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Zoologisch-Botanische Datenbank/Zoological-Botanical Database

Digitale Literatur/Digital Literature

Zeitschrift/Journal: Herpetozoa

Jahr/Year: 2017

Band/Volume: 29_3_4

Autor(en)/Author(s): Eroglu Ali Ihsan, Bülbül Ufuk, Kurnaz Muammer

Artikel/Article: Age structure and growth in a Turkish population of the Crimean Wall Lizard, Podarcis tauricus (PALLAS, 1814) (Squamata: Sauria: Lacertidae) 125-133 Buelbuel_etal_Podarcis_tauricus_skeletochronology:heRPeTozoa.qxd 08.02.2017 15:32 seite 1

heRPeTozoa 29 (3/4): 125 - 133 125 Wien, 30. Jänner 2017

age structure and growth in a Turkish population of the Crimean Wall Lizard, Podarcis tauricus (PaLLas , 1814) (squamata: sauria: Lacertidae)

altersstruktur und Wachstum in einer türkischen Population der Taurischen eidechse, Podarcis tauricus (PaLLas , 1814) (squamata: sauria: Lacertidae)

aLi İhsaN eRoğLU & U fUK BüLBüL & m UammeR KURNaz

KURzfassUNG

Wachstums- und entwicklungsmerkmale von Podarcis tauricus (PaLLas , 1814) wurden mit skelettochrono- logischen arbeitsmethoden untersucht. das alter in der adulten stichprobe (20 männchen, 20 Weibchen aus sergen,Westtürkei) betrug für beide Geschlechter gemeinsam betrachtet 4-10 (  = 6,68), für männchen 5-10 (  = 7,2) und Weibchen 4-10 (  = 6,15) Jahre und differierte zwischen den Geschlechtern signifikant. aus der anord- nung der Linien verlangsamten Wachstums (LaGs) ergab sich ein eintrittsalter der Geschlechtsreife von 2 bis 3 Jahren. die Kopf-Rumpflänge der echsen war positiv mit der anzahl der LaGs in den zehenknochen korreliert. der geschlechtsbedingte Größendimorphismus zugunsten der männchen war gering (sdi = -0.005) , der Wachstumskoeffizient (k) bei männchen kleiner als bei Weibchen (k ± Konfidenzintervall; männchen: 0.37 ± 0.16; Weibchen: 0.67 ± 0.22). die Wachstumsraten der Geschlechter unterschieden sich nicht voneinander. aBsTRaCT

Life-history traits of a Turkish population of Podarcis tauricus (PaLLas , 1814) were studied using the meth - ods of skeletochronology. in the adult sample (20 males and 20 females from the population of the village of sergen), the age ranged from 4-10 (  = 6.68) years for both sexes collectively, 5-10 (  = 7.2) years in males, 4-10 ( = 6.15) years in females and differed significantly between sexes. derived from the LaG (Lines of arrested Growth) configuration, the age at sexual maturity was 2-3 years in both males and females. There was a positive correlation between the lizards’ body size (sVL) and number of LaGs counted in the toe bones. sexual size dimor - phism was weakly expressed by the slightly bigger snout-vent-length of the males (sdi = -0.005). The growth coefficient (k) was lower in males than in females (k ± Confidence interval; males: 0.37 ± 0.16; females: 0.67 ± 0.22). There was no difference in growth rate between sexes. Key WoRds Reptilia: squamata: sauria; Lacertidae; Podarcis tauricus ; skeletochronology, LaG, Lines of arrested Growth, von Bertalanffy’s growth model, age structure, growth, longevity, body size, population ecology, Turkey

iNTRodUCTioN

The Crimean Wall Lizard, Podarcis CasTaNeT 1994; G UaRiNo et al. 2010; Kim tauricus (PaLLas , 1814), is found at alti - et al. 2010; aRaKeLyaN et al. 2013; KURiTa tudes of 0 to 2,350 m a.s.l. , from the Balkan & T oda 2013) and various growth parame - Peninsula in the west to the Crimean Pen- ters ( CasTaNeT & B aez 1988; h aLLiday & insula in the east, with its Turkish range area VeRReLL 1988; R oiTBeRG & s miRiNa 2006; restricted to the extreme northwest of the Kim et al. 2010). skeletochronological age country ( KaBisCh 1986; B öhme et al. 2009). estimates in lizards are more accurate and The iUCN Red List of Threatened animals timesaving as compared to age estimates classifies the species at the LC (Least using a mark-recapture approach ( smiRiNa & Concern) category since 2009 ( Böhme et al. TseLLaRiUs 1996). This is why this method 2009). was applied here to study life-history traits skeletochronology is a widely recog - such as age and size structure, longevity and nized tool for the study of the age structure age at maturation in a Turkish population of within reptile populations (J ames 1991; Podarcis tauricus (PaLLas , 1814). Buelbuel_etal_Podarcis_tauricus_skeletochronology:heRPeTozoa.qxd 08.02.2017 15:32 seite 2

126 a. İ. e RoğLU & U. B üLBüL & m. K URNaz

maTeRiaLs aNd meThods

forty adult lizards (20 females, 20 deparaffinized (at 60-70 °C for an hour, there - males) from the population at the Village of after rinsed with xylene for 7-10 minutes) sergen (41°42’20” N, 27°43’40” e, 450 m and stained with haematoxylin were put on a.s.l., Province of Kırklareli, european microscope slides and closed using entellan ® Turkey) were collected on may 09 and 10, for observation under a light microscope. during the breeding season of 2015. The age was estimated using skeletochro - specimens were caught by hand and sexed by nological methods ( CasTaNeT & s miRiNa visual examination of secondary sex charac - 1990; C asTaNeT 1994). The numbers of ters (e.g., presence of dark blue spots on the Lines of arrested Growth (LaGs) visible on margins of ventral plates and the more con - the phalangeal bone cross-sections were spicuous orange colored belly in males only). counted independently by all three authors; The active period for lizards lasts from early the observers’ results were harmonized fol - april to early october in the sergen area; the lowing discussions. in the bone, arrival at average air temperature by day was about 20 sexual maturity was assumed to be indicated °C during the sampling period. The habitat by an obvious decrease in spacing between consists of rocky ground, sparsely vegetated two subsequent LaGs ( RyseR 1998; yiLmaz by thorny plants, which provide refuges. The et al. 2005; özdemiR et al. 2012). lizard species Podarcis tauricus , P. muralis since age classes and body measure - (L aUReNTi , 1768) , Lacerta trilineata BedRi - ments (sVL) were normally distributed aGa , 1886 and Lacerta viridis (L aUReNTi , (one-sample Kolmogorov-smirnov test), 1768) live in sympatry at the study locality. parametric tests were used for comparison snout-vent length (sVL) was meas - of means (independent sample t-test) and ured to the nearest 0.01 mm using a digital correlations (spearman’s correlations test). caliper; sexual size dimorphism was quanti - all tests were processed with the iBm soft - fied by the sexual dimorphism index (sdi) ware package sPss 21.0 for Windows with as described by the formula: sdi = (mean the level of significance set at P < 0.01. sVL of the larger sex / mean sVL of the according to von Bertalanffy’s (1938) smaller sex) ± 1. The value +1 is used if growth model, the authors computed growth males are larger than females and -1 if the curves as was done in other studies (e. g., opposite is true. The result is arbitrarily James 1991; W aPsTRa et al. 2001; RoiTBeRG defined as positive if the females are larger & s miRiNa 2006; G UaRiNo et al. 2010). The and negative if the males are larger ( LoViCh authors used the general formula of the von – k (t-t ) & G iBBoNs 1992). Bertalanffy equation, Lt = L∞ (1 – e 0 ), from each lizard, the second phalange where Lt is the sVL at the age t, L∞ the of the longest (4th) toe was clipped and pre - asymptotic maximum sVL, e the base of the served in 10 % formalin solution for subse - natural logarithm, k a growth coefficient and quent histological analyses. after toe-clip - t0 the age at hatching, which is the starting ping, the lizards were released back to their point of the growth interval described in the natural habitats. The specimens were treat - present study. Because of lack of data on ed in accordance with the guidelines of the hatching size in the studied population, 26.7 ethics committee of the Karadeniz Technical mm was taken for Lt0, which is the mean University (KTü.53488718-651/2014/56). value for the species provided by iN deN after peeling the skin from the toes, BosCh & B oUT (1998). The parameters L∞ they were preserved in 10 % formalin for 2.5 (asymptotic sVL) and k, and their asymp - hours and thereafter transferred into 5 % totic confidence intervals (Ci), were esti - nitric acid solution for decalcification of the mated using the non-linear regression pro - bone. Later, the toe samples passed a tissue cedure in sPss 21.0. The growth rate ( R, processing system (Leica Tissue Processor mm/yr) was calculated as R = k (L∞ – Lt). TP1020) before they were transferred into a Growth curves were considered significant - paraffin embedding workstation (Thermo ly different if the 95 % confidence in tervals shandon B64100010). Phalangeal cross-sec - did not overlap ( James 1991; W aP sTRa et al. tions (15 µm) cut with a rotary micro tome, 2001). Buelbuel_etal_Podarcis_tauricus_skeletochronology:heRPeTozoa.qxd 08.02.2017 15:32 seite 3

Life-history traits of Podarcis tauricus (PaLLas , 1814) 127 s s u u ResULTs a c i

) r 1 4 u 7

1 a . t 0 8

8

3

7 4 - 1

7 a series of growth zones, each fol - s R . .

i 5 , 1 2 c 0 s r . lowed by a thin hematoxylinophilic line a a 0 L d

L corresponding to a winter line of arrested s o a e

P l P r

a growth were present in the phalangeal cross ( e f

t l s o m

0 u 5 8 e sections of all 40 (100 %) adult individuals

a n 0

c 1 2 6 f 6 6 / i . . - o 4

r + i 0 6

4 (fig. 1). The resorption zone did not reach e t s u a g e a l l t the first (innermost) LaG in all specimens a

u a s p i m o

c of P. tauricus , clearly seemed to be out of r p 7

a 9 n endosteal bone in all preparations and never d . e

o 8 6

g 6 L P 6 r caused difficulties for age determination. 1 0

5 . - e . V 4 n 1 0 s 0 s

o double lines were observed in 13 (32.5 %) 6 7

. v e

1 h ] specimens. The oldest female and male were 5 t a

/ f 10 years old each (fig. 2). age at maturity o m

] m r was not correlated with sex, two years in 32 , y / R

[ specimens (17 males, 15 females; 80 %) while m

e

t m

3 it was three years in 8 (3 males, 5 females;

a , r 8 . s R 5 6

2 20 %) specimens. [ 0 9 - m

6 R . . 5 u e 1 0 t t The means of sVL [mm], age [yr] and 0 s a . r h 0

c growth rate [mm/yr] were 61.16 ± 0.56, h a . t s

W 6.68 ± 0.25 and 2.48 ± 1.70, respectively for w e r

t o e r r s t r e e l e all individuals of P. tauricus (61.81 ± 0.75; g l

h 0 4 5

w a a c l 0

1 l 3 1 6 5 i / . . - a 7.20 ± 0.34; 1.78 ± 1.18 in males and 60.50 2 l e

m t r 0 6 4 u e t e h i n g

f ± 0.81; 6.15 ± 0.34; 0.96 ± 1.05 in females, ä n j m a a

d s respectively) (Table 1). d 2 e n n 1 d u .

a age was ranged from 5-10 years in

r ] 7

0

1 ] L e a 6 5 l r 0 [ 8 . - the male and 4-10 years in the female sam -

. V h y 2 0 0 r [ 0 e s 6 e 7

f t . ple. The mean age was significantly higher e l d 1 r g a 5 a a in males (independent samples t-test; t = ]

d , n ] m

a 2.210, df = 38, P = 0.033). intersexual dif - t m m

s ,

m ference in body size (snout-vent-length)

. – L , )

L n V

e was slightly male-biased (sdi = -0.005). a 1 s s V e [

7

, . s The mean sVL ( t = 1.811, df = 38, P = 8 8

[ e m

g 3

1 7 - g 6 n R . . e h 8 n a 0.245), did however not differ significantly t 1 1 h f 5 ä t g . l

f n m f 0 between sexes. There was a significant pos - e p u o l

n - r m

t itive correlation between age and sVL for e o r u n b r e e r t R o

l males (spearman’s correlation coefficient r s - e v r

f

- 0 e 4 0 p a t l d p 0

1 3 2 h 7 6 r / a u . . = 0.599, P < 0.01), females ( r = 0.499, P < - o 2

c a o 0 7 5 i e m t K d n

g 0.05) and all individuals together ( r = 0.576, s n s n

a a f t o –

s P < 0.01). Growth pattern estimated by von o v

7 N – s n

9

c Bertalanffy’s growth equation was in good . e

. i

e ) t 8 k 1

i 5 i L s s 6 8 t

i

0 fit with the data plot of age versus sVL e 7 . - , t s . V 2 k i 1 4 e a r 0 t s t z 6 6 obtained from the individuals of the study a ü s i . t

s 4 T

s e ( 5

e sample (fig. 3). for both sexes, the estimated

v l e i n t p v e i

p asymptotic sVL was lower than the maxi - t i g m r r p a i e c s

r mum sVL recorded (sVL ± Ci; males:

s asym s k

e – e

s

t

t n i t 63.77 ± 10.05 mm; females: 61.16 ± 13.40 r d e r o N e

e

e d

v .

:

w w ) mm). Growth coefficient was lower in males

w l 1 n n : l 4 a

o n 1 e 1 i e

d t than in females (k ± Ci, males: 0.37 ± 0.16; a t l . t 8 o i p a b b 1 l

s a a females: 0.67 ± 0.22). There was no differ - , m u

m

T T

/ s / p n

/

a

a e o ence in growth rate between sexes (inde - e i L n g P d L a d

n r e o e a pendent samples t-test; t = 1.300, df = 10, P

a e e P m d N m m R s ( = 0.223) (fig. 4). Buelbuel_etal_Podarcis_tauricus_skeletochronology:heRPeTozoa.qxd 08.02.2017 15:32 seite 4

128 a. İ. e RoğLU & U. B üLBüL & m. K URNaz

fig. 1: Cross section (15µm thick) of a toe bone of a five-year-old female (58.98 mm sVL) of Podarcis tauricus (PaLLas , 1814) from the sergen population in Turkey. The age was derived from the presence of five Lines of arrested Growth (LaGs 1-5) surrounding the resorption line. mC – marrow cavity; eB – endosteal bone; RL – resorption line; dL – double line; P – periphery. abb. 1: Querschnitt (15 µm dick) eines zehenknochens eines fünf Jahre alten Weibchens (58,98 mm Kopf- Rumpflänge) von Podarcis tauricus (PaLLas , 1814) aus der Population von sergen (Türkei). das alter wurde aus der anzahl von fünf Linien verlangsamten Wachstums (1-5) abgeleitet. mC – markhöhle; eB – endostaler Knochen; RL – Resorptionslinie, dL – doppelline, P – Peripherie.

N

age (yr) / alter (a)

fig. 2: frequency distribution of the age in 20 male (black) and 20 female (white) Podarcis tauricus (PaLLas , 1814), from the sergen population (Turkey). N - Number of individuals. abb. 2: die häufigkeitsverteilung des alters bei 20 männchen (schwarz) und 20 Weibchen (white) von Podarcis tauricus (PaLLas , 1814) in der Population von sergen (Türkei). N - anzahl der individuen. Buelbuel_etal_Podarcis_tauricus_skeletochronology:heRPeTozoa.qxd 08.02.2017 15:32 seite 5

Life-history traits of Podarcis tauricus (PaLLas , 1814) 129 ) m m ( f L R K

/

) m m m (

L V s

age (yr) / alter (a)

fig. 3: Von Bertalanffy growth curves for males (open square, solid m-line), females (solid square, solid f-line) and all specimens (dotted line) of Podarcis tauricus (PaLLas , 1814). solid circle and arrow point to the mean sVL of the lizards at hatching (26.7 mm acording to iN deN BosCh & B oUT 1998). abb. 3: Von Bertalanffy Wachstumskurven für männchen (offenes Quadrat, durchgezogene m-Linie), Weibchen (gefülltes Quadrat, durchgezogene f- Linie) und beide Geschlechter (Punktlinie) von Podarcis tauricus (PaLLas , 1814). Voller Kreis und Pfeil weisen auf die mittlere Kopf- Rumpflänge der eidechsen beim schlupf (26,7 mm nach iN deN BosCh & B oUT 1998) . ) 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 (yr) / alter (a)

fig. 4: male (f), female (f) and adult (a) annual growth rates (mm/yr) of Podarcis tauricus (PaLLas , 1814), from the sergen population (Turkey). abb. 4: die Wachstumsraten (mm/a) von männchen (m), Weibchen (f) und allen erwachsenen individuen (a) von Podarcis tauricus (PaLLas , 1814) in der Population von sergen (Türkei). Buelbuel_etal_Podarcis_tauricus_skeletochronology:heRPeTozoa.qxd 08.02.2017 15:32 seite 6

130 a. İ. e RoğLU & U. B üLBüL & m. K URNaz

disCUssioN

in populations living under harsh eco - and environmental factors ( GüL et al. 2014; logical conditions (e. g., unusual dry period, BüLBüL et al. 2016) on the other hand, lack of food), double lines are found in an TaRKhNishViLi & G oKheLashViLi (1996) increased proportion of specimens com - suggested that longevity might be related to pared to populations living under less stress - the type of locality rather than climate. The ful conditions. for instance, BüLBüL et al. high maximum age of ten years for males (2016) found 54.5-57.8 % in populations of and females of the sergen population of P. Darevskia parvula (L aNTz & C yRéN , 1913) tauricus is very likely due to favorable con - and aLTUNişiK et al. (2013) reported 66 % in ditions and paralleled by the findings of Eremias strauchi KessLeR , 1878. The study üzüm et al. (2014) who reported the high site at sergen is characterized by its moder - maximum longevities of nine (males) and ate climate and rich insect food resources, seven (females) years in a lowland popula - which is in accordance with the observed tion of A. boskianus . low percentage (32.5 %) of individuals in the P. tauricus lowland population showing double lines. studied, mean sVL did not differ signifi - Climate conditions as well as daily cantly between sexes, which was also ob- and annual activity can affect endosteal served in a population of D. rudis at 700 m resorption ( smiRiNa 1972; hemeLaaR 1988; a.s.l. (G üL et al. 2014). This is obviously a esTeBaN 1990; L eCLaiR 1990, a UGeRT 1992; species-specific trait in some small lacertids esTeBaN et al. 1999). Congruently, endosteal from which sexual size differences are oth - resorption was not observed in the studied erwise definitely known, e.g., A. boskianus population which inhabits a rather low ele - (üzüm et al. 2014). The significant positive vation site. a converse trend was reported correlation between age and sVL found for in Darevskia lizard species ( aRaKeLyaN et male and female P. tauricus was also al. 2013; GüL et al. 2014). observed in the sexes of A. boskianus an increase in mean age with increas - (ü züm et al. 2014) whereas, a converse ing altitude was reported for populations of trend was reported in D. rudis by GüL et al. various lizard species ( RoiTBeRG & s miRiNa (2014). 2006). although the altitude of the study according to studies by BeeBee & site was comparatively low (450 m a.s.l.), GRiffiThs (2000) and oLssoN & m adseN mean age was high (6.68 years) in the P. (2001), male lizards mature earlier than tauricus sample studied. similarly, the high females in some species. This was, howev - mean age of 6.82 years was identified in a er not observed in the studied population of low altitude (445 m a.s.l.) population of the P. tauricus. similarly, GaLáN (1996) and lacertid Acanthodactylus boskianus (d aU- RoTGeR et al. (2016) reported the absence of diN , 1802) (üzüm et al. 2014). however, sexual difference in age at maturity in Pod - also low altitude populations can show low arcis bocagei (seoaNe , 1884) and Podarcis mean age values: 2.21 years were found in lilfordi (GüNTheR , 1874). age at the arrival an island population of Podarcis siculus of sexual maturity was the second and third (RafiNesQUe -s ChmaLTz , 1810) ( Raia et al. calendar year of life in the studied popular - 2010) and 4.12 years in a lowland popula - tion of P. tauricus whereas, KaBisCh (1986) tion (50 m a.s.l) of Eremias argus PeTeRs , reported one and a half years referring to 1869 ( Kim et al. 2010). The present study NöLLeRT (1983). revealed that the mean age in the male and The adult body size depends on many female samples of P. tauricus differed sig - factors including age at maturity and long- nificantly. üzüm et al. (2014) received sim - evity ( özdemiR et al. 2012). accordingly, ilar results for A. boskianus whereas, GüL et similar age at maturity (2-3 years) and al. (2014) did not find such differences in longevity (10 years) resulted in similar adult Darevskia rudis (BedRiaGa , 1886). sVL (61.81 ± 0.75 mm; 61.50 ± 0.81 mm) Longevity depends on the duration of in both sexes of P. tauricus . the annual activity period, which is a func - in many lacertid species and other tion of altitude, latitude and other climatic saurian families, males are larger in body Buelbuel_etal_Podarcis_tauricus_skeletochronology:heRPeTozoa.qxd 08.02.2017 15:32 seite 7

Life-history traits of Podarcis tauricus (PaLLas , 1814) 131

size than females ( KaLioNTzoPoULoU et al. ronmental temperatures at higher eleva - 2007). also, in his compilation of literature tions. as expected, ssd was low in the data, KaBisCh (1986) reports male P. tauri - studied lowland population of P. tauricus . cus to attain longer sVLs than females (79.3 on the other hand, longevity and age at first mm versus 70.5 mm). in the present study reproduction were identified as the main the intersexual differences in body size determinants of ssd at an intra-specific (sVL) were insignificant, albeit slightly level ( Liao & L U 2010; L yaPKoV et al. male-biased (sdi = -0.005). also, üzüm et 2010; Liao et al. 2013, 2015). Congruently, al. (2014) and GüL et al. (2014) found male- longevity and age at sexual maturation were biased (sdi = -0.02) intersexual differences similar between the sexes in the present in body size in A. boskianus and D. rudis . a study. in many lizard species, adult ssd similar observation was made in P. siculus arises from sexual differences in the growth (VoGRiN 2005), whereas, ŽaGaR et al. (2012) rates, the larger sex growing faster than the reported female-biased ssd in P. muralis . smaller sex ( JohN -a dLeR & C ox 2007; sexual size dimorphism (ssd) may be KoLaRoV et al. 2010; üzüm et al. 2014). in affected by climate (R oiTBeRG 2007) . GüL accordance to this, the present study found et al. (2014) reported increased ssd values no significant difference between growth in a highland population (2,137 m a.s.l.) of rates for both sexes and the observed low- D. rudis , probably as a result of colder envi - level male-biased ssd.

aCKNoWLedGmeNTs

This study was supported financially by the issued by the ministry of forest and Water affairs Karadeniz Technical University scientific Researches under the number 72784983-488.04-94286. Unit (fdK-2015-5215). Capture permission was

RefeReNCes

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daTe of sUBmissioN: January 11, 2016 Corresponding editor: heinz Grillitsch

aUThoRs: ali İhsan eRoğLU < [email protected] >; Ufuk BüLBüL (Corresponding author < ufukb@ ktu.edu.tr >); muammer KURNaz < [email protected] > − Karadeniz Technical University, faculty of science, department of Biology, 61080 Trabzon, Turkey.