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Lynx Lynx lynx returned to a semi-natural, human dominated landscape in the Jura Mountains in France and Switzerland after reintroductions in the early 1970s. Controversy has resulted from lynx attacking sheep and preying on game species such as roe deer Capreolus capreolus and chamois Rupicapra rupicapra. We review the history of the lynx, the transition of the landscape and fauna in the Jura Mountains, and recent findings from long-term field studies on the species. Possible threats to the survival of the population are assessed. The ecological conditions for the existence of the lynx in the Jura Mountains have improved since the species was eradicated in the 19th century. Both habitat and prey base are suitable for maintaining the population. Immediate threats include traffic accidents and illegal killings. Long-term threats include small population size and genetic impoverishment as a result of the post-reintroduction bottleneck. We recommend conservation and management that involve local people and cooperation at national and international scales. Fragmentation of the habitat and the management system should be avoided, and landscape linkages from the Jura Mountains to adjacent mountain ranges should be established to promote a lynx metapopulation.
Lynx were reintroduced to the Jura Mountains in the mid-1970s. A first retrospective update of the situation in France and Switzerland was undertaken 10 years later. Since then, real-time collection of occasional observations has been going on in both countries. The monitoring methods have been standardised since the beginning of the 1990s using, among other methods, a network of observers. During 1972-2001, 862 observations were collected in Switzerland and in this paper, these are used to describe the colonisation of the Swiss Jura Mountains and the present distribution of lynx. A comparison with the results of French researchers shows that during the first years of recolonisation, most of the observations were collected in the Swiss part of the Jura Mountains. Today, the French territory has become the core area of the population and includes about ⅔ of the population. The population has recovered after a period of reduced presence at the beginning of the 1990s. The range occupied permanently by lynx in the Swiss Jura Mountains is estimated to be 2,100 km2, representing a potential population of 17-23 resident individuals. The range occupied permanently by lynx in the Jura Mountains on both sides of the border is estimated to be 7,100 km2. Depending on the degree of saturation in the population, this represents a potential population of 56-78 resident individuals. Compared to existing habitat suitability models, most of the suitable habitat has already been occupied. In the future more importance will be given to the exchange of information across the international border and the participation of local people in the survey. I recommend that active monitoring using camera-traps be carried out periodically as a supplement to the ongoing passive surveillance system.
A total of 18 Eurasian lynx Lynx lynx were radio-tagged between March 1988 and June 1998 in the Swiss Jura Mountains, and during 1995-1997 eight animals were radio-tagged on the French side of the mountain chain. Adult males occupied larger long-term home ranges than adult females (283 km2 vs 185 km2). Neighbouring males shared 7.3% of their home ranges and females 0.2%. The mean distance between males and females living in the same area for fixes taken the same day was 10.94 ± 8.61 km, underlining the solitary character of the species. Consecutive individual annual home ranges overlapped 71.7 ± 7.3% for females and 77.5 ± 7.9% for males, indicating high spatial stability over time. In the Swiss study area, two adult animals were followed for seven and nine years, respectively, and another two lynx were observed in the study area for nine years. Range size did not vary across three distinct periods, P1-P3, but the sex ratio did. Generally, males covered the ranges of 1-2 females, but during the second period, P2, the range of a single male overlapped with those of six females. Dead females were all immediately replaced, but dead males were not. Two poached males were only replaced after three and five years, respectively. Population density, ranging within 0.7-0.8 adult resident lynx/100 km2, did not vary significantly over time in Switzerland. Including kittens and subadults, the density was 1.1-1.6 lynx/100 km2. Our study in the Jura Mountains indicated that there is long-term stability in the social and spatial structure of the lynx population, but this stability was temporarily disturbed by the lack of adult resident males.
We radio-collared a total of 29 lynx in the Swiss Jura Mountains and collected data on demography during 1988-1998. We were able to observe 10 2-14 year-old females for 32 female years. Lynx kittens were born between 12 May and 13 June (26 May ± 9 days). One female gave birth to a litter on 26 August after she had lost her first litter born in May. The average litter size was 2.00 ± 0.75 kittens (range: 1-3). The sex ratio of 1.67 females:1 male did not differ significantly from the expected ratio of 1:1. On average, 81% of adult females reproduced each year. The overall reproduction rate was 1.67 kittens/female year. Of 49 kittens, 43-49% survived until independence at the age of about 10 months. The survival rate of subadults and adults was 53 and 76%, respectively. Litter size and survival of kittens and subadults varied considerably between three observation periods, i.e. P1-P3 (P1: 1988-1991, P2: 1992-1994, P3: 1995-1997), characterised by substantial changes in the social structure of the population. During P2 only one resident male was present in the main study area, and during this period, the average litter size was lowest with 1.50 kittens only. The survival of kittens was with 37-44% lowest as well, mainly because of the high proportion of litters where all kittens were lost. However, the survival of subadults was highest during P2, when observed animals survived to adulthood. During 1974-2002, 124 mortalities were documented for the whole of the Jura Mountains. Human related mortalities were responsible for 70% of known losses. An estimation based on the radio-collared lynx suggested that poaching may have been responsible for as much as 32% of total mortality. In contrast to other study areas, traffic accidents were very important in the Jura Mountains. They might represent as much as 29% of mortality. During our 10-year study period, no significant spatial expansion of the population was recorded in Switzerland. We suppose that the high human-related mortalities limited the population, which also would explain the lack of a numerical response to an increase in prey abundance observed during the same period in the study area in Switzerland.
Anja Molinari-Jobin, Fridolin Zimmermann, Andreas Ryser, Christine Breitenmoser-Würsten, Simon Capt, Urs Breitenmoser, Paolo Molinari, Heinrich Haller, Roman Eyholzer
To analyse the factors responsible for the interplay of Eurasian lynx Lynx lynx predation and home-range size, we reviewed patterns of lynx predation in Switzerland by comparing the prey spectrum of lynx in five studies performed in the following study areas: the northwestern Alps, where lynx were studied both in the 1980s and 1990s, the central Alps, the Jura Mountains, and northeastern Switzerland. We then compared home-range size of female lynx with two indirect measures of prey abundance, roe deer Capreolus capreolus and chamois Rupicapra rupicapra harvested per km2 and habitat suitability for roe deer and chamois as derived from a GIS model. Lynx diets were similar among sites. Roe deer and chamois made up 90% of prey items in all five studies. Comparing the proportion of roe deer and chamois in the diet with availability, Manly's preference indices indicated selective predation in all studies. Roe deer were preferred over chamois in all areas except in the Jura Mountains where relatively few chamois were present. Predation was least selective in northeastern Switzerland, where the initial phase of recolonisation by lynx was studied. Variation in prey availability is often identified as an important factor explaining intraspecific variation in home-range size. Due to differences in roe deer and chamois abundance from one study area to another, we expected female lynx home ranges to decrease with increasing prey abundance. The predictors for Minimum Convex Polygon (MCP) and Kernel home-range estimators differed. MCP home-range sizes were best explained by the interactions of study with the number of locations per lynx, roe deer harvested per km2, and good roe deer habitat, whereas Kernel home-range sizes were best explained by the interactions of study with good roe deer habitat, good chamois habitat, and the interaction of good roe deer and chamois habitat plus an additive effect of the study. Contrary to our expectations, there was no simple correlation of prime roe deer and chamois habitat nor between the number of roe deer and chamois harvested per km2 and the size of female lynx home ranges. The comparison of the five studies suggested that this expectation may only be valid if lynx populations are close to carrying capacity (e.g. the Jura Mountains and the northwestern Alps in the 1990s). For predictions of home-range size both habitat (spatial factor) and the status and dynamic of the predator/prey populations (temporal factor) need to be taken into account.
To estimate the potential population size of the Eurasian lynx Lynx lynx in the Jura Mountains and to assess possible corridors between this population and adjacent areas (the Vosges Mountains, the Black Forest and the Alps), we adapted a previously developed Geographic Information system (GIS) probability model for lynx distribution and extrapolated it over the entire mountain range. The model was based on knowledge of the habitat use and land tenure system of resident animals from the central part of the Jura Mountains, where lynx were followed by means of radio-telemetry. Corridors were computed in the GIS using a friction grid and a cost distance function. The friction value attributed to each land use variable was assessed from our observations of lynx dispersal. Our model predicts a breeding population in the Jura Mountains of 74-101 individuals and 51-79 individuals when continuous habitat patches of < 50 km2 are disregarded. The Jura population lies within the range of a viable population if only demographic aspects are taken into account, but is rather small from a genetic point of view. Genetic viability would be assured if the Jura lynx population were part of a larger metapopulation. Potential corridors exist from the Jura Mountains to the Vosges Mountains, the Black Forest and the Alps (Chartreuse and Salève, respectively). The length of these corridors range within 7.3-37.3 km, and their costs are all within the range of radio-collared lynx roaming outside their prime habitat. The best corridor leads south to the Chartreuse, an isolated part of the French Alps, which is itself connected to the rest of the Alps by two corridors of 4.5 and 6.5 km, respectively. Observations in the Chartreuse indicate that lynx may have immigrated from the Jura Mountains, but there is no evidence for the use of northern corridors, as the species has not yet completely occupied this area. We conclude that the monitoring of the population size, its spatial expansion, and the genetic surveillance in the Jura Mountains must be continued, as the status of the population is still critical. Only good surveillance would allow the necessary conservation measures to be initiated in time.
We retrospectively investigated characteristics of den structures and den sites used by female Eurasian lynx Lynx lynx in the Jura Mountains and the northwestern Alps of Switzerland. During 1983-2000, we discovered 30 natal and 40 maternal dens belonging to 26 females. Important den structures were closed, i.e. provided good shelter, had few entrances, and measured 1 m2. Dens were found in rocky places, caves and wooden surroundings. Most dens were located in mixed forests with relatively open vegetation allowing for a visibility of 10-20 m. Contrary to our expectations, natal and maternal dens were equally exposed to human disturbance and were found in terrain which could be dangerous for the kittens. Overall, the two den types barely differed. While concealment did not seem to play a very important part at natal dens and while natal dens were almost never open structures, maternal dens were surrounded by a large number of hiding places and the dens and surroundings were rich in visual contrasts providing good camouflage options. Dens in the Alps and in the Jura Mountains were located in steeper terrain than available on average. The quality of the den sites did not seem to affect the survival of young lynx. Well-suited den sites are so abundant in the Alps and the Jura Mountains that females obviously have no problems finding good den sites.
Modelling species distribution is an important aspect of conservation ecology. Empirical models are most commonly used. However, collecting data for these models is time-consuming and expensive. Expert models may be a good alternative method, though previous studies have found mixed results. The purpose of our study was first to create an expert model and evaluate it with independent lynx data, and second to use two discrete types of experts to control for prior radio-tracking experience. Two habitat suitability expert models (scientific and local experts) were constructed in a Geographical Information System using the Analytical Hierarchy Process and Compromise Programming. The models were evaluated with lynx data, taken from the study area in the northwestern Swiss Alps, using Resource Selection Index and Spearman correlation. The correlations showed that both models fitted the data well. However, the local expert model was better (rs = 0.964, P < 0.001) than the scientific expert model (rs = 0.833, P < 0.001). The models were also evaluated in the Jura Mountains to test the local nature of the models. It was found that the local expert model performed less well (rs = 0.939, P < 0.001) than the scientific expert model (rs = 0.967, P < 0.001) as expected. Comparison between weights for each expert group revealed some interesting differences, which showed the local nature of answers and how personal experience and theoretical knowledge can lead to different answers. Our study shows that expert knowledge, and especially local knowledge, can be employed to create a good habitat suitability model. This has important implications for conservation and science because it shows not only that expert knowledge may be used when no other data exist, but also that local ‘ground workers’ should be employed more often in the development of habitat suitability models or conservation plans. However, there are limitations to the models and, as expert models are relatively new in ecology, more research is needed. Nevertheless, in a climate where there is pressure to keep up with human exploitation of natural resources and to adopt a more strategic approach to conservation, the findings of our study are encouraging.
John D. C. Linnell, John Odden, Reidar Andersen, Henrik Brøseth, Henrik Andrén, Olof Liberg, Per Ahlqvist, Pål Moa, Tor Kvam, Peter Segerström, Krzysztof Schmidt, Włodzimierz Jędrzejewski, Henryk Okarma
Monitoring of lynx Lynx lynx populations in Scandinavia is largely based around unreplicated minimum counts of family groups, i.e. adult females with dependent kittens. When observations cannot be separated from each other on the basis of back-tracking in the snow it is desirable to use a distance rule to separate observations of groups that are so far apart that they are unlikely to be derived from the same group. We have analysed radio-telemetry derived movement data from five study areas, corresponding to three ecologically different regions, in Scandinavia, and included data from Poland for comparison. We derive examples of two different distance rules; one is a static rule based on home-range size and length which is suitable for observations accumulated during a whole winter, and the other is a dynamic rule suitable for observations collected within 1-7 days of each other. Because of inter-study area variation in home-range size and movement there is a need to use different rules in different regions. Within Scandinavia, average maximum home-range lengths varied from 28 to 54 km, and average maximum daily distances travelled varied from eight to 16 km in the three regions. This implies that locally collected movement data are a prerequisite for application of this type of methodology.
Counts of tracks that passively accumulate on a suitable substrate are commonly used to derive indices of large carnivore abundance. In this study we evaluate the suitability of a similar survey using multiple 3-km long transect-lines to detect changes in population size for Eurasian lynx Lynx lynx in central Norway. We used GIS methods to simulate the crossing of transect lines by lynx using real telemetry data from the study area. We compared the effect of transect-line placement (deliberate vs random), transect-line density, and the number of nights over which tracks can accumulate in the snow. For each scenario we evaluated both the probability of detecting lynx that are present in the survey area, and the power of the index to detect changes between consecutive surveys. Deliberately placed lines performed significantly better than randomly placed lines, and as expected, increases in line density and the period of track accumulation improved the outcome. Using three nights of track accumulation and the highest density of deliberately placed lines that we simulated (1/38 km2) indicated that both the probability of detecting individual lynx present within the survey area, and the power to detect a 33% change in population size between two surveys, were > 80%.
Data on the age, sex and condition of roe deer Capreolus capreolus killed by Eurasian lynx Lynx lynx and human hunters were collected in two study areas in the counties of Nord-Trøndelag and Hedmark in central and southeastern Norway, respectively. Data on the age and sex structure of the standing population were also collected. No differences in the age or sex structure of roe deer killed by lynx were found between the two study areas or between years with differing snow depths which was expected to affect age-class vulnerability. The profile of 151 lynx-killed roe deer (24% adult males, 44% adult females, 11% male fawns and 21% female fawns) was not statistically different from that of the standing population. In contrast, hunters killed a significantly larger proportion of adult animals, especially males (44% adult males, 28% adult females, 15% male fawns and 13% female fawns). The detailed age structure of lynx killed and hunter killed animals was not different within the adult age class. The body condition of roe deer declined during late winter (more so for males than for females), however, there was no difference in the condition of roe deer killed by lynx or hunters during early winter (the hunting season ends on 24 December). The overall picture is typical of a stalking predator that has few options to select individuals based on condition, age or sex in a prey species with no sexual dimorphism and which is relatively small in relation to the size of the predator (lynx are 50-70% the size of an adult roe deer). Human hunters, on the other hand, are clearly harvesting a non-random section of the population. The result is that hunting does not replicate natural predation, although lynx predation is likely to have a higher per capita impact on roe deer growth rates.
We studied the diet of red fox Vulpes vulpes in the presence of lynx Lynx lynx in the southern boreal forest of Sweden during 1999-2002 using scat analyses. Diet composition and food niche breadth was compared to an identical study conducted in the same area during 1974-1979 prior to the recolonisation by lynx. The consumption of roe deer Capreolus capreolus was higher in the presence of lynx, even compared to the particularly harsh winter in 1976/77 when roe deer suffered serious starvation related mortality. Scavenging on roe deer replaced feeding on presumably less profitable food such as fish and reptiles, invertebrates and vegetables. The winter food niche was narrowed. The frequent consumption of roe deer could have been a combined effect of a stable supply of deer carrion and a low abundance of other prey species. We thus conclude that presence of lynx feeding on roe deer creates an important, stable food supply for red foxes.
Home-range size of Eurasian lynx Lynx lynx was studied in two different mountain ranges of the Polish Carpathians during 1999-2004. We live-trapped and radio-collared five adult lynx (two males and three females). One female was probably killed only 10 days after being radio-collared. Home-range size of radio-collared individuals calculated for the whole period of radio-tracking (MCP with 100% observations) was 164 km2 for male M1 from the Beskid Sądecki Mountains, and it was 124 and 190 km2, respectively, for the two females F1 and F2 and 199 km2 for male M2 in the Bieszczady Mountains.
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