2

, e

[ e 1 e g b

n . g h

n found three to four years in a high alpine h o 0 2 7 t n m t a -

v ä 2 3 f u g f

l 3 5 t r 0 0 population of another green lizard, Lacerta f n n o . . r 0 m

e p e r 0 0 0 e l u -

o iNNaeUS 0 h agilis L , 1758, from italy. Their re - n m t . W r t

e

u r n 0 r s b e e r

e sults were below those reported by

e o - v t d d r f - r

o r t t p p e rOiTBerG & S MiriNa (2006) for populations a n t u e h s o l e d o n 6 c e 3 7 d i n k a n h

8 of the same species in the republic of

t 4 l

1 0 a s ) c / 4 . - t S 1 n

i

a .

s 9 f 7 r e o e

0 dagestan in the russian Federation. These ( – o z v g

– M

e e s e N a t n

b authors determined maximum longevities of

c

a e . S i 8 ) r

) t k

, i 8 i s r

. six years for males and five for females in 6 e t h e i 5 ( t t 7 s z 3

k 8 i i a r 1 e 2 L t w t s populations from lowland and submontane 7 t 4 1

ü a 1 s o a 2 V . - t 1

e r r T l 6 0 5 S S e ( s g regions (up to 600 m. a.s.l.) and seven years .

p

1 3 v 1 e . i n m n 1 t m i v 7 e

u for males and six for females in high alti - i p a t 9 t g i s s N r r

p

h i e

c tude populations (above 960 m. a.s.l). f – r c s S o k a

e e N t s LSSON HiNe e n However , O & S (1996) reported

i t d e

W r u o e .

l e r

v d 6 a

: higher longevity (11 years for males and 12

w

e

8 w v 1 n n : d l

8

o n 1 e e i for females) in L. agilis from Sweden. as a i e

1 t a t e l .

t h , i p a b b r T B l a general rule, individuals from northern lati - S a a e ) ) u

M t G T 1 1 T

/ p

e / a

i tudes and high altitudes live longer than e o n m r g P a a )

d n r r 1 individuals who live in southern latitudes e

e a e a e M d N r S P B and low altitudes ( WaPSTra et al. 2001; Odabas_Buelbuel_etal_Lacerta_trilineata_Age_Growth_TR:HERPETOZOA.qxd 12.02.2019 14:56 Seite 7

Life-history traits of Lacerta trilineata BedriaGa , 1886, from Sergen, Turkey 189 ) a / m m (

e t a r s m u t s h c a W

/

) r y / m m (

e t a r

h t w o r G age (yrs) / alter (a)

Fig. 4: Male (M) and female (F) annual growth rates (mm/yr) of Lacerta trilineata BedriaGa , 1886, from the Sergen population (Turkey). abb. 4: die Wachstumsraten (mm/a) bei Männchen (M) und Weibchen (F) von Lacerta trilineata BedriaGa , 1886 der Population von Sergen (Türkei).

rOiTBerG & S MiriNa 2006; G UariNO et al. strigata were two to three years old. as 2010; SearS & a NGiLLeTTa 2004). This observed by GUariNO et al. (2010) for L. was shown in the study by kaLaYCi et al. agilis and most likely by kaLaYCi et al. (2018) who found seven year-old individu - (2018) for L. trilineata , there was no signifi - als in Turkish L. trilineata from Bolu (1,250 cant difference in mean age of the sexes in m a.s.l.). Other longevity studies on lizards the L. trilineata sample of the present study. of the genus Lacerta reported five to six in the specimens of the Sergen popu - years for Lacerta bilineata daUdiN , 1802 lation, the age when maturity was achieved (SaiNT GirONS et al. 1989) and five years was three years (i.e., after their third hiber - for Lacerta strigata eiCHWaLd , 1831 (rOiT - nation) in both males and females of L. tri - BerG & S MiriNa 2006), which is also clear - lineata , which is in accordance with ly below the present maximum age results kaLaYCi et al. (2018) who found immatures for L. trilineata . to be one or two and the young est matures in the present study, the mean age was three years old . rOiTBerG & S MiriNa determined as 9.07 years in males and 8.42 (2006) reported that males of L. agilis attain in females of L. trilineata. kaLaYCi et al. maturity after the second, females after the (2018) reported the much lower values of third year of life. an age of maturity at 22 4.60 and 4.86 years for the males and 4.00 to 23 months was previously reported for L. and 5.44 years for the females of edirne and agilis and L. strigata (MUSkHeLiSHViLi Bolu specimens, respectively. in the study 1970; k HONYakiNa 1970, 1972; d areVSkiJ by GUariNO et al. (2010), the mean age was 1984; T erTYSHNikOV 2002), except by found as 2.4 years in males and 2.5 years in SHaMMakOV (1981) who demonstrated the females for L. agilis . rOiTBerG & S MiriNa reproduction of yearlings in L. strigata from (2006) reported that most individuals of L. south-western Turkmenistan. These results Odabas_Buelbuel_etal_Lacerta_trilineata_Age_Growth_TR:HERPETOZOA.qxd 12.02.2019 14:56 Seite 8

190 Y. O daBaş & U. B üLBüL & a. İ. e rOğLU & H. k Oç & M. k UrNaz & B. k UTrUP

indicate that the age upon arrival at sexual their earlier maturation as compared to maturity may vary among species, as well males, and the male bias in SSd appears as their sub-populations according to lati - more pronounced in low-altitude popula - tude and altitude of their geographical range tions ( rOiTBerG 2007). area. adOLPH & P OrTer (1996) suggested in the Sergen sample, the correlation that the phase of slower body growth in between age and SVL was weak for both females (associated with the first reproduc - females and males ( r = -0.675 and 0.318, tion) relative to males is shifted to younger respectively), whereas kaLaYCi et al. (2018) ages in the lowland populations, because the found significant positive correlations ( r = warmer climate accelerates growth and mat - 0.94 and 0.8) in their Bolu and edirne sam - uration. accordingly, the growth rate may ples. also, GUariNO et al. (2010) reported a be related to the age when sexual maturity is statistically significant positive relationship attained. equal age at maturity (three years) between age and SVL in L. agilis . BaU- of males and females might explain similar WeNS (1999) pointed to inconsistencies be - growth rates in both sexes of the Sergen tween increase in body size and age which population. as reported in various studies, may explain the above differences. sexual size dimorphism in many adult although the mean SVL was found lizards arises from sexual differences in the slightly higher in males (110.12 mm) than growth rates ( JOHN -a Lder & C Ox 2007; females (108.51 mm) in the Sergen sample, TOMašeVić -k OLarOV et al. 2010). Since this difference was not significantly differ - any bias in SSd was not significant, growth ent between sexes. Similarly, kaLaYCi et al. rates were not different between the sexes of (2018) observed mean SVLs of 109.57 mm L. trilineata from Sergen. and 102.5 mm in the males and 108.49 mm SMiriNa (1972) suggested that end - and 98.33 mm in the females from edirne osteal resorption may be affected by envi - and Bolu respectively. Comparative data ronmental conditions so that elevation may from Greece by Meiri (2007) indicated influence endosteal resorption. al though higher mean SVL values in an island eSTeBaN et al. (1999) found that high-alti - (Spetsai) and a mainland (Peloponesus) tude populations of Pelophylax saharicus population of L. trilineata (129.7 mm and (BOULeNGer , 1913), exhibited less resorp - 155.2 mm for males, 122.8 mm and 117.0 tion than lowland populations, CaeTaNO & for females, respectively). CaSTaNeT (1993) observed less resorption Sexual size dimorphism (SSd) is an in low-altitude populations. Similar to the important feature in the lacertid family latter findings, L. trilineata from the rather (ar riBaS 1996). That the success of large low altitude site at Sergen showed a low males over small ones in combats favors endosteal resorption rate (15.4 %). the evolution of large males may be a Unpredictable environmental events prevalent reason for male-biased SSd of ecological relevance (e.g., abnormal (a NderSON & V iTT 1990; C Ox et al. 2003). drought, humidity or changes in food avail - Most species of the Lacertidae have a male- ability) may cause unexpected double lines biased sexual size dimorphism ( kaLiONT - (JakOB et al. 2002; GUariNO & e riş Miş zOPOULOU et al. 2007). although a clear 2008; ö zdeMir et al. 2012). although the male biased sexual size dimorphism is Sergen population studied lives under stable reported for L. trilineata in the literature and moderate climate conditions and can (HerreL et al. 1996; SCHarF & M eiri utilize ample food resources, a high rate 2013) , it was insignificantly expressed (65.4 %) of double line formation was (SSd = 0.014) in the sample from Sergen, observed. However, erOğLU et al. (2017) which is in accordance with kaLaYCi et al. reported 32.5 % of individuals with double (2018) who reported insignificantly longer lines in Podarcis tauricus (P aLLaS , 1814) SVLs in males than females. rOiTBerG & from the Sergen population . it is not known SMiriNa (2006) stated that the inhibitory what factor caused the formation of double effect of reproduction on growth is stronger lines in the L. trilineata individuals of the in females than males. also, the smaller Sergen population. body size of females could be the effect of Odabas_Buelbuel_etal_Lacerta_trilineata_Age_Growth_TR:HERPETOZOA.qxd 12.02.2019 14:56 Seite 9

Life-history traits of Lacerta trilineata BedriaGa , 1886, from Sergen, Turkey 191

aCkNOWLedGMeNTS

This study was supported financially by the Unit (FYL-2017-6945). karadeniz Technical University Scientific researches

reFereNCeS

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daTe OF SUBMiSSiON: april 22, 2018 Corresponding editor: Heinz Grillitsch

aUTHOrS: Yasemin OdaBaş < [email protected] > 1) ; Ufuk BüLBüL (Corresponding author < [email protected] >) 1) ; ali İhsan erOğLU < [email protected] > 1) ; Halime kOç < koc.halime@gmail. com > 1) ; Muammer kUrNaz < [email protected]> 1) & Bilal kUTrUP < [email protected] > 1) . 1) karadeniz Technical University, Faculty of Science, department of Biology, 61080 Trabzon, Turkey.