NATURA SLOVENIAE 18(2): 47-62 SCIENTIFIC PAPER Prejeto / Received: 2.11.2016 Sprejeto / Accepted: 15.12.2016 Altitudinal distribution and habitat use of the common wall lizard Podarcis muralis (Linnaeus, 1768) and the Horvath's rock lizard Iberolacerta horvathi (Mehely, 1904) in the Kočevsko region (S Slovenia) Anamarija ŽAGAR CIBIO, Centro de Investigado em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de Vairao, 4485-661 Vairao, Portugal; E-mail: anamarija.zagar@gmail.com Abstract. The study reports on the distribution and habitat use of two lizard species in the Kočevsko region: Horvath's rock lizard and common wall lizard. Extensive sampling across an altitudinal span of 200 to 1,100 m a.s.l. in the study area revealed 62 localities with populations of both or either species. At 11 of these localities (18%) species occurred in syntopy, at 42 locations (68%) only common wall lizards were found, while at 9 locations (14%) only Horvath's rock lizards were recorded. Both species occurred across the entire altitudinal span but exhibited an opposite pattern of relative abundances and frequencies, which increased with increasing altitude in Horvath's rock lizard and with decreasing altitude in common wall lizard. The habitat use of common wall lizard was more general (it was found in seven habitat types) than Horvath's rock lizard that was registered only in three habitat types with rocks. Key words: Podarcis muralis, Iberolacerta horvathi, altitudinal gradient, habitat use, Kočevsko region, Slovenia Izvleček. Višinska razširjenost in raba prostora pozidne kuščarice Podarcis muralis (Linnaeus, 1758) in velebitske kuščarice Iberolacerta horvathi (Mehely, 1904) na Kočevskem (J Slovenija) - V raziskavi smo pridobili skupno 62 novih podatkov o razširjenosti dveh vrst kuščaric na Kočevskem: za pozidno kuščarico (Podarcis muralis) in velebitsko kuščarico (Iberolacerta horvathi). Od vseh 62 lokacij sta se vrsti pojavljali sintopično na 11 lokacijah (18 %), na 42 lokacijah (68 %) je bila zabeležena le pozidna kuščarica, na 9 lokacijah (14 %) pa izključno velebitska kuščarica. Obe vrsti sta na Kočevskem razširjeni čez celotni višinski gradient, ki se razteza med 200 m n.m. v dolini reke Kolpe do 1100 m n.m. na najvišjih vrhovih planot. Vendar pa se vrsti pojavljata v višjih relativnih gostotah na različnih nadmorskih višinah, in sicer je velebitska kuščarica pogostejša v višjih legah, pozidna kuščarica pa v nižjih. Kar zadeva rabo prostora, smo ugotovili, da je pozidna kuščarica nagnjena bolj k splošni rabi prostora kot velebitska kuščarica. Pozidna kuščarica je bila najdena v sedmih različnih habitatnih tipih, medtem ko je bila velebitska kuščarica najdena le v treh, in sicer: v naravnih in umetnih ostenjih in presvetljenem gozdu. Ključne besede: Podarcis muralis, Iberolacerta horvathi, višinski gradient, raba prostora, Kočevsko, Slovenija Biotehniška fakulteta Univerze v Ljubljani in Nacionalni inštitut za biologijo, Ljubljana, 2016 48 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER Introduction Horvath's rock lizard (Iberolacerta horvathi (Méhely, 1904)) and the common wall lizard (Podareis muraiis (Laurenti, 1768)) are small lacertid lizards that exhibit a sympatric distribution, where the distribution range of I. horvathi overlaps completely with the range of P. muralis (Sillero et al. 2014) and have similar life-history traits and ecology (heliothermy, diet, habitat use, activity, etc. (review in Žagar 2016)). Horvath's rock lizard is one of the eight species currently recognized in the genus Iberolacerta Arribas, 1997 (Mayer & Arribas 1996, Odierna et al. 1996, Arribas 1999a, 1999b, Almeida et al., 2002, Mayer & Arribas, 2003, Arribas & Carranza 2004, Carranza et al. 2004, Crochet et al. 2004, Arribas et al. 2006, Arnold et al. 2007, Galán et al. 2007, Mayer & Pavlicev 2007). Seven of these species live in the Pyrenees and in the northern and central mountains of the Iberian Peninsula, while one, Horvath's rock lizard, occurs in Central and South-eastern Europe (Gasc et al. 1997, Arnold et al. 2007, Sillero et al. 2014). Today, the distribution of Horvath's rock lizard is restricted to a relatively small range extending across the eastern Alps, pre-Alps and northern Dinaric Mountains (Bischoff 1984, Sillero et al. 2014, Žagar et al. 2014). It occurs in at least four countries: Italy (Lapini & Dolce 1983, De Luca 1989, Lapini et al. 1993, 2004, Lapini & Dal Farra 1994, Rassati 2010), Austria (Grillitsch & Tiedemann 1986, De Luca 1989, Tiedemann 1992, Grillitsch et al. 2001, Cabela et al. 2002, 2004, 2007), Slovenia (Brelih 1954, Brelih & Džukic 1974, De Luca 1989, Tome 1996, Mršic 1997, Tome 2001, Žagar et al. 2007, 2013, Žagar 2008a, 2008b, Krofel et al. 2009, Cafuta 2010) and Croatia (Méhely 1904, Karaman 1921, Arnold 1987, De Luca 1989, Tvrtkovic & Veen 2006, Kryštufek et al. 2008, Jelic 2014). It likely occurs also in Bosnia and Hercegovina, but has not been discovered there yet (Žagar et al. 2014). The report on the population found in Karwendel Gebirge in south Germany (Capula & Luiselli 1990) was strongly disputed (Bischoff 1991, Faberl & Faberl 1991, Tiedemann 1992, Capula & Luiselli 1993, Franzen et al. 1993, Schmidtler & Schmidtler 1996) and has not been re-confirmed (Cabela et al. 2004). The common wall lizard has the largest distributional range of all species of the genus Podarcis Wagler, 1830 (Gasc et al. 1997, Sillero et al. 2014). Previous studies revealed that this species originated from multiple glacial refugia (Gassert et al. 2013, Salvi et al. 2013), and multiple lineages were identified within three Mediterranean peninsulas (Iberian, Apennine and Balkan; Salvi et al. 2013). Its widespread distribution expands across most of Central Europe, the northern part of Iberian Peninsula, large parts of the Apennine and the Balkan Peninsulas and stretches to the east into North Turkey (Gasc et al. 1997, Sillero et al. 2014). The northernmost native distribution is probably still unresolved because results from a recent genetic study suggested that the population on Jersey (Channel Islands, UK) and in the Chausey archipelago may be of native origin (Michaelides et al. 2015), while in the past it has been believed that the species distribution does not extend beyond the Netherlands and that common wall lizards found in UK were introduced (Arnold 1995). In Slovenia, it is relatively common and widespread (Tome 1996, Mršic 1997, Tome 2001, Krofel et al. 2009). Syntopic populations of I. horvathi and P. muraiis have been most frequently found at low and middle altitudes in Slovenia (Brelih 1954, Žagar et al. 2007, Žagar 2008a) as well as elsewhere (Bischoff 1984, Arnold 1987, De Luca 1989, Lapini et al. 1993, Richard & Lapini 1993, Grillitsch et al. 2001, Cabela et al. 2002, 2007, Lapini et al. 2004, Rassati 2010). NATURA SLOVENIAE 18(2): 47-62 49 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER The species tandem studied here is not unique, since other Iberolacerta-Podareis species pairs with completely or partly attitudinally segregated distributional patterns have been observed also in the Iberian Peninsula, where other species of Iberolacerta occur (e.g. Moreira et al. 1999, Arribas et al. 2006, Monasterio et al. 2010). In several parts of the species range, I. horvathi populations tend to be denser at higher altitudes (e.g. De Luca 1989), while density in P. muraiis follows an opposite trend (e.g. Krofel et al. 2009, Žagar et al. 2013). In general, both species are found on rocky substrates with sparse vegetation (Arnold 1987; Arnold & Ovenden, 2004; Arnold et al. 2007; Cabela et al. 2007; Žagar et al. 2013), except that Horvath's rock lizards are more associated with rocks, while common wall lizards occur in a wider variety of different habitats (Arnold & Ovenden 2004). In this study, an extensive sampling across an altitudinal span of 200 to 1,100 m a.s.l. was conducted in the Kočevsko region in order to comprehensively recognise syntopic and allotopic occurrence, altitudinal distribution and habitat use of Horvath's rock lizard and common wall lizard. Materials and methods The study was limited to the Kočevsko region, where we collected data on the presence and relative abundances (using transect line counts) of the study species in the period between 2006 and 2015 (Fig. 1, Annex 1). Part of the data collected in the 2006-2008 period was obtained within the framework of a diploma thesis (Žagar 2008a) and was published in a study of habitat use of reptile community in the Kočevsko region (Žagar et al. 2013). Specifically, in that work we included information on the altitude, exposition, vegetation cover and habitat type of 10 reptile community members, from which we included for I. horvathi and P. murais finds from 33 localities that are also included in this analysis. The data of the 2009-2015 period was collected within the framework of a PhD thesis (Žagar 2016). Species recognition was done by either coming very close to the lizard or photographing it, to inspect the position of scales on the head or colouration of the throat region. The species can readily be identified upon either of these characteristics (Tome 1999, Arnold & Ovenden 2004). We did not distinguish sex or age of individuals in this data set (Annex 1). Locations were described as allotopic, when all visits of that location confirmed the presence of only one species, and syntopic, when both species were found at least once during the same visit. Transect line counts (Buckland et al. 1993) were conducted in one or up to three replicates (Annex 1). We summed all observations per transect and corrected for the number of times that we walked that transect (divided by number of replicates) to calculate the frequency of individuals recorded on each transect. We grouped transects into five altitudinal belts, each encompassing 200 m of altitude (Tab. 1). Thereupon, we determined the relative abundances for each altitudinal belt by summing up frequencies of individuals and dividing it by the summed length of transects inside each altitudinal belt. NATURA SLOVENIAE 18(2): 47-62 50 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER Figure 1. Map of the Kočevsko region with localities where one or both studied species, Horvath's rock lizard (Iberolacerta horvathi) and common wall lizard (Podarcis muralii), were found (N = 62) in the 2006-2015 period (see Annex 1). Slika 1. Karta razširjenosti lokacij (N = 62) na Kočevskem, kjer je bila najdena ena ali obe preučevani vrsti, velebitska kuščarica (Iberolacerta horvathi) in pozidna kuščarica (Podarcis murališ), v obdobju 2006-2015 (glej Prilogo 1). Table 1. Distribution of transects across five altitudinal belts with corresponding frequencies corrected for the replicated transect visits (No. of ind.) and calculated relative abundances of Horvath's rock lizard (Iberolacerta horvathi) and common wall lizard (Podarcis muralis) in the Kočevsko region. Tabela 1. Razporeditev transektov v petih višinskih razredih in pripadajoče frekvence osebkov z upoštevanjem števila pregledov posameznega transekta (No. of ind.) in preračunane relativne gostote (ind./km) za vrsti velebitska kuščarica (Iberolacerta horvathi) in pozidna kuščarica (Podarcis muralis) na Kočevskem. ALTITUDINAL No. of Total P. muralis I. horvathi BELT (m a.s.l.) transects distance of No. of Relative No. of Relative transects ind. abundance ind. abundance (m) (ind./km) (ind./km) 100-299 20 5430 50 9.21 3 0.55 300-499 10 3960 30 7.57 1 0.25 500-699 16 5470 26 4.75 4 0.73 700-899 6 2720 19 6.96 6 2.21 900-1099 10 2910 12 4.12 37 12.71 SUM 62 20490 NATURA SLOVENIAE 18(2): 47-62 51 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER We compared the observed frequencies of lizards (corrected for replicated transect visits, Tab. 1) in five altitudinal belts with expected frequencies (if species were equally distributed across the altitudinal span in the study area corrected for the total distance of surveyed transects in each altitudinal belt) using the Chi square test. For assessing habitat use, one of the seven different habitat types were assigned to each transect occupied by study species: (i) natural rock area, (ii) urban area, (iii) agricultural land, (iv) water bank, (v) road, (vi) artificial rock area, and (vii) open forest (Annex 1). Habitat types describe the typical areas where transects were located in the study area. Natural rock areas were naturally occurring rock cliffs and screes, urban area included backyards, cemeteries and house ruins, agricultural land included grasslands, pastures and crop fields, water banks were banks of rivers, streams or lakes, roads were gravel or asphalt roads, artificial rock areas comprised of any rocky ground or walls originating from human activities, and open forest were located in forests with <85% crown coverage (Žagar et al. 2013). We calculated the relative proportion of allotopic and syntopic populations in each habitat type to present it graphically (Fig. 2). Results Syntopic and allotopic occurrence Results represent a dataset of 62 localities, at which one or both study species, Horvath's rock lizard and common wall lizard, were found in the Kočevsko region within the 2008-2015 period (Fig. 1, Annex 1). Both species were found to occur in syntopy at 10 localities (16%), common wall lizard was allotopic at 43 locations (69.5%) and Horvath's rock lizard at 9 locations (14.5%) (Fig. 1). Syntopic populations were found across the whole altitudinal span but with the majority of them located at middle altitudes (average altitude of syntopic populations (N = 10) was 620 m a.s.l., lower quartile range = 512 m a.s.l., upper quartile range = 813 m a.s.l., Annex 1). The lowest syntopic population was found at the entrance to Bilpa cave at 200 m a.s.l. and the highest at Kameni zid at 1,061 m a.s.l. (Annex 1). Altitudinal distribution The highest relative abundances of Horvath's rock lizard were determined for the highest altitudinal belt (900-1099 m a.s.l.) and relative abundances decreased with decreasing altitude (Tab. 1, Fig. 2). The opposite pattern was observed for the common wall lizard; relative abundance was highest at the two lowest altitudinal belts (100-299 and 300-499 m a.s.l.) and decreased with increasing altitude (Tab. 1, Fig. 2). Results of the Chi square test to compare observed frequencies of lizards (Tab. 1) in five altitudinal belts with expected frequencies (if species were equally distributed across the altitudinal span, see also Methods) showed significant differences between expected and observed frequencies for both species (for Horvath's rock lizard: x2 = 145.27, df = 4, P < 0.0001); for common wall lizard: x2 = 11.31, df = 4, P = 0.0233). Results of comparing altitudes from all finds of both species also showed a statistically significant difference between the species (Horvath's rock lizard, N = 71, NATURA SLOVENIAE 18(2): 47-62 52 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER median = 948 m a.s.l., and common wall lizard, N Whitney U tests: U = 1425, Z = 10.10, P < 0.0001). 205, median = 430 m a.s.l., Mann- Figure 2. Relative abundances of the studied species across five altitudinal belts. Slika 2. Relativne gostote preučevanih vrst v petih razredih nadmorskih višin. Habitat use The study species were found in seven different habitat types; Horvath's rock lizard in three and common wall lizard in seven of them (Fig. 3, Annex 1). Allotopic populations of common wall lizard were found in all seven habitat types, syntopic populations in all three habitat types where Horvath's rock lizard was found: in natural and artificial rocky habitats and in open forests (Fig. 2). These three habitat types occurred throughout the altitudinal range (artificial rock: 203-1058 m a.s.l. (min-max), natural rock: 208-1055 m a.s.l. (min-max), open forest: 336-1118 m a.s.l. (min-max); Annex 1). On the other hand, four habitat types exclusively occupied by common wall lizard (agricultural land, road, urban area, and water banks) were mostly limited to middle and lower altitudes (agricultural land: 588 a.s.l. (one location), roads: 204-612 a.s.l. (min-max), urban area: 229-959 a.s.l. (min-max), water banks: 204-274 m a.s.l. (min-max); Annex 1). NATURA SLOVENIAE 18(2): 47-62 53 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER □ Allotopic P. muralis ■Allotopic I. horvathi nSyntopic 30,0 -, 25,0 - 20,0 - 15,0 - O 10,0 - 5,0 - ■o C « 75 k_ 3 3 O U) IS Figure 3. Relative proportion of allotopic and syntopic populations of Horvath's rock lizard (Iberolacerta horvathi) and common wall lizard (Podarcis muralis in seven different habitat types in the Kočevsko region. Slika 3. Relativni delež alotopičnih in sintopičnih populacij za vrsti velebitska kuščarica (Iberolacerta horvathi) in pozidna kuščarica (Podarcis muralis) v sedmih habitatnih tipih na Kočevskem. To check whether the changes in relative abundance with altitude can be explained by observed differences in habitat use between the species due to changes in habitat availability across the altitude, we decided to repeat the comparison of altitudes between the species by including findings in only three habitat types that occurred throughout the altitudinal range and were used by both species (artificial and natural rock areas and open forest). Results showed that in these habitat types, too, Horvath's rock lizard was found at significantly higher altitudes (N = 71, median = 948 m a.s.l.) than common wall lizard (N = 113, median = 507 m a.s.l..; Mann-Whitney U tests: U = 815, Z = 9.09, P < 0.0001). Discussion In conclusion, we have found that in the Kočevsko region, Horvath's rock lizard and the common wall lizard - two lizard species, which exhibit a high resemblance in overall body plan and many ecological characteristics - occurred across the entire altitudinal span but exhibited an opposite pattern of relative abundances and frequencies, which increased with increasing altitude in Horvath's rock lizard and with decreasing altitude in common wall lizard. The observed pattern of habitat use suggests that the common wall lizard occupies here a more diverse array of habitat types than Horvath's rock lizard. NATURA SLOVENIAE 18(2): 47-62 54 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER Jointly, the opposite pattern in relative abundances across the altitudinal span and wider use of habitat types of the common wall lizard compared to Horvath's rock lizard suggest that the species segregate to some extent in their spatial distribution and spatial niches in the Kočevsko region. However, compared to other studies of distribution of these two species (see introduction), our results showed an interestingly high altitudinal overlap in the distribution of the two species and relatively high proportion of syntopic populations (16%), as well as an overlap in three habitat types. Previous studies reported that syntopic populations of studied species occurred only in a limited zone of middle altitudes, while Horvath's rock lizard was found in allotopic populations at higher altitudes and the common wall lizard in allotopic populations at lower altitudes (De Luca 1989, Lapini et al. 1993, Richard & Lapini 1993, Lapini et al. 2004, Cabela et al. 2007, Rassati 2010). So far, this is the first observation of syntopic populations found across the entire altitudinal span of an area for these two species. This may be due to the specific topography of the Kočevsko region where altitudes do not exceed 1,100 m a.s.l. (Perko & Orožen Adamič 1998), whereas other study areas had higher altitude ranges (over 2,000 m a.s.l. in the Alpine region or up to 1,757 m a.s.l. at Velebit). Horvath's rock lizard was found there in places up to the highest peaks in Velebit (De Luca 1989) or up to 2,000 m a.s.l. in the Alps (De Luca 1989, Lapini et al. 1993, Richard and Lapini 1993, Lapini et al. 2004, Cabela et al. 2007, Rassati 2010). The found between-species differences in altitudinal distribution, not only in the Kočevsko region but elsewhere, reinforce that Horvath's rock lizard is a high-altitude species that can also occur in lowlands but on rarer occasions, while the common wall lizards' populations are most abundant in lowlands and become less dense at higher altitudes. Recent research revealed that both species also exhibit differences in physiological characteristics and that Horvath's rock lizard has adaptations that are potentially advantageous in high-altitude areas that are climatically thermally more restrict (lower yearly average air temperatures and shorter activity periods for lizards) compared to lowlands. For example, the study species differ in seasonal variation of their preferred body temperatures in terms that Horvath's rock lizard exhibits a more accurate thermoregulation across the seasons than the common wall lizard (Osojnik et al. 2013). The differences between the species were also observed on the cellular level where Horvath's rock lizard had higher potential metabolic activity than the common wall lizard, which may be advantageous in thermally restrictive environment together with more precise thermoregulatory behaviour as exhibited by Horvath's rock lizard (Žagar et al. 2015). Acknowledgements The work of AŽ was funded through a PhD grant (SFRH/BD/81324/2011) supported by Fundado para a Ciencia e Tecnologia (FCT) doctoral fellowships under the Programa Operacional Potencial Humano - Quadro de Referencia Estratégico Nacional funds from the European Social Fund and Portuguese Ministerio da Educado e Ciencia and has in 2016 received a grant of the program »Women in Science« from L'Oreal Slovenia d.o.o. and Slovenian national committee for UNESCO. I would greatly like to thank supervisors of my PhD thesis, Dr Miguel A. Careterro and Dr Al Vrezec, the reviewer of my PhD thesis Dr Boris Kryštufek and two anonymous reviewers of this article for their constructive comments that improved this work. I also greatly thank for assistance in the field to: M. Krofel, M. A. Careterro, C. Rato, N. Sillero, V. Gomes, N. Osojnik, K. Bitenc, K. Drašler, D. Marguč, V. Cafuta, L. Jamnik, M. Gojkič, students from the reptile group at RTŠB Kočevje 2014 and Medvedjak 2012, P. Bizjan, Dominiku, A. Pekolj, J. Vidmar, D. Stanojevič, J. Pintar, N. Pajk, D. Vlačic, V. Petkovska, T Delič, M. Jelenčič, N. Ražen, T. Rezek., T. Krofel, R. Portas, F. Alvares. NATURA SLOVENIAE 18(2): 47-62 55 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER References Almeida A., Rosa H., Paulo O., Crespo E. (2002): Genetic differentiation of populations of Iberian rock' lizards Iberolacerta (Iberolacerta) sensu Arribas (1999). J. Zool. Syst. Evol. Res. 40(2): 57-64. Arnold E.N. (1987): Resource partition among lacertid lizards in southern Europe. J. Zool. Ser. B 1: 739-782. Arnold H.R. (1995). Atlas of amphibians and reptiles in Britain. HMSO, London, 40 pp. Arnold E.N., Arribas O., Carranza S. (2007): Systematics of the Palaearctic and Oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae), with descriptions of eight new genera. Zootaxa 1430: 1-86. Arnold E.N., Ovenden D. (2004): A field guide to the reptiles and amphibians of Britain and Europe. Collins, London, 288 pp. Arribas O.J. (1999a): Taxonomic revision of the Iberian "Archaeolacertae" II: Diagnosis, morphology and geographic variation of 'Lacerta' aurelioiArribas, 1994. Herpetozoa 11(3/4): 155-180. Arribas O.J. (1999b): Phylogeny and relationships of the mountain lizards of Europe and Near East (Archaeolacerta Mertens, 1921, sensu lato) and their relationships among the Eurasian lacertid radiation. Russ. J. Herpetol. 6(1): 1-22. Arribas O., Carranza S. (2004): Morphological and genetic evidence of the full species status of Iberolacerta cyrenimartinezricai(Arribas, 1996). Zootaxa 634: 1-24. Arribas O., Carranza S., Odierna G. (2006): Description of a new endemic species of mountain lizard from Northwestern Spain: Iberolacerta galani sp. nov. (Squamata: Lacertidae). Zootaxa 1240: 1-55. Bischoff W. (1984): Lacerta horvathi Mehely, 1904-Kroatische Gebirgseidechse. In: Bohme W. (Ed.), Handbuch der Reptilien und Amphibien Europas 2, Aula-Verlag, Wiesbaden, pp. 265-289. Bischoff W. (1991): Lacertiden-Neunachweise für Italien und Deutschland. Die Eidechse 3: 17-19. Brelih S. (1954): Prispevek k poznavanju favne plazilcev slovenskega ozemlja. Biol. vest. 3: 128-131. Brelih S., Dzukic G. (1974): Catalogus Faunae Jugoslaviae. Ljubljana, Academia Scientarum et Artium Slovenica, 32 pp. Buckland S.T., Anderson D.R., Burnham K.P., Laake J.L. (1993): Distance sampling: estimating abundance of biological populations. Chapman and Hall, London, 446 pp. Cabela A., Grillitsch H., Tiedemann F. (2002): New records of Lacerta horvathi Mehely, 1904, in Carinthia (Austria). Herpetozoa 15: 190-192. Cabela A., Grillitsch H., Tiedemann F. (2004): Lacerta horvathi (Mehely, 1904) in the Tyrol south of the Central Alps. Herpetozoa 16(3/4): 175-176. Cabela A., Grillitsch H., Tiedemann F. (2007): Habitatpraferenzen von Podarcis muralis (Laurenti, 1768) und Iberolacerta horvathi (Mehely, 1904) bei gemeinsamem Vorkommen. Herpetozoa 19: 149-160. NATURA SLOVENIAE 18(2): 47-62 32 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER Cafuta V. (2010): First record of Hor/ath's rock lizard (Iberolacerta hor/athi) in the Kara/anke Mts., Slo/enia. Nat. Slo/. 12(1): 127-128. Capula M., Luiselli L. (1990): Notes on the occurrence and distribution of Lacerta hor/athi Méhely, 1904 in Federal Republic of Germany. Herpetol. J. 1(11): 535-536. Capula M., Luiselli L. (1993): Northernmost Hor/ath's rock lizard populations, the Vipera xanthina complex and the meaning of a correct herpetology: within what limits should authors of science act? Die Eidechse 8: 8-14. Carranza S., Arnold E.N., Amat F. (2004): DNA phylogeny of Lacerta (Iberolacerta) and other lacertine lizards (Reptilia: Lacertidae): did competition cause long-term mountain restriction? Syst. Biodi/. 2(1): 57-77. Crochet P.A., Chaline O., Surget-Groba Y., Debain C., Cheylan, M. (2004): Speciation in mountains: phylogeography and phylogeny of the rock lizards genus Iberolacerta (Reptilia: Lacertidae). Mol. Phylogenet. E/ol. 30: 860-866. De Luca N. (1989): Taxonomic and biogeographic characteristics of Hor/ath's rock lizard (Lacerta hor/athi Méhely, 1904, Lacertidae, Reptilia) in Yugoslavia. Scopolia 18: 1-48. Faberl F., Faberl H. (1991): Zwischenbericht zur Ausbreitung des Archelacerta-hor/athi- Komplexes in Bayern und seinen Nachbarländern. Die Eidechse 2(4): 8-12. Franzen M., Gruber H.-J., Heckes U. (1993): Untersuchungen zum Vorkommen der Kroatischen Gebirgseidechse (Lacerta hor/athi) im Karwendel- und Mangfallgebirge.- München. Bericht im Auftrag des Bayerischen Landesamtes für Umweltschutz. Ökokart, München, 11 pp. Galán P., Vila M., Remón N., Naveira H.F. (2007): Caracterización de las poblaciones de Iberolacerta monticola en el Noroeste ibérico mediante la combinación de datos morfológicos, ecológicos y genéticos. Munibe 25: 34-43. Gasc J.P., Cabela A., Crnobrnja-Isailovic J., Dolmen D., Grossenbacher K., Haffner P., Lescure J., Martens H., Martínez Rica J.P., Maurin H., Oliveira M.E., Sofiandiou T.S., Veith M., Zuiderwijk A. (1997): Atlas of amphibians and reptiles in Europe. Paris, Collection Patrimoines Naturels 29, Societas Europaea Herpetologica, Muséum National d'Histoire Naturelle & Service du Patrimoine Naturel, 496 pp. Gassert F., Schulte U., Husemann M., Ulrich W., Rödder D., Hochkirch A., Engel E., Meyer J., Habel J.C. (2013): From southern refugia to the northern range margin: genetic population structure of the common wall lizard, Podarcis muralis. J. Biogeogr. 40(8): 1475-1489. Grillitsch H., Tiedemann F. (1986): Lacerta hor/athi Méhely 1904 - Erstnachweis für Österreich. Ann. Naturhistor. Mus. Wien 88/89(B): 357-359. Grillitsch H., Cabela A., Tiedemann F. (2001): Lacerta hor/athi Méhely, 1904. Kroatische Gebigseidechse. In: Cabela A., Grillitsch H., Tiedemann F. (Eds.), Atlas zur Verbreitung und Ökologie der Amphibien und Reptilien in Österreich. Umwelfbundesamt-Naturhistorisches Museum Wien, Wien, pp. 481-488. Jelic D. (2014): Checklist of Croatian amphibians and reptiles with bibliography of 250 years of research. Nat. Slo/. 16(2): 17-72. Karaman S. (1921): Beiträge zur Herpetologie /on Jugoslawien. Glas. Hr. naravosl. društ. 33: 194-209. Krofel M., Cafuta V., Planinc G., Sopotnik M., Šalamun A., Tome S., Vamberger M., Žagar A. (2009): Distribution of reptiles in Slo/enia: a rewew of data collected until 2009. Nat. Slo/. 11(2): 61-99. NATURA SLOVENIAE 18(2): 47-62 57 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER Kryštufek B., Janžekovič F., Donev N.R. (2008): Elevational diversity of reptiles on two Dinaric mountains. J. Nat. Hist. 42(5-8): 399-408. Lapini L., Dolce S. (1983): Lacerta (Archaeolacerta) horvathi Méhely, 1904 in Italia; nuove stazioni per le Alpi Carniche e Giulie. Atti. Mus. Friul. Stor. Nat. 4(1982): 213-225. Lapini L., Dal Farra A. (1994): Lacerta horvathi Méhely, 1904 sulle Dolomiti (Reptilia, Lacertidae). Boll. Mus. Civ. Stor. Natur. 43: 205-208. Lapini L., Richard J., Dall'Asta A. (1993): Distribution and ecology of Lacerta horvathi Méhely, 1904 (Reptilia, Lacertidae) in North-Eastern Italy. Atti. Mus. Civ. Stor. Nat. Trieste 14: 213-230. Lapini L., Dall'Asta A., Luiselli L., Nardi P. (2004): Lacerta horvathi in Italy: a review with new data on distribution, spacing strategy and territoriality (Reptilia, Lacertidae). Boll. Zool. 71: 145-151. Mayer W., Arribas O.J. (1996): Allozyme differentiation and relationship among the Iberian-Pyrenean Mountain Lizards (Squamata: Sauria: Lacertidae). Herpetozoa 9(1/2): 57-61. Mayer W., Arribas O. (2003): Phylogenetic relationships of the European lacertid genera Archaeolacerta and Iberolacerta and their relationships to some other 'Archaeolacertae' (sensu lato) from Near East, derived from mitochondrial DNA sequences. J. Zool. Syst. Evol. Res. 41(3): 157-161. Mayer W., Pavlicev M. (2007): The phylogeny of the family Lacertidae (Reptilia) based on nuclear DNA sequences: convergent adaptations to arid habitats within the subfamily Eremiainae. Mol. Phylogenet. Evol. 44(3): 1155-1163. Méhely L. (1904): Eine neue Lacerta aus Ungarn. Ann. Hist.-Nat. Mus. Natio. Hung. 2: 362-367. Michaelides S., Cornish N., Griffiths R., Groombridge J., Zajac N., Walters G.J., Aubert F., While G.M., Uller T. (2015): Phylogeography and conservation genetics of the common wall lizard, Podarcis muralis, on islands at its northern range. PLoS ONE 10(2): e0117113. Monasterio C., Salvador A., Diaz J.A. (2010): Competition with wall lizards does not explain the alpine confinement of Iberian rock lizards: an experimental approach. Zool. 113: 275-282. Moreira P.L., Almeida A., Rosa H., Paulo O., Crespo E. (1999): Bases para a conservacao da lagartixa-da-montanha, Lacerta monticola. Estudos de Biologia e Conservagao da Natureza no 25. Grupo de Herpetologia do Centro de Biologia Ambiental, Faculdade de Ciencias da Universidade de Lisboá, Lisbon, XX pp. Mršic N. (1997): Plazilci (Reptilia) Slovenije. Zavod Republike Slovenije za šolstvo, Ljubljana, 167 pp. Odierna G., Aprea G., Arribas O.J., Capriglione T., Caputo V., Olmo E. (1996): The karyology of the Iberian rock lizards. Herpetologica 52(4): 542-550. Osojnik N., Žagar A., Carretero M.A., García-Muñoz E., Vrezec A. (2013): Ecophysiological dissimilarities of two sympatric lizards. Herpetologica 69: 445-454. Perko D., Orožen Adamič M. (1998): Slovenija: Pokrajine in ljudje. Mladinska knjiga, Ljubljana, 735 pp. Rassati G. (2010): Contributo alla conoscenza della distribuzione della Lucertola di Horvath Iberolacerta horvathi e della Lucertola dei muri Podarcis muralis in Friuli Venezia Giulia e in Veneto. Atti. Mus. Civ. Stor. Nat. Trieste 54: 133-146. Richard J., Lapini L. (1993): Trophic niche overlap in syntopic populations of Lacerta horvathi and Podarcis muralis (Reptilia, Lacertidae). Atti. Mus. Civ. Stor. Nat. Trieste 45: 151-157. NATURA SLOVENIAE 18(2): 47-62 58 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER Salvi D., Harris D.J., Kaliontzopoulou A., Carretero M.A., Pinho C. (2013): Persistence across Pleistocene ice ages in Mediterranean and extra-Mediterranean refugia: phylogeographic insights from the common wall lizard. BMC Evol. Biol. 13(1): 147. Schmidtler H., Schmidtler J. (1996): Zur Reptilienfauna der Nördlichen Kalkalpen zwischen Isar und Inn (Bayern/Tirol). Mitt. Landesverband Amphibien und Reptilienschutz (LARS) in Bayern 15(1): 1-36. Sillero N., Bonardi A., Corti C., Creemers R., Crochet P., Ficetola G.F., Kuzmin S., Lymberakis P., Pous P.D., Sindaco R., Speybroeck J., Toxopeus B., Vieites D.R., Vences M. (2014): Updated distribution and biogeography of amphibians and reptiles of Europe. Amphibia-Reptilia 35: 1-31. Tiedemann F. (1992): Zur Verbreitung der Kroatischen Gebirgseidechse, Lacerta horvathi Mehely, 1904, in Osterreich (Squamata: Sauria: Lacertidae). Herpetozoa 5(1/2): 67-69. Tome S. (1996): Pregled razširjenosti plazilcev v Sloveniji. Annales 9/'96 - Anali za istrske in mediteranske študije, Series historia naturalis 3: 217-228. Tome S. (1999): Razred: Plazilci, Reptilia. In: Kryštufek B., Janžekovič F. (Eds.), Ključ za določanje vretenčarjev Slovenije, DZS, Ljubljana, pp. 284-305. Tome S. (2001): Plazilci. In: Kryštufek B. (Ed.), Raziskava razširjenosti evropsko pomembnih vrst v Sloveniji, Končno poročilo, Prirodoslovni muzej Slovenije, Ljubljana, pp. 480-545. Tvrtkovic N., Veen P. (2006): The Dinaric Alps rare habitats and species. A nature conservation project in Croatia. Part A. Hrvatski prirodoslovni muzej, Zagreb, Royal Dutch Society for Nature Conservation, 67 pp. Žagar A. (2008a): Importance of open areas in forest landscape for reptiles (Reptilia) (in Slovene). Graduation thesis, University of Ljubljana, Ljubljana, 80 pp. Žagar A. (2008b): The lowest altitudinal record of Horvath's Rock Lizard (Iberolacerta horvathi) in Slovenia. Nat. Slov. 10(2): 59-61. Žagar A. (2016): Interspecific competition between the Common wall lizard (Podarcis muralis [Laurenti 1768]) and the Horvath's rock lizard (Iberolacerta horvathi [Mehely 1904]). PhD Doctoral dissertation, University of Ljubljana, Ljubljana, 123 pp. Žagar A., Planinc G., Krofel M. (2007): Records of Horvath's Rock Lizard (Iberolacerta horvathi) from the Notranjsko podolje region (central Slovenia). Nat. Slov. 9(2): 43-44. Žagar A., Kos I., Vrezec, A. (2013): Habitat segregation patterns of reptiles in Northern Dinaric Mountains (Slovenia). Amphibia-Reptilia 34: 263-268. Žagar A., Carretero M.A., Krofel M., Lužnik M., Podnar M., Tvrtkovic N. (2014): Reptile survey in Dinara Mountain (Croatia) revealed the southernmost known population of Horvath's rock lizard (Iberolacerta horvathi). Nat. Cro. 23(1): 235-240. Žagar A., Simčič T., Carretero M.A., Vrezec A. (2015): The role of metabolism in understanding the altitudinal segregation pattern of two potentially interacting lizards. Comp. Biochem. Physiol. Part A 179: 1-6. NATURA SLOVENIAE 18(2): 47-62 Annex 1. Locality descriptions with presence of studied species that were surveyed in the period between 2006 and 2015 in Kočevsko region and sorted by altitude. Surveys done in spring (before 30th June of the calendar year) are labelled with (1), while (2) corresponds to surveys done in summer (after 1st July of the calendar year). During each survey, we either made counts of individuals on a predetermined transect route or we only noted presence (P) or absence (/) of studied species. Maximum numbers of individuals counted are in bold. In cases when the maximum number of individuals was found more than once per location, only the first such count is in bold. Abbreviations: Alt. - average altitude, GKY and GKX = Y and X coordinates of the Gauss Kriiger coordinate system, Sp. pres. - species presence, P.m. = Common wall lizard (Podarcis mura/is) and I.h. = Horvath's rock lizard (Iberolacerta horvath!), A = Allotopic, S = Syntopic. Priloga 1. Opisi lokacij v Kočevski regiji, Iger sta bili zabeleženi ena ali obe preučevani vrsti v obdobju raziskave med leti 2006 in 2015. Lokacije so razporejene po nadmorski višini. Najdbe, ki so bile zabeležene na popisih pred 30. junijem v koledarskem letu, so označene z (1) in tiste pol. juliju v koledarskem letu z (2). Na posameznem obisku lokacije smo prešteli osebke na predhodno določenem transektu (N) ali pa smo zabeležili le prisotnost vrste (P = vrsta prisotna, / = vrsta ni prisotna). Največje število osebkov iz transektnih popisov na posamezni lokaciji je poudarjeno (v primeru ponavljanja je poudarjeno le časovno prej zabeleženo največje število osebkov). Okrajšave: Alt. - povprečna nadmorska višina, GKY in GKX = Y in X koordinate Gauss Kriigerjevega koordinatnega sistema, Sp.pres. -prisotnost vrste, P.m. = pozidna kuščarica (Podarcis muralis) in I.h. = velebitska kuščarica (Iberolacerta horvath!), A = alotopična populacija, S = sintopična populacija. g 3 Locality Alt. GKX Habitat Sp. CN CN CN N N iH O N ë o: CN CO iH O CN ë o: CN (m) GKY type pres. IB o o N ë o: iH iH CN iH CN iH iH N \P.m. 2012 (] iH N co iH O CN ë o: iH CN iH CN \P.m. 2015 (] iH IS o o CN ■ti >-! o o CN ë o: O O o CN ■ti O o CN ë o: O O O CN ■ti >-! o o CN ë o: CO o o CN ■ti >-! o o CN ë o: CO O o CN ■ti >-! iH o CN ë o: O iH O CN ■ti iH O CN ë o: iH iH O CN ■ti iH O CN ë o: iH iH O N ■ti N iH O N ■ti >-! N iH O N ■ti CO iH O CN ■ti CO iH O CN ■ti iH O CN ë o: iH O CN ■ti >-! iH O CN ë o: t iH O CN ■ti >-! m iH o CN ■ti >-! med Žlebi in Grgeljem 203 495705 42336 artificial rock A- P.m. 5 0 0 0 5 0 med Bilpo in Lazami 204 497228 41041 road A- P.m. 12 0 p / P / pri Žlebih 204 494406 42644 water bank A- P.m. 0 0 1 0 0 0 pri Gorenji Zagi 207 4936854 artificial A- 4 0 0 0 3 0 P / 1801 rock P.m. Bilpa 208 497441 40858 natural rock S 6 3 p P p P P P med Gorenjo Zago in Žlebi 215 493751 41851 road A- P.m. i 0 0 0 1 0 0 0 pri Gorenji žagi 215 493920 41899 artificial rock A- P.m. 8 0 5 0 3 0 P / Žlebi 215 494412 42675 artificial rock A- P.m. 1 0 1 0 2 0 P / med Kobilno jamo in Gorenjo Zago 219 494125 42501 artificial rock A- P.m. 1 0 0 0 2 0 P / med Sapnikom in Brsnikom 226 493567 38681 artificial rock A- P.m. 2 0 1 0 4 0 P / med Lobičem in Žlebi 228 494419 42673 natural rock A- P.m. 2 0 0 0 5 0 P / Maverc 229 493728 40033 urban area A- P.m. 2 0 2 0 2 0 P / Sapnik 230 493652 38810 urban area A- P.m. 0 0 0 0 2 0 60 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER I.h. 2015 (1) D m Oni C MA - - - - - P.m. 2015 (1) I.h. 2014 (2) D m Oni A. f1\ P.m. 2014 (2) I.h. 2014 (1) D m Oni A (A ~\ P.m. 2014 (1) I.h. 2013 (2) D »m Om "2 P.m. 2013 (2) I.h. 2013 (1) D m Oni O HA P.m. 2013 (1) I.h. 2012 (2) D »M om O AOA P.m. 2012 (2) I.h. 2012 (1) D m Oni O HA - - - P.m. 2012 (1) I.h. 2011 (2) D m Oni 1 f OA P.m. 2011 (2) I.h. 2011 (1) D m Om 1 HA - P.m. 2011 (1) I.h. 2010 (1) d m oni n fi A P.m. 2010 (1) I.h. 2008 (2) D m mno 0 0 0 0 0 o P.m. 2008 (2) I.h. 2008 (1) d m onna f-iA 0 0 0 0 0 0 0 0 0 O o o o iH o o O iH o o P.m. 2008 (1) I.h. 2007 (2) d m 0nn*7 (OA 0 0 0 0 0 0 0 O o o iH O o o O o o P.m. 200/ (2) I.h. 200/ (1) d m onn*7 0 0 ri 0 o o P.m. 200/ (1) I.h. 2006 (1) P.m. 2006 (1) 0 m 0 iH 0 iH iH O O o (N o O O iH o o o o O O o 10 Locality Alt. GKX Habitat Sp. (m) GKY type pres. pokopališče v Fari 238 491217 urban A - 37083 area P.m. med Grivacom in Gladloko 245 486880 road A - 35251 P.m. pri Gladloki 245 486880 artificial A - 35251 rock P.m. Mirtoviči 260 483050 urban A - 40666 area P.m. Srobotnik 266 484664 urban A - 39915 area P.m. Mirtoviški potok 274 482871 water A - 41106 bank P.m. nad cerkvijo v Fari 288 491251 road A - 37067 P.m. pokopališče sv. Štefan 310 491452 urban A - 37337 area P.m. pri Srobotniku 336 484894 open A - 39571 forest P.m. Kostel 348 492939 urban A - 40372 area P.m. nad vasjo Planina zapuščen 378 489201 artificial A -kamnolom 37430 rock P.m. kamnolom v Podstenah 384 491440 artificial A - 42191 rock P.m. od Planine na Planinsko steno 430 489253 road A - 37450 P.m. med Friškovo grabo in Dolenjim 435 489983 road A -Potokom 42358 P.m. pri Dolenjem Potoku 435 490044 artificial A - 42240 rock P.m. pot na Krempo 459 482701 open A - 41016 forest P.m. od vasi Podstene do sten 499 492289 artificial S 42271 rock pod Podstenami 501 490457 road A - 43051 P.m. od Planine na Planinsko steno 506 489309 road A - 37690 P.m. NATURA SLOVENIAE 18(2): 47-62 61 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... / SCIENTIFIC PAPER I.h. 2015 (1) D m oni C Q_ Q_ - - P.m. 2015 (1) I.h. 2014 (2) D m Oni A. f1\ - - P.m. 2014 (2) I.h. 2014 (1) D m oni A f 1 A - Q_ P.m. 2014 (1) I.h. 2013 (2) D »m Om "2 - Q_ P.m. 2013 (2) I.h. 2013 (1) D m oni 7 AlA - P.m. 2013 (1) I.h. 2012 (2) D »M om O AOA - Q_ P.m. 2012 (2) I.h. 2012 (1) D m oni O fl A - o o - o Q_ P.m. 2012 (1) I.h. 2011 (2) D m oni 1 fOA O P.m. 2011 (2) I.h. 2011 (1) D m Oni 1 HA - o iH - Q_ P.m. 2011 (1) I.h. 2010 (1) d on 1 n ha iH P.m. 2010 (1) I.h. 2008 (2) d m onna (o A 0 o o o iH P.m. 2008 (2) I.h. 2008 (1) d m onna f-iA 0 t 0 iH O iH o o o o o o iH O P.m. 2008 (1) I.h. 2007 (2) d m onn-7 f OA O 0 iH iH O o o o O iH P.m. 200/ (2) I.h. 200/ (1) d m onn-7 o o o P.m. 200/ (1) I.h. 2006 (1) P.m. 2006 (1) 0 M Q_ P.m. 2011 (1) I.h. 2010 (1) P m 9f)1 n m Q Q. P.m. 2010 (1) I.h. 2008 (2) n--- ^nno O M iH o o P.m. 2008 (2) I.h. 2008 (1) d m onna ha 10 O m o iH P.m. 2008 (1) I.h. 2007 (2) d m inn7 fi\ P.m. 200/ (2) I.h. 200/ (1) d m inn7 HA P.m. 200/ (1) I.h. 2006 (1) P.m. 2006 (1) m 0 iH o iH O m o iH O Locality Alt. GKX Habitat Sp. (m) GKY type pres. pred Žurgarsko steno 899 477181 artificial A - 49089 rock I.h. na Žurgarski steni 2 948 477075 natural A - 48768 rock I.h. Fridrikštajn 959 488966 urban A - 52243 area P.m. Male Bele stene 976 476572 artificial A - 61464 rock I.h. Taborska stena 991 478628 natural A - 48293 rock I.h. ob cesti pod Velikimi Belimi stenami 996 477087 artificial A - 59976 rock I.h. Kameni zid 1028 479672 natural S 52082 rock and open forest na Taborski steni 1 1029 478193 natural A - 48482 rock I.h. na Taborski steni 2 1055 478891 natural A - 48207 rock I.h. na ovinku pred Velikimi Belimi 1058 477753 artificial A -stenami 59090 rock I.h. Velike Bele stene na robu sten 1118 47744 natural A - 59648 rock and I.h. open forest NATURA SLOVENIAE 18(2): 47-62