Biologia, Bratislava, 60/2: 189—195, 2005
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Morphological differentiation of an isolated population of the Italian wall lizard (Podarcis sicula) of the southeastern Adriatic coast Katarina Ljubisavljevic1, Stasa Tome3, Georg Dzukic1 & Milos L. Kalezic1,2 1
Institute for Biological Research “Sinisa Stankovic”, 29. Novembra 142, 11060 Belgrade, Serbia & Montenegro; e-mail:
[email protected] 2 Institute of Zoology, Faculty of Biology, Studentski trg 16, 11000 Belgrade, Serbia & Montenegro 3 Trnovska 8, SI-1000 Ljubljana, Slovenia
Abstract: Variation in a number of morphometric, meristic and qualitative characters of the Italian wall lizard (Podarcis sicula) of eastern Adriatic coast was analysed using multivariate statistics. The analysis included samples from the contiguous portion of the range of P. s. campestris and an isolated southern population from Kotor (P. s. cattaroi). All three sets of parameters revealed a similar pattern of variation in the extent of divergence of the disjunct population. It was characterised by large overall body size, while particular meristic and qualitative traits varied differently in the direction of small or great values and percentages. The fluctuating asymmetry (FA) indexes of several meristic traits in Kotor sample appeared to be significantly higher than in other samples. Key words: Morphology, Italian wall lizard, isolated population.
Introduction As an autochthon species, the Italian wall lizard (Podarcis sicula Rafinesque-Schmaltz, 1810) inhabits the Apennine Peninsula, the eastern Adriatic coast, as well as a large number of islands and islets in the Tyrrhenian and Adriatic Sea. Introduced isolated populations have been reported in Turkey, Spain, France, Switzerland, Germany, Great Britain, Greece, Libya, Tunisia, Cyprus, Israel and USA (Henle & Klaver, 1986; Corti et. al., 1997; Arnold & Ovenden, 2002). The present status of some of these colonies is uncertain (see Henle & Klaver, 1986; Arnold & Ovenden, 2002). Furthermore, there are doubts about the autochthon origin of some island populations in the Tyrrhenian Sea (Capula, 1992; Corti et al., 1997). Morphologically, the Italian wall lizard is also known as a highly inter and often intra-populationally variable polytypic species. The number of reported nominal taxa is in a state of flux (Mertens & Wermuth, 1960; Henle & Klaver, 1986; Corti et al., 1997; Arnold & Ovenden, 2002). Morphological and biochemical studies have called into question the taxonomic status of many island subspecies of P. sicula (Gorman et al., 1975; Tome, 1995). Podarcis sicula occurs along the eastern Adriatic coast mainland from the Istrian Peninsula to Neum (Veith, 1991; Schmidtler, 1999), in a more or less continuous distribution. Southwards, isolated populations have been
recorded in Dubrovnik (Croatia) and Kotor (Montenegro). The Neum – Dubrovnik and Dubrovnik-Kotor “sicula free zones” cover approximately 70 and 100 km, respectively, in a coastal line. According to previous studies (Mertens & Wermuth, 1960; Henle & Klaver, 1986), three subspecies have been identified on the Adriatic coast mainland: P. s. campestris Betta, 1857, P. s. ragusae Wettstein, 1931 (Dubrovnik) and P. s. cattaroi Taddei, 1950 (Kotor). To the best of our knowledge, comparative morphological analyses of populations from the Adriatic coast mainland are rather descriptive (Cvitanic, 1976) and usually based on few characters (Wettstein, 1926; Kramer & Mertens, 1938a, b; Taddei, 1950; Radovanovic, 1960). Moreover, the southern Adriatic distribution border, the taxonomic status, the origins of the Dubrovnik and Kotor populations and their relationship to northern populations remained unclear (Henle & Klaver, 1986). Attempts have been made to explain the presence and the origin of P. sicula populations along the Adriatic coast and on the Adriatic islands in a number of hypotheses (Wettstein, 1926, 1949; Radovanovic, 1960 and references therein; Brelih, 1963; Witte, 1965; Cvitanic, 1971; Gorman et al., 1975; Henle & Klaver, 1986; Mayer & Podnar, 2002a, 2003). The authors mainly agree about the origin of both the Istrian and Dalmatian mainland populations, assigning them to P. s campestris and stating that in this area
190 P. sicula is a relatively recent invader from the northeastern part of Italy around the Adriatic. However, there is lot of uncertainty concerning the origin of the two isolated Kotor and Dubrovnik populations. Some authors (Radovanovic, l.c.; Witte, l.c.; Henle & Klaver, l.c.) presumed their colonization was facilitated by humans, mainly on the basis of the existence of a large “sicula-free zone” between them, as well as between the Dubrovnik and mid Dalmatian populations. They excluded the hypothesis that the Dubrovnik population could be the remains of a former continuous population, which extended over the “land bridge” from southern Italy (Monte-Gargano) to mid Dalmatia during the Pleistocene (Bognar, 1987), assuming that in that case the distribution of P. sicula would be more frequent on the mid Dalmatian islands. On the basis of geological data, other authors (Wettstein, 1926; Cvitanic, 1971) supported the idea of the “land-bridge” origin of the Dubrovnik population. Cvitanic (l.c.) argued that the presence of P. sicula on the Pleistocene southern Adriatic land mass had not been strictly continuous, but rather mosaic, alternating with P. melisellensis. Over time, this resulted in the presence of the two species on different islands in the southern Adriatic. Further assumptions were that the southern populations of P. sicula were older than those in the north (Cvitanic, l.c.; Clover, 1979). In this paper, we analysed the relationship between the isolated population from Kotor, the mid Dalmatian and the northern population samples, on the basis of multivariate statistics of a number of qualitative, meristic and morphometric characters. An attempt is made to check the possible “small isolated population effect” on intra- population variability in the Kotor sample.
Material and methods Samples and measurements A total of 71 specimens from four sites along the east Adriatic coast were analysed: 1 – Kotor (Montenegro, 42◦ 26 N, 18◦ 46 E, 8 females + 1 juvenile female, 6 males + 0 juvenile males); 2 – Filipjakov (Croatia, 43◦ 58 N, 15◦ 26 E, 6+2, 8+4); 3 – Bokanjac (Croatia, 44◦ 08 N, 15◦ 15 E, 11+1, 7+0); 4 – Rovinj (Croatia, 45◦ 05 N, 13◦ 38 E, 8+0, 9+0). Females larger than 50 mm in snout-vent length were considered as mature (e.g. HENLE, 1988). The smallest adult male measured was 57 mm in body length. Specimens preserved in 70% ethanol came from the Herpetological Collection of the Slovenian Museum of Natural History, Ljubljana (sample Nos 2, 3, 4) and from dr Georg Dzukic’s herpetological collection, Institute for Biological Research, Belgrade (No. 1). Twenty-six morphometric, 21 meristic and 11 qualitative characters were examined (see Appendix). Dimensions of pileus plates and counts were taken under a stereoscopic microscope, while the body and head dimensions were taken with digital calipers to the nearest 0.01 mm. Symmetrical characteristics were taken from either side of the body, and the data processing concerned the mean of the right and
K. Ljubisavljevic et al. left values. For estimates of the fluctuating asymmetry (FA), counts for the right and left hand sides were used separately. Statistical analyses Descriptive statistics (mean, standard error, range) for quantitative traits, and percentages of states for each qualitative trait were calculated. Data are available from the first author on request. For morphometric characters, statistical analyses included only mature individuals, and the data were analysed separately for females and males. To reduce the influence of body size, the analysis was performed on residuals from a regression of each trait on Lcor. For analysis of scalation, adults and immature individuals were pooled since meristic characters do not show ontogenic variation (e.g. SARRE & DEARN, 1991), while the sexes were considered separately. A Mann-Whitney U test was used to identify differences between the sexes within population samples, while the least significant difference (LSD) test was used for average comparisons between samples for morphometric traits. Preliminary analyses revealed no significant age or sex related variations in the frequencies of qualitative traits (Yates corrected X 2 test, P > 0.05, for all comparisons). Therefore, qualitative data from all specimens of the same sample were pooled for further analyses. Multivariate statistics included separate Canonical Discriminant Analysis (CDA) for morphometric and square-root transformed (SOKAL & ROHLF, 1981) meristic traits and Correspondence Analysis (CA) for qualitative traits. The described statistics were used to identify the generalized patterns of geographic variation in these traits. In order to examine the relative levels of possible developmental instability within the isolated population of Kotor, we used the fluctuating asymmetry (FA) index calculated for meristic characters (according to procedure given in SARRE & DEARN, 1991). The unsigned differences between the left and right hand side values tested for sex related withinsample variation, appeared non-significant (Wilcoxon signed ranks test, P > 0.05, for all comparisons). Therefore, the between-sample comparisons for FA were done on pooled samples of males and females. Characters that showed significant differences were then tested for concordance with each other, by the Friedman ANOVA ranks test (SOKAL & ROHLF, 1981). The analyses were carried out using the STATISTICA (StatSoft, Inc., 1997) program package.
Results Morphometric characteristics Pronounced intersexual differences were found for a number of morphometric characters (Mann-Whitney U test, P < 0.05, in 14–22 out of 26 measures) in all samples. All measures (excluding distance between the limbs) showed higher values in males. Paired testing between the samples revealed the greatest differences for Kotor vs. Bokanjac and Rovinj sample pairs in males and Kotor vs. Rovinj in females, respectively (LSD test, P < 0.05 in 12–16 variables, respectively). In both genders no significant differences were found between two mid Dalmatian samples. The limb measurements showed the greatest differences across the paired-comparisons of samples in males (in 4–5 out of 6 pairs), while the head height was the most variable
Morphological differentiation. . .
191
Figs 1, 2. Two-dimensional scatterplot on the first and the second canonical axes (CAN) of morphometric characters in adult males (1) and females (2) of Podarcis sicula.
character in females (4 pairs). In both genders the first canonical axis discriminated the Kotor sample from all others, while the second axis separated Rovinj from the Filipjakov and Bokanjac samples (Figs 1, 2). This tendency was apparent especially in females. The isolated population sample from Kotor was discriminated on the basis of large length dimensions of the head and limbs, in both sexes (the table of pooled within-group correlations between variables and discriminant functions is available from the first author upon request). In both genders the influence of site was very high, since there were no misclassifications among samples (correctness of 100%). Meristic characters Sexual dimorphism in meristic characters appeared low and was limited to particular traits. The females from all samples had significantly higher number of ventral scales. The greatest differences were found between the Kotor and Bokanjac sample for females (MannWhitney U test, Z = −3.80 to 2.38, P < 0.05 for 9 variables) and between the Kotor and Rovinj sample for males (Mann-Whitney U test, Z = −3.06 to 2.71, P < 0.05 for 6 variables), while non-significant differences were observed between two mid Dalmatian female samples. The number of supraciliary granules, supratemporals and subdigitals revealed the greatest differences across the paired-comparisons of female samples (3–4 out of 6 pairs), while supraciliary granules and number of femoral pores were the most variable characters in males (3 pairs). The CDA correctly classified 89% of female specimens. The correctness of 100% was recorded only in the Kotor sample. In females, similarly to the CDA on morphometric variables, the first axis separated the Kotor sample from others, while the second axis separated Rovinj from mid Dalmatian samples (Fig. 3). Females from Kotor differed from the remainder in high average values for subdigital, colar and
caudal scales, and low values for supraciliary granules, supratemporal, ventral and gular scales. On the second axis, segregation between the Rovinj and mid Dalmatian samples was caused by significant differences in the subdigital and vertebral scales counts (low for the first, and higher for the second). The table of pooled withingroup correlations between the variables and discriminant functions is available from the first author upon request. There were no misclassifications among the male samples. The Kotor sample was distinctive by low counts for supraciliary granules, supratemporal and guilar scales but high mean number of femural, collar and caudal scales (Fig. 4). However, there was also a segregation of the Filipjakov sample from the other two Dalmatian samples in the upper left quadrant, mainly on the basis of low counts for femural scales. Levels of FA were significantly higher in isolated population samples from Kotor than the other ones for four characters (supraciliar, temporal, supratemporal nd femural scales; Mann-Whitney U test; Z = 1.96 to 2.32; P < 0.05). These characters were significantly correlated with each other (X 2 = 93.46, P 0.001, 3 df; Freedman’s ANOVA). Qualitative traits There was a tendency towards an increase in the percentage of indistinct massetericum (VI1, VI2), deformed occipital plate (IX1) southwards, with the decrease in the Kotor sample. The percentage of symmetrical dividing of the preocular scale (II2) increased northwards. An incomplete row of supraciliary granules (XI1, XI2) and the asymmetrical presence of indistinct massetericum (VI1) was recorded only in the Kotor sample. This traits and to a lesser extent the presence of additional pileus plates and supralabials in largest percentage (VIII1, IV1, IV2, I1, X1), caused a segregation of this sample on the second correspondence
192
K. Ljubisavljevic et al.
Figs 3, 4. Two-dimensional scatterplot on the first and the second canonical axes (CAN) of meristic characters in females (3) and males (4) of Podarcis sicula.
Fig. 5. Plot of population samples (solid circles) and states of qualitative characters (crosses) in the space of the first and the second correspondence axes (DIM) for Podarcis sicula. For numbers of samples see “Material and methods”. Abbreviations of characters are given in the Appendix.
axis (Fig. 5). Distinction of the other three samples on the opposite end of the first axis was influenced by the presence of symmetrical polymeric states of preoculars, submaxilars and supralabials (opposite to this, % II2, V2, X2 = 0 in the Kotor sample). Slight discrimination of the Bokanjac sample on the second axis is caused by the exclusive presence of additional postnasal and frenal scales on both sides of the head (III2, VII2), while only one specimen from Rovinj sample had the VII1 character state. Discussion All three sets of parameters revealed a similar pattern of variation in the extent of the divergence of the isolated southern population of Kotor. In our analysis, a tendency for larger body measurements appeared in both genders of this population, which was already found for P. sicula (Radovanovic, 1960; Henle &
Klaver, 1986). This could also be a consequence of the so-called “effect of small island populations” found in other lacertid species (e.g. Bejakovic et al., 1996). The “continental isolate” of the Kotor population acted differently from the “island isolates” of P. sicula. The Adriatic island populations of this species appeared to be smaller in overall body size in relation to mainland populations from the contiguous part of the range (Kramer & Mertens, 1938a; Radovanovic, 1960). Our results were somewhat different (in terms of absolute measure values) from those given for P. s. cattaroi (Taddei, 1950) probably due to different samples or sample sizes. However, observations about a greater number of dorsalia, and larger body proportions in the Kotor sample compared to the northern populations, are similar. Additionally, we noticed that the most conspicuous feature of lizards from the Kotor sample was the reduction of supraciliary granules. Similar trends in the extent of the larger body measurements,
Morphological differentiation. . . greater number of dorsal and temporal scales, presence of some asymmetrical states and deformities of qualitative traits and additional plates of head scutelation, were observed in the isolated population of Dubrovnik. However, since an insufficient number of individuals from this site was available to us, we did not include this sample in analyses and restrain ourselves from definite conclusions. One of the possible explanations for the differences between the isolated population (Kotor) and the populations further up north, could be the more ancient origin of the southern populations (Cvitanic, 1971; Clover, 1979). During a later period, the Kotor (and Dubrovnik) populations could have differentiated through shrinking of the range, isolation and/or adaptation to their local environment. However, bearing in mind that P. sicula is easily transported by humans (Capula, 1992; Corti et al., 1999; Ugurtas & Yildirmhan, 2000), further investigation of mainland populations might reveal alternative explanations (Thorpe, 1980). It also needs to be noted that Kotor was a major trading seaport for many centuries, and that this too could be a possible explanation for the presence of P. sicula exclusively in this city along the southern Adriatic coast. Cvitanic (1976) excluded the possibility that P. sicula could have come to the southern Adriatic mainland through expansion from the north. He believed that the climatic conditions (a high July isotherm of 26–27 ◦C) southwards from Split were not appropriate for the spread of the Italian Wall Lizard. However, records of P. sicula in Makar, Podgora (Schmidtler, 1999) and Neum (Veith, 1991), support the assumption that the “sicula free zone” towards Dubrovnik could be much smaller than previously thought. Another indication of a more southern distribution of P. sicula are high heterozygosity records for the central Dalmatian (Trogir) population (Gorman et al., 1975), which is not in agreement with the authors’ hypothesis about the Trogir population being at the periphery, “presumably near the moving front of an invasion”. They also believed that the gap towards Dubrovnik, was not a result of extinction, but of a separate invasion from Italy, which probably accounted for the southern populations. However, the distributional gap between Dubrovnik and Kotor seems to be real. Within the last ten years we have made a thorough check of the coast of the Boka Kotorska bay, but we could not confirm Hales’ (1988) data concerning the presence of P. sicula in the vicinity of Zelenika. Whatever the origin of the Kotor population, it is isolated on the southern Adriatic border of the P. sicula range. Great evolutionary potential is often attributed to periphery isolates (e.g. Nevo, 1989; Lessica & Allendorf, 1995; Garcia-Ramos & Kirkpatrick, 1997). Small disjunct lizard populations on the mainland appeared to be similar to island populations, showing indices of genetic diversity smaller than the contiguous part of areal (Mac Cullogh et al., 1997). A
193 lower genetic variability can result from the loss of variability in a relict population or from the “bottle-neck” of a founder event (Mac Cullogh et al., 1997). FA has been shown to vary inversely with heterozygosity (Sarre & Dearn, 1991) and various fitness components (Møller, 1997), and is more likely to reflect changes in developmental stability before major genetic disruption or severe environmental stress causes fitness problems within a population. In our case, higher FA indexes of several meristic traits in Kotor population may indicate that populations were exposed to either environmental or genetic stresses. However, P. sicula is believed to be an antropochorous species (Corti et al., 1999) favoured by drastic habitat alteration caused by man over the last century (e.g. deforestation, fire, agricultural activities) (Capula, 1992). As better adapted to open environments, it seemed to have competed successfully with the native congeneric species, reducing their range (e.g. P. tiliguerta in Corsica and Sardinia), or replacing them through competitive exclusion (e.g. P. muralis in the Tuscan Archipelago, Capula, 1992). Evaluations of competitive interaction between P. sicula and P. melisellensis were different (Radovanovic 1959, 1960; Raynor, 1989; Stamenkovic, 1992). Although we found a clear differentiation of the Kotor sample on the basis of three sets of morphological characteristics, molecular and biochemical analyses are needed to verify the subspecific status of this isolate. Comparative studies on a larger number of samples, both from the eastern and western Adriatic coast, together with a detailed investigation of paleological data, are also needed to clarify the picture of the historical biogeography of this species. Acknowledgements We thank Mr. S. BRELIH who collected most of the specimens from the Croatian coast. Thanks to G. SOKORAC for linguistic help. This research was supported by the Serbian Ministry of Science, Technologies and Development (grant No. 1623, “Integrative Study of Amphibians and Reptiles from the Central Balkans”). References ARNOLD, E.N. & OVENDEN, D. 2002. Field guide of reptiles and amphibians of Britain and Europe. Second Edition. Collins, London, 288 pp. BEJAKOVIC, D., ALEKSIC, I., TARASJEV, A., CRNOBRNJAISAILOVIC, J., DZUKIC, G. & KALEZIC, M.L. 1996. Lifehistory variation in a community of lacertid lizards from the Lake Skadar region (Montenegro). Herpetol. J. 6: 125–132. BOGNAR, A. 1987. Reljef i geomorfoloske osobine Jugoslavije, pp. 13–21. In: BERTIC, J. (ed.) Veliki geografski atlas Jugoslavije, SNL, Zagreb. BRELIH, S. 1963. Prispevek k poznavanju kvarnerskih kuscaric (Reptilia, Lacertidae). Biol. Vest., Ljubljana 11: 107–113. CAPULA, M. 1992. Competitive exclusion between Podarcis lizards from Tyrrhenian islands: Inference from comparative species distributions, pp. 89–93. In: KORSOS, Z. & KISS, I. (eds), Proc. Sixth Ord. Gen. meet. S. E. H., Budapest 1991.
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Appendix. Quantitative and qualitative characters used for analyses. Morphometric characters: Lcor – snout-vent length (body length from tip of snout to vent), Lcap – head length (from tip of snout to posterior margin of ear opening), Altcap – head height (at position of parietal plates), Ltcap – head width (maximum transverse distance), Lfo – mouth length (from tip of snout to corner of mouth), Ltfo – mouth width (between corners of mouth), Lpan – forelimb length (from axilla to tip of longest finger), Lpp – hindlimb length (from groin to tip of the longest toe), Ldq – length of fourth toe on hindlimb (from basis to tip), Lpil – pileus length (from tip of snout to posterior margin of occipital plate), Ltpil – pileus width (at parietal plates), Lin – length of internasal plate, Ltin – width of internasal plate, Lfr – length of frontal plate, Ltfr – width of frontal plate, Lfp – length of frontoparietal plates, Ltfp – width of frontoparietal plates, Lip – length of interparietal plate, Ltip – width of interparietal plate, Lpa – length of parietal plates, Ltpa – width of parietal plates, Lan – length of anal plate, Ltan – width of anal plate, Doa – orbit to ear distance (from posterior margin of eye to anterior edge of ear opening), Pap – distance between fore and hind limbs (from posterior margin of forelimb insertion to anterior margin of hindlimb insertion), Lspf – length of the row of femoral pores. Meristic characters: soc – supraocular scales, cil – supraciliary scales, gra – supraciliary granules, poc-postocular scales, tmp – temporal scales, tsp – scales between masseteric and supratemporals, stm – supratemporal scales, slb – supralabial scales, san – supralabial scales anterior to subocular, sub – sublabial scales, smx – submaxilar scales, gul – gular scales along the throat midline, col – large scales in the collar, vnt – inner ventral scales counted longitudinally, dor – dorsal scales around mid-body, pra – praeanal scales
surrounding anteriorly the anal plate, fpo – femoral pores, fem – femural scales, sdg – lamellar scales under the fourth toe, ver – dorsal scales counted longitudinally along the occipital stripe from occipital plate to the base of the tail, scd – caudal scales in the 13th coil counting from the vent. Qualitative characters: (I) PF – additional plate between the praefrontals: 0 – absent, 1 – present; (II) PRO – praeocular scale: 0 – undivided, 1 – divided on one head side, 2 – divided on both the right and left sides; (III) FF – additional scale between frenal and frenoocular scale: 0 – absent, 1 – present on one head side, 2 – present on both the right and left sides; (IV) PS – additional scale between the first postocular (1POC) and the last supraocular (4SOC) scale: 0 – absent, 1 – present on one head side, 2 – present on both the right and left sides; (V) SM – polymeric states of submaxilars (having more elements than in the typical pattern, i.e. adding, partition or insertion of scales): 0 – absent, 1 – present on one side, 2 – present on both the right and left sides; (VI) MS – masseteric plate: 0 – distinct, 1 – indistinct on one head side, 2 – indistinct on both the right and left sides; (VII) PN – postnasal scale: 0 – undivided, 1 – divided on one head side, 2 – divided on both the right and left sides; (VIII) IN – insertion of additional plates in internasal: 0 – absent, 1 – present; (IX) OC – occipital plate: 0 – normal; 1 – deformed; (X) SL – polymeric states of supralabials (having more elements than in the typical pattern, i.e. adding, partition or insertion of scales): 0 – absent, 1 – present on one head side, 2 – present on both the right and left sides; (XI) GR – row of supraciliary granules: 0 – complete; 1 – incomplete on one head side, 2 – incomplete on both the right and left sides.
