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1 NATURA LOVENIAE 8(): 7-6 rejeto Received:..6 CIENTIFIC AER prejeto Accepted: 5..6 Altitudinal distribution and habitat use of the common wall lizard odarcis muralis (Linnaeus, 768) and the Horvath s lizard Iberolacerta horvathi (Méhely, 9) in the Kočevsko region ( lovenia) Anamarija ŽAGAR CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do orto, Campus Agrário de Vairão, Vairão, ortugal; anamarija.zagar@gmail.com Abstract. The study reports on the distribution and habitat use of two lizard species in the Kočevsko region: Horvath s lizard and common wall lizard. Extensive sampling across an altitudinal span of to, m a.s.l. in the study area revealed 6 localities with populations of both or either species. At of these localities (8%) species occurred in syntopy, at locations (68%) only common wall lizards were found, while at 9 locations (%) only Horvath s 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 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 lizard that was registered only in three habitat types with s. Key words: odarcis muralis, Iberolacerta horvathi, altitudinal gradient, habitat use, Kočevsko region, lovenia Izvleček. Višinska razširjenost in raba prostora pozidne kuščarice odarcis muralis (Linnaeus, 758) in velebitske kuščarice Iberolacerta horvathi (Méhely, 9) na Kočevskem (J lovenija) V raziskavi smo pridobili skupno 6 novih podatkov o razširjenosti dveh vrst kuščaric na Kočevskem: za pozidno kuščarico (odarcis muralis) in velebitsko kuščarico (Iberolacerta horvathi). Od vseh 6 lokacij sta se vrsti pojavljali sintopično na lokacijah (8 %), na lokacijah (68 %) je bila zabeležena le pozidna kuščarica, na 9 lokacijah ( %) pa izključno velebitska kuščarica. Obe vrsti sta na Kočevskem razširjeni čez celotni višinski gradient, ki se razteza med m n.m. v dolini reke Kolpe do 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. ozidna 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: odarcis muralis, Iberolacerta horvathi, višinski gradient, raba prostora, Kočevsko, lovenija Biotehniška fakulteta Univerze v Ljubljani in Nacionalni inštitut za biologijo, Ljubljana, 6

2 8 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER Introduction Horvath s lizard (Iberolacerta horvathi (Méhely, 9)) and the common wall lizard (odarcis muralis (Laurenti, 768)) are small lacertid lizards that exhibit a sympatric distribution, where the distribution range of I. horvathi overlaps completely with the range of. muralis (illero et al. ) and have similar life-history traits and ecology (heliothermy, diet, habitat use, activity, etc. (review in Žagar 6)). Horvath s lizard is one of the eight species currently recognized in the genus Iberolacerta Arribas, 997 (Mayer & Arribas 996, Odierna et al. 996, Arribas 999a, 999b, Almeida et al.,, Mayer & Arribas,, Arribas & Carranza, Carranza et al., Crochet et al., Arribas et al. 6, Arnold et al. 7, Galán et al. 7, Mayer & avlicev 7). even of these species live in the yrenees and in the northern and central mountains of the Iberian eninsula, while one, Horvath s lizard, occurs in Central and outh-eastern Europe (Gasc et al. 997, Arnold et al. 7, illero et al. ). Today, the distribution of Horvath s lizard is restricted to a relatively small range extending across the eastern Alps, pre-alps and northern Dinaric Mountains (Bischoff 98, illero et al., Žagar et al. ). It occurs in at least four countries: Italy (Lapini & Dolce 98, De Luca 989, Lapini et al. 99,, Lapini & Dal Farra 99, Rassati ), Austria (Grillitsch & Tiedemann 986, De Luca 989, Tiedemann 99, Grillitsch et al., Cabela et al.,, 7), lovenia (Brelih 95, Brelih & Džukić 97, De Luca 989, Tome 996, Mršić 997, Tome, Žagar et al. 7,, Žagar 8a, 8b, Krofel et al. 9, Cafuta ) and Croatia (Méhely 9, Karaman 9, Arnold 987, De Luca 989, Tvrtković & Veen 6, Kryštufek et al. 8, Jelić ). It likely occurs also in Bosnia and Hercegovina, but has not been discovered there yet (Žagar et al. ). The report on the population found in Karwendel Gebirge in south Germany (Capula & Luiselli 99) was strongly disputed (Bischoff 99, Faberl & Faberl 99, Tiedemann 99, Capula & Luiselli 99, Franzen et al. 99, chmidtler & chmidtler 996) and has not been re-confirmed (Cabela et al. ). The common wall lizard has the largest distributional range of all species of the genus odarcis Wagler, 8 (Gasc et al. 997, illero et al. ). revious studies revealed that this species originated from multiple glacial refugia (Gassert et al., alvi et al. ), and multiple lineages were identified within three Mediterranean peninsulas (Iberian, Apennine and Balkan; alvi et al. ). Its widespread distribution expands across most of Central Europe, the northern part of Iberian eninsula, large parts of the Apennine and the Balkan eninsulas and stretches to the east into North Turkey (Gasc et al. 997, illero et al. ). 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. 5), 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 995). In lovenia, it is relatively common and widespread (Tome 996, Mršić 997, Tome, Krofel et al. 9). yntopic populations of I. horvathi and. muralis have been most frequently found at low and middle altitudes in lovenia (Brelih 95, Žagar et al. 7, Žagar 8a) as well as elsewhere (Bischoff 98, Arnold 987, De Luca 989, Lapini et al. 99, Richard & Lapini 99, Grillitsch et al., Cabela et al., 7, Lapini et al., Rassati ). NATURA LOVENIAE 8(): 7-6

3 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER 9 The species tandem studied here is not unique, since other Iberolacerta-odarcis species pairs with completely or partly attitudinally segregated distributional patterns have been observed also in the Iberian eninsula, where other species of Iberolacerta occur (e.g. Moreira et al. 999, Arribas et al. 6, Monasterio et al. ). In several parts of the species range, I. horvathi populations tend to be denser at higher altitudes (e.g. De Luca 989), while density in. muralis follows an opposite trend (e.g. Krofel et al. 9, Žagar et al. ). In general, both species are found on y substrates with sparse vegetation (Arnold 987; Arnold & Ovenden, ; Arnold et al. 7; Cabela et al. 7; Žagar et al. ), except that Horvath s lizards are more associated with s, while common wall lizards occur in a wider variety of different habitats (Arnold & Ovenden ). In this study, an extensive sampling across an altitudinal span of to, 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 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 6 and 5 (Fig., Annex ). art of the data collected in the 6 8 period was obtained within the framework of a diploma thesis (Žagar 8a) and was published in a study of habitat use of reptile community in the Kočevsko region (Žagar et al. ). pecifically, in that work we included information on the altitude, exposition, vegetation cover and habitat type of reptile community members, from which we included for I. horvathi and. muralis finds from localities that are also included in this analysis. The data of the 9 5 period was collected within the framework of a hd thesis (Žagar 6). pecies 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 999, Arnold & Ovenden ). We did not distinguish sex or age of individuals in this data set (Annex ). 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. 99) were conducted in one or up to three replicates (Annex ). 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 m of altitude (Tab. ). 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 LOVENIAE 8(): 7-6

4 5 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER Figure. Map of the Kočevsko region with localities where one or both studied species, Horvath s lizard (Iberolacerta horvathi) and common wall lizard (odarcis muralis), were found (N = 6) in the 6 5 period (see Annex ). lika. Karta razširjenosti lokacij (N = 6) na Kočevskem, kjer je bila najdena ena ali obe preučevani vrsti, velebitska kuščarica (Iberolacerta horvathi) in pozidna kuščarica (odarcis muralis), v obdobju 6 5 (glej rilogo ). Table. 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 lizard (Iberolacerta horvathi) and common wall lizard (odarcis muralis) in the Kočevsko region. Tabela. 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 (odarcis muralis) na Kočevskem. ALTITUDINAL BELT (m a.s.l.) No. of transects Total distance of transects (m) No. of ind.. muralis I. horvathi Relative No. of abundance ind. (ind.km) Relative abundance (ind.km) UM 6 9 NATURA LOVENIAE 8(): 7-6

5 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER 5 We compared the observed frequencies of lizards (corrected for replicated transect visits, Tab. ) 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) area, (ii) urban area, (iii) agricultural land, (iv) water bank, (v) road, (vi) area, and (vii) open forest (Annex ). Habitat types describe the typical areas where transects were located in the study area. Natural areas were ly occurring 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, areas comprised of any y ground or walls originating from human activities, and open forest were located in forests with <85% crown coverage (Žagar et al. ). We calculated the relative proportion of allotopic and syntopic populations in each habitat type to present it graphically (Fig. ). Results yntopic and allotopic occurrence Results represent a dataset of 6 localities, at which one or both study species, Horvath s lizard and common wall lizard, were found in the Kočevsko region within the 8 5 period (Fig., Annex ). Both species were found to occur in syntopy at localities (6%), common wall lizard was allotopic at locations (69.5%) and Horvath s lizard at 9 locations (.5%) (Fig. ). yntopic populations were found across the whole altitudinal span but with the majority of them located at middle altitudes (average altitude of syntopic populations (N = ) was 6 m a.s.l., lower quartile range = 5 m a.s.l., upper quartile range = 8 m a.s.l., Annex ). The lowest syntopic population was found at the entrance to Bilpa cave at m a.s.l. and the highest at Kameni zid at,6 m a.s.l. (Annex ). Altitudinal distribution The highest relative abundances of Horvath s lizard were determined for the highest altitudinal belt (9 99 m a.s.l.) and relative abundances decreased with decreasing altitude (Tab., Fig. ). The opposite pattern was observed for the common wall lizard; relative abundance was highest at the two lowest altitudinal belts ( 99 and 99 m a.s.l.) and decreased with increasing altitude (Tab., Fig. ). Results of the Chi square test to compare observed frequencies of lizards (Tab. ) 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 lizard: χ = 5.7, df =, <.); for common wall lizard: χ =., df =, =.). Results of comparing altitudes from all finds of both species also showed a statistically significant difference between the species (Horvath s lizard, N = 7, NATURA LOVENIAE 8(): 7-6

6 5 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER median = 98 m a.s.l., and common wall lizard, N = 5, median = m a.s.l., Mann- Whitney U tests: U = 5, Z =., <.). Figure. Relative abundances of the studied species across five altitudinal belts. lika. Relativne gostote preučevanih vrst v petih razredih nadmorskih višin. Habitat use The study species were found in seven different habitat types; Horvath s lizard in three and common wall lizard in seven of them (Fig., Annex ). Allotopic populations of common wall lizard were found in all seven habitat types, syntopic populations in all three habitat types where Horvath s lizard was found: in and y habitats and in open forests (Fig. ). These three habitat types occurred throughout the altitudinal range ( : 58 m a.s.l. (min max), : 8 55 m a.s.l. (min max), open forest: 6 8 m a.s.l. (min max); Annex ). 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: 6 a.s.l. (min max), urban area: a.s.l. (min max), water banks: 7 m a.s.l. (min max); Annex ). NATURA LOVENIAE 8(): 7-6

7 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER 5 Figure. Relative proportion of allotopic and syntopic populations of Horvath s lizard (Iberolacerta horvathi) and common wall lizard (odarcis muralis) in seven different habitat types in the Kočevsko region. lika. Relativni delež alotopičnih in sintopičnih populacij za vrsti velebitska kuščarica (Iberolacerta horvathi) in pozidna kuščarica (odarcis 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 ( and areas and open forest). Results showed that in these habitat types, too, Horvath s lizard was found at significantly higher altitudes (N = 7, median = 98 m a.s.l.) than common wall lizard (N =, median = 57 m a.s.l..; Mann-Whitney U tests: U = 85, Z = 9.9, <.). Discussion In conclusion, we have found that in the Kočevsko region, Horvath s 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 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 lizard. NATURA LOVENIAE 8(): 7-6

8 5 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER 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 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 (6%), as well as an overlap in three habitat types. revious studies reported that syntopic populations of studied species occurred only in a limited zone of middle altitudes, while Horvath s lizard was found in allotopic populations at higher altitudes and the common wall lizard in allotopic populations at lower altitudes (De Luca 989, Lapini et al. 99, Richard & Lapini 99, Lapini et al., Cabela et al. 7, Rassati ). o 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, m a.s.l. (erko & Orožen Adamič 998), whereas other study areas had higher altitude ranges (over, m a.s.l. in the Alpine region or up to,757 m a.s.l. at Velebit). Horvath s lizard was found there in places up to the highest peaks in Velebit (De Luca 989) or up to, m a.s.l. in the Alps (De Luca 989, Lapini et al. 99, Richard and Lapini 99, Lapini et al., Cabela et al. 7, Rassati ). The found between-species differences in altitudinal distribution, not only in the Kočevsko region but elsewhere, reinforce that Horvath s 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 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 lizard exhibits a more accurate thermoregulation across the seasons than the common wall lizard (Osojnik et al. ). The differences between the species were also observed on the cellular level where Horvath s 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 lizard (Žagar et al. 5). Acknowledgements The work of AŽ was funded through a hd grant (FRHBD8) supported by Fundação para a Ciência e Tecnologia (FCT) doctoral fellowships under the rograma Operacional otencial Humano Quadro de Referência Estratégico Nacional funds from the European ocial Fund and ortuguese Ministério da Educação e Ciência and has in 6 received a grant of the program»women in cience«from L'Oreal lovenia d.o.o. and lovenian national committee for UNECO. I would greatly like to thank supervisors of my hd thesis, Dr Miguel A. Careterro and Dr Al Vrezec, the reviewer of my hd 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. illero, V. Gomes, N. Osojnik, K. Bitenc, K. Drašler, D. Marguč, V. Cafuta, L. Jamnik, M. Gojkič, students from the reptile group at RTŠB Kočevje and Medvedjak,. Bizjan, Dominiku, A. ekolj, J. Vidmar, D. tanojevič, J. intar, N. ajk, D. Vlačić, V. etkovska, T Delič, M. Jelenčič, N. Ražen, T. Rezek., T. Krofel, R. ortas, F. Alvares. NATURA LOVENIAE 8(): 7-6

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10 56 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER Cafuta V. (): First record of Horvath's lizard (Iberolacerta horvathi) in the Karavanke Mts., lovenia. Nat. lov. (): 7-8. Capula M., Luiselli L. (99): Notes on the occurrence and distribution of Lacerta horvathi Méhely, 9 in Federal Republic of Germany. Herpetol. J. (): Capula M., Luiselli L. (99): Northernmost Horvath's 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-. Carranza., Arnold E.N., Amat F. (): DNA phylogeny of Lacerta (Iberolacerta) and other lacertine lizards (Reptilia: Lacertidae): did competition cause long-term mountain restriction? yst. Biodiv. (): Crochet.A., Chaline O., urget-groba Y., Debain C., Cheylan, M. (): peciation in mountains: phylogeography and phylogeny of the lizards genus Iberolacerta (Reptilia: Lacertidae). Mol. hylogenet. Evol. : De Luca N. (989): Taxonomic and biogeographic characteristics of Horvath's lizard (Lacerta horvathi Méhely, 9, Lacertidae, Reptilia) in Yugoslavia. copolia 8: -8. Faberl F., Faberl H. (99): Zwischenbericht zur Ausbreitung des Archelacerta-horvathi- Komplexes in Bayern und seinen Nachbarländern. Die Eidechse (): 8-. Franzen M., Gruber H.-J., Heckes U. (99): Untersuchungen zum Vorkommen der Kroatischen Gebirgseidechse (Lacerta horvathi) im Karwendel- und Mangfallgebirge.- München. Bericht im Auftrag des Bayerischen Landesamtes für Umweltschutz. Ökokart, München, pp. Galán., Vila M., Remón N., Naveira H.F. (7): Caracterización de las poblaciones de Iberolacerta monticola en el Noroeste ibérico mediante la combinación de datos morfológicos, ecológicos y genéticos. Munibe 5: -. Gasc J.., Cabela A., Crnobrnja-Isailović J., Dolmen D., Grossenbacher K., Haffner., Lescure J., Martens H., Martínez Rica J.., Maurin H., Oliveira M.E., ofiandiou T.., Veith M., Zuiderwijk A. (997): Atlas of amphibians and reptiles in Europe. aris, Collection atrimoines Naturels 9, ocietas Europaea Herpetologica, Muséum National d'histoire Naturelle & ervice du atrimoine Naturel, 96 pp. Gassert F., chulte U., Husemann M., Ulrich W., Rödder D., Hochkirch A., Engel E., Meyer J., Habel J.C. (): From southern refugia to the northern range margin: genetic population structure of the common wall lizard, odarcis muralis. J. Biogeogr. (8): Grillitsch H., Tiedemann F. (986): Lacerta horvathi Méhely 9 - Erstnachweis für Österreich. Ann. Naturhistor. Mus. Wien 8889(B): Grillitsch H., Cabela A., Tiedemann F. (): Lacerta horvathi Méhely, 9. 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 Jelić D. (): Checklist of Croatian amphibians and reptiles with bibliography of 5 years of research. Nat. lov. 6(): 7-7. Karaman. (9): Beiträge zur Herpetologie von Jugoslawien. Glas. Hr. naravosl. društ. : 9-9. Krofel M., Cafuta V., laninc G., opotnik M., Šalamun A., Tome., Vamberger M., Žagar A. (9): Distribution of reptiles in lovenia: a review of data collected until 9. Nat. lov. (): NATURA LOVENIAE 8(): 7-6

11 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER 57 Kryštufek B., Janžekovič F., Donev N.R. (8): Elevational diversity of reptiles on two Dinaric mountains. J. Nat. Hist. (5-8): Lapini L., Dolce. (98): Lacerta (Archaeolacerta) horvathi Méhely, 9 in Italia; nuove stazioni per le Alpi Carniche e Giulie. Atti. Mus. Friul. tor. Nat. (98): -5. Lapini L., Dal Farra A. (99): Lacerta horvathi Méhely, 9 sulle Dolomiti (Reptilia, Lacertidae). Boll. Mus. Civ. tor. Natur. : 5-8. Lapini L., Richard J., Dall'Asta A. (99): Distribution and ecology of Lacerta horvathi Méhely, 9 (Reptilia, Lacertidae) in North-Eastern Italy. Atti. Mus. Civ. tor. Nat. Trieste : -. Lapini L., Dall'Asta A., Luiselli L., Nardi. (): Lacerta horvathi in Italy: a review with new data on distribution, spacing strategy and territoriality (Reptilia, Lacertidae). Boll. Zool. 7: 5-5. Mayer W., Arribas O.J. (996): Allozyme differentiation and relationship among the Iberian- yrenean Mountain Lizards (quamata: auria: Lacertidae). Herpetozoa 9(): Mayer W., Arribas O. (): hylogenetic 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. yst. Evol. Res. (): Mayer W., avlicev M. (7): The phylogeny of the family Lacertidae (Reptilia) based on nuclear DNA sequences: convergent adaptations to arid habitats within the subfamily Eremiainae. Mol. hylogenet. Evol. (): Méhely L. (9): Eine neue Lacerta aus Ungarn. Ann. Hist.-Nat. Mus. Natio. Hung. : Michaelides., Cornish N., Griffiths R., Groombridge J., Zajac N., Walters G.J., Aubert F., While G.M., Uller T. (5): hylogeography and conservation genetics of the common wall lizard, odarcis muralis, on islands at its northern range. Lo ONE (): e7. Monasterio C., alvador A., Diaz J.A. (): Competition with wall lizards does not explain the alpine confinement of Iberian lizards: an experimental approach. Zool. : Moreira.L., Almeida A., Rosa H., aulo O., Crespo E. (999): Bases para a conservacao da lagartixa-da-montanha, Lacerta monticola. Estudos de Biologia e Conservação da Natureza no 5. Grupo de Herpetologia do Centro de Biologia Ambiental, Faculdade de Ciências da Universidade de Lisboá, Lisbon, XX pp. Mršić N. (997): lazilci (Reptilia) lovenije. Zavod Republike lovenije za šolstvo, Ljubljana, 67 pp. Odierna G., Aprea G., Arribas O.J., Capriglione T., Caputo V., Olmo E. (996): The karyology of the Iberian lizards. Herpetologica 5(): Osojnik N., Žagar A., Carretero M.A., García-Muñoz E., Vrezec A. (): Ecophysiological dissimilarities of two sympatric lizards. Herpetologica 69: 5-5. erko D., Orožen Adamič M. (998): lovenija: okrajine in ljudje. Mladinska knjiga, Ljubljana, 75 pp. Rassati G. (): Contributo alla conoscenza della distribuzione della Lucertola di Horvath Iberolacerta horvathi e della Lucertola dei muri odarcis muralis in Friuli Venezia Giulia e in Veneto. Atti. Mus. Civ. tor. Nat. Trieste 5: -6. Richard J., Lapini L. (99): Trophic niche overlap in syntopic populations of Lacerta horvathi and odarcis muralis (Reptilia, Lacertidae). Atti. Mus. Civ. tor. Nat. Trieste 5: NATURA LOVENIAE 8(): 7-6

12 58 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER alvi D., Harris D.J., Kaliontzopoulou A., Carretero M.A., inho C. (): ersistence across leistocene ice ages in Mediterranean and extra-mediterranean refugia: phylogeographic insights from the common wall lizard. BMC Evol. Biol. (): 7. chmidtler H., chmidtler J. (996): Zur Reptilienfauna der Nördlichen Kalkalpen zwischen Isar und Inn (BayernTirol). Mitt. Landesverband Amphibien und Reptilienschutz (LAR) in Bayern 5(): -6. illero N., Bonardi A., Corti C., Creemers R., Crochet., Ficetola G.F., Kuzmin., Lymberakis., ous.d., indaco R., peybroeck J., Toxopeus B., Vieites D.R., Vences M. (): Updated distribution and biogeography of amphibians and reptiles of Europe. Amphibia-Reptilia 5: -. Tiedemann F. (99): Zur Verbreitung der Kroatischen Gebirgseidechse, Lacerta horvathi Méhely, 9, in Osterreich (quamata: auria: Lacertidae). Herpetozoa 5(): Tome. (996): regled razširjenosti plazilcev v loveniji. Annales 9'96 Anali za istrske in mediteranske študije, eries historia is : 7-8. Tome. (999): Razred: lazilci, Reptilia. In: Kryštufek B., Janžekovič F. (Eds.), Ključ za določanje vretenčarjev lovenije, DZ, Ljubljana, pp Tome. (): lazilci. In: Kryštufek B. (Ed.), Raziskava razširjenosti evropsko pomembnih vrst v loveniji, Končno poročilo, rirodoslovni muzej lovenije, Ljubljana, pp Tvrtković N., Veen. (6): The Dinaric Alps rare habitats and species. A nature conservation project in Croatia. art A. Hrvatski prirodoslovni muzej, Zagreb, Royal Dutch ociety for Nature Conservation, 67 pp. Žagar A. (8a): Importance of open areas in forest landscape for reptiles (Reptilia) (in lovene). Graduation thesis, University of Ljubljana, Ljubljana, 8 pp. Žagar A. (8b): The lowest altitudinal record of Horvath's Rock Lizard (Iberolacerta horvathi) in lovenia. Nat. lov. (): Žagar A. (6): Interspecific competition between the Common wall lizard (odarcis muralis [Laurenti 768]) and the Horvath's lizard (Iberolacerta horvathi [Méhely 9]). hd Doctoral dissertation, University of Ljubljana, Ljubljana, pp. Žagar A., laninc G., Krofel M. (7): Records of Horvath's Rock Lizard (Iberolacerta horvathi) from the Notranjsko podolje region (central lovenia). Nat. lov. 9(): -. Žagar A., Kos I., Vrezec, A. (): Habitat segregation patterns of reptiles in Northern Dinaric Mountains (lovenia). Amphibia-Reptilia : Žagar A., Carretero M.A., Krofel M., Lužnik M., odnar M., Tvrtković N. (): Reptile survey in Dinara Mountain (Croatia) revealed the southernmost known population of Horvath's lizard (Iberolacerta horvathi). Nat. Cro. (): 5-. Žagar A., imčič T., Carretero M.A., Vrezec A. (5): The role of metabolism in understanding the altitudinal segregation pattern of two potentially interacting lizards. Comp. Biochem. hysiol. art A 79: -6. NATURA LOVENIAE 8(): 7-6

13 Annex. Locality descriptions with presence of studied species that were surveyed in the period between 6 and 5 in Kočevsko region and sorted by altitude. urveys done in spring (before th June of the calendar year) are labelled with (), while () corresponds to surveys done in summer (after st July of the calendar year). During each survey, we either made counts of individuals on a predetermined transect route or we only noted presence () 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 = Common wall lizard (odarcis muralis) and = Horvath s lizard (Iberolacerta horvathi), A = Allotopic, = yntopic. riloga. Opisi lokacij v Kočevski regiji, kjer sta bili zabeleženi ena ali obe preučevani vrsti v obdobju raziskave med leti 6 in 5. Lokacije so razporejene po nadmorski višini. Najdbe, ki so bile zabeležene na popisih pred. junijem v koledarskem letu, so označene z () in tiste po. juliju v koledarskem letu z (). Na posameznem obisku lokacije smo prešteli osebke na predhodno določenem transektu (N) ali pa smo zabeležili le prisotnost vrste ( = 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 Krügerjevega koordinatnega sistema, p.pres. prisotnost vrste,.m. = pozidna kuščarica (odarcis muralis) in = velebitska kuščarica (Iberolacerta horvathi), A = alotopična populacija, = sintopična populacija. Locality med Žlebi in Grgeljem med Bilpo in Lazami pri Žlebih pri Gorenji Žagi Bilpa med Gorenjo Žago in Žlebi pri Gorenji žagi Žlebi med Kobilno jamo in Gorenjo Žago med apnikom in Brsnikom med Lobičem in Žlebi Maverc apnik Alt. (m) GKX GKY road urban area 965 urban 88 area Habitat type road water bank p. pres Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER 59.m..m..m..m..m..m..m..m..m..m..m..m..m. 6 () 6 ().m. 7 () 7 ().m. 7 () 7 ().m. 8 () 8 ().m. 8 () 8 ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. 5 () 5 () count is in bold. Abbreviations: Alt. average altitude, GKY and GKX = Y and X coordinates of the Gauss Krüger coordinate system, p. pres. species presence,.m. NATURA LOVENIAE 8(): 7-6

14 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER 5 ().m. 5 () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () 8 ().m. 8 () 8 ().m. 8 () 7 ().m. 7 () 7 ().m. 7 () 6 ().m. 6 ().m..m..m..m..m..m..m..m..m..m..m..m..m..m..m..m..m..m. Locality pokopališče v Fari med Grivacom in Gladloko pri Gladloki Mirtoviči robotnik Mirtoviški potok nad cerkvijo v Fari pokopališče sv. Štefan pri robotniku Kostel nad vasjo lanina zapuščen kamnolom kamnolom v odstenah od lanine na laninsko steno med Friškovo grabo in Dolenjim otokom pri Dolenjem otoku pot na Krempo od vasi odstene do sten pod odstenami od lanine na laninsko steno Alt. (m) Annex riloga. Continued. Nadaljevanje GKX GKY Habitat type urban area road urban area urban area water bank road urban area open forest urban area road road open forest road road p. pres. 6 NATURA LOVENIAE 8(): 7-6

15 5 5 Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER 6 5 ().m. 5 () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () 8 ().m. 8 () 8 ().m. 8 () 7 ().m. 7 () 7 ().m. 7 () 6 ().m. 6 ().m..m..m..m..m..m..m..m..m..m..m..m. Locality pri robotniku na odstenah pod laninsko steno na odstenah na odstenah od vasi odstene do sten kamnolom pri Kočevski Reki pot na Krempo od lanine na laninsko steno na planinski poti na Krempo med Gornjo Brigo in Borovcem pri Kočevski Reki laninska stena na planinski poti na Krempo območje tevniki na Mali gori zahodno od Špičastega vrha na Mali gori Gornje niše na Mali gori pod Kuželjsko steno Kuželjska stena na Žurgarski steni Alt. (m) Annex riloga. Continued. Nadaljevanje. GKX GKY Habitat type open forest agricultu -ral land road open forest open forest p. pres. NATURA LOVENIAE 8(): 7-6

16 p. pres Anamarija ŽAGAR: Altitudinal distribution and habitat use of the common wall lizard... CIENTIFIC AER 5 ().m. 5 () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () ().m. () 8 ().m. 8 () 8 ().m. 8 () 7 ().m. 7 () 7 ().m. 7 () 6 ().m. 6 ().m. Locality pred Žurgarsko steno na Žurgarski steni Fridrikštajn Male Bele stene Taborska stena ob cesti pod Velikimi Belimi stenami Kameni zid na Taborski steni na Taborski steni na ovinku pred Velikimi Belimi stenami Velike Bele stene na robu sten Annex riloga. Continued. Nadaljevanje. Alt. (m) GKX GKY Habitat type urban area and open forest and open forest NATURA LOVENIAE 8(): 7-6