Little is known on the ecology and behaviour of the alpine mountain hare
(Lepus timidus). Between 1996 and 1997, we analysed by radiotracking the pattern of space
use of 8 mountain hares from the Swiss Alps. We estimated home range size using both the
kernel density estimator and the minimum convex polygon. We found smaller ranges (38
ha) compared to those reported for the species in boreal or arctic habitats, but similar to
ranges in Scotland. Hares did not use a centre of major activity (core area) and showed high
home range overlap, confirming their non-territorial behaviour. Smaller ranges were used
during winter compared to the other seasons, whilst no difference in size was found
between sexes

Bisi, F.

Gamboni, A.-S.G.

Martinoli, A.

Masseroni, E.

Nodari, M.

Preatoni, D.G.

Tosi, G.

Wauters, L.A.

ABSTRACT- Little is known on the ecology and behaviour of the alpine mountain hare (Lepus timidus). Between 1996 and 1997, we analysed by radiotracking the pattern of space use of 8 mountain hares from the Swiss Alps. We estimated home range size using both the kernel density estimator and the minimum convex polygon. We found smaller ranges (38 ha) compared to those reported for the species in boreal or arctic habitats, but similar to ranges in Scotland. Hares did not use a centre of major activity (core area) and showed high home range overlap, confirming their non-territorial behaviour. Smaller ranges were used during winter compared to the other seasons, whilst no difference in size was found between sexes

Anne-sophie Genini Gamboni

Francesco Bisi

Elisa Masseroni

Mosè Nodari

Damiano G. Preatoni

Lucas A. Wauters

Adriano Martinoli

Guido Tosi

CiteSeerX

HOME RANGE DYNAMICS OF MOUNTAIN HARES (LEPUS TIMIDUS) IN THE SWISS ALPS

Little is known on the ecology and behaviour of the alpine mountain hare
(Lepus timidus). Between 1996 and 1997, we analysed by radiotracking the pattern of space
use of 8 mountain hares from the Swiss Alps. We estimated home range size using both the
kernel density estimator and the minimum convex polygon. We found smaller ranges (38
ha) compared to those reported for the species in boreal or arctic habitats, but similar to
ranges in Scotland. Hares did not use a centre of major activity (core area) and showed high
home range overlap, confirming their non-territorial behaviour. Smaller ranges were used
during winter compared to the other seasons, whilst no difference in size was found
between sexes

Gamboni, A. S. G.

NODARI, MOSE'

PREATONI, DAMIANO

WAUTERS, LUCAS ARMAND

MARTINOLI, ADRIANO

TOSI, GUIDO

Archivio istituzionale della ricerca - Università dell'Insubria

Hystrix It. J. Mamm. (n.s.) 19 (2) (2008): 77-83  HOME RANGE DYNAMICS OF MOUNTAIN HARES (LEPUS TIMIDUS) IN THE SWISS ALPS  ANNE-SOPHIE GENINI GAMBONI1, FRANCESCO BISI2,  ELISA MASSERONI2,3, MOSÈ NODARI2, DAMIANO G. PREATONI2, LUCAS A. WAUTERS2, ADRIANO MARTINOLI2, GUIDO TOSI2  16714, Semione, Switzerland 2Department Environment-Health-Safety, Universiy of Insubria Varese, Via J.H. Dunant 3, 21100 Varese, Italy 3Institute Oikos NGO Via Crescenzago 1, 20134 Milano, Italy  Received 19 August 2008; accepted 24 November 2008  ABSTRACT - Little is known on the ecology and behaviour of the alpine mountain hare (Lepus timidus). Between 1996 and 1997, we analysed by radiotracking the pattern of space use of 8 mountain hares from the Swiss Alps. We estimated home range size using both the kernel density estimator and the minimum convex polygon. We found smaller ranges (38 ha) compared to those reported for the species in boreal or arctic habitats, but similar to ranges in Scotland. Hares did not use a centre of major activity (core area) and showed high home range overlap, confirming their non-territorial behaviour. Smaller ranges were used during winter compared to the other seasons, whilst no difference in size was found between sexes.  Key words: Lepus timidus, mountain hare, home range, core area, alpine ecosystem  RIASSUNTO - Dinamica dell’uso dello spazio della Lepre bianca (Lepus timidus) nelle Alpi Svizzere. Le informazioni relative all’ecologia e al comportamento della lepre alpina (Lepus timidus) sono ad oggi scarse. In questo studio abbiamo analizzato l’utilizzo dello spazio di una popolazione di lepre bianca sulle Alpi Svizzere. Tra il 1996 e il 1997 sono stati marcati con radio collare 8 individui di lepre alpina. L’home range è stato calcolato utilizzando lo stimatore di densità kernel (KDE) e il metodo del minimo poligono convesso (MCP). L’ampiezza degli home range (38 ha) è risultata inferiore a quella riportata per la specie in habitat boreali e artici, ma simile a quella riscontrata in Scozia. All’interno dell’home range non è stato rilevato alcun centro di maggiore attività (core area) ed è stata evidenziata una notevole sovrapposizione tra gli stessi, confermando la non territorialità della specie. Le aree frequentate in inverno sono risultate più piccole rispetto alle altre stagioni e non sono state riscontrate differenze tra i sessi.  Parole chiave: Lepus timidus, Lepre alpina, home range, core area, ecosistema alpino  INTRODUCTION  The mountain hare (Lepus timidus Linnaeus, 1758) is a Palearctic relict species with a wide distribution area, from Ireland and Scandinavia in the west, across Siberia, Mongolia and China to the northern islands of Japan in the east (Angerbjörn and Flux, 1995). In Europe, isolated alpine 157ystrix It. J. a . 19(2) 2008: 157-163Genini Gamboni et al.  populations of Lepus timidus occur above 1300 m a.s.l. in the mountains of France (Couturier, 1964), Germany, Switzerland, Austria, Slovenia (Isa-kovic, 1970) and Italy (Mitchell-Jones et al., 1999). Although the ranging and spacing behaviours of the mountain hare have been studied in Scotland (Hewson, 1976; Hewson and Hinge, 1990; Hulbert et al., 1996) and Scandinavia (Hiltunen et al., 2004; Dahl, 2005a, b; Kauhala et al., 2005; Hiltunen, 2006), little is known about the social structure and home range characteristics of alpine mountain hare Lepus timidus varronis. In contrast with the large, contiguous boreal or arctic habitats occupied by mountain hares in the northern part of their range, the distribution of suitable habitats in the Alps (sub-alpine coniferous forest, alpine shrub and meadows) is heterogeneous and patchy (our unpubl. data). This difference in landscape structure is likely to affect home-range size and overlap. Here we analyse the pattern of space-use of eight mountain hares that were radio-tracked in the Swiss Alps in 1996, comparing data on home-range size and overlap with the spacing patterns described for populations from Scotland and Scandinavia.  STUDY AREA AND METHODS  Mountain hares were studied in the Blenio Valley (46°28’ N, 8°53’ E) - Ticino Canton, southern Switzerland -, between 1500 and 2050 m a.s.l. The habitat is characterised by typical alpine vegetation with alpine grasslands (59%), clearings and landslide cones (19%) and Norway spruce forest (22%). Between November 1995 and October 1996, eight mountain hares were trapped using hand-made traps of three different kinds: wooden traps (62 x 58 x 125 cm) and aluminium traps (60 x 60 x 125 cm and 60 x 40 x 120 cm). Hares were radio-collared with adjustable necklace transmitters (L-I/ER3, article S212, A. Wagener, Köln, Germany) weighing 35g. Radio-tagged hares were located using a Yaesu receiver (148-149 MHz, FT-290R II., Yaesu Musen Co., Ltd., Tokyo, Japan) and a three-elements directional Yagi antenna. The animals were localised by triangulation (White and Garrott, 1990) once a week, taking one fix every 15 minutes for two hours, in order to cover periodically the whole 24 hours. At each fix, hares’ location and activity (1 = active, 2 = not active) were recorded. We did not test for independence of subsequent fixes, since, using a fixed study period, concerns regarding autocorrelation are relatively unimportant compared to obtaining a representative sample of fixes to reliably estimate home range size (Otis and White, 1999; Fieberg, 2007). For example, to subsample observations that are equally spaced throughout the study period would only decrease the precision of home range and the estimates of the utilisation distribution (UD) (Fieberg, 2007).  We estimated home ranges at a seasonal level: winter (December-March, ground continuously covered with snow), spring-summer (April-August, corresponding to hare “breeding season” and vegetation growth), and autumn (September-November, post-breeding season). We used the R software (R Development Core Team, 2007) for all analyses. We calculated the fixed kernel density estimator (KDE; Worton, 1989; Seaman and Powell, 1996) with adjusted smoothing factor (KDE with hadj), following the method proposed by Wauters et al. (2007) to calculate hadj. For comparison with 158Home range of alpine mountain hare  previous studies, we also calculated home range size with the minimum convex polygon (MCP) using the recalculated arithmetic mean and compared the two methods – KDE and MCP -, by the Wilcoxon matched-pairs signed-ranks test. Internal home range structure was explored for a core area of high activity (Kenward, 2001), examining the UD (range size included in 5% incremental isopleths containing between 20% and 95% of fixes) of the fixed kernel density estimator (KDE; Kenward, 2001). A one-way ANOVA was applied to define the core area, with the size of hares’ range included in a given isopleth as dependent variable and the 5% incremental isopleths as class. Home range overlap was expressed as the percentage of a hare's range overlapped by those of all other hares (e.g. Wauters and Dhondt, 1992; Lurz et al., 2000). Values are presented as mean ± standard deviation (S.D.). We investigated the effect of season (1= winter, 2 = spring-summer, 3 = autumn) and sex on home range size using the 95% KDE estimator (Kenward et al., 2001; Wauters et al., 2005, 2007; Di Pierro et al., 2008) with a two-way ANOVA (Proc mixed; SAS, 1999). When necessary, home range data were ln-transformed to meet the assumptions of normality (Shapiro-Wilk’s test W >0.93, P >0.3). To account for pseudo-replication of repeated measures, “individual” was added as a repeated factor. Models with simple correlation structure of the residual correlation matrix, corresponding to no effect of repeated measures on the individual, had the smallest values of Schwarz’s Bayesian Information Criterion (BIC), indicating the best fit (Verbeke and Molenberghs, 2000).  RESULTS AND DISCUSSION  We used 13 seasonal home-range estimates from eight different hares, six males (3 sub-adults, 3 adults) and two adult females. For each individual, 95% KDE estimates of home-range size were larger than 95% MCP (Wilcoxon matched-pairs signed-ranks test: P = 0.0005, 95% KDE mean = 38.1 ± 23.7 ha, min-max = 11.9-77.2 ha; 95% MCP mean = 26.7 ±ҏ 21.6 ha, min-max = 2.5-64.1 ha, Tab. 1). Home range size (95% KDE) did not differ between the two sexes (Sex-effect: F1,9 = 0.27, P = 0.61), but was smaller in winter than in the other seasons (Season-effect: F2,9 = 7.02, P = 0.015, Fig. 1, 2). Due to the small sample size, we cannot draw any general conclusion about sex-specific space use patterns in alpine habitats. The use of small home ranges in winter might be a strategy to reduce energy expenditure during a season with critical weather condition, such as low temperatures (mean monthly T < 0°C) and deep snow cover. In comparison with other studies, home range sizes in the Swiss Alps were smaller than in Scandinavia (min-max = 71-280 ha; Hiltunen et al., 2004; Dahl, 2005a, b; Dahl and Willebrand, 2005; Kauhala et al., 2005; Hiltunen, 2006), but similar to those documented in Scotland (min-max = 15-113 ha; Hewson and Hinge, 1990; Hulbert et al., 1996, Rao et al, 2003). In Scotland (moorlands) and alpine environments (mountain pine or other forests) a homogeneous distri-bution of food resources in sufficiently large patches might explain the reduced movements when searching for food or hiding places, and, consequently, the use of relatively small home ranges. We found only one other study, in Austria, on the ecology of alpine populations of this species (Slotta-Bachmayr, 1998), that reported average home ranges of 435 ha (95% 159Genini Gamboni et al.  Table 1 - Seasonal home range size in hectares calculated for each hare by the kernel density estimator (KDE) and the minimum convex polygon (MCP).  Hare code Season No. locations 95% KDE 95% MCP 9001 Autumn 99 26.9 7.7 9001 Winter 245 15.9 11.6 9002 Winter 693 16.8 12.4 9003 Winter 950 29.4 20.4 9003 Breeding 58 27.6 2.5 9004 Winter 264 11.9 8.4 9005 Winter 1045 33.2 19.7 9005 Breeding 1353 48.5 49.5 9006 Breeding 2178 77.0 60.6 9006 Autumn 954 46.1 40.1 9007 Winter 566 15.1 8.3 9009 Breeding 1881 77.2 64.1 9009 Autumn 557 70.0 41.3                        Figure 1 - Seasonal home range size in hectares (mean ± S.E.): winter (4 males, 2 females), breeding season (3 males, 1 female), autumn (3 males). 160Home range of alpine mountain hare  MCP) and 551 ha (95% KDE), hence, much larger than in our study. However, the period over which each hare were monitored was not clearly defined (entire study period from November 1992 to March 1996), and hares made exploratory movements over wide areas (Slotta-Bachmayr, 1998) that could introduce bias in home range estimation. Exploring the use of a core area, no significant increase in range area was observed in any pair wise comparisons between consecutive contours (Tukey’s test on differences between means, all P >0.05). Hence, no area of con-centrated use could be detected. Accordingly, day-time radiotracking showed that each animal used several forms, widely spaced throughout its home range (our unpubl. data). The average percentage of home rangeoverlap between mountain hares was 51±ҏ22% (N = 44, min-max: 14-100%). Mean overlap between male ranges was 52±24% (N = 17), of a male by females was 49±17% (N = 7) and of a female by males was 47±22% (N = 8). The large home range overlap found in this study supports the hypothesis that the mountain hare is a non-territorial species, as documented in several study areas of northern Europe (Flux, 1970; Hewson, 1976; Hewson and Hinge, 1990; Dahl, 2005b). Ecosystems are rapidly changing due to human activities and global warming (Stenseth et al., 2002; Walther et al., 2005; Thomas et al., 2006). Studies on a wider range of populations, living in different habitat types, of the alpine mountain hare are necessary to better understand the social organisation and habitat use of this keystone species for alpine ecosystems.                      Figure 2 - Home ranges of two mountain hares (male no. 1: continuous lines and female no. 5: dashed lines) in different seasons. Thin lines: winter home range; thick lines: autumn and breeding season for male and female respectively. 161Genini Gamboni et al.  ACKNOWLEDGEMENTS  We are grateful to Marco Salvioni and the Hunting and Fishing Office of Can-ton Ticino, Switzerland. This research was part of the Degree Thesis of A-S. G. G. at Neuchâtel University.  REFERENCES  Angerbjörn A. and Flux J. E. C. 1995. Lepus timidus. Mammal. Spec., 495: 1-11. Couturier M. 1964. Le gibier des montagnes françaises. 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Home range dynamics of mountain hares (Lepus timidus) in the Swiss Alps.

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Home range dynamics of mountain hares (Lepus timidus) in the Swiss Alps.

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