ࡱ> G +bjbjَ +,]8TLA@(i k k k - l@$A5dd@i i  I i $@4 ֓a 11 August 1997 Josip Kusak Biology Department, Veterinary Faculty of the University of Zagreb Heinzelova 55 10000 Zagreb Croatia Phone: 385-1-239-0141 Fax: 385-1-214-697 email: kusak@mavef.vef.hr or josip.kusak@alf.tel.hr RH: Brown bear habitat in Croatia ( Kusak and Huber BROWN BEAR HABITAT QUALITY IN GORSKI KOTAR, CROATIA JOSIP KUSAK, Biology Department, Veterinary Faculty of the University of Zagreb, Heinzelova 55, 10000 Zagreb, Croatia, email:kusak@mavef.vef.hr or josip.kusak@alf.tel.hr DJURO HUBER, Biology Department, Veterinary Faculty of the University of Zagreb, Heinzelova 55, 10000 Zagreb, Croatia, email:huber@mavef.vef.hr Abstract: The brown bear (Ursus arctos) population in Croatia is a possible source of bears for reintroducing and augmenting disappearing European brown bear populations in western Europe. For successful reintroduction, knowledge about bear habitat qualities of both source and target areas is necessary. A habitat suitability index (HSI) model was developed to assess habitat quality of European brown bear in Gorski kotar, Croatia. We identified natural components of bear habitat, most of which had been deteriorated by human activities. Food sources were available year-round. Human influence was manifested through a relatively high density of roads (1.91 km/km2), including forest roads. Habitat fragmentation did not occur within the study area, but a highway under construction was a possible threat. Habitat in Gorski kotar can be improved with changes in management practice. Changes that could have an immediate effect include closing forest roads, seasonally avoiding of logging in denning areas, and changing laws that prevent smallscale bear management. Proposed measures with long-term effects include modifying forest management to improve natural forest cover and educating residents about garbage management. Int. Conf. Bear Res. and Manage. 10:000-000 Key words: Croatia, European brown bear, habitat, suitability, index, HSI, quality, Ursus arctos. __________________________________________________________ One of the largest remaining brown bear populations in Europe occurs in the Dinara, ara, and Pindus mountains (approximately 60,000 km2) spanning Slovenia, Croatia, Bosnia and Herzegovina, Monte Negro, Macedonia (Huber and Frkovic 1993), Albania and Greece (Servheen 1990). Within the Croatian portion of this range, about 400 bears inhabit approximately 9,800 km2 (Huber and Frkovic 1993), and this number has been stable since the early 1980s (Frkovic et al. 1987). Mitochondrial DNA testing has demonstrated that the brown bears in the Dinara Mountains have stronger genetic similarities to the small, isolated populations in the Alps, Apennines, and Pyrenees than to other European populations (Taberlet and Bouvet 1994, Randi et al. 1994). Consequently, they may be the best source to reintroduce or augment bear populations in western Europe. Projects to reintroduce bears from the Dinara Mountains are in progress, with 3 (2 F, 1 M) bears already translocated to the Austrian Alps and another 3 (2 F, 1 M) to French Pyrenees. Maintenance of suitable bear habitat is central to the continued presence of the species in Europe (Servheen 1990). Knowledge of brown bear habitat relationships is needed to enhance management of existing European brown bear populations and to improve chances for successful reintroduction efforts. We developed a habitat suitability index model for brown bear in southern Europe, following the U.S. Fish and Wildlife Service's (1981) habitat evaluation procedures. We used the HSI to evaluate brown bear habitat in the Gorski kotar region of Croatia; we recommend management changes based on that analysis. This study was funded by the Croatian Ministry of Science, Wildbiologische Geselschaft - Mnchen, John Sheldon Bevins Memorial Foundation, and Stiftung Europische Naturerbe (EURONATUR). We thank A. Frkovi, D. Majnari, and B. Pleae for logistic field support. Special thanks are due to the reviewers for improving the style and quality of the manuscript. BROWN BEAR HABITAT SUITABILITY INDEX MODEL FOR SOUTHERN EUROPE HABITAT USE INFORMATION General Bear management in southern Europe includes feeding with carrion and corn on permanent feeding sites and a regulated hunting season. The most intensive feeding is during hunting season (1 Oct to 30 Apr). Annual hunting quotas allow taking 10% of estimated number of bears (Huber and Frkovic 1993). Hunting law (Narodne novine 1994) allows leasing of hunting areas to hunting clubs and individuals. Food Requirements Food habits and habitat use of brown bear in Croatia were analyzed by Cicnjak (1991). In spring and early summer, bears were found at lower elevations where food was available. Main food sources included broad-leaved garlic (Allium ursinum), lords and ladies (Arum maculatum), and food at permanent feeding sites (carrion and corn). Bears also visited meadows where they consumed grasses (Gramineae spp.), clovers (Trifolium spp.), and sorrels (Rumex spp.). During the summer wild angelica (Angelica silvestris) and stinking aposeris (Aposeris foetida), were the main plant foods eaten although bears occasionally fed in and substantially damaged oat fields (Huber and Mori 1989). By late summer, fruits ripened in meadows, abandoned orchards, along fields and roads, and in the forest; the most important bear foods were raspberry (Rubus idaeus), bramble (Rubus fruticosus), common buckthorn (Rhamnus cathartica), and blueberry (Vaccinium myrtillus). During fall, bears spent the most time in forests where beech (Fagus sp.) nuts were abundant. Similar patterns were found in Cantabrian mountains, Spain (Clevenger et al. 1992a,b). Depredation on domestic animals was lower than in other European countries because of availability of natural and supplemental food available (Cicnjak 1991). During feeding on natural food, the average distance of bears from settlements and roads was not statistically different from the randomly distributed points (Cicnjak 1991). Water Requirements Brown bear distribution and abundance has not been associated with water availability in Croatia (Cicnjak 1991). Cover Requirements Resting sites (day beds) had the shortest horizontal visibility ( EMBED Equation.2 =8.4 m, range=3-46 m), followed by denning sites ( EMBED Equation.2 =12 m, range=1-18 m) and feeding sites ( EMBED Equation.2 =36, range=6-186 m) (Cicnjak 1991). European brown bears do not rely on protective cover to access food, they have developed crepuscular and nocturnal activity patterns (Roth and Huber 1986) when protection cover is less important. The closest distance to which man can cautiously approach a European brown bear is about 30m (pers. observ.). Bears approached closely may abandon some feeding places, but this may not be of great importance in habitat with good food sources elsewhere. Protective cover may be more important for selection of suitable daybeds or dens. Reproductive requirements Bears in Croatia den in forested areas at elevations from 450 m to 1370 m( EMBED Equation.2 =863m, N=28). Average distance from den to road was 486 m (range=39-1500 m) and from den to settlement was 1435 m (range=200-4000 m) (Huber and Roth 1995). Feeding on energetically rich beech nuts, which were the most important food source for bears in Croatia in autumn (Cicnjak 1991), could be of great importance for successful wintering and giving birth (Schooley et al. 1994). Beech trees start producing nuts at 40-50 years. Maximum crop production ( EMBED Equation.2 =1.38 kg/tree) can be expected every 7th to 12th year; in other years production is 0.11 kg/tree. Cicle of beech productivity occures in different parts of forest in different years (Regent 1980). Interspersion requirements Arangement of different habitat types follows the changes in elevations, geology and other ecological factors. Seasonal movements of bears reflect changes of food availability in different habitat types. In a continuous habitat as Gorski kotar, home range size of bears for both sexes did not vary between spring, summer, and fall ( EMBED Equation.2 =28km2, range=1-102km2, N=32), although winter range was significantly smaller ( EMBED Equation.2 =4km2, range=0-18 km2, N=5) (Huber and Roth 1993). Special Considerations Forest management in the mountains of Croatia involves building a dense network of forest roads. These roads are not closed which increases human access. Increased traffic on local and regional roads and railroads can increase traffic mortality of bears. Crossing sites on roads and railroads were recognized as important parts of migrating corridors (Huber et al. This Volume). Telemetry studies in Slovenia (Kaczensky et al. In Press) showed that highways are a physical border for movements of females, whereas males occasionally cross during the mating season. HSI MODEL Model Applicability Geographic Area This model describes habitat suitability in Gorski kotar, Croatia. It can be applied to similar mountainous areas in Europe, like Alps, Pyrenees, Apenines, and Carpathians. Season This model is designed to produce HSI values for year-round habitat needs of the brown bear. We assume that carrying capacity is restricted by food availability, and that a weak compensatory relationship exists for food availability among seasons. To stress importance of fall food, we weighted it by a factor of 2. Cover Types The model can be applied on areas where beech, mixed beech-fir (Abies sp.), and spruce (Picea sp.) forests prevails. Minimum Habitat Area Home range size of bears for both sexesin foraging season averages28km2, range=1-102km2 (Huber and Roth 1993). The minimum habitat area was assumed to be 30 km2. Verification level This model has not been tested under field conditions. Model description A habitat suitability index (HSI) is a numerical index that represents the ability of a given habitat to support a specific species. An index of 0.00 represents unsuitable habitat, and maximum HSI1.00 represents optimal habitat. For HSI to be between 0.00 and 1.00 the assumed linear relationship between chosen habitat components and carrying capacity must be met (U.S. Fish and Wildlife Service 1981). Graphic Overview We defined food, cover, and human impact as the main bear habitat components (Fig. 2). Van Manen and Pelton (1993) defined ecological, physical, and cultural habitat components for American black bear (Ursus americanus). Aste (1993) listed the following bear habitat components for Austria: food type, food availability, availability of remote and safe places, and effect of limiting factors. Clevenger et al. (1992a) ranked bear habitat in Cantabrian mountains (Spain) by quality of forest cover, elevation, distance to nearest village, and distance to nearest paved road. Model relationships The contribution of each habitat component to the HSI was evaluated by measuring 1 or more variables (Fig. 2). A suitability index (SI) can be calculated for each variable (Fig. 3). based on our knowledge of bear biology, expert opinion (A. Frkovi, Croatian Forests, Delnice, Croatia, pers. commun. 1993) and through comparison to other authors (Aste 1993; Van Manen and Pelton 1993, 1995; Clevenger et al. 1995). Life Requisite Components Food We considered only plants that were present in bear diets with a frequency (5% and volume (0.5% (Cicnjak 1991). Total cover of 10% for 1 or several plants was taken as the threshold of optimal index (Fig. 3 A,C). When total cover was <10% (V1 or V3<10%) calculation for spring and summer was SIV1 or V3 = 1/10 V1 or V3. When total cover was ( 10% (V1 or V3 ( 10%), the value was SIV1orV3=1.00 (Fig. 3 A,C). Beech cover, because it produced the most important fall food, changed SI lineary from 0% to 100%, (Fig. 3 E). This relationship was chosen because productivity cycles of beech trees varies in time for a different areas. Not all beech trees have maximum crop in the same year (Regent 1980). The same model was used by Aste (1993). When a single significant bear plant was found in spring or summer, the area was assigned an SIV2 or V4 = 0.50. If ( 2 plants were found, we assigned SIV2orV4=1.00) (Fig. 3 B,D). When < 40% of beech trees produced nuts (V6<40%) SI was calculated as SIV6=(1/40)V6. For V6>40%, SIV6 = 1.00 (Fig. 3 F). Cover We evaluated hiding cover separately for the foraging and denning period. If<25% of the sites in a foraging area had adequate cover (V7<25%), the SIV7 was calculated by SIV7=(1/25)V7. When ( 25% of the sites had adequate cover, SIV7was assigned the value of 1.00 (Fig. 3 G). Suitable denning areas in forested areas had SIV8a=1.00; denning areas outside forests had for SIV8a=0.00 (Fig. 3H). Denning areas > 500 m from roads (V8b > 500 m) were assigned 1.00 for SIV8b, whereas closer distances (less suitable) were assigned SIV8b = 0.00 (Fig. 3 I). Denning areas < 1500 m from human settlements also were considered unsuitable for denning (SIV8c=0.00), but areas >1500 m were considered suitable bear denning habitat (SIV8c=1.00) (Fig. 3 J). Areas suitable for denning (V8) were determined by the factor with the lowest value, because we assumed that factors would be limited (V8=min.(V8a, V8b, V8c)). It means that part of the habitat was suitable for denning if it was in the forest, and >500 or >1500 m away from the road or settlement, respectively. Habitat section should have >25% suitable denning sites for SIV8to be >0.00 (SIV8=(1/75)V8(1/3)). The same model was used by Aste (1993) for calculating availability of remote areas. Human Impact We evaluated human impact by summing total length (V9, km/km2) of all roads, including forest roads that are open and accessible to the public, and by searching for possible habitat fragmentation. We assumed that habitat quality decreased as the density of roads increased, where areas with >1.5 km/km2 roads received SIV9=0.00 (Fig. 3 L). We used the same model developed by Aste (1993). Part of the habitat was considered isolated if it was surrounded by 500 m of unforested area. We did not consider habitat as isolated, even if it contained railroads or road, if there were places for crossing (tunnel, viaduct, "green bridge"). Habitat fragmentation was calculated from colored, 1:25,000 topographic maps. Areas <10 km2 (V10<10km2) received an SIV10 of 0.00, whereas SIV10 for areas of 10 to 2,500 km2 were calculated from SIV10=(1/2490)V10-(1/249) (Fig. 3 M). The same model was used by Aste (1993). Equations Suitability indexes of all habitat components were combined according to importance, value, and compensatory or limiting nature. To emphasize the importance of fall food, we squared average of SIV5 and SIV6. Also, for overall HSI, SIFOOD was multiplied by 2 because of its greater importance compared to other habitat components. The following equations were used to calculate HSI: SIFOOD=(((SIV1+SIV2)/2)*((SIV3+SIV4)/2)*(((SIV5+SIV6)/2)2))1/4 (1) SICOVER=(SIV7+SIV8)/2 (2) SIHUMAN IMPACT=(SIV9+SIV10)/2 (3) HSI=(2SIFOOD+SICOVER+SIHUMAN IMPACT)/4 (4) All statistical analyses were performed using SAS software (SAS Institute, Inc. 1994) Application of the Model This model can be applied to areas with plant communities defined by Braun-Blanquet's method. Other areas would reqire field sampling to determine presence and amount of bear related plants. Sources of Other Models Existing models for evaluating bear habitat suitability (Aste 1993, Van Manen and Pelton 1994, Clevenger et al. 1995) were not found to be appropriate for European southeastern mixed deciduous-conifer forests, but were used as a starting point for construction of the model. APPLICATION OF THE HSI IN GORSKI KOTAR, CROATIA Study Area Gorski kotar (1,796 km2) is situated in Croatia in the narrowest part of the Dinarids (Fig. 1). It comprises the northwest end of the Dinara Mountains, which divide Mediterranean from the continental part of Croatia and the Adriatic Sea from the Black Sea drainage. The mountains gmainly consist of carbonate rocks with older silicates that are not water permeable (Horvat 1962). The highest mountains are Bjelolasica (1,533 m) and Risnjak (1,528 m). Gorski kotar has a moderately cold climate (yearly average about 8C) with a relatively large amount of precipitation (up to 377 cm/year) and high snow cover, which averages 139 days/year (Penzar 1959). The cover types used by brown bears in Croatia are defined by several plant communities. Forests cover 66% (1,191 km2) of the total area, of which 91% is managed. Vegetation shows clear altitudinal differentiation, and Horvat (1962) described 3 main vegetation belts. Deviations from the main zonality are varied by different exposure, slope, wind, and other ecological factors. Elevations from 0 m to 850 m by the Adriatic Sea are covered with sub-Mediterranean thermophil forests belonging to Ostryo-Carpinion orientalis group. Deciduous and mixed forests of Fagion illyricum group cover the largest part of Gorski kotar stretching from 900 m (600 m on continental side) to 1400 m above sea level. Shrublike forest of dwarf mountain pine (Pinetum mugi croaticum) is found above 1400 m (Horvat 1962). Gorski kotar area is divided in 4 political units: abar, Delnice, Vrbovsko, and Rijeka (Fig. 1). These are also forest management units and our study area sections. The area is inhabited by 236,000 people in 308 settlements, but the majority (74%) live in 3 towns on the Adriatic coast (i.e., in the 10% of land area that is marginal bear habitat). The rest (90%) is mountainous area inhabited by 24 humans/km2. A majority of local people work in forestry (9.5%) and related industries (48%). Only 1.4% work in agriculture on 331 km2 of pastures, meadows, fields, gardens, and orchards that comprise 18% of total study area (Dr~avni zavod za statistiku 1993). Data Sources We calculated availability of plant food using phytocenological data (Horvat 1938; Trinajsti 1972, 1995; Pavleti et al. 1982; Vukeli 1985) for specific plant communities for each season except winter. Data of location and size of each phytocenosis came from Delnice Forest Manage. (1994), the agency that manages the forest in the study area. Calculation for percent of beech nuts producing trees was based on information that classifies crop producers (K.Potenjak, Dipl ing. Jastrebarslo, Croatia, pers. commun., 1994) and on number of trees/hectare in each perimeter class for each plant community (Cestar et al. 1986). We estimated density of understory vegetation on random plots of 450 m2 (N=20 for each plant community) by measuring the percentage of a bear silhouette (average natural size - 95x130 cm) covered at 15 m distance from east, west, north, and south. This method combined those of Cicnjak (1991) and Van Manen and Pelton (1995). The mean of these 4 measurements was used as a percent cover for each site. For each plant community we determinated how many random points had cover >60% of the bear silhouette. Areas suitable for denning were obtained from 1:25,000 topographic maps. For total lengths of main, regional, and local roads, we used data from the State Department for Statistic (Dr~avni zavod za statistiku 1993). For forest roads lenghts, we used data from Delnice Forest Manage. (1994). Results and Discussion Plants belonging to genera Allium, Arum, and Rumex represented spring food availability and provided an overall 4.94% cover (SIV1=0.48; Table 1). The minimum cover of spring food of 1.38% was recorded for habitat facing the Adriatic Sea (Rijeka), whereas maximum cover (8.77%) was found at lower elevations on the continental side (Vrbovsko). In each section, only lords and ladies were present with 10% coverage, resulting in SIV2=0.50 (Table 1) for the entire study area for spring food diversity. Summer food covered >10% of each habitat section ( EMBED Equation.2 =25.21%) giving an overall SIV3=1.00 (Table 1). The lowest abundance of food was again found near seacoast compared to the higher continental areas. In all habitat sections we found >1 (max.=10) summer food, plants resulting in SIV4=1.00 for summer food diversity (Table 1). Beech, the only important producer of fall food in the study area, covered 55.92% of forested area, resulting in a fall food availability of SIV5=0.56 (Table 1). An average 11.04% of all beech trees were capable of producing nuts, for a suitability index for fall food productivity of SIV6=0.28 (Table 1). Private forests (20% of all forests), which are situated mostly around villages, lack mature beech trees and conifers. There were no significant differences among study area sections in food availability in different seasons (P > 0.05). Bear movements are related to food availability (Craighead et al. 1982, Cicnjak 1991, Hamer et al. 1991). Home range sizes of brown bears in Croatia do not differ among seasons except winter (Huber and Roth 1993). This may be explained by year-round balanced food availability in our study area. Spring food had the lowest abundance in Rijeka section, near the Adriatic Sea, whereas lower continental areas provided the best spring habitat. Fall food availability was good through the entire study area because beech trees covered about 50% of all forested surfaces. The limiting factor seemed to be beech productivity. Management practices have resulted in 11.04% of beech trees at productive ages, decreasing suitability of bear habitat. Clearcuts, which are sometimes made as circular openings or clearcuts of entire mountain slopes, may provide good summer foods during early succession stages. Replanting such areas with spruce (Picea abies), which suppresses most understory vegetation, will not promote good bear habitat for the future. Permanent bear feeding sites, which offered carrion and slaughterhouse refuse are visited mostly in spring (A. Frkovi, Croatian Forests, Delnice, Croatia, pers. commun. 1993), but there is no data on quantities of food eaten by bears. Only 2.87%of study area had horizontal protective cover with a density >60% which resulted in SIV7=0.14 (Table 1). All areas with low horizontal visibility were found in beech forests growing on a rugged terrain with steep slopes and with abundant running water. Low visibility was related more to terrain configuration than vegetation density. The overall area providing suitable den sites was 19.56%, SIV8=0 (Table 1). Number and size of available sites varied within areas. The Delnice section had 41 suitable areas totaling 71.85 km2 for SIV8=0.00 (Table 1). The Rijeka section had 88.76 km2, (37.09%) of suitable denning habitat which differed significantly (P=0.00046, ANOVA [analysis of variance]) from other sections. For Rijeka SIV8=0.16 (Table 1); it was the only section with SIV8>0. The Vrbovsko section had 25.46 km2 of suitable denning habitat resulting in SIV8=0.00 (Table 1). abar also had a SIV8=0.00 (Table 1) for 40.39km2, (17.52%) of suitable denning habitat. Risnjak National Park (16.90 km2), and the section of Rijeka (31.55 km2), that bordered to the national park, were the 2 biggest continuous denning areas in the northwestern part of study area. Availability of suitable denning sites was limited mainly by presence of roads, especially forest roads. The number and size of suitable denning areas can indicate possible habitat fragmentation in the future. Bears in Croatia den outside areas that we indicate as suitable, but there is a higher possibility of disturbance. Mild winters and early springs with little or no snow cover are regular periods for logging beech trees. At least 3 maternal dens in 1993-1994 have been disturbed due to logging and hunting. Areas identified as suitable for denning should be excluded from logging during winter until late April if maternal dens need to be protected. The density of all roads in study area was 1.91 km/km2, resulting in SIV9=0.0 (Table 1). The mean density of all roads in Delnice, abar, Rijeka, and Vrbovsko was 2.07, 1.95, 1.74, and 1.89 km/km2, respectively, resulting in SIV9=0.0 for all sections (Table 1). This density does not prevent brown bears from inhabiting the area. In the Cantabrian mountains of Spain, areas inhabited by brown bears have an average roads density of 0.71 km/km2 while areas uninhabited by bears have a mean road density of 0.85 km/km2 (Clevenger et al. 1995). The road density indicates possible human access into bear habitat, affecting the bears security (i.e., hunting, forestry operations, traffic caused mortality, tourism, and other activities). By excluding forest roads, density of main, regional, and local roads dropped to 0.83 km/km2. The highest density of non-forest roads was found in Rijeka, (1.31 km/km2), followed by Vrbovsko (0.72km/km2), Delnice (0.71km/km2), and abar (0.59km/km2). Closing forest roads would increase SIV9 to 0.45, improving SIHUMAN IMPACT to 0.73, overall HSI to 0.48, and increase the area of suitable denning sites. Habitat fragmentation did not occur neither on any section or on the 1,796-km2 of study area. The SIV10 values for each of study area sections and for entire study area were limited only by the sizes of the areas. Gorski kotar is connected with the rest of bear range in Croatia (9,800 km2), which is a part of continuous brown bear range stretching from Slovenian thorough Croatian, Bosnian, and Monte Negro parts of the Dinara Mountains (Huber and Frkovi 1993). Because this range is part of a much larger area, we assigned SIV10=1.00 (Table 1). The main road stretched through the middle of study area, and the majority of settlements were located along it. The 5-km belt stretching along the main road still had 66% forest vegetation. At least 3 bear crossing sites were identified on the road (Huber et al. This Volume) as part of travel corridors. A new highway under construction will connect the Adriatic Sea coast with the continental part of Croatia, through the middle of Gorski kotar. If all proposed measures (D. Huber, Veterinary Faculty, Zagreb, Croatia, unpubl. data.) are implemented, existing bear corridors will not be cut. The final SI values for the food, cover and human impact habitat components were 0.55, 0.09, and 0.50, respectively, resulting in an overall HSI of 0.42 (Table 2). The results of the HSI evaluation indicate good food base, especially during summer, protective cover below average, and average impact of humans resulting in an average quality of bear habitat in Gorski kotar, Croatia. We did not evaluate garbage management and brown bear management in this study. It can be assumed that each settlement (N=308) has a garbage dump located usualy in the forest. New possibilities of leasing hunting areas to hunting clubs and individuals can harm the bear population. Trophy hunting is the easiest way to compensate for leasing expenses. We conclude that brown bear habitat in Gorski kotar is still in good condition and connected with the remaining bear range on Dinara mountain. 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Rasprave i prikazi MZH-a, 4. Zagreb, Croatia. (In Croatian). Randi, E., L. Gentile, G. Boscagli, D. Huber, and H.U. Roth. 1994. Mitochondrial DNA sequence divergence among some west European brown bear (Ursus arctos) populations. Lessons for conservation. Heredity 73:480-489. Regent, B. 1980. umsko sjemenarstvo. Dokumentacija za tehniku i tehnologiju u umarstvu. Broj:79. JPC, Beograd, Yugoslavia. (In Serbian). Roth, H.U., and D. Huber. 1986. Diel activity of brown bears in Plitvice Lakes National Park, Yugoslavia. Int. Conf. Bear Res. and Manage. 6:177-181. Sas Institute, Inc. 1994. SAS(r) Propriatery Software Release 6.10 TS019. Licenced to SRCE, Site 0082452003. SAS/Stat( Users Guide Version 6, Fourth Edition, Volume 1, Cary, NC: SAS Institute Inc; 1989., 943pp. Schooley, R.L., C.R. McLaughlin, G.J. Matula, Jr., and W.B. Krohn. 1994. Denning chronology of female black bears: effects of food, weather, and reproduction. J. Mammal. 75(2):466-477. Servheen, C. 1990. The status and conservation of the bears of the world. Int. Conf. Bear Res. and Manage. Monogr. Series No. 2. 32pp. Taberlet, P., and J. Bouvet. 1994. Mitochondrial DNA polymorphism, phylogeography, and conservation genetics of the brown bear Ursus arctos in Europe. Proc. R. Soc. Lond. B. 255:195-200. Trinajsti, I. 1972. Fitocenoloaka istra~ivanja bukovih auma Gorskog kotara. Acta Bot. Croat. 31:173-180. (In Croatian). ------, 1995. Virgin forest, natural forest and exploited forest. A comparision of the floristical structure. Sauteria 6:109-132. U.S. Fish and Wildlife Service. 1981. Standards for the development of habitat suitability index models 103 EMS. U.S. Dep. Inter. Fish and Wildl. Serv., Div. Ecol. Serv., Washington, D.C. 162pp. Van Manen, F., and M. Pelton. 1993. Data-based modelling of black bear habitat using GIS. in I. Thompson, ed. Proceedings of the 21st. International Union of Game Biologists Congr. (1):323329. ------, and ------. 1994. Habitat evaluation to study the feasibility of reestablishing a black bear population. Int. Conf. Bear Res. and Manage. 9(1):438-457. Vukeli, J. 1985. Doprinos fotointerpretacijske analize vegetacijskom istra~ivanju `umskih zajednica Nacionalnog parka Risnjak. Glasnik za aum. pokuse 23:95-140. (In Croatian). Table 1. Suitability indices (SI) of brown bear habitat variables for Gorski kotar, Croatia, 1995. Suitability indices (SI)SectionSIV1aSIV2 bSIV3 cSIV4 dSIV5 eSIV6 fSIV7 gSIV8 hSIV9 iSIV10 jDelnice0.470.501.001.000.450.270.190.000.000.28 abar0.440.501.001.000.640.270.060.000.000.11Rijeka0.120.501.001.000.570.310.200.160.000.21Vrbovsko0.880.501.001.000.590.250.100.000.000.11OVERALL0.480.501.001.000.560.280.140.040.001.00 Legend: a spring food availability b spring food diversity c summer food availability d summer food diversity e autumn food availability f autumn food productivity g protective cover in foraging period h protective cover in denning period i density of all roads j habitat fragmentation Table 2. Component indices and overall brown bear habitat suitability index (HSI) for Gorski kotar, Croatia, 1995. SectionSIFOOD1SICOVER2SIHUMAN IMPACT3HSI4Delnice0.500.100.140.31 abar0.560.030.060.30Rijeka0.490.180.110.32Vrbovsko0.590.050.060.32OVERALL0.550.090.500.42 Figure captions Fig. 1 Location of Croatia (top), the Gorski kotar study area (middle), and the study area sections (bottom). Fig. 2 Structure of the brown bear habitat suitability index (HSI) model for the Gorski kotar, Croatia. Fig. 3 Relationships of suitability indices (SI) and 13 habitat variables used in the brown bear HSI model. (A) Spring food availability. (B) Spring food diversity. (C) Summer food availability. (D). Summer food diversity. (E) Autumn food availability. (F) Autumn food productivity. (G) Protective cover in foraging period. (H) Sites for denning. (I) Distance of denning areas from road. (J) Distance of denning areas from settlement. (K) Protective cover in denning period. (L) Density of all roads. (M) Habitat fragmentation. 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