Splitting hairs: How to tell hair of hares apart for predator diet studies

Mammalian Biology 89 (2018) 84–89

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Original investigation

Splitting hairs: How to tell hair of hares apart for predator diet studies Niccolò Fattorini a , Lucia Burrini a , Giovanni Morao b , Francesco Ferretti a , Giorgia Romeo c , Emiliano Mori a,∗ a

Department of Life Sciences, University of Siena, Via P.A. Mattioli 4, 53100 Siena, Italy Dipartimento di Agronomia Animali Alimenti Risorse Naturali e Ambiente – DAFNAE, Università degli Studi di Padova – Viale dell’università 16, 35020, Legnaro-Padova, Italy c Tuscan Regional Council, Via Pietro Micca 39, 58100 Grosseto, Italy b

a r t i c l e

i n f o

Article history: Received 21 September 2017 Accepted 13 January 2018 Handled by Mauro Lucherini Keywords: Hair determination Italian hares Lepus Macroscopical analysis Microscopical analysis

a b s t r a c t Hares are a major prey for many carnivorous vertebrates worldwide. Their occurrence in the diet of predators is mostly assessed through the analysis of indigested remains (especially hair) in faeces or pellets. In Italy, four hare species are present, locally occurring in sympatry, and several studies confirmed they are preyed upon by 15 carnivorous vertebrates, overall. A reliable identification of hare species in their diet is only possible if specific diagnostic keys of their hair are available. To provide diagnostic features of the four Italian hare species, we collected 218 hair samples from 37 individuals belonging to 13 hare populations. Samples were measured and analysed at the microscope; five indices were assessed. Hair indices of morphology differed significantly across the four species, both in the cortex and in medulla structure. Species discrimination through hair may be crucial especially if the range overlap among hare species will increase, due to environmental/climatic changes and/or human management actions (e.g. restocking). ¨ Saugetierkunde. ¨ © 2018 Deutsche Gesellschaft fur Published by Elsevier GmbH. All rights reserved.

Introduction Food habits of vertebrate species are mainly assessed through the analysis of food remains from faeces (i.e., seeds, stems, hair and bones: e.g. Putman, 1984; Reynolds and Aebischer, 1991; Klare et al., 2011) or egested pellets (Yalden and Yalden, 1985; Johnstone et al., 1990; Votier et al., 2003). Food remains can be determined through specific reference atlases (e.g. Mayer, 1952; Teerink, 1991; De Marinis and Agnelli, 1993; Oli, 1993; Nappi, 2001; De Marinis and Asprea, 2006) or ad hoc reference collections. Mandibular structures and hair of prey of carnivores often show species-specific diagnostic features (Hausman, 1920; Faliu et al., 1980; Teerink, 1991). When the size of the prey is relatively large with respect to that of predator, teeth are rarely found in faeces (Nappi, 2001), and, thus, researchers have to rely on hair analysis; accordingly, macroscopical and microscopical analyses of mammal hair may represent a valuable tool for ecological research, including food habits of carnivores and raptors (e.g. Yalden and Yalden, 1985; Oli, 1993; Redpath et al., 2002; Kerley et al., 2015).

∗ Corresponding author. E-mail address: [email protected] (E. Mori).

Hares Lepus spp. are widespread worldwide, but for Antarctica, and are important prey species for many vertebrates (e.g. felids: Lovari et al., 2013; Apostolico et al., 2016; canids: Hayward et al., 2014; Pagh et al., 2015; Newsome et al., 2016; mustelids: Macdonald et al., 2000; Posłuszny et al., 2007; raptors: Kopij, 2016; Rehnus et al., 2016). In the specific case of Italy, it has been reported that hare species are present in the diet of 7 species of mammalian carnivores, 6 species of raptors, the wild boar Sus scrofa and the carrion crow Corvus corone have been reported to feed on hare species (Appendix I in Supplementary material). Effective species determination from undigested hairs is needed to estimate the actual role of hares in the diet of predators (Temple and Terry, 2009), as well as to obtain sound information on mortality/limiting factors for conservation/management of wild hare populations. De Marinis and Agnelli (1993) reported that hair of all species belonging to the genus Lepus show a multicellular column-shaped medulla in the shield, a concave cross section and a pale bar. Furthermore, hairs of Lepus species show a diagnostic pale (light brown to red) bar near the tip, which allow researchers to distinguish them from those of other mammal species (Teerink, 1991). Diagnostic keys are required to identify hair at the specific level when two or more hare species co-occur. Italy represents a useful case study for setting up speciesspecific diagnostic keys to identify hair of hare species. In this

https://doi.org/10.1016/j.mambio.2018.01.005 ¨ Saugetierkunde. ¨ Published by Elsevier GmbH. All rights reserved. 1616-5047/© 2018 Deutsche Gesellschaft fur

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Table 1 Percent overlap in the Italian range of Lepus europaeus with L. corsicanus and L. timidus. SPECIES 1 → SPECIES 2 ↓

L. europaeus

L. corsicanus

L. timidus

L. europaeus L. corsicanus L. timidus

– 64.1 47.4

40.0 – 0.0

9.3 0.0 –

country, 4 out of 6 European hare species are present (Aulagnier et al., 2010). Mainly due to restocking events for hunting purposes (Santilli, 2007; Ferretti et al., 2010), the European brown hare L. europaeus is the most widespread species, and it is the only one coexisting with other species (Table 1). The Sardinian hare (L. capensis mediterraneus, hereafter L. capensis: Slimen et al., 2005) has been released in historical times to Sardinia from North Africa. Concerning the other species, the Apennine hare L. corsicanus is endemic to central and southern Italy (Scalera and Angelici, 2003), whereas the mountain hare L. timidus is widely distributed throughout the northern Palearctic and the Alps (Aulagnier et al., 2010). The latter species is listed within the European Union Habitat Directive, Annex V. Thus, both the Apennine hare and the mountain hare are species of conservation concern (Amori et al., 2008) and their distribution ranges partially overlap with that of the brown hare (Aulagnier et al., 2010; www.iucnredlist.org, accessed on 04th May 2017: Table 1). A morphological characterization of hair is only available for the mountain- and European brown hares (Teerink, 1991). Rugge et al. (2009) found that coat colouration is an effective feature for discriminating between L. corsicanus and L. europaeus in Southern Italy, suggesting that differences in hair morphology may also occur. Thus, our aim was to identify diagnostic features to distinguish the hairs of the four hare species, as to provide researchers with an effective key for this genus. Material and methods Collection and measurements of hair samples We collected guard hair samples from 13 populations (L. corsicanus, N = 3; L. timidus, N = 3; L. capensis, N = 3; L. europaeus, N = 4) of all the 4 hare species, from a total of 37 individuals (L. corsicanus, N = 8; L. timidus, N = 8; L. capensis, N = 6; L. europaeus, N = 15). Hairs were taken from individuals of both sexes. Coat colouration and morphology (i.e. hair length) do not change throughout the year for most species adapted to live in areas with limited annual climatic fluctuations (Stoner et al., 2003). We decided to use only hair collected between September and November to avoid the winter-early spring white coat of the mountain hare. Furthermore, most captures for behavioural and parasitological studies, as well as hunting, mainly occur in these months or early after (e.g. Santilli, 2007; Ferretti et al., 2010; Zaccaroni et al., 2013), therefore increasing the success of hair collection. We assumed that no age difference in hair ultrastructure would occur in hares (cf. Faliu et al., 1980; Teerink, 1991). We also have no evidence of any age class being more susceptible to predation. Since most restocked European hares are adult (Amori et al., 2008; Sokos et al., 2015), we only sampled hairs of subadult/adult individuals. Hairs were taken from killed and live captured individuals, from both the rump and hind legs. A total of 218 hair samples was analysed, between 4 and 6 hairs per individual. The total hair length, pale bar width (hereafter “bar width”), and distance between the bar and hair tip (“bar-tip distance”: Fig. 1) were measured by a precision calliper (0.1 mm in sensitivity: © DIN 862, Würth, Germany). In turn, we used five indices of hair morphology, some of which can

Fig. 1. General structure of hair of hare species in Italy.

be obtained even from hairs partially destroyed by digestive processes: i) hair length; ii) bar width; iii) bar-tip distance; iv) “bar width: bar-tip distance” ratio; v) “bar width: hair length” ratio. Hairs were prepared through standard protocols (Mori et al., 2016). The analysis of the cortex and cuticula was carried out by placing each hair on a glass slide, on a layer of nail polish. When polish was completely dry, the hair was removed, and the mould observed at the microscope and described according to the categories reported by Teerink (1991). The analysis of the medulla was carried out with the hair longitudinally sectioned and wet with cedar oil at the section level. Slides were observed under the microscope to compare the structure of the medulla with patterns reported by Teerink (1991). Statistical analyses We used Fisher’s linear discriminant analysis (LDA: Rencher, 1995) to assess the separation among species hair morphology. LDA replaced the values of original variables (i.e. hair indices) with the scores of three discriminant functions, i.e. linear combinations of the variables providing the greatest separation between species. We applied a non-parametric adaptation of the multivariate analysis of variances based on Euclidean distances between LDA scores to test for the statistical significance of differences between species (NPMANOVA; Anderson, 2001). The significance was computed by class permutations, using 99,999 replicates. The pairwise comparison of species was used as a post-hoc test using Bonferroni’s correction. These statistical analyses were performed through the software PAST (Hammer et al., 2001). We evaluated whether indices of hair morphology differed among species through generalised linear mixed models (GLMMs; Zuur et al., 2009). Being continuous, positive variables, indices were modelled through an inverse Gaussian error distribution and an

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Fig. 2. Hair morphological differences among hare species in Italy (N = number of measurements) through LDA. The ellipses containing 95% observations within each species are also shown. The x and y axes represent the scores of first and second discriminant function, respectively, which mainly contributed (c. 90%, see Section Results) to species separation.

identity link function, with species as predictors. Lepus capensis was arbitrarily set as the reference category and we estimated the model parameters for each of the other species. Wald’s test was used to assess the significance of predictors (Bolker et al., 2009). To account for repeated measures, individual was inserted as random effect, nested within each species. Crossed random effects across species/individuals were allowed to account for the body position where the hair sample was taken (i.e. from the back or the hind thig). Tukey’s HSD tests were used for post-hoc pairwise comparisons between species (by correcting p-values for multiple comparisons through the single-step method: Hothorn et al., 2008). GLMMs and post hoc tests were performed with the R packages lme4 (Bates et al., 2015) and multcomp (Hothorn et al., 2008), respectively. Validation of the statistical methods are reported in Appendix II in Supplementary material. Results Microscopical analyses of hair were insufficient for species determination, but medullar structure allowed the distinction of sympatric hare species (Table 2). Accordingly, two species (i.e. L. timidus and L. corsicanus) showed straight medullar margins, the other two (i.e. L. europeaus and L. capensis) had scalloped margins. The contribution to species separation provided by the first two discriminant functions was about 90% (first discriminant function: 60.39%; second discriminant function: 29.17%). The four species were significantly distinguished by their indices of hair morphology, both overall (NPMANOVA test: F: 281.7; p = 0.0001; Fig. 1) and pairwise (L. capensis vs L. corsicanus: p = 0.0006; L. capensis vs L. europaeus: p = 0.0006; L. capensis vs L. timidus: p = 0.0006; L. corsicanus vs L. europaeus: p = 0.0006; L. corsicanus vs L. timidus: p = 0.0006; L. europaeus vs L. timidus: p = 0.0006: Fig. 2). Hair length was the greatest in L. timidus and the lowest in both L. capensis and L. corsicanus (Tables 3a and 4a ; Fig. 3A). Bar width was the greatest in L. timidus and the lowest in L. capensis (Tables 3b and 4b; Fig. 3B). L. timidus showed the greatest distance between bar and hair tip, which was the lowest in both L. europaeus and L. corsicanus (Tables 3c and 4c; Fig. 3C). The ratio “bar width: bartip distance” was the highest in L. corsicanus and the lowest in L. capensis (Tables 3d and 4d; Fig. 3D). Lepus corsicanus showed the greatest ratio between bar width and hair length, which was the lowest in both L. capensis and L. europaeus (Tables 3e and 4e; Fig. 3E). Bar width is the only index providing an unequivocal separation amongst all four species. Furthermore, hairs of L. europaeus can be

Fig. 3. Indices of hair morphology across hare species in Italy. Boxplots delineate the median (thick horizontal line), 25%–75% quartiles (box), upper/lower range (short horizontal lines) and outliers (circles). All the pairwise comparisons were significant (Tables 2 and 3), except for those indicated with ns (not significant).

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Table 2 Summary of the main macroscopical and microscopical features of the hair hare species in Italy.

N subpopulations N individuals N hair Bar width (cm) Hair length (cm) Distance bar-tip (cm) Scale position (cortex) Scale margin (cortex) Dist. between scale-margin Scale pattern (cortex) Width composition (medulla) Medulla structure Medulla margin

Lepus timidus

Lepus europaeus

Lepus corsicanus

Lepus capensis

3 8 39 0.50 (0.48–0.56) 3.70 (3.68–3.71) 0.49 (0.47–0.53) transversal smooth near regular wave multicellular (rows) filled straight

4 15 86 0.38 (0.37–0.40) 3.20 (3.10–3.40) 0.39 (0.38–0.41) transversal smooth near regular wave multicellular (rows) filled scalloped

3 8 60 0.44 (0.42–0.46) 2.70 (2.60–2.80) 0.38 (0.37–0.39) transversal smooth near regular wave multicellular (rows) filled straight

3 8 38 0.35 (0.34–0.36) 2.90 (2.80–2.93) 0.43 (0.42–0.44) transversal smooth near regular wave multicellular (rows) filled scalloped

Table 3 Parameters estimated from GLMMs: coefficients (␤), standard error (SE) and significance (p). For each response variable, the variance of random effects is also shown (random factors, in parentheses). Lepus capensis was arbitrarily set as reference. Index of hair morphology

Predictor

␤

SE

p

(a) Hair length (Species/Invidual) = 0.635 (Hair position) = 0.172

intercept L. corsicanus L. europaeus L. timidus intercept L. corsicanus L. europaeus L. timidus intercept L. corsicanus L. europaeus L. timidus intercept L. corsicanus L. europaeus L. timidus intercept L. corsicanus L. europaeus L. timidus

27.581 −1.345 3.914 8.605 3.518 0.850 0.359 1.655 4.308 −0.538 −0.417 0.657 0.818 0.348 0.177 0.228 0.128 0.037 −0.003 0.017

1.271 0.950 0.782 0.840 0.109 0.141 0.127 0.138 0.063 0.082 0.074 0.087 0.022 0.029 0.026 0.029 0.006 0.006 0.005 0.006

<0.001 0.157 <0.001 <0.001 <0.001 <0.001 0.005 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 0.625 0.004

(b) Bar width (Species/Invidual) = 0.017 (Hair position) <0.001 (c) Bar-tip distance (Species/Invidual) = 0.005 (Hair position) < 0.001 (d) “Bar width: bar-tipdistance” ratio (Species/Invidual) < 0.001 (Hair position) < 0.001 (e) “Bar width: hair length” ratio (Species/Invidual) < 0.001 (Hair position) < 0.001

Table 4 Post hoc comparisons between species assessed through general linear hypothesis/Tukey’s HSD tests: coefficients (␤), standard error (SE) and adjusted significance (adjusted p). Index of hair morphology

Post hoc comparison

␤

SE

adjusted p

(a) Hair length

L. corsicanus vs L. capensis L. europaeus vs L. capensis L. timidus vs L. capensis L. europaeus vs L. corsicanus L. timidus vs L. corsicanus L. timidus vs L. europaeus L. corsicanus vs L. capensis L. europaeus vs L. capensis L. timidus vs L. capensis L. europaeus vs L. corsicanus L. timidus vs L. corsicanus L. timidus vs L. europaeus L. corsicanus vs L. capensis L. europaeus vs L. capensis L. timidus vs L. capensis L. europaeus vs L. corsicanus L. timidus vs L. corsicanus L. timidus vs L. europaeus L. corsicanus vs L. capensis L. europaeus vs L. capensis L. timidus vs L. capensis L. europaeus vs L. corsicanus L. timidus vs L. corsicanus L. timidus vs L. europaeus L. corsicanus vs L. capensis L. europaeus vs L. capensis L. timidus vs L. capensis L. europaeus vs L. corsicanus L. timidus vs L. corsicanus L. timidus vs L. europaeus

−1.345 3.914 8.605 5.259 9.950 4.691 0.850 0.359 1.655 −0.491 0.805 1.296 −0.538 −0.417 0.657 0.121 1.195 1.075 0.348 0.177 0.228 −0.165 −0.114 0.051 0.037 −0.003 0.017 −0.040 −0.020 0.019

0.950 0.782 0.840 0.777 0.835 0.645 0.141 0.127 0.138 0.111 0.128 0.108 0.082 0.074 0.087 0.065 0.080 0.071 0.029 0.026 0.029 0.023 0.026 0.022 0.006 0.005 0.006 0.005 0.005 0.005

0.484 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 0.025 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 0.243 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 0.103 <0.001 0.961 0.023 <0.001 <0.001 <0.001

(b) Bar width

(c) Bar-tip distance

(d) “Bar width: bar-tip distance” ratio

(e) “Bar width: hair length” ratio

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distinguished from those of L. capensis through all indices, with the exception of the ratio bar width: hair length. Lepus europaeus and L. corsicanus can be discriminated through all the measures, except for the bar-tip distance. All indices (but the ratio bar width: bartip distance) may discriminate between hairs of L. europaeus and L. timidus. Lepus capensis and L. corsicanus can be distinguished by all the indices, with the exception of the hair length. Hairs of L. capensis and L. timidus and hairs of L. corsicanus and L. timidus are discriminated by all indices.

Discussion The coloration of the pelage of lagomorph species is known to be matched to habitat type, geographical region and altitude above sea level (Stoner et al., 2003; Rugge et al., 2009). With the present work, we provided a first key for the macroscopical and microscopical identification of hair of all hare species living in Italy. Our results showed that hare species in Italy can be identified through hair analyses. The mountain hare, Sardinian hare and Apennine hare showed different hair morphologies (with the exception of the medullar pattern, which is similar among Sardinian and Apennine hares), probably because these species are adapted to live in different, non-overlapping habitats (Stoner et al., 2003; Rugge et al., 2009). Hair morphology of the European brown hare showed clear differences from that of mountain hare, but a partial overlap with Apennine and Sardinian hare was found. Hairs of the mountain hare were the longest amongst the Italian hares, a result consistent with the spatial ecology of this species, adapted to live in high mountains (Hewson and Hinge, 1990; Thulin, 2003). The largest width of the pale bar depends on the overall pale coloration of this hare species. Both the Apennine hare and the Sardinian hare showed short hair, being adapted to live in warm, mostly plain areas of the warmest Italian regions (Amori et al., 2008). The former is distributed in Mediterranean “macchia” habitats (i.e. sclerophillic scrubwood), as well as in warm oakwoods (Angelici and Luiselli, 2007; Mori et al., 2014), whereas the latter is mainly present in dense, thick shrubwoods (Murgia et al., 2003). Accordingly, the Apennine hare showed a wider pale bar with respect to the Sardinian species, being more adapted to open habitats (Amori et al., 2008). Medullar margin type is sufficient to differentiate between sympatric hare species in dietary studies of predators. Despite preferring open habitats (i.e. fallows and cultivated fields: Lewandowski and Nowakowski, 1993), the European brown hare occurs from plain areas to high mountains (up to 2412 m a.s.l. in the Italian Alps: Patriarca and Debernardi, 2014), also due to frequent releases. Therefore, it may cover a wide range of habitat types (wooded areas: e.g. Zaccaroni et al., 2013; farmlands: e.g. Santilli et al., 2014; mountain meadows: Bisi et al., 2015), despite no difference in hair morphology was observed among different sites. Range overlap between L. europaeus and L. timidus, as well as between L. europaeus and L. corsicanus is known to occur in Europe (Thulin, 2003; Fulgione et al., 2009; Bisi et al., 2015). Furthermore, range overlap between restocked and naturally occurring hare specie may further increase, because of continuous releases of European brown hares (Angelici and Luiselli, 2007; Fusco et al., 2007; Bisi et al., 2015). Thus, given the importance of hares in the diet of many species (Appendix I in Supplementary material), an effective identification of hair of hares should be required for food habit studies, through macroscopical and microscopical (i.e. medullar pattern) analyses, to obtain reliable species determination. We are aware that a very well-preserved hair sample (i.e. hairs which preserve the real length) is hard to be found in scats, because hair pass through a digestive process, which is somehow destructive. At the light of this assertion, we suggest to use ratios between

hair characteristics, more than hair characteristics themselves, for hair discrimination. Acknowledgements We are grateful to M. Calosi, A. Canu, P. Di Luzio, F. Lucati and P. Di Bari, who provided us with hair samples. EM had the idea of this project, carried out hair measurements, worked out data as well as wrote all drafts. NF performed statistical analyses and participated in writing up all drafts. LB prepared hair samples for microscopical analyses. GM provided a high amount of hair samples. FF participated in writing and contributed critically to the drafts. GR provided several hair samples and contributed with comments to the final draft of the paper. All the authors gave final approval for publication. Authors declare no conflict of interests. Valentina Mancino helped us in slide preparation. Two anonymous reviewers, Mauro Lucherini and Sandro Lovari greatly improved our first draft with their comments. Vasco Sfondrini kindly took the time to read and correct the English grammar and syntax, markedly improving the readability of the manuscript. Appendix A. Supplementary data Supplementary data associated with this article can be found, in the online version, at https://doi.org/10.1016/j.mambio.2018.01. 005. References Amori, G., Contoli, L., Nappi, A., 2008. Fauna d’Italia: Mammalia II, Erinaceomorpha, Soricomorpha, Lagomorpha, Rodentia. Calderini – Il Sole 24 ore, Bologna, IT. Anderson, M.J., 2001. A new method for non-parametric multivariate analysis of variance. Austral Ecol. 26, 32–46. Angelici, F.M., Luiselli, L., 2007. Range, dynamic biogeography and ecological interactions of two species: Lepus corsicanus and Lepus europaeus in Italy. Wildl. Biol. 13, 251–257. Apostolico, F., Vercillo, F., La Porta, G., Ragni, B., 2016. Long-term changes in diet and trophic niche of the European wildcat (Felis silvestris silvestris) in Italy. Mammal Res. 61 (2), 109–119. Aulagnier, S., Haffner, P., Mitchell-Jones, A.J., Moutou, F., Zima, J., 2010. Guide des Mammifères d’Europe, d’Afrique du Nord et du Moyen-Orient. Delachaux and Niestlè SA, Paris, France. Bates, D., Maechler, M., Bolker, B., Walker, S., 2015. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48. Bisi, F., Wauters, L.A., Preatoni, D.G., Martinoli, A., 2015. Interspecific competition mediated by climate change: which interaction between brown and mountain hare in the Alps? Mammal. Biol. 80, 424–430. Bolker, B.M., Brooks, M.E., Clark, C.J., Geange, S.W., Poulsen, J.R., Stevens, M.H.H., White, J.S.S., 2009. Generalized linear mixed models: a practical guide for ecology and evolution. Trends Ecol. Evol. 24, 127–135. De Marinis, A.M., Agnelli, P., 1993. Guide to the microscope analysis of Italian mammals hairs Insectivora, Rodentia and Lagomorpha. Ital. J. Zool. 60, 225–232. De Marinis, A.M., Asprea, A., 2006. Hair identification key of wild and domestic ungulates from southern Europe. Wildl. Biol. 12, 305–320. Faliu, L., Lignereux, Y., Barrat, J., 1980. Identification des poils des mammifères ˜ Acta Vert. 1, 125–212. pyreneens. Donana Ferretti, M., Paci, G., Porrini, S., Galardi, L., Bagliacca, M., 2010. Habitat use and home range traits of resident and relocated hares (Lepus europaeus, Pallas). Ital. J. Anim. Sci. 9, e54. Fulgione, D., Maselli, V., Pavarese, G., Rippa, D., Rastogi, R.K., 2009. Landscape fragmentation and habitat suitability in endangered Italian hare (Lepus corsicanus) and European hare (Lepus europaeus) populations. Eur. J. Wildl. Res. 55, 385–396. Fusco, L., Vaccaro, L., Troisi, S.R., Accardo, Y., Caliendo, M.F., Gabriele, F., 2007. Segregazione ambientale tra popolazioni simpatriche di Lepus corsicanus e L. europaeus nel Parco Nazionale del Cilento e Vallo di Diano. Conservazione di Lepus corsicanus De Winton, 1898 e stato delle conoscenze. Pubblicazioni IGF, pp. 119–122. Hammer, Ø., Harper, D.A.T., Ryan, P.D., 2001. PAST: paleontological statistics software package for education and data analysis. Palaeontol. Electronica 4, 1–9. Hausman, L.A., 1920. Structural characteristics of the hair of mammals. Am. Nat. 54, 496–523. Hayward, M.W., Lyngdoh, S., Habib, B., 2014. Diet and prey preferences of dholes (Cuon alpinus): dietary competition within Asia’s apex predator guild. J. Zool. (Lond.) 294, 255–266.

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