Seasonal migration of male red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) in southern Sweden and consequences for management

Red deer Cervus elaphus is a highly appreciated and intensively managed game species throughout Europe. A common management objective is a sustainable harvest of large trophies. In southern Sweden, management has mainly aimed at preserving the nominate subspecies C. elaphus elaphus. Seasonal migration of red deer males may, however, complicate both harvest management as well as conservation efforts. I used individually identified male red deer in southern Sweden to observe distance travelled from rutting areas to areas used by males in summer and winter. Adult males were identified by antler shape and photo-documented during rut. Photos from the rut were compared to trophies of deer harvested or found dead, to found cast antlers and to stags photographed during summer. From 1969 to 2007, a distance between rutting ground and summer/winter quarters was established for 96 identified stags. An average distance of 14 km and a maximum distance of 47 km were recorded between rut and summer/winter observations. The seasonal migration of males increases the risk of overexploitation of males with harvest in both rutting areas and wintering areas. Harvest management and conservation efforts may fail if males seasonally migrate outside the management unit. The results suggest that seasonal migration must be considered in harvest management and conservation and that there is a need for a regulation of male harvest. Furthermore, the study stresses that the success in deer management of single hunting units, may be largely dependent on the harvest policies in the near surroundings as well as in areas tenths of kilometres away, suggesting that a successful management must rely on co-operation and co-ordination on a landscape scale.     

Effects of immunocontraception on behavior in fallow deer (Dama dama)

Immunocontraceptives can control growth in wild and captive populations; however, in doing so, they should not disrupt species‐typical behavior patterns. The presence of treated females could disrupt social interactions in a population; yet, few studies have examined effects of immunocontraception on behavior. The goal of this project was to determine whether behavior in a fallow deer (Dama dama) population vaccinated with porcine anti‐zona pellucida (pZP)‐immunocontraceptives differed from behavior in a population of unvaccinated deer. We predicted that pZP‐treated females would spend a higher percentage of time exhibiting mating and dominance behavior than nontreated females, and that males interacting with pZP‐treated females would spend a higher percentage of time demonstrating mating and aggressive behavior than males interacting with nontreated females. We recorded activity budgets of males and females in two fallow deer populations, one pZP‐treated and the other untreated, before, during, and after rut. Females did not differ in time spent displaying mating or dominance behavior compared to nontreated females. Males coexisting with nontreated females spent more time exhibiting aggressive behavior during rut than males living with pZP‐treated females, but males did not differ in time spent in mating behavior. Thus, immunocontraception did not seem to affect behavior adversely. However, sample sizes, living conditions, and sex ratios may have affected the results. Given these limitations, future research is needed to confirm our findings. Zoo Biol 27:49–61, 2008. © 2007 Wiley‐Liss, Inc.     

Age-specific feeding cessation in male red deer during rut

Tactics for resource-use involve both using stored reserves (capital breeding) and feeding while reproducing (income breeding). In polygynous ungulates, males often use an income breeding strategy when young and shift to a capital breeding tactic at prime age. Little is known regarding why prime-aged males stop or largely reduce eating during rut but still remain inactive for as much time as before rut. A detailed exploration of how rumen content correlates with age and date may shed light on the ultimate causes of why some males stop eating. We provide quantitative empirical data on rumen content from red deer Cervus elaphus during the rutting season in Norway. In male red deer, rumen content declined with age, up to around 6 years of age. Above this age, rumen content was low and stable. A time-of-year effect on rumen content was best described with a second-order polynomial term, as rumen fill was lowest during mid-October when the mass loss of males and the ovulation rate of females both peak. We present two new hypotheses related to why males reduce eating ( physical rest and parasite hypotheses). Two related patterns need to be explained and better documented: (1) why are resting times stable before, after and during rut, and (2) why do non-prime-aged males eat more between rutting activities?     

Red deer females collect on male clumps at mating areas

Mating strategies in mammalian herbivores are adapted to thedispersion of females, and female dispersion is mainly determinedby resource dispersion, although it is frequently unclear whetherfemales may also be influenced by the location of males. Inthe red deer (Cervus elaphus) the distribution of females beforethe rut predicts the places were males should establish territoriesand even their relative success. However, the number of femalesusing the mating areas in Doñana increases during therut. We observed 20 areas of meadows, used by grazing femalesbefore the rut. At the onset of the rut, the number of females increasedin some of these areas and decreased in others, and the opposite patternwas found after the rutting period. Changes in the vegetationat mating and nonmating areas could not account for the changesin female distribution; even some of the highest quality meadowswere vacated by females during rut. In selecting the matingareas, females avoided isolated small meadows within the scrubarea and preferred larger meadows where a number of neighboringrutting males could be found. Females also avoided those areas heavilyused by fallow deer (Dama dama), a competing sympatric species.We found that females suffered less sexual harassment when inlarger harems and when their harem was surrounded by other harems.Our results, together with those in the literature about thispopulation, indicate that red deer females collect during theearly rut in mating areas containing several rutting males,although once there they may select particular sites based on availabilityof food rather than based on the presence of a particular male. Byjoining harems in large meadows they are less harassed, andat the same time they probably increase their chances of matingwith highly competitive males. The results from Doñanasupport the idea that harassment avoidance may lead to femalemovements to areas with male territories without lek breedingor female comparison of male phenotypes and may bring an insightinto those factors leading to clumps of male territories andleks.     

Population dynamics and resource use of red deer after release from harvesting in New?Zealand

Despite periods of extensive government-funded control, fluctuating commercial exploitation and ongoing recreational hunting, little is known about how red deer (Cervus elaphus scoticus L?nnberg) in New?Zealand respond to the cessation of harvesting in terms of population growth rate and resource use. We describe the population dynamics and resource use of red deer in a montane catchment over 5?years (1962?67) following cessation of intensive government-funded control in 1961. Locations and sex?age classes of deer were observed monthly along a fixed route in the Harper-Avoca catchment, inland Canterbury. A total of 2036 red deer groups were observed. The number of groups observed annually increased during the study but no trends in median (2 or 3) or modal (1 or 2) group sizes were found. Population growth rates (r) of deer were extraordinarily high in the first two years (e.g. 2.33 ? 0.22 for adult females and 1.61 ? 0.23 for adult males), but decreased in subsequent years and were not biologically possible without substantial immigration and/or changes in detectability of deer. Sexual segregation and selection of vegetation types (alpine grassland, montane grassland, and forest) and 10 topographic landforms showed stronger intra-annual than inter-annual patterns. Segregation was greatest in spring and summer, least in the rut, and variable in winter. In all seasons, sexual segregation was greatest at 25- and 50-ha scales, moderate at 100-ha, and absent at the 500- and 1000-ha scales. Selection of vegetation types also varied seasonally, with deer of both sexes preferring montane grasslands in spring and summer and alpine grasslands in the rut. Backslopes were preferred landforms in spring and summer, spurs during spring and the rut, and hollows during the rut. Our results highlight the need to consider spatial scale, immigration, and detectability in the design of red deer culling and harvesting programmes. Studies of home-range size and use, migration patterns, dispersal rates and distances are required to better understand the impacts of red deer on New?Zealand ecosystems and the effects of management on red deer populations.     

Individual variation in migratory and exploratory movements and habitat use by adult red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis> L.) in a mountainous temperate forest

Using data from a global positioning system (GPS), seven adult red deer (Cervus elaphus L.) were tracked in the Parc National des Cévennes, southern France, between November 1998 and December 2000 to assess the factors affecting large-range movement patterns and habitat use. The home range varied from a single compact area for females to three distinct seasonal ranges for males, which used alternative migratory strategies (i.e. non-, downward- and upward-migrants). The migrants used mainly southerly and easterly aspects, and wintered in areas having steeper slopes than were used during summer or the rut season. For males, the time of rut migration was mid-September and they finally entered wintering ranges from mid-December to the beginning of January. Exploratory behaviour (i.e. individuals found outside the limits of their familiar area but returning to it a few days later) occurred in both sexes and for all individuals monitored during at least a 6-month period. Velocity and efficiency of exploratory movements were higher than usual movements. During these exploratory movements, hinds may have used different landscape attributes (elevation, slope, canopy cover) while stags did not. These results provide new empirical information that could be used for building and applying broad-scale spatial and landscape use models in ecological research.     

Male red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) dispersal during the breeding season

Breeding dispersal can be of significant ecological and evolutionary importance. Yet, it is seldom considered in mammals. I present data on male red deer (Cervus elaphus) movements between sub-populations in southern Sweden during the rut. I investigated whether these movements could be breeding dispersal driven by mate competition. During the ruts of 1998–2009, I recorded 91 movements of males. The longest movement distance was 18.5 km. Dispersal was not restricted to yearlings or sub-adults, but also observed among adult stags. Of 91 movements observed, 7 were made by yearlings, 46 by sub-adults and 38 by adults. There was a significant move among yearlings and sub-adults towards areas with a higher ratio of females/adult males and towards areas with more females. The movements between rutting areas thereby seemed driven by sexual competition.     

Season effect on genitalia and epididymal sperm from Iberian red deer, roe deer and Cantabrian chamois

Seasonality deeply affects the physiology and behavior of many species, and must be taken into account when biological resource banks (BRBs) are established. We have studied the effect of seasonality on many reproductive parameters of free-ranging Iberian red deer, roe deer and Cantabrian chamois, living in Spain. Testicles from hunted animals were collected and sent to our laboratory at different times during the year. We recorded the weight and volume of testis, the weight of the epididymis and its separate parts (caput, corpus, and cauda), the weight of the sperm sample collected from the cauda epididymis, and several sperm parameters (sperm concentration, spermatozoa recovered, motility, HOS test reactivity, acrosomal status, and viability). We studied the data according to several periods, defined accordingly to each species. For red deer, we defined rut (mid-September to mid-October), post-rut (mid-October to mid-December), and non-breeding season (February). For roe deer, they were pre-rut (June), rut (July), post-rut (first fortnight of August), and non-breeding season (September). For chamois: non-breeding season (June to mid-September) and breeding season (October-November). The rut/breeding season yielded significantly higher numbers for almost all parameters. However, in the case of red deer, sperm quality was higher in the post-rut. For roe deer, testicular weight was similar in the pre-rut and in the rut, and sperm quality did not differ significantly between these two periods, although we noticed higher values in the rut. In the case of chamois, sperm quality did not differ significantly from the breeding season, but data distribution suggested that in the non-breeding season there are less males with sperm of good quality. On the whole, we find these results of interest for BRB planning. The best season to collect sperm in this species would be the breeding season. However, post-rut in red deer, pre-rut in roe deer, and non-breeding season in chamois could be used too, because of the acceptable sperm quality, despite the lower quantity salvaged. More in-depth research needs to be carried out on the quality of sperm salvaged at different times of the year in order to confirm these findings.     

Seasonal behavioral patterns of captive alpine musk deer (<Emphasis Type="Italic">Moschus sifanicus</Emphasis>): Rut and pre-rut comparisons

Alpine musk deer (Moschus sifanicus), well-known for their musk production, are endemic to western China. Due to historical unrestricted illegal hunting and habitat loss, captive farming has been employed as a means of conserving this endangered species and developing sustainable musk harvesting techniques. For captive animals, an understanding of behavioral characteristics is vital to improve management practices. This study addressed a lack of information regarding the behavioral characteristics of alpine musk deer; specifically daily activity patterns in respect to gender and reproductive season. From August 2002 to January 2003, focal sampling was employed to observe 32 adult captive alpine musk deer (13 females and 19 males), at Xinglongshan Musk Deer Farm (XMDF), located at Xinglongshan National Nature Reserve, Gansu Province, China. Results indicated that the general behavioral patterns were similar between female and male captive alpine musk deer throughout both reproductive and non-reproductive seasons (rut and pre-rut season). Both male and female alpine musk deer demonstrated tail-pasting behavior during rut season, a previously male-only behavior trait. Female musk deer also rested comparatively more than males during pre-rut season.     

Experimental evidence for density-dependence of home-range size in roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis> L.): a comparison of two long-term studies

The effect of experimental manipulation of population density on home-range size was investigated in two free-ranging roe deer (Capreolus capreolus) populations under contrasting environmental conditions. In these two long-term monitoring studies, one in Bogesund, Sweden (12 years) and one in Dourdan, France (10 years), deer density varied fourfold through varying culling pressure. Home-range data were collected by radio-tracking across the periods of contrasting density of the studies. We predicted that home-range size for females should vary in relation to the level of feeding competition, while for males, competition for mating opportunities should also influence range size, at least in summer when roe bucks are territorial. We found a highly consistent pattern over the two populations, with strong effects of deer density on home-range size, as well as significant differences between winter and summer ranges and between the sexes. Home ranges were consistently smaller at high density compared to low density. Males had larger ranges than females and this was particularly so during summer. Lastly, winter ranges were generally larger than summer ranges, particularly among females, although males at Dourdan had larger summer ranges compared to winter ranges. We suggest that the reduction of range size at high deer density during winter, as well as summer, is linked to the solitary behaviour and territorial social system of roe deer, with possible effects of dominance rank, even outside the mating season.     

Reproductive effort of bison bulls (<Emphasis Type="Italic">Bison bison</Emphasis>) in semi-natural conditions

Reproductive effort (RE) is defined as the investment in any current reproductive act. In male ungulates, variation in RE can be explained by differences in age, dominance, phase in the breeding season and number of oestrus females simultaneously present. We investigated the relation between these factors and RE in American bison males in semi-natural conditions. We looked for differences between males in RE, measured by tending behaviour, and maintenance activities (feeding/resting) during the rut. We found no effect of age on RE. During early rut there was no difference in RE between males of different rank. Dominant males had a higher RE during peak rut, and in late rut both the first and second ranking males showed more RE than lower ranking bulls. For all males, there was a clear increase in RE towards peak rut and when more females were attractive. Maintenance behaviours decreased when more females were synchronously in oestrus. The most dominant bulls spent less time grazing than lower ranking males, and all males decreased feeding during peak rut. Resting decreased with age. We conclude that dominance, phase of rut and the number of females synchronously in oestrus are the factors that best explain variation in RE.     

Female Sexual Preferences Toward Conspecific and Hybrid Male Mating Calls in Two Species of Polygynous Deer, <Emphasis Type="Italic">Cervus elaphu</Emphasis>s and <Emphasis Type="Italic">C. nippon</Emphasis>

The behavioral processes at the basis of hybridization and introgression are understudied in terrestrial mammals. We use a unique model to test the role of sexual signals as a reproductive barrier to introgression by investigating behavioral responses to male sexual calls in estrous females of two naturally allopatric but reproductively compatible deer species, red deer and sika deer. Previous studies demonstrated asymmetries in acoustic species discrimination between these species: most but not all female red deer prefer conspecific over sika deer male calls while female sika deer exhibit no preference differences. Here, we extend this examination of acoustic species discrimination to the role of male sexual calls in introgression between parent species and hybrids. Using two-speaker playback experiments, we compared the preference responses of estrous female red and sika deer to male sexual calls from conspecifics versus red × sika hybrids. These playbacks simulate early secondary contact between previously allopatric species after hybridization has occurred. Based on previous conspecific versus heterospecific playbacks, we predicted that most female red deer would prefer conspecific calls while female sika deer would show no difference in their preference behaviors toward conspecific and hybrid calls. However, results show that previous asymmetries did not persist as neither species exhibited more preferences for conspecific over hybrid calls. Thus, vocal behavior is not likely to deter introgression between these species during the early stages of sympatry. On a wider scale, weak discrimination against hybrid sexual signals could substantially contribute to this important evolutionary process in mammals and other taxa.     

Early life expenditure in sexual competition is associated with increased reproductive senescence in male red deer

The evolutionary theories of senescence predict that investment in reproduction in early life should come at the cost of reduced somatic maintenance, and thus earlier or more rapid senescence. There is now growing support for such trade-offs in wild vertebrates, but these exclusively come from females. Here, we test this prediction in male red deer (Cervus elaphus) using detailed longitudinal data collected over a 40-year field study. We show that males which had larger harems and thereby allocated more resources to reproduction during early adulthood experienced higher rates of senescence in both harem size and rut duration. Males that carried antlers with more points during early life did not show more pronounced declines in reproductive traits in later life. Overall, we demonstrate that sexual competition shapes male reproductive senescence in wild red deer populations and provide rare empirical support for the disposable soma theory of ageing in males of polygynous vertebrate species.     

Solar Radiation Determines Site Occupancy of Coexisting Tropical and Temperate Deer Species Introduced to New Zealand Forests

Assemblages of introduced taxa provide an opportunity to understand how abiotic and biotic factors shape habitat use by coexisting species. We tested hypotheses about habitat selection by two deer species recently introduced to New Zealand’s temperate rainforests. We hypothesised that, due to different thermoregulatory abilities, rusa deer (Cervus timorensis; a tropical species) would prefer warmer locations in winter than red deer (Cervus elaphus scoticus; a temperate species). Since adult male rusa deer are aggressive in winter (the rut), we also hypothesised that rusa deer and red deer would not use the same winter locations. Finally, we hypothesised that in summer both species would prefer locations with fertile soils that supported more plant species preferred as food. We used a 250 × 250 m grid of 25 remote cameras to collect images in a 100-ha montane study area over two winters and summers. Plant composition, solar radiation, and soil fertility were also determined for each camera location. Multiseason occupancy models revealed that direct solar radiation was the best predictor of occupancy and detection probabilities for rusa deer in winter. Multistate, multiseason occupancy models provided strong evidence that the detection probability of adult male rusa deer was greater in winter and when other rusa deer were present at a location. Red deer mostly vacated the study area in winter. For the one season that had sufficient camera images of both species (summer 2011) to allow two-species occupancy models to be fitted, the detection probability of rusa deer also increased with solar radiation. Detection probability also varied with plant composition for both deer species. We conclude that habitat use by coexisting tropical and temperate deer species in New Zealand likely depends on the interplay between the thermoregulatory and behavioural traits of the deer and the abiotic and biotic features of the habitat.     

Age- and density-dependent reproductive effort in male red deer

Reproductive effort in female ungulates originates from gestation and lactation and has been studied extensively; however, no comparable studies of reproductive effort in males (due to fighting for access to mates) have, to our knowledge, previously been reported. Here, we report on weight loss of male red deer during the annual mating season--a direct measure of male reproductive effort (or somatic reproductive costs). The 'terminal investment' hypothesis predicts that reproductive effort should increase with age, given that costs remain stable. We also propose the 'mating strategy-effort' hypothesis, which predicts that reproductive effort peaks in prime-aged males, since they are most often the harem holders. Consistent with the mating strategy-effort hypothesis, relative weight loss during the rutting season peaked at prime age and was lower in younger and senescent males. Weight loss during the rut was relatively smaller as density increased and more so for older males. This is probably primarily due to males (particularly senescent males) starting their rut in poorer condition at high density. The pattern of reproductive effort in males with regard to age and density therefore differs markedly from the pattern reported for females.     

Relative Effect of Food Supplementation and Natural Resources on Female Red Deer Distribution in a Mediterranean Ecosystem

ABSTRACT Supplementary feeding is a widespread game management practice in several red deer (Cervus elaphus) populations, with important potential consequences on the biology of this species. In Mediterranean ecosystems food supplementation occurs in the rutting period, when it may change mating system characteristics. We studied the role of food supplementation relative to natural resources in the spatial distribution, aggregation, and mean harem size of females in Iberian red deer (Cervus elaphus hispanicus) during the rut. We studied 30 red deer populations of southwestern Spain, 63% of which experienced supplementary feeding. Using multivariate spatial analyses we found that food supplementation affected distribution of females in 95% of the populations in which it occurred. Green meadows present during the mating season acted as an important natural resource influencing female distribution. Additionally, the level of female aggregation and mean harem size were significantly higher in those populations in which food supplementation determined female distribution than in populations in which female distribution did not depend on supplementary feeding. Because female aggregation and mean harem size are key elements in sexual selection, supplementary feeding may constitute an important anthropogenic element with potential evolutionary implications for populations of Iberian red deer.     

Sex differences in weather sensitivity can cause habitat segregation: red deer as an example

Sex differences in habitat use (habitat segregation) are widespread in sexually dimorphic ungulates. A possible cause is that males are more sensitive to weather than females, leading to sex differences in sheltering behaviour (the 'weather sensitivity hypothesis'). However, this hypothesis has never been tested. We considered the allometric rates of net energy gain during times of cold weather and food shortage in a model. We argue that the higher absolute heat losses relative to intake rates of larger ungulates should indeed lead to higher weather sensitivity in males than in females. Furthermore, we tested the weather sensitivity hypothesis empirically in red deer, Cervus elaphus, on the Isle of Rum, U.K. We predicted that (1) use of relatively exposed, high-quality forage habitat should be negatively influenced by bad weather; and (2) this influence should be stronger in males. We found that bad weather (strong wind, low temperature, heavy rain) in winter and spring influenced use of high-quality forage habitat negatively in all deer; that adult males responded more strongly to low temperature and strong wind than did females; and that adult males foraged on windy days at better sheltered sites than did females. Thus, the weather sensitivity hypothesis is supported both theoretically and empirically. We suggest that the weather sensitivity hypothesis can potentially explain winter habitat segregation in a large number of ungulate species. Copyright 2000 The Association for the Study of Animal Behaviour.     

Evaluation of Shifts in White-Tailed Deer Winter Yards in the Adirondack Region of New York

ABSTRACT In the Adirondack region of northern New York, USA, severe weather and deep snow typically force white-tailed deer (Odocoileus virginianus) to congregate in areas of dense coniferous cover and along watercourses at lower elevations. We examined 16 yards in the Adirondacks and explored the observation that deer have changed their movement behavior to incorporate residential communities within their wintering areas. We compared locations of deer herds in 2003 and 2004 to deer wintering areas mapped during the 1960s and 1970s. Deer were predominantly absent in 9 of 16 historical yards but were present in residential communities within the same drainage. Yarding areas to which deer shifted contained more residential, deciduous, and mixed cover than yards where no shift occurred, indicating that deer in residential areas were using conifer and mixed cover at a finer scale than deer in nonresidential areas. Smaller winter ranges and core areas of marked deer in a residential winter yard further imply greater concentration of resources available in these areas. Marked deer demonstrated flexibility in core winter range fidelity, a behavior that allows for more permanent shifts as habitat and food resources change or as new areas with appropriate resources are encountered. Our study suggests that low-density residential areas in lowland conifer forests may provide an energetic advantage for deer during winter due to the assemblage of quality habitat interspersed with open areas and a variety of potential food sources in environments where movement is typically constrained by deep snow. Managers should consider the potential for changes in use of deer wintering areas prior to land conservation efforts and may need to adapt management strategies to reduce conflicts in communities occupied by deer during winter.     

To Feed or Not to Feed? Testing Different Hypotheses on Rut‐Induced Hypophagia in a Mountain Ungulate

In many ruminant species, males dramatically reduce forage intake during the rut. To date, different hypotheses have been suggested to explain this rut‐induced hypophagia. To assess the predictions of the main hypotheses, we analysed Alpine ibex (Capra ibex) activity budget and compared the behaviour of males and females before, during, and after the rut. Only males spent significantly less time foraging during the rut than outside of it, whereas females allocated a similar proportion of time to foraging before, during, and after the rut. Our results showed that during the rut males also reduced lying time, while the ratio of time spent feeding to time spent lying did not change for males among periods. In conclusion, during the breeding season males maximized energy intake when not actively engaged in mating activities and rut‐induced hypophagia appeared to result from time budget constraints generated by mating‐related activities. Accordingly, the foraging constraint hypothesis seems appropriate to explain this phenomenon in Alpine ibex males.     

Male Alpine Chamois Occupy Territories at Hotspots Before the Mating Season

Ungulate mating systems vary broadly both between and within species. Studies on mating systems in different habitats can provide clues to the ecological factors determining this diversity. Despite its abundance in the European Alps and its importance as a game species, surprisingly little is known about the mating system of Alpine chamois Rupicapra rupicapra rupicapra. We tested the hypothesis that adult males first defend mating territories in late spring, when females segregate from males and well before the Nov. rut. In the Gran Paradiso National Park (north‐western Italian Alps), adult males shared a winter range but occupied individual ranges in summer and early autumn. Males were more aggressive to each other in the summer than in the spring. A strong site fidelity from one year to the next was found for the summer and early autumn months. Those males that occupied the same territories both in the summer and during the rut (Nov.) appeared to be at hotspots, attractive to females during the rut because of reduced snow cover. Other males appeared to cluster around these hotspots during the rut. Territories that were first occupied during the summer were visited by more females than those that were not established until the rut began. Our results suggest that the mating system of this population of Alpine chamois consists of the early occupation of clustered mating territories. The early establishment of mating territories in areas frequented by females during the rut may lead to reproductive benefits for male chamois.