Arch. Biol. Sci., Belgrade, 60 (3), 459-468, 2008 DOI:10.2298/ABS0803459C BREEDING SITE TRAITS OF EUROPEAN NEWTS (TRITURUS MACEDONICUS, LISSOTRITON VULGARIS, AND MESOTRITON ALPESTRIS: SALAMANDRIDAE) IN THE MONTENEGRIN KARST REGION RUŽA ĆIROVIĆ1, D. RADOVIĆ2, and TANJA D. VUKOV3 1Nature Protection Institute of Montenegro, 81000 Podgorica, Montenegro 2Institute of Zoology, Faculty of Biology, University of Belgrade, 11000 Belgrade, Serbia 3Siniša Stanković Institute for Biological Research, 11060 Begrade, Serbia Abstract — We recorded the occurrence of three European newt species - the smooth newt (Lissotriton vulgaris), the eastern alpine crested newt (Triturus macedonicus), and the alpine newt (Mesotriton alpestris) - in the Montenegrin karst, as well as their breeding site characteristics. In terms of long-lasting breeding site numbers and occupation rate, the most common species is the smooth newt, followed by the alpine newt and the crested newt. The e��������������������������xamined water bodies with- out newts showed no significant differences of aquatic habitat characteristics compared to water bodies with newts. The factors that explained most of the observed variation in newt breeding site traits were the habitat category and habitat origin. The alpine newt primarily inhabits natural lakes, while the crested newt inhabits artificial breeding sites such as lithotelma and ubao. The smooth newt is less choosy and occurs in different types of natural and artificial habitats. The aquatic requirements of Montenegrin newt species do not differ substantially in many respects from requirements of the core species range populations. Key words: Smooth newt, alpine newt, crested newt, holokarst, allotopy, syntopy, conservation, Montenegro Udc 597.9(4:497.16) INTRODUCTION B u s k i r k , 2003). Temporary ponds provide a rich (in terms of food resources) but transient environ- European newts (Lissotriton sp., Triturus spp., and ment for amphibians (G r i f f i t h s , 1997) and are Mesotriton sp.) use a variety of water bodies that a major breeding habitat for amphibians in the differ in an array of abiotic factors (e.g., pond water Mediterranean region (D i a z - P a n i a g u a , 1990). depth, water chemistry, pond isolation), as well as However, amphibian breeding sites with a mostly biotic factors (e.g., presence of competitors and stable hydroperiod, which are by far the predomi- predators and effects of their interactions, potential nant type of water bodies in the Montenegrin karst mate presence, etc; J a k o b et al., 2003). The qual- region, have escaped investigations so far. ity and quantity of these characteristics partially determine the presence of newt populations to the Available data on newts’ habitat selection pri- extent that some features can be isolated as being of marily refer to populations from Western, Northern, particular diagnostic importance and used as a basis and Central Europe (C o o k e and F r a z e r, 1976; for determining the likelihood of newts’ occurrence S w a n and O l d h a m , 1994; M a r n e l l , 1998; (O l d h a m et al., 2000). S z t a t e c s n y et al., 2004; S k e i et al., 2006).2006�������). Much less is known about populations from Eastern Europe Freshwater lentic habitats range from ephem- (B a b i k and R a f i n s k i , 2001) and the Western eral puddles to large permanent lakes, with con- Mediterranean (P a v i g n a n o et al., 1990; I l d o s comitant gradients in many features that covary and A n c o n a , 1994; D e n o ë l and L e h m a n n , with size and permanence of the water body (Va n 2006), and only anecdotal data are available on 459 460 RUŽA ĆIROVIĆ et al. ones from Southeaste Europe, including the Balkan The aims of this paper are twofold. To begin Peninsula (B o u s b o u r a s and I o a n n i d i s , 1997; with, we present for the first time an account of D e n o ë l , 2004, K a t i et al., 2007). Objective meth- breeding site characteristics of Montenegrin karst ods of measuring and monitoring newt populations area newts. Secondly, based of this analysis, we have never been applied in the Dinaric karst regions. sought to answer (1) whether some of these breed- The current information on newt breeding sites in ing site characteristics shape the observed distribu- this area is scarce and limited to narrow territories tion pattern of European newts in the Montenegrin (D ž u k i ć , 1981; K a l e z i ć and D ž u k i ć , 1985; karst region, and (2) do the newt species differ much D ž u k i ć and K a l e z i ć , 1988; B r u n o , 1988). in their aquatic requirements when they inhabit What is more, no studies so far have examined fringe areas of the taxon's distribution? simultaneously the factors that affect the distribu- tion of newts in the Dinaric karst regions. MATERIALS AND METHODS Our primary objective was to assess patterns of Study area variation in European newt breeding site character- istics in the Montenegrin karst area and search for The segment of the Dinaric Alps which forms the possible association(s) between aquatic habitat traits terrain of Montenegro is predominantly built of and the presence of particular species. We studied all limestone and dolomite sediments that have been newt species living in this region: the smooth newt exposed to various forms of karstification (C v i j i ć , (Lissotriton vulgaris), the eastern alpine crested newt 1989). The karst zoning of Montenegro is very (Triturus macedonicus, hereinafter referred to simply complex (R a d u l o v i ć anandd R a d u l o v i ć , 1997). as the crested newt), and the alpine newt (Mesotriton However, for the sake of simplicity with regard to alpestris). The Montenegrin karst area is relevant in our investigation, the karst region can be divided several ways for this study. Firstly, as noted below, into two zones separated mainly by the spacious this region is inhabited by endemic or subendemic Zeta depression, where flysch deposits are found. taxa and/or phylogenetic clades of each of the newt The landscape of the northern Upper Zone is an groups, whose breeding site characteristics have exclusively mountainous region, covered with for- never been assessed systematically. Secondly, t��he ests and grasslands, with little bare karst. Short examined region is on the range margins of all three spring-fed watercourses are relatively common, but newt groups (G r i f f i t h s , 1996), thus harboring lakes of various sizes, mostly of glacial origin, as well geographically (and most likely ecologically) periph- as different kinds of ponds, are the main amphib- eral populations (L e s i c a and A l l e n d o r f , 1995; ian (for newts almost the only) breeding sites in A r a ú j o and Wi l l i a m s , 2001). this zone. On the whole, these water bodies are not densely distributed. The landscape of the southern In a previous study of ours, we determined Lower Zone is a less elevated region with bare karst European newt distribution patterns in the (called holokarst) over its largest part. Most of this Montenegrin karst (Ć i r o v i ć et al., submitted). is exposed karst with scanty plant growth in the Briefly, the smooth newt occurred throughout the form of crack dwellers. The Lower Zone is a flow- Montenegrin karst region, except in the most ele- ing-water-free karst region. The limestone base of vated parts. The alpine newt was restricted mostly the zone, being highly porous, drains much of the to the Upper Zone, while the crested newt was rainfall, causing extreme aridity during the sum- represented the least, occupying only a part of the mer months. Relatively scarce surface water bodies, Lower Zone (Figs. 1 and 2). In addition, �����������using land- usually at the sites of sinkholes and potholes with scape traits obtained by combining remote sensing impermeable bottoms, in which water comes almost and geographic information systems, we found that exclusively from rainfall and rarely from boiling geographic position (latitude and longitude) and springs, are the only ones available for amphibian (especially) elevation have a great impact on newt reproduction. These bodies have been used since occurrence. a very distant past as cattle drinking water or even Breeding site traits of European newts in montenegro 461 Fig. 1. Range of distribution of L. vulgaris in Montenegro Fig. 2. Ranges of distribution of T. macedonicus (light gray) (shaded areas) and geographic position of L. vulgaris vulgar- and M. alpestris (dark gray) and geographic position of their is and L. vulgaris tomasinii breeding sites used in this study. breeding sites in Montenegro. Dashed line delimits the Upper Dashed line delimits the Upper and Lower Zones. and Lower Zones. for human consumption. As water was constantly vegetation types near the water bodies, the extent in short supply, people have always taken great care of karstification, and, concomitantly, the extent of of these ponds. By building dams or walls, as in the population connectivity. case of wells, they have enabled these ponds to last a Studied newt taxa and distribution patterns long time. Thus, the holokarst is characterized by for the most part permanent ponds, the majority being Four newt taxa inhabit the Montenegrin artificial or partly artificial.