Time Allocation during the Reproductive Season in Vicuñas

Vicuñas (Vicugna vicugna) are wild South American camelids that live in high-altitude grasslands (Puna). Their social organization is based on family and bachelor groups. The amount of time they allocated to walking, running, lying, grazing and alert behaviour was studied in Abrapampa, northwestern Argentina, during the 1988–1990 reproductive seasons. Females and yearlings spent significantly more time grazing, and less time walking, running and alert than territorial males and bachelors. This result suggests that females maximized the time spent foraging to fulfil the energetic demands of reproduction. The alertness of the territorial males increased and the grazing decreased with the number of females in the families, suggesting that males that incurred a higher cost of lost feeding time through vigilance activities increased their mating success.     

青藏铁路对雄性藏羚行为时间分配的影响

行为时间分配是动物对资源获取和风险防御的权衡,受到诸多因素影响。本研究采用目标动物观察法记录了可可西里冬季（交配季）雄性藏羚的行为表现,并将行为分为觅食、警戒、卧息、移动和“其他”5种类型。首先,本文通过对比青藏铁路运营前后两个交配季（2003-2004年建设期和2017-2018年运行期）之间雄性藏羚的行为时间分配差异,探讨青藏铁路的出现对雄性藏羚行为的影响。随后,基于2017-2018年交配季雄性藏羚的行为数据,分析了年龄对其行为时间分配的影响。结果表明：与建设期相比,雄性藏羚的觅食和“其他”行为时间比例明显增加,而警戒和卧息的行为时间比例显著降低,这说明铁路稳定运营后对雄性藏羚的影响降低。交配群中成年雄性藏羚的警戒、移动,以及“其他”行为中的繁殖行为的时间比例显著高于亚成体,而觅食和卧息行为时间比例显著低于亚成体, 这与交配群中不同年龄雄性个体所处的地位等级有关。成年雄性藏羚在交配群中占据主导地位,拥有更多配偶资源,增加警戒和移动,减少觅食和卧息有助于其维持交配群的稳定。     

Fighting behaviour in territorial male roe deer <Emphasis Type="Italic">Capreolus capreolus</Emphasis>: the effects of antler size and residence

In all areas where they have been studied, male roe deer are believed to have a territorial mating system, although few quantitative studies have been conducted and there remains considerable debate about the function of male roe deer territories. We observed 139 aggressive interactions between male roe deer in Storfosna Island (Norway) during one territorial season (March–August). We recognised seven rank levels of escalation according to the potential danger of the behaviour. On the basis of the number of escalation levels included in the interactions, the complexity of the fights was also scored. We recorded the presence of other individuals during the interaction, the age, the antler size, the territorial status and the residency status of the two contestants and tested how these variables affected escalation, complexity and outcome of the fights. Most of the interactions ended with low levels of escalation, and physical contact occurred only in fights between two territorial bucks. The escalation was also affected by the difference in antler size index (the bucks escalated more when the difference in antler size was smaller) and increased with an increasing number of female deer present during the interaction. The resident buck won in 81% of the fights. When it drew or lost, it was generally both inferior in age and antler size, and the duration and escalation of the interactions were higher. However, even when a fight was lost, no territory loss occurred. These results are consistent with the evolutionary game theory and the proposed low risk–low gain strategy of roe deer bucks.     

Home range dynamics of male roe deerCapreolus capreolus in a mountainous habitat

Male spacing behaviour of roe deer Capreolus capreolus (Linnaeus, 1758) was studied in a wooded mountainous habitat in the Casentinesi Forest National Park, Italy. Data were collected using radio-tracking techniques from March 1997 to February 1998. Annual, seasonal, and bimonthly home ranges were analysed. Different factors may influence male spacing behaviour throughout the year. Winter home range sizes may be dependent on environmental conditions, while social factors could determine a high level of individual variability during the territorial and reproductive period. Prime age males showed great spatial stability, in contrast to the ranging movements of yearlings.     

To scan or not to scan? Occurrence of the group‐size effect in a seasonally nongregarious forager

Group‐living requires a compromise between safety and direct/indirect costs for individuals. The larger is the group, the greater is the collective vigilance, leading to a greater net food intake per forager because of the time saved individually from scanning behaviour. In turn, individual alertness usually decreases with increasing group size (“group‐size effect”). Information on the occurrence of group‐size effect is still unclear. Previous studies have shown that it may fail to occur or even reverse, for example when costs of interference between conspecifics are high. In turn, assessing whether the group‐size effect would occur in weakly or seasonally gregarious species may help to understand its drivers. We evaluated the occurrence and the extent of group‐size effect in a seasonally nongregarious herbivore, the roe deer Capreolus capreolus. We examined the roles of sex/age class and season as drivers of vigilance behaviour. In roe deer, the group‐size effect did not depend on sex/age class: time spent foraging increased with increasing group size; in turn, vigilance increased with decreasing group size, in all sex/age classes. Females with fawns were the most vigilant sex/age class, thus revealing the cost of offspring protection. Accordingly, the higher spring vigilance levels of does could be related to reproductive costs (e.g., defence of newborn fawns). Conversely, the greater summer vigilance of bucks could result from patrolling/defence of territories. Both adult males and females also showed the higher vigilance in winter, likely because of an increase in the perception of predation risk and/or, possibly, hormones linked to an increase in intolerance of conspecifics, in males. However, the group‐size effect occurred in all the seasons, for adult males and females. Our findings suggest that foraging benefits provided by the group‐size effect may have overcome costs of group‐living, even in a weakly gregarious forager.     

Mutual benefits of associations between breeding and non-breeding White-fronted Geese Anser albifrons

The relationships between yearlings and adult pairs of White-fronted Geese Anser albifrons were studied during pre-nesting, laying and early incubation in the central Canadian Arctic. Prior to nesting, females of lone pairs spent 75–81% of their time feeding, while males spent only 42–47% of time feeding and 46–50% alert. In pairs with one or more associated yearlings, both females and males fed significantly more and spent less time vigilant. Yearlings spent significantly less time (59%) feeding when alone compared with 71–76% when with pairs. Associations between yearlings and paired adults were most frequent before adult females began prospecting for nest sites. No prospecting pairs were associated with yearlings. After the egg-laying period, groups of geese, predominantly yearlings, made distraction flights over humans and terrestrial predators approaching nests, in contrast to the more cryptic behaviour of nesting pairs. The presence of groups of geese associated with some nest sites suggests that continuing parent-offspring relationships may involve assistance with nest defence.     

MOLTING PHENOLOGY OF HARBOR SEALS ON TUGIDAK ISLAND, ALASKA

We documented the progression and timing of the annual molt of harbor seals on Tugidak Island, Alaska, from 1997 to 1999. In all years the timing of molting differed among age-sex classes. Yearlings molted first, subadults second, adult females third, and lastly adult males. Timing of molting was nearly identical in 1997–1998, whereas in 1999 molting occurred three to six days later for all age-sex classes except yearlings. Estimated dates when peak proportions of each age-sex class were molting ranged from 2 August (yearlings) to 2 September (adult males). The number of seals hauled out was positively related to the proportion of seals in the molt and negatively related to the proportion of seals in the postmolt. Population trend estimates, based on aerial counts conducted during a narrow window within the molting period, are likely biased toward certain age-sex classes. Statistical models used to estimate trend include covariates to help account for within-year variation in seal numbers, but do not account for compositional changes that occur during molting. Population modeling may elucidate the effects of within-year population structure on trend estimates. Monitoring molting phenology at additional sites is necessary to determine the extent of geographic variation in molting.     

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.     

Habitat quality influences relative antler size and hunters’ selectivity in roe deer

Understanding factors affecting antler size, and the extent to which harvesting is selective for these traits, is important in order to address management strategies aimed to minimize the risk of negative evolutionary consequences. In an Alpine study area, we compared the phenotypic quality and the antler size of 2,725 male roe deer hunted in two regions differing for winter harshness and habitat quality, and evaluated whether the selective behaviour of recreational hunters was influenced by phenotypic quality and antler size. Antler length and antler circumference relative to both body mass and jaw length were larger in the region with more favourable climate and habitat conditions, indicating that here roe deer were able to allocate more resources to antler growth. The analysis of the temporal trends of harvest bags suggested that hunters did not select roe deer for their body mass or size, but instead for antler size. This resulted also in a preference for sub-adult and adult age classes, while yearlings were culled reluctantly, especially in the region where antlers were smaller. Our results indicate that environmental heterogeneity may influence the relative investment in antler growth. In this way, it may interact with the hunters’ preferences increasing the risk that recreational hunting of roe deer, which is a widespread practice in many European countries, might result in alteration of male age structure and possibly in directional artificial selection.     

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.     

鄱阳湖围垦区藕塘越冬白鹤的时间分配与行为节律

2016年12月—2017年3月,采用瞬时扫描法研究了鄱阳湖围垦区藕塘生境中白鹤Grus leucogeranus越冬期的行为,共扫描2560次,23219只次,包括18031只次成鹤和5188只次幼鹤。结果表明,觅食(41.78%)、警戒(25.02%)、修整(15.00%)和休息(10.84%)是白鹤越冬期的主要行为。与自然生境相比,藕塘生境中白鹤主要采取多休息和修整的策略来节省能量支出。成鹤的觅食行为(35.29%)显著低于幼鹤(62.42%)(F_(1,12)=45.977,P0.01),警戒行为(28.66%)则显著高于幼鹤(10.26%)(F_(1,12)=38.975,P0.01)。家庭群成鹤觅食行为(43.96%)极显著高于非家庭群成鹤(27.04%)(F_(1,12)=60.169,P0.01)。家庭群成鹤需要喂食幼鹤,它们花费更多的时间觅食弥补能量的消耗。行为节律上,白鹤各时段觅食行为占总行为的比例均较高,11:00—11:59出现明显高峰,占总行为的48.64%。警戒行为无明显的低谷和高峰。成鹤在各时段的觅食行为比例均明显低于幼鹤,成鹤和幼鹤的觅食曲线变化趋势相似,即觅食比例的升高和降低比较同步。成鹤在各时段的警戒行为和修整行为比例均明显高于幼鹤,成幼鹤的休息行为比例在各时段差别不大,均在14:00—14:59有个明显的高峰。家庭群成鹤的觅食行为比例几乎在各时段均明显高于非家庭群,家庭群成鹤的修整行为和休息行为在大部分时段均低于非家庭群成鹤。因此家庭群成鹤采取多取食,少休息和修整的策略提高自身的适合度,同时保证对后代的抚育。     

Territory Establishment and Antler Cycle in Male Roe Deer

Territory establishment and antler cycle in male roe deer, Capreolus capreolus, were studied in two areas in south-eastern Sweden during 1989–1993. Initial territorial behaviour was observed in early Mar., when overlap between male ranges decreased from 66 % to 0 % over a 3–6-wk period, as determined from telemetry data. The decrease was associated with an increased number of agonistic interactions. Most territories were established at the end of Mar. to early Apr., with a tendency for males re-establishing old territories to be first. In all years, territories were taken up 1–3 wk before males shed velvet from their antlers. The distance between geometric centres of neighbouring males in winter and after territory establishment increased from 164 to 364 m. Former territory holders moved the geometric centres of their winter range and subsequent territory a shorter distance than first-year territorials, 129 and 267 m respectively. Territory defence ceased after the rut in late Aug., and the spatial relations between neighbours reverted to the situation before the territorial season. Neither velvet shedding nor antler casting was correlated with male age (yearlings excluded), which contradicts earlier statements. Neither were there any associations between timing of antler casting and subsequent velvet shedding, nor vice versa. Hence, a male casting antlers early one year derives no advantage by early shedding the next season. Furthermore, to be in hard antlers is not a prerequisite for territory take-up, whereas antlers probably are necessary for maintaining the area throughout the season. Functional aspects on roe buck territoriality are also discussed.     

Territorial cooperation and social monogamy: factors affecting intersexual behaviours in pair-living snapping shrimp

Among the factors that may contribute to the evolution of social monogamy are selection for extended mate guarding of females and selection for territorial ‘cooperation’. Many socially monogamous taxa are also territorial, with ‘partners’ sharing a single territory, suggesting that one or both partners may benefit by sharing territorial maintenance. Snapping shrimp (genus Alpheus) are socially monogamous and territorial, living in excavated burrows or with host organisms, with females performing all parental care. The territorial cooperation hypothesis predicts that male and female partners share (1) territorial defence, resulting in a reduction in the risk of eviction from the burrow, (2) burrow construction duties, such that individuals in pairs spend less time in burrow construction relative to solitary individuals, and/or (3) foraging duties, by returning food to the burrow, where it is consumed by both partners. UsingA. angulatus as a model species, a territorial defence experiment revealed that females in pairs were significantly less likely than solitary females to be evicted by female intruders, but males in pairs were not significantly less likely than solitary males to be evicted by male intruders. A subsequent experiment revealed that paired males were significantly less likely to be evicted by an intruding male if paired with sexually receptive females than if paired with nonreceptive females. Another experiment revealed that (1) paired females spent significantly more time in burrow construction than paired males, and (2) both males and females consistently returned food items to the burrow, perhaps incidentally provisioning their mates. These data suggest that social monogamy may have been selected for in part because of the advantages of territorial cooperation, as both males and females are likely to benefit by dividing the labour of territorial defence and maintenance. These tests of the territorial cooperation hypothesis are synthesized with data from tests of the extended mate-guarding hypothesis to place snapping shrimp pairing behaviour into a larger construct incorporating both the influence of ecological pressures (territoriality) and mating interactions between the sexes. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Space use in a multi-male group of the group-living lizard

The aim of the study was to determine the space use of individuals in a multi-male group of the armadillo lizard Cordylus cataphractus . Specific focus was on the adult males and females of the group, because, ultimately, the manner in which adult males and females utilize space determines the mating system. A two-dimensional (2D) grid pattern was delineated at the crevice of a free-living, multi-male group to facilitate the recording of the positions of individuals during observations. All marked individuals ( n =55) that were visible were scanned through a telescope, and their respective positions were recorded at 30-min intervals over several days during the breeding season. Arcview 3.2 Geographical Information Software was used to create a computerized replicate of the 2D grid pattern at the crevice and to depict graphically the recorded positions for each individual using a minimum convex polygon procedure. From the polygons generated by this procedure, it was clear that the space use of adult males overlapped greatly with the space use of adult females, but that there was no overlap in space-use among adult males. Space use overlapped greatly among subadults and juveniles and also among these size classes and adult males and females. Adult male C. cataphractus are territorial, defending specific sites at the crevice. The territory of each male in the group incorporated at least one female. The space use of several females overlapped with the space use of two or more males. Males were observed to mate with one or more females, and one female was observed to mate with two different males on more than one occasion. Cordylus cataphractus males display typical territorial polygyny whereas females may be promiscuous.     

A one night stand? Reproductive excursions of female roe deer as a breeding dispersal tactic

Breeding dispersal, defined as the net movement between successive breeding sites, remains a poorly understood and seldom reported phenomenon in mammals, despite its importance for population dynamics and genetics. In large herbivores, females may be more mobile during the breeding season, undertaking short-term trips (excursions) outside their normal home range. If fertilisation occurs, leading to gene flow of the male genome, this behaviour could be considered a form of breeding dispersal from a genetic point of view. Here, we investigated ranging behaviour of 235 adult roe deer using intensive GPS monitoring in six populations across Europe within the EURODEER initiative. We show that excursions are common from June to August among females, with 41.8 % (vs. 18.1 % of males) making at least one excursion. Most individuals performed only one excursion per season and departure dates for females were concentrated in time, centred on the rutting period, suggesting a link with reproduction. The distance females travelled during excursions was significantly greater than the site-specific average diameter of a male home range, while travel speed decreased once they progressed beyond this diameter, indicating search behaviour or interaction with other male(s) outside the resident male’s territory. Because adults are normally highly sedentary, the potential for mating with relatives is substantial; hence, we conclude that rut excursions could be an alternative tactic enabling females to avoid mating with a closely related male. To understand better the ultimate drivers at play, it will be crucial to explore the genetic causes and consequences of this behaviour.     

Differences in size and energy content affect the territorial status and mating success of a neotropical dragonfly

In male odonates, both size and fat content are related to territory defence and mating success. Males that are larger and have higher energy reserves win relatively more disputes for territory and attract more females. Wing colour has also been regarded as a mechanism that influences agonistic behaviour between males, as wing pigmentation might be regarded as a sign of male quality. In this study, we analysed whether a set of male physical (body size and wing colour), physiological (body fat content) and behavioural (disputes between males) characteristics were involved in the territory defence and mating behaviour of the neotropical dragonfly Zenithoptera lanei Santos, 1941 (Anisoptera: Libellulidae). Males were characterised as territorial whenever they warded‐off other males and remained in the same place within the pond for two consecutive days. In general, these territorial males were larger and had more abdominal and thoracic fat, engaged in pursuits more frequently, spent more time on sexual behaviour and female guarding, and mated more in comparison to subordinate males. By evaluating whether the percentage of wing area covered by black ink influenced male behaviour, we found that territorial males tended to act aggressively towards other males whose wings were partially painted, and sexually towards females irrespective of wing area painted. In Z. lanei, both body size and fat content play a role in defining territoriality. By subduing competitors and dominating preferred locations within high‐quality sites, these males are likely to be visited by females and engage in mating.     

Metabolic cost of first reproduction in young female European roe deerCapreolus capreolus

In female roe deerCapreolus capreolus (Linnaeus, 1758), like in several deer species, first reproduction occurs before they have reached their full body size. This study quantifies, in young females, the energetic costs induced by the contemporaneously occurring events: growth and first reproduction. Resting metabolic rate and body mass were measured in young primiparous females from first mating to 4 months after parturition, and compared to values measured in fully-grown adult multiparous females. Throughout the 10-month period from mating to fawning, body mass increased in yearlings. Prior to the blastocyst implantation (month −5 before parturition) young females were lighter than adult females whereas after parturition they had the same body mass. Our results suggest that body growth was reinitiated during pregnancy in primiparous females. From mating to fawning, except in the first part of pregnancy, mass-specific metabolism was higher in primiparous females than in multiparous individuals indicating the occurrence of an additional cost due to growth in young females. The depressed level observed at the beginning of gestation could allow the resumption of growth at lower cost. Thus, the allocation of resources to both reproduction and growth was not detrimental to first reproduction in young female roe deer under experimental conditions withad lib feeding.     

Do male roe deer clump together during the rut?

Movement s an d habitat use of 7 male and 7 female roe deerCapreolus capreolus (Linnaeus, 1758) were studied by radioteleme try from March 1999 to February 2001. Annual and bimonthly home ranges of males were small (ca 10 ha, 95% kernel), with large overlap among individuals throughout the year. Exclusive core areas (ca 0.4 ha, 25% kernel) were concentrated in the forest, a limited and sought-after resource in the study area. The difference in overlap between male exclusive core areas and female home ranges in the pre-rut and rut periods suggested that females made excursions to search for territorial males during the rut Our results support the mating strategy hypothesis of territorial behaviour Different space use patterns occurred between the sexes, with females apparently playing an active role in mate choice by visiting males at clumps of core areas in the forest.     

Foraging strategies and seasonal diet optimization of muskoxen in West Greenland

Various aspects of optimal foraging and seasonal diet composition of bulls (bachelor and dominant), cows, subadults, and yearlings of muskoxen Ovibos moschatus were investigated in West Greenland during the following seasons: calving, post-calving, summer, rut and mid-winter. The following hypotheses were tested: (1) muskoxen maximize daily energy intake during spring and summer, (2) dominant bulls monopolizing cows during the rutting season shift from an energy maximizing to a time minimizing foraging strategy in order to maximize the time available for reproductive activities, and (3) muskoxen employ a time minimizing foraging strategy during winter to conserve energy. As forage quality changed throughout the short Arctic growing season, muskoxen responded by changing the proportions of daily time spent feeding on graminoids (Cyperaceae, Poaceae) and dicots (Salix, Betula), respectively. This seasonal variation in the relative proportion of daily feeding time spent ingesting graminoids followed approximately the energy maximization prediction over the periods calving to rut. Neither time minimizing nor random foraging could explain the observed diets in this period, thus confirming hypothesis 1. Dominant bulls did not shift to the time minimizing strategy as predicted by hypothesis 2. However, during the pre-rutting and rutting seasons bulls deviated from the other sex/age classes by failing to obtain the daily maximum energy predicted by the model, as a result of a higher proportion of time allocated to agonistic and sexual behaviour. During winter, none of the sex/age classes employed a time minimizing strategy, so rejecting hypothesis 3. Instead, muskoxen were found to maximize Na intake, indicating that Na is of major importance for winter survival. The results emerging from a linear programming model with constraint settings varying over seasons confirm that the constraint parameters applied are indeed important limiting factors for muskoxen in natural populations.