Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.2011 11:46 S
HeRPeTozoa 24 (1/2): 73 - 88 73 Wien, 30. Juli 2011
external morphological and osteological features of Turkish populations of Laudakia stellio (Linnaeus , 1758) (squamata: sauria: agamidae)
eidonomische und osteologische charakteristika türkischer Populationen von Laudakia stellio (Linnaeus , 1758) (squamata: sauria: agamidae)
çiğdeM GüL & M uRaT TosunoğLu
kuRzFassunG die studie vergleicht äußere morphologische und osteologische Merkmale der schleuderschwanzagame, Laudakia stellio (Linnaeus , 1758) aus sechs ihrer türkischen vorkommensgebiete. dazu wurden Merkmals- ausprägungen der Färbung und zeichnung, Beschuppung, und körperproportion von 173 individuen sowie osteo- logische Merkmalsausprägungen bei 40 Repräsentanten dieser Populationsgruppen im Hinblick auf signifikante unterschiede untersucht. nach den vorliegenden Befunden hebt sich die Population von Hatay von allen übrigen in ihrer äußeren Morphologie und osteologisch ab. aBsTRacT The authors conducted a comparative study of the morphology and osteology of six Turkish populations of the agamid lizard Laudakia stellio (L innaeus , 1758). color and pattern characteristics, pholidosis features and body measurements of 173 specimens were examined for morphological differences between the populations. Besides the analysis of external morphological traits, 40 specimens from five localities/populations were examined osteologically. in conclusion, the Hatay population differed morphologically and osteologically from the other pop - ulations. keyWoRds Reptilia: squamata: sauria: agamidae, Laudakia stellio , external morphology, osteology, subspecific vari - ation, distribution, Turkey
inTRoducTion
Laudakia stellio (Linnaeus , 1758) , et al. 2001). color-pattern, pholidosis, mor - the Hardun or Roughtail Rock agama, is phometry and molecular traits were utilized distributed in the east Mediterranean area to distinguish subspecific units ( daan from Greece across the Levant as far as 1967; B euTLeR & F RÖR 1980; G ÖçMen et al. egypt, including neighboring countries of 2003; aLMoG et al. 2005; ÖzdeMiR et al. the near east ( sindaco & J eReM čenko 2011). 2008 ). it is found in almost all of anatolia on the basis of color-pattern, pholido - except for Thrace and the Black sea regions sis and morphometric studies, all L. stellio (BaRan & a TaTüR 1998; G üL et al. 2010). populations of anatolia were previously studies of particular anatolian popu - treated as representatives of the nominate lations dealt with taxonomy ( BaRan & Ö z subspecies ( daan 1967; B aiG 1992; L evi- 1985; B aRan et al. 1989; GÖçMen et al. Ton et al. 1992). in the following years they 2003; aLMoG et al. 2005), distribution ( Ba - were assigned to two subspecies, viz. L. s. Ran 1980; M uLdeR 1995; B udak et al. stellio and L. stellio daani (BeuTLeR & 1998; Tok 1999; u ğuRTaş et al. 2000; kuM- FRÖR , 1980), again in terms of color-pattern, LuTaş et al. 2004), feeding biology ( düşen morphometry and pholidosis ( BeuTLeR & & Ö z 2001) and enteric protozoa ( üçüncü FRÖR 1980; BaRan & Ö z 1985; B aRan & Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.2011 11:46 S
74 ç. G üL & M. T osuno ğLu
Fig. 1: sampling regions of Laudakia stellio (Linnaeus , 1758) in this study. 1 - West anatolian population (Balıkesir, çanakkale, İzmir, Muğla, Manisa), 2 - south anatolian population (antalya, Mersin, adana), 3 - central anatolian population (karaman, kayseri), 4 - north anatolian population (Tokat, amasya, sinop), 5 - Hatay population, 6 - southeast anatolian population (kahramanmaraş, Gaziantep, adıyaman, şanlıurfa). abb. 1: die Gebiete, in denen die in der vorliegenden untersuchung verwendeten exemplare von Laudakia stellio (Linnaeus , 1758) gesammelt wurden. 1 - Westanatolische Population (Balıkesir, çanakkale, İzmir, Muğla, Manisa), 2 - südanatolische Population (antalya, Mersin, adana), 3 - zentralanatolische Population (karaman, kayseri), 4 - nordanatolische Population (Tokat, amasya, sinop), 5 - Hatay Population, 6 - südostanatolische Population (kahramanmaraş, Gaziantep, adıyaman, şanlıurfa).
aTaTüR 1998; G ÖçMen et al. 2003; keTe & males, and 16 to 21 in females of L. s. stel - yiLMaz 2006). in one of the latest studies lio , whereas, the number of preanal pores (aLMoG et al. 2005), however, the authors was more than 30 and the number of sub - concluded that, according to external mor - digital lamellae under the fourth toe of the phology traits, all the anatolian populations hind limb ranged from 16 to 23 in males, represented L. s. daani . and from 17 to 22 in females in L. s. daani . Regarding the color-pattern, L. s. stel - as a result of pholidosis and morphometric lio was said to have a yellow-red temporal analyses, keTe & y iLMaz (2006) detected and occipital region, in contrast with the that the number of subdigital lamellae was brown body, and an inconspicuous gular smaller and the snout-vent length longer in area displaying a dark striped pattern. in L. L. s. daani than in L. s. stellio . s. daani , however, the top and posterior knowledge about the osteology of L. portions of the head were reported to be stellio is still fragmentary. in his Phd thesis blackish grey like the rest of the body, and on the “ stellio group” within the family the gular region bears large black spots in agamidae, BaiG (1992) included some oste - most specimens ( daan 1967; B euTLeR & ological information. However, the only FRÖR 1980; a LMoG et al. 2005). aLMoG et detailed osteology of the species L. stellio al. (2005) and BeuTLeR & F RÖR (1980) was the study by eL-T ouBi (1947) about declared that there were no blue colored specimens from two localities in egypt scales present in L. s. stellio populations, (Bourg el-arab and the south of the sina whereas blue elements could be found in L. Peninsula), in which the most important s. daani . characteristics were provided. Regarding the pholidosis, aLMoG et our study aims to present an external al. (2005) stated that the number of preanal morphological and osteological examina - pores was less than 30 and the number of tion of the Laudakia stellio populations dis - subdigital lamellae underneath the fourth tributed in anatolia and to describe the vari - toe of the hind limb ranged from 17 to 20 in ation among the populations. Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.2011 11:46 S
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external morphological and osteological features of anatolian Laudakia stellio 77
MaTeRiaLs and MeTHods
in this study, 173 (77/61/35 male/ Pholidosis.- From each specimen female/juvenile) specimens were collected col lected, eight meristic pholidosis charac - from different localities of Turkey in april ters (numbers of sub- and supralabials, ven - and august, between 2006 and 2009. sexes trals, subdigital lamellae underneath the were distinguished by presence/absence of fourth toe of the hindlimb, subdigital lamel - hemipenes and callous glandular scales in lae un derneath the third finger of the fore - pre-anal and midbody position. The speci - limb, scales on the 5th whirl of the tail as of mens were generally caught by hand on, its beginning, ventral glandular scales and under or between the rocks at sunny hours preanal glandular scales) were examined and kept alive in cloth bags until the color- and subjected to interpopulation compari - pattern characteristics were registered in son. The characters that varied among pop - detail, and color photographs were taken. ulations were: number of sublabialia, num - Later, the lizards were anesthetized with ber of subdigital lamellae underneath the ether; 96% alcohol was injected into the body fourth toe of the hind limb, number of sub - cavity, and the specimens were transferred digital lamellae underneath the third finger into 70% ethanol for permanent preservation. of the fore limb and number of scales on the The specimens used in this study were regis - 5th whirl of the tail as of its beginning. tered i n the collection of the zoology section Morphometry.- using a digital at the Biology department in the Faculty of caliper sensitive to 0.01 mm for the mor - science and Letters at çanakkale onsekiz phological measurements, four body meas - Mart uni versity, çanakkale, Turkey. urements were obtained for each specimen each of the 173 specimens was as- (TL - Tail Length, svL - snout-vent Length, signed to one out of six populations/regions HL - Head Length and HW - Head Width). according to where it was caught, namely, From these the following ratios and indexes West anatolia (21/18/8 male/female/juve - were computed: TL / svL, svL / HL, HL / nile), south anatolia (14/11/15), central HW, Head index (Hi) [100 x HL / HW], anatolia (8/5/2), north anatolia (9/8/5), Head Length index (HLi) [100 x HL / svL]. Hatay (12/9/4 and southeast anatolia (13/10/1) (Fig. 1). The inventory numbers osteological examinations and GPs based coordinates and altitudes of the specimens’ record localities are present - There are many osteological studies in ed in Table 1. which the double skeletal staining method was used ( davis & G oRe 1936; G Reen 1952; Morphological examinations McLeod 1980; s unay 2005). Transparent stained skeletons formed the essence of the color-pattern.- From each speci - present study as well. double staining was men collected, 18 characters of color and applied to the transparent stained total skele - pattern (dorsal and ventral ground color; tons, i.e. cartilage and bone. after the fresh - ventral pattern and its range; color of dorsal ly euthanized specimens were skinned their and head scales; gular ground color, pattern internal organs were extracted, and the car - type and its range; coloration of the sides of cass was placed in 95% ethanol for five days the head; color of sublabialia and supralabi - and in standard laboratory acetone for two alia; symmetry, number and location of ver - days. The preparations were then left in the tebral dots; expression of transverse ledges stain prepared from a mixture of alizarin in the vertebral dots; coloration of arms and Red-s and alcian Blue 8Gs for three days legs; coloration of hands and feet) were and went through a glycerin series for one qualitatively examined . The color-pattern week each. after the cartilages were stained characteristics varying among populations blue, the bones red and the stain remnants were: dorsal ground color, ventral ground completely removed from the muscles using color, pattern of throat, symmetry and shape a mixture of glycerin and potassium hydrox - of the vertebral dots and the extent of blue ide, the preparations were preserved in coloration on the femur. 100% glycerin ( McLeod 1980). Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.2011 11:46 S
78 ç. G üL & M. T osuno ğLu
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external morphological and osteological features of anatolian Laudakia stellio 79
Fig. 3: The hyoid apparatus of anatolian Laudakia stellio (Linnaeus , 1758). The terminal portion Fig. 2: Gular pattern types in anatolian Laudakia (pharyngobranchial) of the first branchial arc was stellio (Linnaeus , 1758). a - striped pattern, withdrawn (a) in the majority of populations and B - mosaic pattern as found in the Hatay bent outwards (b) in the Hatay population. population only. abb. 3: das zungenbein anatolischer Laudakia abb. 2: kehlzeichnungstypen anatolischer Laudakia stellio (Linnaeus , 1758). das endstück (Pharyngo- stellio (Linnaeus , 1758). a – streifenmuster, branchiale) des ersten Branchialbogens wies bei der B – Mosaikmuster wie es nur in der Hatay Mehrzahl der Populationen nach hinten (a), bei der Population vorkam. Population von Hatay jedoch nach auswärts.
The authors intended to detect skeletal ments and proportions. Whether there were variation among the populations by examin - differences between male and female speci - ing both qualitative and quantitative charac - mens in terms of the pholidosis counts was teristics of skull and lower jaw (shapes of the detected by applying the non-parametric nasal, frontal, pineal foramen and hyoid ap - two-sample kolmogorov-smirnov test. To paratus; skull length, maximum and mini - understand whether there were sex-related mum skull width, rostrum length, orbit differences in terms of the body and osteo - length, frontal length and width, nasal length logical measurements, indexes and ratios, and width, mandibula length, dentale length, the parametric independent two-sample and retroarticulare length) and limb bones t test was executed. To compare the pholi - (lengths of humerus, radius, ulna, third toe of dosis features of the populations, in a first the forelimb, femur, tibia, fibula, fourth toe step the kruskall-Wallis analysis, a non - of the hindlimb). Measurements were done parametric test, was applied. and if there by using a digital caliper sensitive to 0.01 was a difference as a result of this test, the mm. vertebrae, ribs, sternum, pectoral and Mann-Whitney u test was performed to pelvic girdles were qualitatively examined. detect among which populations and at what significance level there was a difference. statistical analysis The comparison of external body and osteo - logical measurements, indexes and ratios The sPss 17.0 software package was among the populations was made by means used in the statistical evaluation of the of discriminant Function analyses. color-pattern, pholidosis, body measure -
ResuLTs and discussion
color-pattern.- The Hatay popu - portion of brownish-grey individuals (40.0 lation differed in the dorsal ground color % ) was higher than elsewhere (14.4 % ). from the other populations in that the pro - cream and orange-cream were regularly ob- Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.201111:46S 8 0
Table 5: Morphological ratios and indexes as found in several anatolian populations of Laudakia stellio (Linnaeus , 1758), according to various sources.
TL / svL – Tail Length / snout-vent Length, Hi – Head index (100•Head Length / Head Width), HLi – Head Length index (100•Head Length / snout-vent Length),
svL / HL – snout-vent Length / Head Length, HL / HW – Head Lenght / Head Width. sexual differences were not considered in various papers. Redundant presen -
tation of both Hi and HL/HW values was made for easier comparison with literature data.
Tab. 5: Morphometrische Quotienten und indizes in verschiedenen anatolischen Populationen von Laudakia stellio (Linnaeus , 1758) nach unterschiedlichen
Quellen. TL / svL – schwanzlänge / kopf-Rumpf-Länge, Hi – kopfindex (100•kopflänge / kopfbreite), HLi – kopflängen-index (100•kopflänge / kopf-Rumpf-
Länge), svL / HL – kopf-Rumpf-Länge / kopflänge, HL / HW – kopflänge / kopfbreite. Geschlechtsunterschiede wurden in zahlreichen arbeiten nicht berücksichtigt.
die redundante Präsentation von Hi und HL/HW - Werten erfolgte zum einfacheren vergleich mit den angaben in der Literatur.
Reference/Quelle Region / Population TL/svL Hi HLi svL/HL HL/HW
Present study/diese arbeit W anatolia GG 1.48 EE 1.39 GG 117.20 EE 120.15 GG 30.84 EE 28.69 GG 3.24 EE 3.48 GG 1.17 EE 1.20 ç
Present study/diese arbeit s anatolia GG 1.49 EE 1.42 GG 109.55 EE 113.56 GG 30.92 EE 29.56 GG 3.23 EE 3.38 GG 1.09 EE 1.13 .
Present study/diese arbeit central anatolia GG 1.37 EE 1.28 GG 119.14 EE 125.34 GG 31.17 EE 27.95 GG 3.20 EE 3.58 GG 1.19 EE 1.25 G ü
Present study/diese arbeit n anatolia GG 1.33 EE 1.29 GG 117.29 EE 123.56 GG 30.74 EE 28.21 GG 3.25 EE 3.55 GG 1.17 EE 1.23 L
Present study/diese arbeit Hatay GG 1.38 EE 1.35 GG 109.81 EE 117.67 GG 31.79 EE 29.44 GG 3.14 EE 3.40 GG 1.09 EE 1.17 &
Present study/diese arbeit se anatolia 1.51 1.29 113.10 118.85 30.72 28.15 3.25 3.57 1.13 1.18
GG EE GG EE GG EE GG EE GG EE M
daan (1967) W anatolia 1.44 (1.37-1.50) - GG 31.8 (30.4-32.8) - - .
T
EE 30.0 (28.8-32.3) o
aan s d (1967) s anatolia 1.43 (1.19-1.57) - GG 31.9 (29.9-33.2) - - u EE 29.5 (28.4-31.0) n o daan (1967) Hatay 1.43 (1.19-1.57) - GG 32.9 (30.8-33.5) - - ğ L
EE 31.3 (30.4-31.9) u BaRan & Ö z (1985) W anatolia 1.48 (1.42-1.56) - 30.17 (27.34-32.38) - - BaRan & Ö z (1985) s anatolia 1.45 (1.36-1.56) - 29.83 (27.16-32.50) - - BaRan & Ö z (1985) Hatay 1.49 (1.41-1.62) - 30.38 (28.95-31.14) - - BaRan & Ö z (1985) urfa 1.34 (1.25-1.45) - 29.59 (27.21-32.16) - - Budak et al. (1998) antalya L. s. daani - - - - 0.75 Tok (1999) - 1.41 - - 3.37 2.09 kuMLuTaş et al. (2004) W Taurus 1.38 - - - - GÖçMen et al. (2003) Hatay 1.42 (1.24-1.50) - 29.89 (24.8-33.5) - - aLMoG et al. (2005) L. s. daani - GG 93.4 - - - aLMoG et al. (2005) L. s. stellio - GG 85.3 - - - keTe & y iLMaz (2006) e amanos 1.44 (1.35-1.56) - 30.90 (27.23-34.04) - GG 1.12 EE 1.23 keTe & y iLMaz (2006) W amanos 1.48 (1.41-1.54) - 31.00 (29.07-32.79) - GG 1.15 EE 1.24 keTe & y iLMaz (2006) Gaziantep 1.41 (1.29-1.51) - 30.82 (28.40-33.72) - GG 1.17 EE 1.22
Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.2011 11:46 S
external morphological and osteological features of anatolian Laudakia stellio 81
served as ventral ground colors in the cent- rocky region. in summary we conclude that ral anatolian population (23.1 % ), whereas color and pattern characters alone are cer - orange-cream was rare (1.9 % ) in the oth - tainly not sufficient to distinguish between ers. a dark mosaic pattern in the gular the subspecies of L. stellio . region was frequently (55%) seen in the Pholidosis.- differences in all Hatay population, whereas longitudinal parameters were observed between males dark stripes on a whitish ground were the and females (kolmogorov-smirnov) and only pattern present in the other populations populations (kruskall-Wallis). The u test (Figure 2). The bright vertebral dots were results (Table 2) show among which popu - generally symmetrical relative to the mid- lations and at what level of significance vertebral line in the west and southeast these differences were. according to this anatolian populations (84.2 %, 76.2 %) but data, but also in the descriptive statistics, the asymmetrical in the others. The blue color numbers of subdigital lamellae were most on the femur was frequently present in west important meristic features to discriminate and central anatolian specimens (52.6 %, between populations in the species L. stel - 69.2 %), but rarer (mean: 38.5 %) in the lio. The Hatay population differed from all other populations. in all populations, some the other populations in these characters, but blue coloration was present on the scales of also from the south and central anatolian the dorsum in adults of both sexes, howev - populations in terms of the sublabialia count er, generally more pronounced in males than (Table 2, p < 0.05), and from the south, north females. and central anatolian populations in the daan (1967 ), BaRan & Ö z (1985) number of scales forming the fifth tail whirl and keTe & y iLMaz (2006) reported some (Table 2, p < 0.05). bleaching of the coloration to occur from The data obtained, as regards to the vividly colored individuals in the west to numbers of subdigital lamellae of the fore paler in the east. according to keTe & and hind limbs, were in agreement with the yiLMaz (2006) the character of having blue minimum and maximum counts by daan scales did not vary in west-eastern direction (1967), BaRan & Ö z (1985), Budak et al. of the range except that blue colored ele - (1998), Tok (1999), kuMLuTaş et al. ments were present in the west of the (2004) and keTe & y iLMaz (2006) for L. amanos Mountains and in Gaziantep, but stellio daani (Table 3). Budak et al. disappeared in parallel with an increase of a (1998) counted 20 subdigital lamellae general bleaching in the east of the amanos underneath the fourth toe of the hind limb Mountains. although having blue scales in the specimens of antalya (=south was reported as a trait of the west anatolian anatolian population in this study) and, populations (see daan 1967; BeuTLeR & thus, included them into the subspecies L. FRÖR 1980), scarce to considerable blue col - stellio daani . in all populations studied, oration was detected in all populations in the number of subdigital lamellae under - our study. kLauseWiTz (1953) observed neath the fourth toe of the hind limb was that various shades occurred in Acantho - higher in our study than indicated by cercus cyanogaster ( RüPPeLL , 1835) [ Agama aLMoG et al. (2005); this discrepancy, cyanogaster atricollis in his paper], depend - however, was not well understood. ing on the intensity of environmental fac - daan (1967) and BaRan & Ö z (1985) tors, such as light and temperature; he reported the Hatay population to have high - reported that the agamas failed to fully er numbers of subdigital lamellae than the return to their pevious colors and that initial - other populations (Table 3) which is fully ly vividly colored specimens became pale. mirrored by our results, however, the high daan (1967) detected that (i) the Harduns numbers of subdigital lamellae observed in from the Limassol region of cyprus were the Hatay population by GÖçMen et al. lead-colored grey, whereas specimens from (2003) surpassed our counts. other regions of cyprus were of lighter col - daan (1967) argued that increased ors and, (ii) that egyptian specimens caught numbers of fore and hind leg subdigital in the sandy region of the sina Peninsula lamellae as found in the Hatay individuals were of lighter color than those caught in the might point to their particular status as a Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.201111:46S 8 2
Table 6: descriptive statistics of nine proportions of the adult skull in five anatolian populations of Laudakia stellio (Linnaeus , 1758). The central anatolian
population was not included due to insufficient sample size. since differences between males and females were not significant, data was pooled. sL - skull Length,
sW - skull Width maximum, sW - skull Width minimum, RL - Rostrum Length, oL – diameter of orbit, FL - Frontal Length, FW - Frontal Width, nL - nasal max min
Length, nW - nasal Width, ML - Mandibula Length, dL - dentale Length, RaL – Retroarticular Length, se – standard error, Min – Minimum, Max – Maximum.
Tab. 6: Beschreibende statistiken von neun Proportionen des adulten schädels bei fünf anatolischen Populationen von Laudakia stellio (Linnaeus , 1758).
zentralanatolische Tiere wurden wegen unzureichender stichprobengröße nicht berücksichtigt. da die Meßwerte der Männchen von denen der Weibchen nicht sig -
nifikant verschieden waren, wurden ihre daten gemeinsam analysiert. sL – schädellänge, sW – maximale schädelbreite, sW – minimale schädelbreite, RL – max min
schnauzenlänge, oL – augenhöhlenlänge, FL – Länge des Frontale, FW – Breite des Frontale, nL – Länge des nasale, nW – Breite des nasale, ML – Länge der
Mandibel, dL – Länge des dentale, RaL – Länge des Retroarticulare, se – standardfehler, Min – Minimum, Max – Maximum.
Population W anatolia ( n = 8) n anatolia ( n = 8) s anatolia ( n = 8)
Proportion Mean se Min Max Mean se Min Max Mean se Min Max ç . sWmax / sL 0.86 0.01 0.82 0.93 0.85 0.01 0.81 0.90 0.83 0.02 0.73 0.88 G ü sWmin / sL 0.42 0.00 0.42 0.43 0.43 0.00 0.42 0.45 0.43 0.00 0.41 0.45 L
RL / sL 0.20 0.00 0.18 0.21 0.23 0.01 0.21 0.27 0.18 0.01 0.16 0.21 &
oL / sL 0.39 0.01 0.37 0.41 0.38 0.00 0.36 0.40 0.39 0.01 0.34 0.41 M
kGmin / sWmax 0.49 0.01 0.45 0.51 0.51 0.00 0.49 0.53 0.52 0.01 0.49 0.59 .
T
dL / ML 0.54 0.01 0.51 0.56 0.54 0.01 0.50 0.57 0.53 0.01 0.51 0.56 o s
RaL / ML 0.18 0.01 0.16 0.20 0.17 0.00 0.16 0.19 0.18 0.00 0.15 0.19 u n
FW / FL 0.27 0.01 0.23 0.32 0.24 0.01 0.20 0.28 0.31 0.01 0.29 0.34 o nW / nL 0.35 0.01 0.31 0.39 0.44 0.03 0.36 0.62 0.38 0.02 0.30 0.43 ğ L u
Population Hatay ( n = 8) se anatolia ( n = 8) Proportion Mean se Min Max Mean se Min Max sWmax / sL 0.85 0.01 0.83 0.89 0.82 0.01 0.79 0.87 sWmin / sL 0.43 0.01 0.41 0.46 0.42 0.00 0.40 0.43 RL / sL 0.20 0.00 0.18 0.22 0.20 0.01 0.17 0.22 oL / sL 0.39 0.01 0.37 0.43 0.38 0.00 0.37 0.41 kGmin / sWmax 0.51 0.01 0.48 0.53 0.51 0.01 0.48 0.55 dL / ML 0.52 0.00 0.50 0.54 0.54 0.00 0.52 0.56 RaL / ML 0.18 0.00 0.17 0.19 0.21 0.01 0.18 0.24 FW / FL 0.29 0.01 0.23 0.33 0.29 0.01 0.23 0.35 nW / nL 0.37 0.02 0.29 0.47 0.38 0.01 0.33 0.42
Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.201111:46S
Table 7: descriptive statistics of the length measurements of limb bones in adults of five anatolian populations of Laudakia stellio (Linnaeus , 1758). The e central anatolian population was not included due to insufficient sample size. since differences between males and females were not significant, data was pooled. x t HL – Humerus Length, RaL – Radius Length, uL – ulna Length, P3L – Length of third toe of the fore limb, FeL – Femur Length, TiL – Tibia Length, FiL – Fibula e r n
Length, P4L – Length of fourth toe of the hind limb, se – standard error, Min – Minimum, Max – Maximum. a l
Tab. 7: Beschreibende statistiken der Länge der extremitätenknochen bei adulten Tieren aus fünf anatolischen Populationen von Laudakia stellio (Linnaeus , m o
1758). zentralanatolische Tiere wurden wegen unzureichender stichprobengröße nicht berücksichtigt. da die Meßwerte der Männchen von denen der Weibchen nicht r p signifikant verschieden waren, wurden ihre daten gemeinsam analysiert. HL – Humeruslänge, RaL – Radiuslänge, uL – ulnalänge, P3L – Länge des dritten Fingers h o l der vorderextremität, FeL – Femurlänge, TiL – Tibialänge, FiL – Fibulalänge, P4L – Länge der vierten zehe der Hinterextremität, se – standardfehler, Min – Minimum, o g
Max – Maximum. i c a l
a n
Population W anatolia ( n = 8) n anatolia ( n = 8) s anatolia ( n = 8) d
Parameter Mean se Min Max Mean se Min Max Mean se Min Max o s t e HL 21.83 0.67 19.76 25.01 18.78 0.58 15.79 20.73 18.97 0.45 17.29 20.58 o l o
RaL 15.18 0.56 12.82 17.33 13.66 0.40 12.10 15.61 13.97 0.56 12.07 16.23 g i uL 16.44 0.72 13.82 20.31 14.31 0.42 12.39 15.63 14.51 0.67 12.43 17.73 c a l
P3L 16.73 0.66 14.07 19.22 14.19 0.56 12.36 16.76 15.03 0.53 12.92 16.99 f e
FeL 27.66 1.10 23.83 31.83 23.86 0.83 20.77 27.34 24.48 0.97 20.85 27.85 a t TiL 21.03 0.77 18.74 24.94 19.02 0.85 16.20 23.40 19.62 0.65 16.89 21.51 u r e
FiL 20.75 0.80 18.29 24.39 18.02 0.55 16.32 20.92 19.14 0.67 16.73 21.08 s
o
P4L 26.93 0.96 24.37 31.60 23.30 0.79 20.81 27.61 24.53 0.55 22.56 26.64 f
a n a t
Population Hatay ( n = 8) se anatolia ( n = 8) o l i
Parameter Mean se Min Max Mean se Min Max a n
L
HL 19.20 0.59 16.55 22.00 18.91 0.81 16.94 21.93 a u
RaL 15.23 0.52 13.28 17.60 13.87 0.60 12.08 16.17 d a
uL 15.34 0.41 13.90 16.86 14.54 0.49 12.90 16.19 k i P3L 15.90 0.32 14.39 16.84 14.97 0.55 13.15 16.73 a
s
FeL 24.78 0.60 22.05 27.29 24.03 1.29 20.40 28.80 t e l
TiL 20.62 0.62 17.51 22.96 19.38 0.94 16.70 22.86 l i FiL 20.32 0.48 18.00 22.07 18.94 0.95 16.13 22.47 o P4L 24.67 0.46 22.52 26.87 24.20 0.97 21.64 28.06 8 3
Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.2011 11:46 S
84 ç. G üL & M. T osuno ğLu
Fig. 4: discriminant function analysis of nine proportions of the skull (see Table 6) in five anatolian populations of Laudakia stellio (Linnaeus , 1758). The central anatolian population was not included due to insufficient sample size. abb 4: diskriminanzanalyse von neun schädelproportionen (siehe Tab. 6) bei fünf anatolischen Populationen von Laudakia stellio (Linnaeus , 1758). zentralanatolische Tiere wurden wegen unzureichender stichprobengröße nicht berücksichtigt.
Fig. 5: discriminant function analysis of eight quantitative parameters of the limb bones (see Table 7) in five anatolian populations of Laudakia stellio (Linnaeus , 1758). The central anatolian population was not included due to insufficient sample size. abb. 5: diskriminanzanalyse von acht quantitativen Merkmalen des extremitätenskeletts (siehe Tab. 7) bei fünf anatolischen Populationen von Laudakia stellio (Linnaeus , 1758). zentralanatolische Tiere wurden wegen unzureichender stichprobengröße nicht berücksichtigt. Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.2011 11:46 S
external morphological and osteological features of anatolian Laudakia stellio 85
subspecies different from daani , rather osteology.- upon the qualitative resembling the populations of Lebanon, examination of the skull, some variation Jordan and israel in these characters (Table was found regarding the shapes of the nasal 3). keTe & y iLMaz (2006) found that the in dorsal aspect, the anterior portion of the west and south anatolian as well as frontal and the pineal foramen, however, Gaziantep (=southeast anatolian) speci - more or less evenly distributed among pop - mens were close to each other in terms of ulations. on the contrary, the shape of the the number of subdigital lamellae, whereas hyoid was found to vary clearly among these values increased in the east of the populations in that the Hatay specimens amanos Mountains to resemble those in differed from all others by their longer first syria, Lebanon and israel where the sub - branchials with their ends (pharyngobran - species L. stellio stellio lived. according to chial) turned outwards (Fig. 3). our findings, the number of subdigital The results of the skull morphometry lamellae of the Hatay population differed are presented in Table 6. The discriminant from those of the west, central, southeast function analysis revealed that the north and and north anatolian populations (Table 2) southeast anatolian populations definitely by their higher values (Table 3). differed from each other in the skull propor - Ratios and indexes calculated tions (Fig. 4). The two first discriminant from morphological measurements.- functions accounted for 100% of the total differences in the Head Length index (HLi) variance at significant p values. The ratios and the ratio snout-vent Length / Head Retroarticular Length / Mandibular Length Length (svL/HL) were found between (RaL/ML) and Rostrum Length / skull males and females in all populations stud - Length (RL/sL) were the characters that ied. individual discriminant function analy - best distinguished the populations in the ses of these characters applied to adult male stepwise analysis. and female specimens did not reveal groups/ Further results of the quantitative populations that significantly differed from examination comprise the observation of each other (Table 4, p > 0.05). individual variation in the numbers of verte - our HLi data resemble the data in brae and oval fenestrae in the sternum. daan (1967 ) and BaRan & Ö z (1985), in- statistics of the limb bone measurements cluding the Hatay values given by GÖçMen are presented in Table 7. The discriminant et al. (2003) (Table 5). BaRan & Ö z (1985) function analysis of the osteological quanti - reported that the Hatay and West anatolian tative data found that the west anatolian and populations were similar in terms of HLi. Hatay populations definitely differed from our information about the ratio TL/ each other (Fig. 5). The two first discrimi - svL was similar to that reported in daan nant functions accounted for 100% of the (1967), T ok (1999), k uMLuTaş et al. (2004) total variance at significant p values. The and GÖçMen et al. (2003) (Table 5). BaRan humerus and fibula lengths were the charac - & Ö z (1985) provided the minimum to max - ters that best distinguished the populations imum intervals of this ratio of four popula - in the stepwise analysis. tions and reported that the Hatay population in summary, both qualitative and resembled the west anatolian population in quantitative osteological examinations this respect. The data we ob tained were revealed that the Hatay population differed within these intervals, except for the Hatay from all other populations studied. population, for which they were strikingly There was no postfrontal bone in the lower than the values given by BaRan & Ö z specimens under examination. BeddaRd (1985) (Table 5). (1905a) reported the absence of this bone The Head index (Hi) values of the in agamid genera, such as Amphi bolurus present study were within the range given and Chlamydosaurus but observed (1905b) for the subspecies L. stellio daani by aLMoG the presence of a very small postfrontal in et al. (2005) and our data of the ratio snout- Uro mastyx aegyptia BeLL , 1825 . eL- vent Length to Head Length (svL/HL) TouBi (1945) reported that Uromastyx did resembled the values given by Tok (1999) not have any postfrontal in some cases and (Table 5). that there was a great tendency towards the Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.2011 11:46 S
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disappearance of the postfrontal in agami - were small and in contact with each along dae. one-third of their length. eL-T ouBi (1945) all L. stellio specimens studied had observed that the basi sphenoid was tightly incisors, canines, and various molar teeth on attached to the pterygoids laterally; however, the upper and lower jaws. sMiTH (1935) the connection was found to be loose in the pointed out that the members of the family specimens we examined. agamidae differed from all the other Range area.- BaşoğLu & B aRan oriental lizards (excluding chameleons) in (1977) reported the west, south, center and that both the acrodont type of tooth attach - southeast of anatolia to represent the dis - ment and the heterodont type of dentition (i. tributional range of L. stellio in Turkey. e. dental polymorphism, such as the simul - Later, BaRan & Ö z (1985) refined their taneous presence of incisors, canines and definition in declaring the islands off the molars) were jointly observed. west anatolian coast, west, south and osteological characteristics of skull, southeast anatolia, and the coastal regions lower jaw, hyoid apparatus, vertebral col - of the south, the range area of the species, umn, ribs, sternum, pectoral and pelvic gir - but did not mention localities from north dles, and anterior and posterior limb bones of anatolia. BaRan et al. (1989) were the L. stellio were qualitatively examined by eL- first to report L. stellio from amasya and TouBi (1947). His results are in good accor - Tokat in the Black sea region, and added dance with most of our observations. some that the presence of the species there was discrepancies are given below. eL-T ouBi not natural, and the population rather (1945) reported the complete sutureless small. a new locality record was given for fusion of frontals and parietals to form a sin - L. stellio from sinop, the northernmost gle bone . However, a long suture was visible point of Turkey ( GüL et al. 2010). The between the frontals and parietals in all pop - locations of sinop, Taşova and erbaa, ulations of L. stellio under examination. eL- where the specimens of the north anatolian TouBi (1945) found the prevomers to be in population were caught belong to the low - contact with each other up to their centers. in land troughs between the Black sea and the the specimens of the presen study, prevomers shrub dry forest section.
concLusions
in conclusion, all anatolian popula - the throat, the number of sublabialia, the tions of L. stellio , with the exception of the numbers of subdigital lamellae of the fore - Hatay population, are within the range of limb and hind limb and the number of scales the subspecies L. stellio daani as defined by on their tail whirls. also osteologically, the aLMoG et al. (2005) in terms of color-pat - Hatay population was found to differ from tern, pholidosis, morphological ratios and the other Turkish populations in qualitative index values. even though the Hatay popu - and quantitative examinations. This study lation is similar to the subspecies L. s. is understood as a signpost for further analy - daani , it differs from the other Turkish pop - ses to be made in order to determine the ulations in some morphological and osteo - taxonomic statuses of the L. stellio popula - logical characters, namely, the pattern on tions in Turkey.
acknoWLedGMenTs
This study (doctoral thesis) was supported by The authors would like to extend their gratitude to the the scientific Research Foundation of the çanakkale foundation. onsekiz Mart university under Project no: 2009/37. Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis_Gül_Tosunoglu_Laudakia_stellio_TR_population_analysis.qxd 01.08.2011 11:46 S
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88 ç. G üL & M. T osuno ğLu
üçüncü , n. & G ÖçMen , B. & ü çüncü , s. yaşayan Protozoonlar ve yapıları.- Türkiye Parazitoloji (2001): dikenli keler Laudakia stellio stellio (Lin- dergisi; izmir, 25 (1): 79-83. naeus , 1758) (Reptilia, Lacertilia) rektumunda
daTe oF suBMission: april 28, 2011 corresponding editor: Heinz Grillitsch
auHoRs: çiğdem GüL (corresponding author) < [email protected] >, < gulcigdem@comu. edu.tr >, Murat TosunoğLu , çanakkale onsekiz Mart university, Faculty of arts and sciences, department of Biology, zoology section, Terzioğlu campus, 17100 çanakkale, Turkey