ࡱ> 7 bZbjbjUU jr7|7|tBlzzzzzzz$^ ^ ^ P $ $(t (* (R R R R $ @ 9';';';';';';'$* >,_'zP R R P P _'xzzR R d(xxxP VzR zR 9'xP 9'x xVe&@zz'R 05+^ ~& '4z(0(&T,$T,'xzzzz Causes of wolf mortality in Croatia during 1986-2001 uro Huber1*, Josip Kusak1, Alojzije Frkovi2, Goran Gu~vica1, and Tomislav Gomer i1 1Biology Department, Faculty of Veterinary Medicine, University of Zagreb, Croatia 2Rijeka, Croatia HUBER, , J. KUSAK, A. FRKOVI, G. GU}VICA, T. GOMER I: Causes of wolf mortality in Croatia during 1986-2001. Vet. Arhiv 71, xxx-xxx, 2001. ABSTRACT We recorded a total of 92 dead grey wolves (Canis lupus) in Croatia during 1986-2001: 60 (65.2%) were shot, 18 (19.6%) died in traffic accidents, the causes of death of 6 (6.5%) wolves remained unknown, 5 (5.4%) had rabies, and one each was killed by axe, by hay-fork, and by other wolves. Among dead wolves the share of females was 56%, the mean age was 1.9 years, and the mean mass of adult wolves was 31.4 kg. We analyzed separately the last 10 years (1986-1995) before, and in the first 6 years (1996-2001) after the legal protection of the species (15 May 1995), when 30 and 62 wolves died, respectively. The yearly mortality grew from 3.0 before protection to 10.3 after protection (3.4 times). This increase in mortality indicates no enforcement of legal protection, as well as the increase of the animosity of local people. It was also partly influenced by a wolf population growth and seems that the recorded mortality is sustainable. We propose to adapt the legislation to allow controlled hunting of wolves. That will likely lead to decreased animosity and will keep the wolf population safe of uncontrolled hunting. Key words: grey wolf, Canis lupus, mortality, Croatia --------- Contact address: Prf. Dr. uro Huber, Biology Department, Faculty of Veterinary Medicine, University of Zagreb, Heinzelova 55, 10000 Zagreb, Croatia, Phone: 385 1 2390141; Fax: 385 1 2441390; E-mail:  HYPERLINK "mailto:huber@vef.hr" huber@vef.hr  Introduction Mortality of individuals within the population is one of the basic parameters of its dynamics (KEETON and GOULD, 1986). Knowledge of mortality rates, its causes and circumstances of occurrence are one of the first steps in saving an endangered population. In addition, the monitoring of mortality over an extended period of time allows the insight into the population trend. Grey wolf (Canis lupus) is the large carnivore species historically spread around the whole Northern hemisphere. It inhabited the entire area of todays Croatia until about 1895. The decrease of range and numbers continued through the 20th century. The low numbers were reached in late 1980s (HUBER et al., 1999). On 15 May 1995 the wolf in Croatia gained full legal protection (ANONYMOUS 1995). The mortality of wolves in Croatia has been recorded for the period 1945 through 1986 (FRKOVIC et al. 1992). We continued this survey with special emphasis on the causes of mortality and change of legal status of wolves. The objectives of this study were to: (1) investigate the wolf mortality rates since 1986, (2) compare the mortality after the species got protection in 1995, (3) determine the main causes of deaths, and (4) evaluate the effects of legal protection and to propose the measures for more effective conservation of wolves in Croatia. Materials and methods Through the network of local experts (foresters, hunters, wolf damage inspectors) we collected the information about cases of wolf deaths in Croatia. Each report has been investigated for confirmation. Locality has been recorded as well as all anecdotal information. Whenever possible we retrieved the body of dead animal. The body has been examined through morphometric, anatomical and pathological procedures. The whole skeleton and the samples of various tissues, the food items from the digestive tract and the parasites found have been collected and stored in the Collection of large carnivores in the Biology Department of the Veterinary Faculty in Zagreb. To comply with legal restrictions concerning the protected species, the retrieval of wolf bodies and the storage of their parts has been approved by the Croatian Ministry of Environmental Protection and Physical Planning. The Croatian Veterinary Institute has determined the cases of rabies in wolves. The bodies of rabid wolves we did not retrieve. Two rabid wolves were found dead, two were shot and one was killed with a stick, but here all were classified as rabies was the cause of death. We are aware that a certain number of dead wolves could have remained unrecorded. Due to our relatively even efforts over the time we believe that the trend of recorded wolf deaths reflects the actual mortality trend. The change of legal status of wolf in 1995 could have created more unrecorded dead wolves, especially the ones that were shot. The bottom line is that our records represent the minimum numbers of dead wolves in Croatia. The data have been analyzed in respect of the trend of mortality over years, territorial distribution, causes of mortality, sex, age and mass distribution of dead animals. Results In the period from 1986 through 2001 we recorded the deaths of 92 wolves. In 33 (35.9%) cases the body has been found, retrieved and thoroughly studied. In 6 (6.5%) cases we recorded the natural deaths causes: 5 (5.4%) cases of rabies and one (1.1%) killed by other wolves. Six (6.5%) death causes remained unrevealed, and the remaining 80 (87.0%) wolves died due to human activities: most were shot (N=60, 65.2%), one was killed by axe (1.1%), one (1.1%) by hay-fork, and the remaining 18 (19.6%) died in traffic. Most of dead wolves were recorded in inland Dalmatia and in Gorski kotar with a few scattered in Lika and other areas (Fig. 1). The sex was known for 50 dead wolves: 28 (56%) were females and 22 (44%) were males. The age estimation could have been done in 36 cases and it ranged from 0.3 to 6.0 years; the mean age was 1.9 and median was 2 years. The mass was measured for 38 wolves and it ranged from 7.0 kg to 47 kg; mean was 28.4 kg. When only adult wolves (1.5 years or more and/or minimum 25 kg) were used for calculation, the mean mass of 27 measured animals was 31.4 kg. There was an increasing trend of mortality during 1990 to 2000 (Fig.2). The mortality was the lowest in the late 1980s and early 1990s, with zero recorded mortality in 1987 and 1989. In 1988 and 1990, each, one dead wolf has been recorded . After 1991 the mortality steadily grew up to 15 in 1998 and 1999. During the last 10 years (1986 to 1995) before protection a total mortality of 30 wolves has been recorded: 3.0 wolves per year. In the first 6 years of full legal protection 62 wolves were found dead; 10.3 per year, i.e.3.4 times more than before protection. While gun was responsible for 29 of 30 wolves (96.7%) killed until 1995, and for 31 wolves (50.0% of 62) since 1996. The only wolf that was not shot prior to 1995 died of rabies, while all other recorded causes of death occurred after 1995. Discussion The main cause of wolf mortality in Croatia in the surveyed period was related to man, comprising mostly intentional killing by gun: 65.2% in the whole period and 50.0% since protection. Similarly, man caused the major part of mortality in the wolf population in Minnesota (USA) even after its protection (1972. 1977.): 59% (FRITS and MECH, 1981), and 80% (FULLER, 1989). Natural causes of wolf mortality in Minnesota totaled 14% combined of 3% by diseases and 11% by other wolves (FULLER, 1989). In another study in Minnesota the intraspecific aggression accounted for 24% mortality (FRITS and MECH, 1981). The highest recorded intraspecific killing among wolves has been recorded in Denali National Park in Alaska (USA) ranging 39%-65% of all mortality causes (MECH et al., 1998). That is a clear sign that the influence of man is neglectable or even absent there. Wolves tend to self-regulate their numbers when other causes of mortality are low. We found only 6.5% of wolves that died of natural causes, and only one intraspecific killing among them, indicating a strong man-caused mortality. Five rabid wolves comprise 83% of all such cases (N=6) recorded in Croatia since 1977 when the rabies outbreak reached this area ( A , personal information). On the population level this disease could not be counted as a major regulatory factor among wolves. The question is how big mortality rate the wolf population can sustain before the population decline begins. One of the variables is the availability of food supply, but when it is not severely limited the reproductive potential of wolves allows relatively high losses. On the Kenai Peninsula (Alaska, USA) the wolf population started to decline after two consecutive years of >40% removal by hunting, but regained numbers when killing was reduced to <35% per year (GASAWAY et al., 1983). FULLER (1989) believes that the wolf population will remain stabile with a total yearly mortality of about 35%. In Croatia in the period 1996-2001 (first full six years of protection) the average yearly mortality was 10.3 wolves. There are indications that the population has been growing in the same period resulting in an estimated maximum of 170 individuals in 1999 (KUSAK, unpublished data). The recorded mortality obviously did not prevent the population growth. Theoretical sustainable loss in the population of 170 animals could be up to 60 wolves per year. However, if fewer wolves are removed from the population one should not expect that the balance means the net population growth. Assuming that the sex ratio in the wild population is 1:1 we expected to find the similar ratio of sexes in dead wolves. However, the share of dead females was 56.0%. We do not know the sex ratio of our living wolves but in some populations it was shown to be higher (64%) on the male side (MECH, 1970). This author believes that the higher aggression of males for food within the pack increases the survival of male pups compared to their sibling sisters. Additionally, he speculates that the prevalence of male wolves may act as an additional mechanism of wolf population self-control. Consequently, the higher female mortality recoded in Croatia might indicate the higher male survival rates. Mean age of the dead wolves of 1.9 years (median = 2 years) indicates the most vulnerable age category. It is known that wolves most commonly disperse from the parental pack in their second year of life (FULLER, 1989; MECH 1970; MECH et al., 1998). The wolves that left the pack have to search for a new territory, for prey and for sexual partner. Often they must go to less favorable areas or do enter into another wolf pack territory. In natural conditions the young dispersers are most commonly killed by other wolves while in man dominated areas they get killed by man (MECH et al., 1998). The recorded range of masses of dead wolves (7-47 kg) and the mean mass of adult wolves 31.4 kg are important data to mitigate misbelieves that average wolves may be over 50 kg. The increase (3.4 fold) of number of dead wolves in the period after 1995, i.e. after the legal protection, indicates the no acceptance (BATH and MAJI, 2000) and no enforcement of positive legislation. Additional concern is the increase of traffic caused wolf mortality: 15 (24.2%) since 1996, and within this period in the last two years (2000 and 2001) even 6 of 12 (50.0%) wolves died in collisions with vehicles. Traditionally, in the period 1945 - 1994 (KUSAK et al., 2000) 3.6% (20 of 560) of total wolf mortality resulted from traffic accidents. FRKOVI et al. (1988) found the average yearly mortality of wolves in Gorski kotar (Croatia) in the period 1945 to 1986 to be 13 individuals (range 3-27) (Fig. 3). In that period the wolves were hunted with no legal limitations, including traps and poisons, and even bounties awarded until early 1970s. It may be assumed that the number of killed wolves reflected the population size and trend in Gorski kotar, as well as in the whole Croatia. In this period the least numbers of wolves were killed in the years: 1945, 1955, 1970 and 1985 (3 to 5 per year), what represents intervals of 15 years. In spite of continuous hunting effort the numbers of killed wolves varied over time what indicates the population size fluctuation, as well as the presence of other factors influencing the population trend aside of man. The most likely factor was the prey availability. This study revealed the next period of low wolf mortality and population size in the years 1986 to 1990 (Fig. 3). The future wolf population trend and fluctuations are hard to predict. We conclude that the 3.4 times high increase of mortality of wolves after the species got legal protection indicates no enforcement of legal protection, as well as the increase of the animosity of local people. The mortality increase was also partly influenced by a wolf population growth but seems that the recorded mortality is sustainable. We propose to adapt the national legislation to allow controlled hunting of wolves and keep it strictly within the sustainable limits. That will likely lead to decreased animosity and will keep the wolf population safe of uncontrolled hunting. ----------- Acknowledgements We wish to thank to numerous local experts who helped in collecting of information and the bodies of dead wolves. Among them are D. Bosiljevac, I. `upe, S. Koki and J. Malnar. We also thank to many students of the Veterinary Faculty for their volunteering work in the field and in the laboratory in Zagreb. The study has been financed by the Croatian Ministry of Science and Technology with additional support by Bernd Thies Foundation and EURONATUR. References ANONYMOUS (1995): Pravilnik o zatiti pojedinih vrsta sisavaca (Mammalia). Narodne novine 31, 968. BATH, A, A. MAJI (2000): Human dimension in wolf management in Croatia. Report by Large Carnivore Initiative for Europe. 166 pp. FRITS, S. H., L. D. MECH (1981): Dynamics, movements, and feeding ecology of a newly protected wolf population in northwestern Minnesota. Wildlife Monographs 80 pp. FRKOVIC, A., R. RUFF, L. CICNJAK, D. HUBER (1992): Wolf mortality during 1946-86 in Gorski kotar, Croatia. IUGB Congress. 18, 353-358. FRKOVI, A., R. L. RUFF, L. CICNJAK, . HUBER (1988): Ulov vuka u Gorskom kotaru u razdoblju od 1945 do 1986 godine, umarski list 112, 519-530. FULLER, T. K. (1989): Population dynamics of wolves in North-Central Minnesota. Wildlife Monographs 105, 41pp. GASAWAY, W. C., R. O. STEPHENSON, J. L. DAVIS, P. E. K. SHEPPERD, O. E. BURRIS (1983): Interrelationship of wolves, prey, and man in interior Alaska. Wildlife Monograph 84, 50 pp. HUBER, ., J. KUSAK, D. KOVA I, A. FRKOVI, J. RADOVI, }. `TAHAN (1999): Privremeni plan gospodarenja vukom u Hrvatskoj. Dr~avna uprava za zaatitu prirode i okoliaa, 76 pp. KEETON, W. T., J. L. GOULD (1986): Biological science. W. W. Norton & Company, Inc., 500 Fifth Avenue, New York, 1175 pp. KUSAK, J., HUBER, D., FRKOVI, A. (2000): The effects of traffic on large carnivore populations in Croatia. Biosphere Conservation 3, 35-39. MECH, L. D. 1970. The wolf. The ecology and behavior of an endangered species. Univ. of Minnesota press, Minneapolis. 384 pp. MECH, L. D., L. G. ADAMS, T. J. MEIER, J. W. BURCH, B. W. DALE (1998): The wolves of Denali. University of Minnesota Press, 227 pp. HUBER, D, J. KUSAK, A. FRKOVI, G. GU}VICA, T. GOMER I: Uzroci smrtnosti vukova u Hrvatskoj tijekom 1986-2001. Vet. Arhiv 71, xxx-xxx, 2001. SA}ETAK Od 1986. do 2001. zabilje~ena je smrt ukupno 92 siva vuka (Canis lupus) u Hrvatskoj: 60 (65.2%) su ustrijeljena, 18 (19.6%) je poginulo u prometnim nezgodama, za 6 (6.5%) vukova je uzrok smrti ostao nepoznat, 5 (5.4%) je imalo bjesnou, a po jedan su ubijeni sjekirom, vilama ili od drugih vukova. Udio ~enki meu mrtvim vukovima je bio 56%, prosje na dob je bila 1.9 godina, a prosje na masa 31.4 kg. Posebno je analizirano posljednjih 10 godina (1986-1995) prije, te prvih 6 godina (1996-2001) nakon zakonske zaatite vrste (15. svibanj 1995.), a u kojim je razdobljima stradalo 30 odnosno 62 vuka. Godianja smrtnost je porasla sa 3.0 prije zaatite na 10.3 nakon zaatite (3.4 puta). Ovaj porast pokazuje da se zakon ne provodi, kao i porast neprihvaanja vuka od lokalnog stanovniatva. Djelom je porast smrtnosti povezan i sa porastom populacije vuka, te se ini da je veli ina zabilje~ene smrtnosti podnoaljiva za populaciju. Predla~e se prihvaanje propisa koji bi omoguavali ograni eni i kontrolirani lov vukova. To bi moglo dovesti i do smanjenja mr~nje prema vuku i posljedi nog smanjenja rizika za populaciju od nekontroliranog ilegalnog ubijanja. Klju ne rije i: sivi vuk, Canis lupus, smrtnost, Hrvatska  Figure 1. The map of Croatia showing the distribution of mortality sites of wolves during 1986 - 2001.  Figure 2. Wolf mortality in Croatia during 1986 2001  EMBED QuattroPro.Chart  Figure 3: Wolf mortality in Croatia during 19452001*. Line indicates the trend shown as smoothed average of the two years before and after.  * Data until 1986 are from FRKOVI et al. 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