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Habitat Selection and Mating Success in a Mustelid

Habitat Selection and Mating Success in a Mustelid Hindawi Publishing Corporation International Journal of Zoology Volume 2011, Article ID 159462, 6 pages doi:10.1155/2011/159462 Research Article Thierry Lode ´ UMR CNRS 6552 ETHOS, Universit´e de Rennes 1, 35042 Rennes, France Correspondence should be addressed to Thierry Lode, thierry.lode@univ-rennes1.fr Received 4 December 2010; Revised 20 February 2011; Accepted 4 March 2011 Academic Editor: Hynek Burda Copyright © 2011 Thierry Lode. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Habitat selection remains a poorly understood ecological process, but relating mating behaviour to pattern of habitat selection constitutes a fundamental issue both in evolutionary ecology and in biological conservation. From radiotelemetry protocol, habitat-induced variations in mating success were investigated in a solitary mustelid carnivore, the European polecat Mustela putorius. Selection for marshy habitat was regarded as adaptive in that mating success was found greater using marches than other habitats. Males consorted with 1.3 females, revealing a low polygyny rate. Pregnant or lactating females selectively shifted to deciduous woods. That some habitat types may favour a good reproduction forms a key factor for species conservation and environmental management. Nevertheless, such as in various vertebrates, habitat requirements seem to be based on simple broad features of habitat, suggesting that habitat avoidance rather than habitat preference can explain polecat habitat predilection. 1. Introduction reproductive sites can be regarded as reflecting habitat selection [8, 12, 13]. It could be predicted that no selection Adaptive significance of a behaviourisrecognized whena occurs when unsuccessful and successful reproductive sites behaviour provides a selective advantage in animal ability to are not discernible from most of their main habitat features, adapt to its environment. Landscape structures are expected but McLoughlin et al. [12] found no variations at the home- to influence especially mating behaviour and numerous range level but differences within the home range level species required heterogeneous habitat composition. underline the importance of spatial scale dependence in Differences in habitat use among animals have been habitat selection [6, 14]. often ascribed to habitat selection although habitat selection Selection of better reproductive sites and habitat features remained a poorly known behavioural ecological process [1– should be indicative of adaptive behavioural preferences but 3]. Patterns of habitat use may reveal specific choice for can often result in increasing competition, habitat selectivity, habitat, which should provide numerous benefits such as and declining population density [4]. Habitat selection and resource availability, shelter, avoidance of predation, and nest-sites choice have been widely studied in birds although reduced interspecific competition [4, 5]. The basic conjecture differences in fitness were not often examined [8, 15, 16]. in habitat selection theory is that animals select habitat However, differences in habitat can have noticeable effects features to maximize their fitness [6–9]. Thus, habitat on reproduction [9]. The relationship between male status selection is a behaviour related to fitness and differs from and mating success was reported from social mammals such habitat use in which habitat preferences are mainly linked to as mongooses and badgers [17–19], but numerous species resources. Habitat selection can lead to a strong fidelity to lived a solitary life, and their social organisation was mainly habitat features in which fitness was previously high [10]. based on spatial pattern [20]. This is especially the case for Numerous species showed partitioning in microhabitat most carnivores, and although habitat features have been preferences, but patent demonstration for habitat selection was rare [11, 12]. Actually, natural selection can occur recognized as major parameters for species conservation (see [21]), the role of habitat selection has still received little when distinct habitat types differ in successful breeding, and putative differences between unsuccessful and successful attention. 2 International Journal of Zoology In solitary carnivores, male and female live indepen- data from radiotracked polecats, we calculated the centre dently over long periods and generally exhibit a mating of activity (arithmetic mean) from fixes, and the minimal system in which the male home range partially overlaps the dispersal distance was assessed comparing minimal distance range of one or several females [22–25]. Because carnivores between telemetry locations. were routine and did not try to avoid poor environmental Habitat use was inferred in male and female radio- conditions, it could be predicted that habitat traits have a tracked polecats based on percentage of fixes found in major influence on their mating success. a considered habitat type and was assigned to two main This study investigates the role of habitat selection on the periods, winter (November–January) and breeding period mating success of a solitary mustelid carnivore, the European (February–April for males, and May-June for females, polecat Mustela putorius. Based on a long-term survey because mating sites may differ with breeding sites in which of radiotracked animals, this study aims at characterising parturitions occurred). Field study for determining habitat habitat features and mating success to assess whether habitat features was performed between July and September from features may affect mating success (habitat selection). 1998 to 2002, considering the minimum convex polygon of each individual and, for random sites, a number of putative locations, on the basis of average number of real fixes recorded, randomly assigned within the study areas. 2. Methods We defined 19 habitat features as deciduous wood, oak Radiotracked polecats (24 males and 25 females, 1992– grove, chestnut grove, hybrid poplar plantation, orchard and 2002) were mainly surveyed from winter to spring in garden, willow grove, flooded herbaceous cover, reed-bed, ◦ ◦ Western France (Brier ` e 47 20 N, 2 10 W, lake of Grand- ditches, river bank and riparian habitat, pond, grassland, ◦ ◦ Lieu and Tenu river 47 05 N, 1 39 W), and data from each pasture, cultivation and vineyard, shrub of bramble, hedges, individual was only used once. The climate of an oceanic road and path, barn and abandoned house, and at last farm type is mild and humid (mean temperature August 20 C, and houses. Habitat diversity was assessed by the index E = ◦ 2 and mean temperature February 5 C, annual precipitation A−1/S−1, where A = 1/Σ Pi [30]and S is the total number of habitat type, considering only 6 major habitat types. An 650 mm, with about 2 snowy days per year). Every animal was surveyed 121–223 days, but numerous individuals were analysis of variance was used to investigate differences in recaptured several times. From a capture-recapture design, habitat features among male sites considering male sites with consortship success, unconsorted male sites, and random density estimates were regarded as moderately high ranging from 0.35 to 0.57 individuals per km in 2002. sites. Because all females consorted, we only discriminated The consortship behaviour consists of a brief association between habitat use from February to April (mating period) between an adult male and an adult female, showing activity and from April to June (period of parturition). Paired t-test within the same restricted area and often resting together in (with Bonferroni adjustment procedure) or Welch alternate the same den during oestrus [26, 27]. Consortship success t-test is used according to the case. was determined by trapping, incidental observations (using night vision optics NONH4U), and radiotracking locations. 3. Results Throughout the reproductive season, trapping sessions were repeatedly conducted using 20 box-traps in every studied site 3.1. Consortship Success. Moving across several female con- in order to detect the presence of other individuals during tiguous ranges, most male polecats (80%, n = 19/24) the mating season. Polecats are usually intolerant towards obtained brief consortships averaging 3.11 days (sd = 1.26, any congeners, exhibiting a strong individual segregation range 1–6 days, and n = 19 males). Males consorted with in the use of space, and associations between individuals 1.33 partners (sd = 0.82, n = 24 males). All females are restricted to the breeding period [24, 28, 29]. Because consorted with at least one male and obtained a mean of copulation induces ovulation, breeding success depends 1.12 consortships (sd = 0.33, n = 25 females; Table 1). The upon the male tenacity, and most of recaptured females consortship success was related to the consortship duration exhibited scratches on the neck and on the back resulting (r = 0.705,P< .0001, and n = 25). While consortship Spearman from the copulation, when the male remained on the female patterns in males characterised a polygynous mating system with a bite often lasting more than one hour [27]. Therefore, with a single male consorting with one to three females, some the consortship behaviour provides a good indication of the females were regarded as polyandrous. mating activity [26] although how consortship behaviour is related to breeding success remains unknown. Most of copulations occurred before Apri, and parturitions, as 3.2. Habitat Selection. During breeding period, polecats revealed by recaptures of lactating females, occurred from mainly used marshes and wooded areas, but habitat use May to June. differed between the sexes, females occupying significantly A consortship was regarded as successful when it has more woods and hedges, while males exploited mainly occurred about 42 days (gestation duration) before the marshes (χ = 42.3, P< .0001, df = 4) (Figure 1). estimated date of parturitions. On a basis of a minimum Actually, habitat preferences seemed to be only based on of six consecutive fixes per day, the minimal consortship superficial features of habitat, and marshy habitat features duration was assessed as a number of days during which and deciduous woods appeared as the most important male and female were diurnally associated in the den. For descriptors of polecat habitat. Habitat use in males was International Journal of Zoology 3 Table 1: Consortship duration and consortship success (number Habitat use of sexual partners) for 24 radiotracked males and 25 females of European polecats Mustela putorius from Western France. Number of Number Successful sexual Consortship of Con- Males partners- duration sexual sortship for (days) partners with female M1 1 F1 1 4 M2 1 F2 1 3 M3 1 F3 1 4 M4 2 F4 1 2 F5 1 1 M5 1 F6 1 1 M6 2 F7 1 5 F8 1 2 M7 1 F9 2 1 0 Males Females M8 1 F10 1 6 M9 3 F11 2 3 F12 2 3 Hedges, shrubs Marshes, ponds Deciduous woods Meadows F13 1 2 Riparian habitats M10 2 F14 1 3 Figure 1: Habitat use in male and female radiotracked polecats F15 1 3 based on percentage of fixes found in a considered habitat. Breeding M11 2 F16 1 4 males F17 1 3 M12 1 F18 1 5 Table 2: Differences among habitats used by 24 male and 25 female polecats and random sites (Anova from radiotracking locations, M13 2 F19 1 3 significant P value in bold). F20 1 4 M14 2 F21 1 5 Mean Degree of Fisher square freedom F F22 1 3 Males M15 2 F23 1 3 Deciduous woods 0.0019 1,39 0.87 .358 F24 1 3 Marshy areas 0.429 1,39 69.25 .0001 M16 1 F25 1 4 Riparian habitat 0.0033 1,39 2.47 0.125 M17 1 F17 1 2 Meadows and cultivation 0.523 1,39 104.38 .0001 M18 1 F18 1 3 Hedges and shrubs 0.021 1,39 12.42 .0011 M19 1 F20 1 2 Females M20 0 0 Deciduous woods 0.102 1,39 16.38 .0001 M21 0 0 Marshy areas 0.114 1,39 18.39 .0001 M22 0 0 Riparian habitat 0.0026 1,39 0.99 .325 M23 0 0 Meadows and cultivation 0.715 1,39 175.6 .0001 M24 0 0 Hedges and shrubs 0.056 1,39 28.9 .0001 Mean = Mean = Mean = 2.33 1.33 1.12 deserted. Moreover, polecats used significantly more shrubs chiefly influenced by willows, reedbeds, flooded herbaceous and hedges than other habitats. Female polecats were very cover, and ditches, and in females, the choice of parturition selective in their habitat preferences using significantly more sites was mainly influenced by ashes, oaks, willows, and reed- deciduous woods, hedges and shrubs, and marshy areas than beds. randomly estimated, and they poorly utilised meadows and Nonetheless, evidence of habitat selection was found cultivation. when differences among habitats, respectively, used by pole- Furthermore, habitat use differences were found accord- cats, and habitats in random sites were tested with ANOVA ing to the consortship success (Figure 2), unconsorted male (Table 2). In male polecats, marshy areas were the dominant polecats using significantly more deciduous woods and less habitat type, while meadows and various cultivations were marshy areas than consorted polecats (Table 3). Because all (%) 4 International Journal of Zoology Habitat shift in female polecats Habitat selection in males June home range Spring home range Consorted Unconsorted Hedges, shrubs Marshes, ponds Meadows Deciduous woods Barns, farms Riparian habitats Riparian habitats Hedges, shrubs Marshes, ponds Deciduous woods Meadows Figure 3: Habitat shift in surveyed female polecats from March (consortship sites) to June (parturition sites) as revealed by Figure 2: Habitat selection according to the consortship success of percentage of fixes recorded in a considered habitat. radiotracked male polecats. (n = 19 consorted polecats and n = 5 unconsorted polecats) as revealed by percentage of fixes recorded in a considered habitat. 4. Discussion Polecats shifted for marshy and wooded areas during their Table 3: Differences among habitat used by consorted (n = 19) and breeding period, but many habitat features did not clearly nonconsorted males (n = 5) (Anova from radiotracking locations, differ from random sites with regard to most of their significant P value in bold). characteristics. Studies of animal distribution in habitats often dealt Mean Degree of Fisher with habitat preferences with no assessment of mating Males P square freedom F success or remained mainly based on indirect approach Deciduous woods 0.089 1,23 4.68 .042 such as variations in density or abundance [3, 15, 31, 32]. Such approaches described habitat features, which could be Marshy areas 0.022 1,23 5.03 .035 poorly related to fitness but rather with resource availability Riparian habitat 0.034 1,23 2.05 .166 [8]. Measurement of habitat features, which were directly associated with breeding activities such as nesting, should Meadows and cultivation 0.018 1,23 0.95 .338 more reflect habitat selection [8, 13, 33, 34]. Hedges and shrubs 0.0008 1,23 0.34 .564 Habitat features associated with consortship success should at least indirectly reflect features that affect fitness. Mating sites differed from random sites only in some rough habitat features, and characteristics of successful mating surveyed female polecats consorted and bred successfully, sites have uncertain ecological meaning. Three nonexclusive it was impossible to run a same analysis. Nevertheless, causes could be evoked for such findings. Firstly, it may be pregnant or lactating females settled in distinct areas in May- alleged that every habitat type induces few differences in June switching for more woody habitats (paired t-test = 6.28, fitness, and thus, further habitat selection does not occur in P< .0001, and df = 23, Figure 3) and riparian habitats polecat. Secondly, habitat use can be regarded as a result of (paired t-test = 2.31, P = .03, and df = 23) whereas the use limited opportunities for animals to optimise when preferred of marshy areas decreased (paired t-test = 9.57, P< .0001, habitats are saturated. Thirdly, searching for the preferred and df = 23). The use of shrubs and hedges increased habitat may result in a relatively low benefit. (paired t-test = 3.58, P< .002, and df = 23) probably Because habitats features differed between random sites related to the search for a quiet parturition place. The index and mating sites, it should be recognised that polecats of habitat diversity significantly differed between space used exhibited at least a slight selection for habitat. Diversity by consorted polecats (E = 0.225, sd = 0.062) and random of habitat used by breeding polecat differed significantly sites (0.428, sd = 0.087) (Welch T-test = 11.2, P< .0001, from random sites, revealing that certain habitat were less df = 53). occupied during reproduction. Nevertheless, it could be (%) (%) International Journal of Zoology 5 suspected that low benefits in mating success resulted from per male, the polygyny rate appeared very low. Because pole- habitat selection. Habitat requirements of most vertebrates cats assiduously exploited some patchy area until resource seem to be based on simple superficial features of habitat depletion, solitary habits were basically proved to be adaptive [21, 34–38]. Thus, the stone marten exhibited a preference [24, 46, 50]. Conversely, such habits result in a low polygyny for “stony” environment and regardless of other habitat rate, suggesting an evolutionary cost induced by this specific features inhabited deciduous woods as well as barns, farms, spatial pattern. or buildings in towns [25, 39]. Anyway, that certain habitat features favour a best mating Following Haila et al. [35], it may be suggested that success constitutes a key factor for polecat conservation and habitat avoidance rather than habitat preference can explain environmental management. such predilection. Polecats were found to avoid open mead- ows and cultivation and seemed to prefer high vegetation Acknowledgments cover or undisturbed areas. Polecats showed very secretive movements in shrubs and along hedges, resting in dense The author thanks all his assistants for help in collecting the vegetation, in burrows or warrens. Habitats were selectively field data and especially Y. Benoit, Y. Biraud, H. Brault, F. exploited, and while polecats often moved sheltered by Cupertino, R. Dagault, F. Dussault, A. Pihuit, M. Pondaven, hedges and shrubs, most open environments were deserted. and J. Y. Robert. Thanks are also due to D. Le Jacques. The use of bramble patches is regarded as a need for protection against predators [40]. Woods, hedges, and shrubs may ensure more protected movements [41], and most References mustelids were also known to avoid openfields [40, 42, 43]. Moreover, as reproductive activity resulting in a reduced [1] R.E.Ricklefs, Ecology,W.H. Freeman,New York,NY, USA, vigilance, mating may be affected by the existence of refuges. At the broader scale of habitat selection, some factors, [2] C. J. Krebs, Ecology, The Experimental Analysis of Distribution chiefly marshy and wooded habitats, were found to influence and Abundance, Harper Collins College, Glenview, Ill, USA, mating success. Although some carnivores could be thought 4th edition, 1994. 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Habitat Selection and Mating Success in a Mustelid

International Journal of Zoology , Volume 2011 – May 17, 2011

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Publisher
Hindawi Publishing Corporation
Copyright
Copyright © 2011 Thierry Lodé. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
ISSN
1687-8477
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1687-8485
DOI
10.1155/2011/159462
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Hindawi Publishing Corporation International Journal of Zoology Volume 2011, Article ID 159462, 6 pages doi:10.1155/2011/159462 Research Article Thierry Lode ´ UMR CNRS 6552 ETHOS, Universit´e de Rennes 1, 35042 Rennes, France Correspondence should be addressed to Thierry Lode, thierry.lode@univ-rennes1.fr Received 4 December 2010; Revised 20 February 2011; Accepted 4 March 2011 Academic Editor: Hynek Burda Copyright © 2011 Thierry Lode. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Habitat selection remains a poorly understood ecological process, but relating mating behaviour to pattern of habitat selection constitutes a fundamental issue both in evolutionary ecology and in biological conservation. From radiotelemetry protocol, habitat-induced variations in mating success were investigated in a solitary mustelid carnivore, the European polecat Mustela putorius. Selection for marshy habitat was regarded as adaptive in that mating success was found greater using marches than other habitats. Males consorted with 1.3 females, revealing a low polygyny rate. Pregnant or lactating females selectively shifted to deciduous woods. That some habitat types may favour a good reproduction forms a key factor for species conservation and environmental management. Nevertheless, such as in various vertebrates, habitat requirements seem to be based on simple broad features of habitat, suggesting that habitat avoidance rather than habitat preference can explain polecat habitat predilection. 1. Introduction reproductive sites can be regarded as reflecting habitat selection [8, 12, 13]. It could be predicted that no selection Adaptive significance of a behaviourisrecognized whena occurs when unsuccessful and successful reproductive sites behaviour provides a selective advantage in animal ability to are not discernible from most of their main habitat features, adapt to its environment. Landscape structures are expected but McLoughlin et al. [12] found no variations at the home- to influence especially mating behaviour and numerous range level but differences within the home range level species required heterogeneous habitat composition. underline the importance of spatial scale dependence in Differences in habitat use among animals have been habitat selection [6, 14]. often ascribed to habitat selection although habitat selection Selection of better reproductive sites and habitat features remained a poorly known behavioural ecological process [1– should be indicative of adaptive behavioural preferences but 3]. Patterns of habitat use may reveal specific choice for can often result in increasing competition, habitat selectivity, habitat, which should provide numerous benefits such as and declining population density [4]. Habitat selection and resource availability, shelter, avoidance of predation, and nest-sites choice have been widely studied in birds although reduced interspecific competition [4, 5]. The basic conjecture differences in fitness were not often examined [8, 15, 16]. in habitat selection theory is that animals select habitat However, differences in habitat can have noticeable effects features to maximize their fitness [6–9]. Thus, habitat on reproduction [9]. The relationship between male status selection is a behaviour related to fitness and differs from and mating success was reported from social mammals such habitat use in which habitat preferences are mainly linked to as mongooses and badgers [17–19], but numerous species resources. Habitat selection can lead to a strong fidelity to lived a solitary life, and their social organisation was mainly habitat features in which fitness was previously high [10]. based on spatial pattern [20]. This is especially the case for Numerous species showed partitioning in microhabitat most carnivores, and although habitat features have been preferences, but patent demonstration for habitat selection was rare [11, 12]. Actually, natural selection can occur recognized as major parameters for species conservation (see [21]), the role of habitat selection has still received little when distinct habitat types differ in successful breeding, and putative differences between unsuccessful and successful attention. 2 International Journal of Zoology In solitary carnivores, male and female live indepen- data from radiotracked polecats, we calculated the centre dently over long periods and generally exhibit a mating of activity (arithmetic mean) from fixes, and the minimal system in which the male home range partially overlaps the dispersal distance was assessed comparing minimal distance range of one or several females [22–25]. Because carnivores between telemetry locations. were routine and did not try to avoid poor environmental Habitat use was inferred in male and female radio- conditions, it could be predicted that habitat traits have a tracked polecats based on percentage of fixes found in major influence on their mating success. a considered habitat type and was assigned to two main This study investigates the role of habitat selection on the periods, winter (November–January) and breeding period mating success of a solitary mustelid carnivore, the European (February–April for males, and May-June for females, polecat Mustela putorius. Based on a long-term survey because mating sites may differ with breeding sites in which of radiotracked animals, this study aims at characterising parturitions occurred). Field study for determining habitat habitat features and mating success to assess whether habitat features was performed between July and September from features may affect mating success (habitat selection). 1998 to 2002, considering the minimum convex polygon of each individual and, for random sites, a number of putative locations, on the basis of average number of real fixes recorded, randomly assigned within the study areas. 2. Methods We defined 19 habitat features as deciduous wood, oak Radiotracked polecats (24 males and 25 females, 1992– grove, chestnut grove, hybrid poplar plantation, orchard and 2002) were mainly surveyed from winter to spring in garden, willow grove, flooded herbaceous cover, reed-bed, ◦ ◦ Western France (Brier ` e 47 20 N, 2 10 W, lake of Grand- ditches, river bank and riparian habitat, pond, grassland, ◦ ◦ Lieu and Tenu river 47 05 N, 1 39 W), and data from each pasture, cultivation and vineyard, shrub of bramble, hedges, individual was only used once. The climate of an oceanic road and path, barn and abandoned house, and at last farm type is mild and humid (mean temperature August 20 C, and houses. Habitat diversity was assessed by the index E = ◦ 2 and mean temperature February 5 C, annual precipitation A−1/S−1, where A = 1/Σ Pi [30]and S is the total number of habitat type, considering only 6 major habitat types. An 650 mm, with about 2 snowy days per year). Every animal was surveyed 121–223 days, but numerous individuals were analysis of variance was used to investigate differences in recaptured several times. From a capture-recapture design, habitat features among male sites considering male sites with consortship success, unconsorted male sites, and random density estimates were regarded as moderately high ranging from 0.35 to 0.57 individuals per km in 2002. sites. Because all females consorted, we only discriminated The consortship behaviour consists of a brief association between habitat use from February to April (mating period) between an adult male and an adult female, showing activity and from April to June (period of parturition). Paired t-test within the same restricted area and often resting together in (with Bonferroni adjustment procedure) or Welch alternate the same den during oestrus [26, 27]. Consortship success t-test is used according to the case. was determined by trapping, incidental observations (using night vision optics NONH4U), and radiotracking locations. 3. Results Throughout the reproductive season, trapping sessions were repeatedly conducted using 20 box-traps in every studied site 3.1. Consortship Success. Moving across several female con- in order to detect the presence of other individuals during tiguous ranges, most male polecats (80%, n = 19/24) the mating season. Polecats are usually intolerant towards obtained brief consortships averaging 3.11 days (sd = 1.26, any congeners, exhibiting a strong individual segregation range 1–6 days, and n = 19 males). Males consorted with in the use of space, and associations between individuals 1.33 partners (sd = 0.82, n = 24 males). All females are restricted to the breeding period [24, 28, 29]. Because consorted with at least one male and obtained a mean of copulation induces ovulation, breeding success depends 1.12 consortships (sd = 0.33, n = 25 females; Table 1). The upon the male tenacity, and most of recaptured females consortship success was related to the consortship duration exhibited scratches on the neck and on the back resulting (r = 0.705,P< .0001, and n = 25). While consortship Spearman from the copulation, when the male remained on the female patterns in males characterised a polygynous mating system with a bite often lasting more than one hour [27]. Therefore, with a single male consorting with one to three females, some the consortship behaviour provides a good indication of the females were regarded as polyandrous. mating activity [26] although how consortship behaviour is related to breeding success remains unknown. Most of copulations occurred before Apri, and parturitions, as 3.2. Habitat Selection. During breeding period, polecats revealed by recaptures of lactating females, occurred from mainly used marshes and wooded areas, but habitat use May to June. differed between the sexes, females occupying significantly A consortship was regarded as successful when it has more woods and hedges, while males exploited mainly occurred about 42 days (gestation duration) before the marshes (χ = 42.3, P< .0001, df = 4) (Figure 1). estimated date of parturitions. On a basis of a minimum Actually, habitat preferences seemed to be only based on of six consecutive fixes per day, the minimal consortship superficial features of habitat, and marshy habitat features duration was assessed as a number of days during which and deciduous woods appeared as the most important male and female were diurnally associated in the den. For descriptors of polecat habitat. Habitat use in males was International Journal of Zoology 3 Table 1: Consortship duration and consortship success (number Habitat use of sexual partners) for 24 radiotracked males and 25 females of European polecats Mustela putorius from Western France. Number of Number Successful sexual Consortship of Con- Males partners- duration sexual sortship for (days) partners with female M1 1 F1 1 4 M2 1 F2 1 3 M3 1 F3 1 4 M4 2 F4 1 2 F5 1 1 M5 1 F6 1 1 M6 2 F7 1 5 F8 1 2 M7 1 F9 2 1 0 Males Females M8 1 F10 1 6 M9 3 F11 2 3 F12 2 3 Hedges, shrubs Marshes, ponds Deciduous woods Meadows F13 1 2 Riparian habitats M10 2 F14 1 3 Figure 1: Habitat use in male and female radiotracked polecats F15 1 3 based on percentage of fixes found in a considered habitat. Breeding M11 2 F16 1 4 males F17 1 3 M12 1 F18 1 5 Table 2: Differences among habitats used by 24 male and 25 female polecats and random sites (Anova from radiotracking locations, M13 2 F19 1 3 significant P value in bold). F20 1 4 M14 2 F21 1 5 Mean Degree of Fisher square freedom F F22 1 3 Males M15 2 F23 1 3 Deciduous woods 0.0019 1,39 0.87 .358 F24 1 3 Marshy areas 0.429 1,39 69.25 .0001 M16 1 F25 1 4 Riparian habitat 0.0033 1,39 2.47 0.125 M17 1 F17 1 2 Meadows and cultivation 0.523 1,39 104.38 .0001 M18 1 F18 1 3 Hedges and shrubs 0.021 1,39 12.42 .0011 M19 1 F20 1 2 Females M20 0 0 Deciduous woods 0.102 1,39 16.38 .0001 M21 0 0 Marshy areas 0.114 1,39 18.39 .0001 M22 0 0 Riparian habitat 0.0026 1,39 0.99 .325 M23 0 0 Meadows and cultivation 0.715 1,39 175.6 .0001 M24 0 0 Hedges and shrubs 0.056 1,39 28.9 .0001 Mean = Mean = Mean = 2.33 1.33 1.12 deserted. Moreover, polecats used significantly more shrubs chiefly influenced by willows, reedbeds, flooded herbaceous and hedges than other habitats. Female polecats were very cover, and ditches, and in females, the choice of parturition selective in their habitat preferences using significantly more sites was mainly influenced by ashes, oaks, willows, and reed- deciduous woods, hedges and shrubs, and marshy areas than beds. randomly estimated, and they poorly utilised meadows and Nonetheless, evidence of habitat selection was found cultivation. when differences among habitats, respectively, used by pole- Furthermore, habitat use differences were found accord- cats, and habitats in random sites were tested with ANOVA ing to the consortship success (Figure 2), unconsorted male (Table 2). In male polecats, marshy areas were the dominant polecats using significantly more deciduous woods and less habitat type, while meadows and various cultivations were marshy areas than consorted polecats (Table 3). Because all (%) 4 International Journal of Zoology Habitat shift in female polecats Habitat selection in males June home range Spring home range Consorted Unconsorted Hedges, shrubs Marshes, ponds Meadows Deciduous woods Barns, farms Riparian habitats Riparian habitats Hedges, shrubs Marshes, ponds Deciduous woods Meadows Figure 3: Habitat shift in surveyed female polecats from March (consortship sites) to June (parturition sites) as revealed by Figure 2: Habitat selection according to the consortship success of percentage of fixes recorded in a considered habitat. radiotracked male polecats. (n = 19 consorted polecats and n = 5 unconsorted polecats) as revealed by percentage of fixes recorded in a considered habitat. 4. Discussion Polecats shifted for marshy and wooded areas during their Table 3: Differences among habitat used by consorted (n = 19) and breeding period, but many habitat features did not clearly nonconsorted males (n = 5) (Anova from radiotracking locations, differ from random sites with regard to most of their significant P value in bold). characteristics. Studies of animal distribution in habitats often dealt Mean Degree of Fisher with habitat preferences with no assessment of mating Males P square freedom F success or remained mainly based on indirect approach Deciduous woods 0.089 1,23 4.68 .042 such as variations in density or abundance [3, 15, 31, 32]. Such approaches described habitat features, which could be Marshy areas 0.022 1,23 5.03 .035 poorly related to fitness but rather with resource availability Riparian habitat 0.034 1,23 2.05 .166 [8]. Measurement of habitat features, which were directly associated with breeding activities such as nesting, should Meadows and cultivation 0.018 1,23 0.95 .338 more reflect habitat selection [8, 13, 33, 34]. Hedges and shrubs 0.0008 1,23 0.34 .564 Habitat features associated with consortship success should at least indirectly reflect features that affect fitness. Mating sites differed from random sites only in some rough habitat features, and characteristics of successful mating surveyed female polecats consorted and bred successfully, sites have uncertain ecological meaning. Three nonexclusive it was impossible to run a same analysis. Nevertheless, causes could be evoked for such findings. Firstly, it may be pregnant or lactating females settled in distinct areas in May- alleged that every habitat type induces few differences in June switching for more woody habitats (paired t-test = 6.28, fitness, and thus, further habitat selection does not occur in P< .0001, and df = 23, Figure 3) and riparian habitats polecat. Secondly, habitat use can be regarded as a result of (paired t-test = 2.31, P = .03, and df = 23) whereas the use limited opportunities for animals to optimise when preferred of marshy areas decreased (paired t-test = 9.57, P< .0001, habitats are saturated. Thirdly, searching for the preferred and df = 23). The use of shrubs and hedges increased habitat may result in a relatively low benefit. (paired t-test = 3.58, P< .002, and df = 23) probably Because habitats features differed between random sites related to the search for a quiet parturition place. The index and mating sites, it should be recognised that polecats of habitat diversity significantly differed between space used exhibited at least a slight selection for habitat. Diversity by consorted polecats (E = 0.225, sd = 0.062) and random of habitat used by breeding polecat differed significantly sites (0.428, sd = 0.087) (Welch T-test = 11.2, P< .0001, from random sites, revealing that certain habitat were less df = 53). occupied during reproduction. Nevertheless, it could be (%) (%) International Journal of Zoology 5 suspected that low benefits in mating success resulted from per male, the polygyny rate appeared very low. Because pole- habitat selection. Habitat requirements of most vertebrates cats assiduously exploited some patchy area until resource seem to be based on simple superficial features of habitat depletion, solitary habits were basically proved to be adaptive [21, 34–38]. Thus, the stone marten exhibited a preference [24, 46, 50]. Conversely, such habits result in a low polygyny for “stony” environment and regardless of other habitat rate, suggesting an evolutionary cost induced by this specific features inhabited deciduous woods as well as barns, farms, spatial pattern. or buildings in towns [25, 39]. Anyway, that certain habitat features favour a best mating Following Haila et al. [35], it may be suggested that success constitutes a key factor for polecat conservation and habitat avoidance rather than habitat preference can explain environmental management. such predilection. Polecats were found to avoid open mead- ows and cultivation and seemed to prefer high vegetation Acknowledgments cover or undisturbed areas. Polecats showed very secretive movements in shrubs and along hedges, resting in dense The author thanks all his assistants for help in collecting the vegetation, in burrows or warrens. Habitats were selectively field data and especially Y. Benoit, Y. Biraud, H. 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