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.     

Social and Spatial Segregation in Alpine Ibex (Capra ibex) in Bargy,French Alps

Outside the rut, a gradual social and spatial segregation was observed between the sexes as male Alpine ibex (Capra ibex) aged. In addition, males joined similar-age peer groups and segregated spatially as the age difference increased. The patterns varied seasonally, influenced by biological and ecological events. During the rut, males and females intermingled and used greatly overlapping ranges. Males ≥ 9 years first left female groups but social and spatial segregation were not complete until May. Younger males shifted to male groups more gradually. Females and males over 2 years of age used almost exclusive ranges in summer and early autumn. Social grouping and range overlap between ibex classes were correlated outside the rut. Grouping and spatial overlap between young and old males were negatively related to association with females.     

Male reproductive pattern in a polygynous ungulate with a slow life-history: the role of age, social status and alternative mating tactics

According to life-history theory age-dependent investments into reproduction are thought to co-vary with survival and growth of animals. In polygynous species, in which size is an important determinant of reproductive success, male reproduction via alternative mating tactics at young age are consequently expected to be the less frequent in species with higher survival. We tested this hypothesis in male Alpine ibex (Capra ibex), a highly sexually dimorphic mountain ungulate whose males have been reported to exhibit extremely high adult survival rates. Using data from two offspring cohorts in a population in the Swiss Alps, the effects of age, dominance and mating tactic on the likelihood of paternity were inferred within a Bayesian framework. In accordance with our hypothesis, reproductive success in male Alpine ibex was heavily biased towards older, dominant males that monopolized access to receptive females by adopting the ‘tending’ tactic, while success among young, subordinate males via the sneaking tactic ‘coursing’ was in general low and rare. In addition, we detected a high reproductive skew in male Alpine ibex, suggesting a large opportunity for selection. Compared with other ungulates with higher mortality rates, reproduction among young male Alpine ibex was much lower and more sporadic. Consistent with that, further examinations on the species level indicated that in polygynous ungulates the significance of early reproduction appears to decrease with increasing survival. Overall, this study supports the theory that survival prospects of males modulate the investments into reproduction via alternative mating tactics early in life. In the case of male Alpine ibex, the results indicate that their life-history strategy targets for long life, slow and prolonged growth and late reproduction.     

不同年龄雄性北山羊发情期时间分配特征

前人研究表明,可多次繁殖的反刍物种其雄性个体在发情期采食时间显著减少。目前有两个假说解释这一现象,即能量摄入最大化假说和能量保存假说。为验证雄性北山羊在不同发情阶段所采取的能量保存策略,作者于2014年10-12月在新疆天山中部采用焦点动物取样法采集数据,采用Kruskal-Wallis检验和Spearman秩相关性检验分析数据。研究发现雄性北山羊成体和亚成体发情期采食时间均显著低于发情前期和发情后期,但二者在不同发情阶段卧息时间无显著变化,发情期采食和卧息时间比亦显著降低。发情期发情行为时间显著高于发情前期和发情后期,非发情时间主要用于采食。发情期采食时间和卧息时间都与发情行为时间呈显著的负相关关系。雌性北山羊发情期采食时间亦显著低于发情前期和发情后期,发情期和发情后期卧息时间显著高于发情前期。本研究结果表明,发情期不同年龄阶段雄羊都主要采取能量摄入最大化策略,但同时也具有部分能量保存策略的特征。     

Group dynamics of the Nubian ibex (Capra ibex nubiana) in the Tuwayiq Canyons, Saudi Arabia

Group dynamics of the Nubian ibex ( Capra ibex nubiana ), a sub-species adapted to arid environments, were studied at the Ibex Reserve, Saudi Arabia. Most of the animals (83%) were seen in mixed herds during the rut and post-rut. Mean group size of all sex-age categories was 7.2, with the majority of the ibex seen in groups of < 10. The sex ratio of adult animals was equal but that of juveniles, older than six months, was skewed towards females. Habitat structure, sparse vegetation cover and a small population size are factors that favour small group size. An equal adult sex ratio and a high number of juveniles indicate that the population has a high reproductive potential.     

Behavioral response of 12 ungulate species in captivity to the presence of humans

Twelve species of captive ungulates were studied to determine behavioral responses to the presence of a zookeeper within the exhibit and in front of the exhibit, with and without zoo visitors present. Significant differences in behavior occurred between species for nearly all behaviors observed. A significantly greater occurrence of vigilance and approach behavior was directed toward the zookeeper while within the exhibit relative to when the zookeeper stood in front of the exhibit. A significantly lower occurrence of eating or drinking occurred when the zookeeper was inside the exhibit. Significant differences occurred across size categories in the occurrence of approaching the zookeeper. The most frequently scored behavior was visual orientation. Statistically significant differences in the occurrence of visual orientation directed by females toward the zookeeper existed across size category, with more vigilance by species with larger body size. A statistically higher occurrence of vigilance toward the zookeeper was directed by female ungulates but not by males when the zoo was closed to the public. Although no statistical significance was found regarding the intraspecific vigilance of males, data on females revealed significant differences across species and across size categories. When data regarding vigilance toward the public were analyzed, statistically significant differences existed between species for females only. Likewise, when data regarding interspecific vigilance were examined, statistically significant differences were found across size categories for females, but not for males. The potential roles of vigilance in the wild are discussed in reference to its role in captivity.     

三种珍稀有蹄类动物的警戒行为数据

警戒行为是野生动物重要的反捕食策略之一, 警戒行为的研究已经成为行为生态学中继动物的采食行为、繁殖行为之外的另一热点研究领域。然而, 尽管研究人员至今已经在数百种鸟类、兽类中开展了各种类型的警戒行为学研究, 但相关研究的原始数据却鲜有共享。本文公开了利用焦点取样法采集的我国3种珍稀有蹄类动物警戒行为数据样本716个, 其中普氏原羚(Procapra przewalskii) 175个、藏原羚(P. picticaudata) 180个、麋鹿(Elaphurus davidiamus) 361个, 并同时公布了采集该行为学数据时的相关变量, 包括性别、群体类型、群体大小、捕食风险、人类干扰等。结果显示在上述3种有蹄类动物中, 均存在明显的集群效应, 且此效应与性别及捕食风险存在交互作用。本文尝试建立标准的行为学数据规范, 以期为未来的行为学研究数据的共享及深度挖掘提供可能。     

Vigilance behavior during the birth and lactation season in naturally occurring ring-tailed lemurs (Lemur catta) at the Beza-Mahafaly Reserve,Madagascar

I examined the vigilance behavior of adult males and females in two groups of ring-tailed lemurs(Lemur catta)during the birth and lactation season at the Beza-Mahafaly Reserve, southwestern Madagascar. I found no sex difference with respect to the rates of overall vigilance, rates of vigilance toward a potential predator or unfamiliar sound, or rates of vigilance toward conspecifics from other social groups, nor were there sex differences in the percentage of time spent vigilant in any of the above categories. Higher-ranking females were vigilant significantly more often toward predators or potential predators than lower-ranking females were. I detected no relationship between vigilance behavior and dominance rank among adult males. The alpha female in each group exhibited significantly more vigilance behavior than all other members of her group. It was predicted that males should exhibit more vigilance behavior than females do, particularly during the birth and lactation season, when predator pressure is high, if they are benefiting females in this respect. I discuss the results in the context of this prediction and in terms of how ring-tailed lemur males benefit females, and why they may be tolerated in social groups.     

Spatial behaviour of adult male Alpine ibex<Emphasis Type="Italic">Capra ibex ibex</Emphasis> in the Gran Paradiso National Park,Italy

During a two year preliminary study, the spatial organization of a group of male Alpine ibexCapra ibex ibex Linnaeus, 1758 was examined in the Gran Paradiso National Park, Western Italian Alps, Italy. From December 1995 to January 1998 we measured annual, seasonal home range and home range during the rut, plus altitudinal migration of 13 radio-collared adult Alpine ibex. The small annual home range size showed a traditional use of space, confirmed by the high overlapping values between home ranges of consecutive years: the ibex used the same places from year to year. This was also true during periods of rut. Home ranges closely overlapped in consecutive ruts, while their size changed from winter to winter. Snow cover limited the movements of the ibex; winter and spring home ranges were smaller than those in summer and autumn. Mean vertical movement patterns were similar in the two years, showing the highest values in summer and the lowest in spring. Space use was never proportional to availability for each altitudinal range.     

Seasonal and sexual variation in vigilance behavior of goitered gazelle (<Emphasis Type="Italic">Gazella subgutturosa</Emphasis>) in western China

Animals receive benefits from social behavior. As part of a group, individuals spend less time having to be vigilant. This phenomenon, called the “group size effect,” is considered the most dominant factor in an animal’s demonstrated level of vigilance. However, in addition to group size, many other social and environmental factors also influence the degree of vigilance, including the season of the year and the sex of the individual. In our study, we examined the vigilant behavior of goitered gazelles in the Xinjiang Province in western China to test whether and how seasons, the yearly breeding cycle, and group size affect vigilance. According to our results, we found that seasonal factors were not a substantial influence on a gazelle’s level of vigilance, while group size had a tangible effect. In comparison, the yearly breeding cycle (a natural phenomenon) was the most powerful factor: it significantly changed the degree of vigilance in females during birthing and males during rut. Anthropogenic factors (unnatural phenomena) were also potential causes of increased vigilance in both sexes during winter.     

青藏高原异地半圈养藏羚警戒行为的适应性研究

行为是反映动物应对环境变化的最直接形式。动物可以根据周围环境条件的变化以及自身的生理状况来调整行为,异地放养是保护珍稀动物的有效方法,但必然会对其行为产生影响。为了探讨藏羚(Pantholops hodgsonii)对异地环境的行为学适应,对异地圈养藏羚的警戒行为进行了不同季节间的比较研究,采用全事件记录法和焦点动物取样法,记录和统计了异地圈养藏羚在冷季和暖季的警戒行为,进而推测其对人类干扰的行为适应性。研究结果表明,雌性和雄性藏羚的警戒时间及警戒比例(警戒时间占全天活动时间的比例)在暖季存在显著差异(警戒时间:Z=4.36,P<0.05;警戒比例:Z=4.559,P<0.05),而在冷季则无差异(警戒时间:Z=0.001,P>0.05;警戒比例:Z=0.0014,P>0.05);而季节差异对雌、雄性藏羚的警戒时间、警戒比例均具有极显著的影响(雄性-警戒时间:F=31.758,P<0.01;警戒比例:F=21.768,P<0.01;雌性-警戒时间:F=14.98,P<0.01;警戒比例:F=11.05,P<0.01);但是季节和性别对藏羚警戒行为的影响没有交互作用(Z=?0.576,P>0.05)。这些结果提示异地圈养藏羚警戒行为的变化可能是对陌生环境适应的结果。     

Social Interactions of Yearling Male Fallow Deer during Rut

Despite a low probability of achieving copulations, yearling male (1.5-year-old) fallow deer Dama dama display behavioural changes during rut, interacting with both females and older males. Juvenile interactions in polygynous ungulates may affect further reproductive success through both motor training for future interactions and/or competition for current resources that affect growth. To test this hypothesis, we intensively studied the behaviour of 12 yearling males in two captive populations during a rutting season. Interactions with females were interpreted as motor training aimed at learning to recognize oestrous females, because yearling males rarely interacted with their mother and because interactions essentially consisted of olfactory investigations focused on oestrous females. Because agonistic interactions with older males never involved physical contact and were mainly initiated by the most dominant buck in each population, we concluded that these interactions resulted from the presence of the yearlings close to oestrous females defended by rutting males. Heavier yearlings tended to be dominant over lighter ones and tended to interact more frequently with other animals, suggesting that early growth may influence the ontogeny of social behaviours.     

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.     

Effects of reproductive status, social rank, sex and group size on vigilance patterns in Przewalski's gazelle

Background

Quantifying vigilance and exploring the underlying mechanisms has been the subject of numerous studies. Less attention has focused on the complex interplay between contributing factors such as reproductive status, social rank, sex and group size. Reproductive status and social rank are of particular interest due to their association with mating behavior. Mating activities in rutting season may interfere with typical patterns of vigilance and possibly interact with social rank. In addition, balancing the tradeoff between vigilance and life maintenance may represent a challenge for gregarious ungulate species rutting under harsh winter conditions. We studied vigilance patterns in the endangered Przewalski''s gazelle (Procapra przewalskii) during both the rutting and non-rutting seasons to examine these issues.

Methodology/Principal Findings

Field observations were carried out with focal sampling during rutting and non-rutting season in 2008–2009. Results indicated a complex interplay between reproductive status, social rank, sex and group size in determining vigilance in this species. Vigilance decreased with group size in female but not in male gazelles. Males scanned more frequently and thus spent more time vigilant than females. Compared to non-rutting season, gazelles increased time spent scanning at the expense of bedding in rutting season. During the rutting season, territorial males spent a large proportion of time on rutting activities and were less vigilant than non-territorial males. Although territorial males may share collective risk detection with harem females, we suggest that they are probably more vulnerable to predation because they seemed reluctant to leave rut stands under threats.

Conclusions/Significance

Vigilance behavior in Przewalski''s gazelle was significantly affected by reproductive status, social rank, sex, group size and their complex interactions. These findings shed light on the mechanisms underlying vigilance patterns and the tradeoff between vigilance and other crucial activities.     

Contrasting Alternative Hypotheses to Explain Rut‐Induced Hypophagia in Territorial Male Chamois

Male ungulates in temperate environments often show a severe reduction in time spent foraging during the mating season. Several hypotheses have been put forward to explain this phenomenon but, so far, no study investigated the proximate mechanisms underlying rut‐induced hypophagia in ungulates using alternative mating tactics (AMTs). Between the pre‐rut and post‐rut of 2011 and 2012, we collected data on activity budgets, parasite burden and androgen levels of territorial and non‐territorial male Alpine chamois Rupicapra r. rupicapra in the Gran Paradiso National Park (Italy). We aimed to investigate whether AMTs showed similar reduction in time spent foraging during the mating period and to test the predictions underlying alternative hypotheses that may explain rut‐induced hypophagia. Only territorial males showed a significant reduction in time spent foraging during the rut; the lack of correlation between proportion of time spent foraging and androgen metabolites or parasite burden did not fully support the physiological and the parasite hypotheses, while the foraging constraint, the energy‐saving and the physical rest hypotheses could not be discounted. Territorial males decreased the time spent lying down from the pre‐rut to the rut, but not their foraging‐to‐lying‐down ratio. During the mating period, we found negative correlations between time spent foraging or lying down and time spent rutting. Our data suggest that territorial males’ behaviour is more consistent with the foraging constraint hypothesis than with the energy‐saving hypothesis previously suggested. Yet, during the rut territorial males did not maximise their foraging time, and the optimisation of their energy balance could rather depend upon feeding on relatively high‐quality plants. This suggestion – possibly named ‘forage quality hypothesis’ – now requires further investigations. This work showed that alternative mating behaviours may underlie different patterns of foraging strategies: we suggest that tests of alternative hypotheses to explain rut‐induced hypophagia within ungulate populations should not ignore the occurrence of AMTs.     

Heterogeneity in male horn growth and longevity in a highly sexually dimorphic ungulate

In sexually dimorphic ungulates, sexual selection favoring rapid horn growth in males may be counterbalanced by a decrease in longevity if horns are costly to produce and maintain. Alternatively, if early horn growth varied with individual quality, it may be positively correlated with longevity. We studied Alpine ibex Capra ibex in the Gran Paradiso National Park, Italy, to test these alternatives by comparing early horn growth and longevity of 383 males that died from natural causes. After accounting for age at death, total horn length after age 5 was positively correlated with horn growth from two to four years. Individuals with the fastest horn growth as young adults also had the longest horns later in life. Annual horn growth increments between two and six years of age were independent of longevity for ibex whose age at death ranged from 8 to 16 years. Our results suggest that growing long horns does not constrain longevity. Of the variability in horn length, 22% could be explained by individual heterogeneity, suggesting persistent differences in phenotypic quality among males. Research on unhunted populations of sexually dimorphic ungulates documents how natural mortality varies according to horn or antler size, and can help reduce the impact of sport hunting on natural processes.     

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.     

Highly Competitive Reindeer Males Control Female Behavior during the Rut

During the rut, female ungulates move among harems or territories, either to sample mates or to avoid harassment. Females may be herded by a male, may stay with a preferred male, or aggregate near a dominant male to avoid harassment from other males. In fission-fusion group dynamics, female movement is best described by the group’s fission probability, instead of inter-harem movement. In this study, we tested whether male herding ability, female mate choice or harassment avoidance influence fission probability. We recorded group dynamics in a herd of reindeer Rangifer tarandus equipped with GPS collars with activity sensors. We found no evidence that the harassment level in the group affected fission probability, or that females sought high rank (i.e. highly competitive and hence successful) males. However, the behavior of high ranked males decreased fission probability. Male herding activity was synchronous with the decrease of fission probability observed during the rut. We concluded that male herding behavior stabilized groups, thereby increasing average group size and consequently the opportunity for sexual selection.     

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.     

Association patterns and kinship in female reindeer (Rangifer tarandus) during rut

Non-random association of females has been reported in many social species, with kinship being the main mechanism in shaping these associations. Understanding the patterns of association during rut is particularly important in polygynous animals, as this may have implications for inbreeding vulnerability and genetic structuring of populations. However, these patterns are highly dynamic and may be influenced by the population sex ratio and male age structure. We examined females' association in reindeer (Rangifer tarandus), a highly polygynous ungulate, to determine whether genetically related females associate more often than non-kin during rut and whether this association was influenced by the aforementioned factors. During three rutting seasons the male age structure and the sex ratio of the herd was manipulated: two extremely female biased sex ratios, one with only adult males and one with only young males, whereas the third being closer to a more even sex ratio and male age structure. We found that genetic relatedness influenced female association in the mixed male age group, but not in the adult and young male age groups. Highly related females (relatedness ??0.5) associated more closely than unrelated females. Further, period of the rut as well as male age structure and sex ratio had a significant effect on female association. Our study suggests that the female reindeer association during rut is not only influenced by preference for kin but also by male?Cmale competition and male age as well.