Could asynchrony in activity between the sexes cause intersexual social segregation in ruminants?

In many sexually dimorphic mammal species, the sexes live outside the mating season in separate social groups (''social segregation''). Social segregation occurs in a wide range of environmental conditions, but its cause in unknown. I suggest that social segregation is caused by a lower level of activity synchrony between individuals in mixed-sex groups than in single-sex groups, owing to sex differences in activity rhythm. As a consequence, mixed-sex groups are more likely to break up than single-sex groups, resulting in a predominance of single-sex groups at equilibrium. To test this hypothesis in red deer (Cervus elaphus L.), I developed an index of activity synchronization and showed that deer in mixed-sex groups were significantly less synchronized in their activity than deer in single-sex groups. Thus, low intersexual synchrony in activity can lead to social segregation. However, a lower level of intrasexual (female-female and male-male) activity synchrony within mixed-sex than within single-sex groups implies that additional factors (other than sex differences in foraging rhythm) contribute to the higher degree of instability of mixed-sex groups.     

Sexual segregation in Tibetan gazelle: a test of the activity budget hypothesis

Tibetan gazelle Procapra picticaudata , a threatened endemic species on the Qinghai-Tibet Plateau, is a sexually dimorphic ungulate. Males are larger than females and both sexes live in different groups, except during the rutting season. In order to test the activity budget hypothesis, which was proposed to explain social segregation in ungulates, we studied the activity budget of male and female Tibetan gazelles during the summers of 2005 and 2006. The activity budget hypothesis predicts that females spend more time feeding and group members synchronize their activities more in the same-sex groups than in the mixed-sex groups. We found that females and males of Tibetan gazelle had different activity budgets; females spent significantly longer time feeding and comparably less time on other activities. Activity synchronization indexes between female groups and male groups were similar (female groups: 0.81±0.17, male groups: 0.80±0.19), whereas both of them were significantly higher than that of mixed-sex groups (0.58±0.29). These results suggest that although sexual segregation might be caused by multiple mechanisms, the activity budget hypothesis about sexual segregation is supported by our studies of Tibetan gazelle on the Qinghai-Tibet Plateau.     

Multiple causes of sexual segregation in European red deer: enlightenments from varying breeding phenology at high and low latitude

Sexual segregation outside the mating season occurs in most species of sexually dimorphic ungulates and has been extensively described in the literature, but the mechanisms causing segregation are still debated. The detailed pattern of sexual segregation throughout the year has rarely been presented for mammals, and no study, to our knowledge, has used latitudinal-related variation in breeding phenology to shed light on the underlying mechanisms. Recent methodological developments have made it possible to quantify separate components of segregation (social, habitat) and activity synchrony in animal groups, but these major improvements have so far been little used. We observed European red deer year round at two widely different latitudes (France and Norway) and tested three different mechanistic hypotheses of segregation related to: (i) predation risk; (ii) body-size-related forage selection; and (iii) activity budget. Habitat segregation peaked during calving in both populations and dropped rapidly after calving. Females with calves were more segregated from males than were females without calves, pointing to a key role of antipredator behaviour even though large predators are absent in France and extremely rare in Norway. However, at both sites individuals also grouped with their own sex within habitat types (i.e. social segregation), and individuals in mixed-sex groups were less synchronized in activity type than individuals in either unisex male or unisex female groups, suggesting that differences in activity budgets are involved. Social segregation peaked during calving and was lowest during the rut (indicating aggregation) in both populations; these activities occurred one month later in the Northern populations, corresponding well with known differences in breeding phenology. We conclude that latitude-dependent breeding phenology shapes the seasonal pattern of sexual segregation and that sexual segregation in ungulates has multiple causes.     

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.     

Unexplained sexual segregation in polygamous ungulates: a defense of an ontogenetic approach

Two processes are generally mixed when considering ungulate sexual segregation: the social segregation which appears to be a rule in polygamous ungulate species and the spatial segregation which is facultative. Early in life, there is evidence that males and females exhibit different levels of activity and patterns of interaction which could lead to behavioural incompatibility. It is proposed that juvenile females, differently motivated than males to interact socially, may soon avoid the pseudo-sexual and agonistic male behavioural components. as body and behavioural dimorphism increase with age, the behavioural incompatibility would be intensified. Since adults often interact agonistically with or avoid them, juveniles will attend to interact with same-sex peers of similar status. Because social experiences are occurring since early stages of development, i.e., during sensitive periods, a social auto-segregation by sex/age classes is expected. Then population would be structured in coherent socio-spatial sets of compatible individuals, sharing similar constraints and properties. Around the birth period, females become asocial promoting a temporal social segregation. Costs linked to gestation and lactation may make females more dependent on water and rich food resources. Female with offspring are also likely to select areas where the risks of predation are reduced, sometimes at detriment to energy intake, while other animals exhibit opportunistic feeding behaviour, resulting in seasonally ecological segregation. Body growth channels a lot of energy for males which are expected to use the best feeding areas. However, indirect competition for the same resources may result in a spatial segregation between the sexes. Sex- and age-specific activity budgets are suspected to favour social and spatial segregations. The social segregation among males and females use exclusive ranges. Whether spatial segregation results in ecological segregation would however depend on habitat heterogeneity, population density, local ecological conditions and the biological cycle periods. It is suggested that spatial segregation will occur not only between the sexes but also among males. If populations are structured in coherent socio-spatial units, spatial segregation among individuals of age and sex classes may result from indirect interactions between sets of individuals (of same age and sex class), that may be translated as an indirect competition between individuals. Studies on ethogenesis are considered as fundamental as they allow to know on what basis social and spatial segregations are set up. In particular, studies on behavioural development could highlight how social segregation between adults may result from a process that originates much earlier in the life of animals. By considering the role of behaviour, we defend the idea that social and spatial segregations may be logically distinguished.     

Establishing baseline values of parameters potentially indicative of chronic stress in red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) from different habitats in western Germany

The use of habitats by wild animals is commonly assumed to be decreasing due to human activities, such as tourism or the installation and use of wind-energy plants. These anthropogenic interferences may subject animals to chronic stress. To be able to objectively characterise the effects on animal populations or on individual animals, the collection of data that might be suitable to monitor such chronic stress is required. In this study of hunted red deer, we report data that are related to adrenal activity and are not affected by the acute stress induced by hunting. Adrenal glands and samples from ileal digesta were collected from 75 hunted deer from seven different habitats in the German Rhineland. The adrenal glands were evaluated histomorphometrically; in the digesta, the concentration of cortisol metabolites, i.e. of 11,17-dioxoandrostanes (11,17-DOA), was measured. Digesta were also examined for parasites. Animals were grouped according to age, sex, habitat, and hunting method. Animals were infected with gastrointestinal helminths and lungworms; examination for liver flukes was negative. Significant differences were not established among the different groups for any of the recorded parameters. For sex, a tendency (P=0.11) towards higher DOA levels was observed in female deer when compared to male deer. The variability of the parameters together with the lack of identifiable influences of hunting indicates that chronic stress might indeed have been a relevant factor. However, none of the parameters analysed can presently be validly used to evaluate habitat quality for red deer since physiological stressors cannot be differentiated.     

Habitat selection and group formation pattern of fallow deer Dama dama in a submediterranean environment

We analyzed the patterns of association between sex and age classes of fallow deer Dama dama in addition to the modification of aggregation patterns as a function or the degree of cover in different habitats in a coastal submediterranean environment in Italy. Deer preferred large open grasslands, small clearings, woods and marshes. Habitat selection varied strongly according to sex, and there were also ecological differences among males of different age classes. Adult females were mostly in family groups, with subadults and fawns. These groups were very stable in time and space. On the contrary bachelor herds were unstable social associations, adapting their size to the local degree of cover. In open grasslands males and females sometimes formed mixed groups with group size similarly being adapted to degree of cover. The possibility that random events fully explained fallow deer aggregation patterns was excluded, while the existence of non-linear mechanisms of aggregation was shown. The persistence of antipredator strategies in this managed population may account for these findings.     

Sexual segregation in ungulates: individual behaviour and the missing link

Previous "explanations" of sexual segregation in ungulates establish no more than a prerequisite for habitat segregation because they do not include a model of competitive habitat selection. Here we provide one based on the ideal free distributions of mutually competing, optimally foraging, individual deer. We parameterised our model using field data collected from a population of fallow deer (Dama dama) in a Mediterranean forest. The predictions of the inter-sex competition model were in full agreement with observational data, but those of single sex distributions (conventional theory) were not. The "conventional" hypothesis, that segregation arises simply from sex differences, predicted no more than moderate (20–40%) levels of segregation, even in optimal conditions. By introducing inter-sex resource competition, the predicted segregation can generally more than double and full segregation becomes possible in some circumstances. The modelling showed segregation to be density-dependent, varying in complicated ways with season and animal density. Sensitivity analysis showed the vulnerability of the "conventional" understanding of environmental variation and uncertainty. Using our competition model we show that as diet difference increases, direct competition between the sexes declines, so that as males increasingly differ from females, segregation declines and the two sexes are more likely to be found mixed (as long as the chosen food is available to both in the same area). Conversely, small differences among male and female deer are amplified by both food depletion and inter-sex competition to give substantial levels of segregation. The theoretical framework on which our model is built strongly suggests that sexual dimorphism in the context of scramble competition may be the fundamental cause of sexual habitat-segregation among ungulates.     

Intersexual niche segregation among three bark-foraging birds of eucalypt forests

All three species of specialized bark-climbing birds in eucalypt forests of north-eastern New South Wales exhibited sexual differences in their foraging behaviour. The degree of these differences was weakest in the least social species and strongest in the most social species, suggesting that intersexual niche segregation is an adaptation to reduce intraspecific competition for food. In the least social White-throated Treecreeper, the slightly longer-billed males foraged on dead branches more than females only during the breeding season when pair members were forced to share the same area. In the group-living Red-browed Treecreeper and Varied Sittella, males foraged lower and used rough-barked trunks and limbs more than females throughout the year. Sexual differences in the foraging substrates of Sittellas may have been related to sexual dimorphism of the bill, but in the monomorphic Red-browed Tree-creeper, male dominance was probably important. Some evidence suggests that further subdivision of the niche may occur in the highly social Sittella along the vertical dimension according to age and/or reproductive status. This report argues that intersexual niche segregation is just as likely to develop under conditions of increased interspecific competition as under reduced competition.     

Patterns of antler consumption reveal osteophagia as a natural mineral resource in key periods for red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>)

Osteophagia is a common behaviour among ungulates. It has been considered as a mineral source for cervids, mainly related to phosphorus deficiency. In this study, we aimed to study the seasonality in antler consumption of Iberian red deer (Cervus elaphus) and to assess the differences among sex/age classes. In a game reserve located in South-Eastern Spain, antlers were offered to free-ranging animals, and their chewing behaviour was recorded with camera traps. Red deer was the species observed with greatest frequency (90.5%), and only red deer was observed chewing the offered antlers. Males showed greatest peak of consumption at the end of antler growth; females showed the greatest peak at the beginning of the lactation and calves after delivery of the newborn. All groups showed an important second peak of consumption in September. We conclude that osteophagia might have a role as a natural mineral supplement for red deer, but not for wild boar or foxes; and that each sex/age class uses this resource at different times according to differential needs in calcium and phosphorus, supporting the “seasonally increased requirements hypothesis”.     

Inter‐ and intrasexual genetic correlations of exaggerated traits and locomotor activity

Exaggerated traits in males can be costly and therefore can negatively affect fitness. Although these costs are thought to be male specific, traits that have a negative effect due to exaggeration are often shared between the sexes as life‐history traits. When there are genetic intersexual correlations for these shared characters, the evolution of the exaggerated traits can impose these costs on nonadorned females through the intersexual correlation. Thus, the exaggerated traits can constrain optimum development of female characters, even if the females lack these exaggerations completely. However, investigation of this pattern has been largely ignored, and thus, it is necessary to investigate genetic architectures of these traits within and across the sexes. Male flour beetles, Gnatocerus cornutus, have enlarged mandibles that are used in male–male competition, but females lack this character completely. Using a traditional full‐sib/half‐sib breeding design, we detected a negative intrasexual genetic correlation between male weapon size and locomotor activity, but not an intersexual genetic correlation for locomotor activity. After subjecting this weapon to 17 generations of bidirectional selection, we found a correlated response to locomotor activity in the male, whereas there was no correlated response in the female. Our results suggest that the costs of exaggerated traits to locomotion are not imposed on females and would be male specific. This is partly explained by genetic decoupling of locomotor activities across the sexes.     

Do Red Deer Stags (Cervus elaphus) Use Roar Fundamental Frequency (F0) to Assess Rivals?

It is well established that in humans, male voices are disproportionately lower pitched than female voices, and recent studies suggest that this dimorphism in fundamental frequency (F0) results from both intrasexual (male competition) and intersexual (female mate choice) selection for lower pitched voices in men. However, comparative investigations indicate that sexual dimorphism in F0 is not universal in terrestrial mammals. In the highly polygynous and sexually dimorphic Scottish red deer Cervus elaphus scoticus, more successful males give sexually-selected calls (roars) with higher minimum F0s, suggesting that high, rather than low F0s advertise quality in this subspecies. While playback experiments demonstrated that oestrous females prefer higher pitched roars, the potential role of roar F0 in male competition remains untested. Here we examined the response of rutting red deer stags to playbacks of re-synthesized male roars with different median F0s. Our results show that stags’ responses (latencies and durations of attention, vocal and approach responses) were not affected by the F0 of the roar. This suggests that intrasexual selection is unlikely to strongly influence the evolution of roar F0 in Scottish red deer stags, and illustrates how the F0 of terrestrial mammal vocal sexual signals may be subject to different selection pressures across species. Further investigations on species characterized by different F0 profiles are needed to provide a comparative background for evolutionary interpretations of sex differences in mammalian vocalizations.     

Boys will be boys: social affinity among males drives social segregation in western grey kangaroos

In many sexually dimorphic and polygynous species, individuals exhibit social segregation by grouping with others of their own sex. Several proximate mechanisms have been proposed to explain social segregation: female avoidance of males, male avoidance of females, avoidance of harassment, male social affinity, female social affinity and activity budget asynchrony. Some of these have been tested in ungulate species, but few previous studies have been able to rigorously test, or distinguish between, these mechanisms because they have failed to examine the fission/fusion dynamics of groups. We tested these proximate mechanisms simultaneously in western grey kangaroos Macropus fuliginosus , by examining whether females, small males or large males instigated segregation by leaving mixed-sex groups or joining individuals of their own sex or size class. Females, small males and large males left mixed-sex groups as if leaving were independent of sex–size class. In contrast, large males joined male-only groups more frequently than expected. These results suggest that the need for males to maintain contact with other males can contribute to the cohesion of male-only groups and promote segregation. As male–male competition occurs in many polygynous species that sexually segregate, a comparable mechanism might be operating in other taxa, and should be examined further.     

Sexual segregation of seasonal foraging habitats in a non-migratory marine mammal

Many animal species segregate by sex. Such segregation may be social in nature, or ecological, or both. Grey seals (Halichoerus grypus), like many large mammals, are sexually size dimorphic. In size dimorphic species, allometric differences in morphology, metabolic rate and reproductive costs are likely. Such differences may require the sexes to use different foraging strategies or different habitats. To investigate sexual segregation of habitat in grey seals, we used satellite tracks from 95 (male 46; female 49) adults breeding at Sable Island, Nova Scotia (44 degrees N, 60 degrees W) collected from 1995 to 2005. Location estimates were made from satellite fixes using a state-space movement model to estimate true locations and regularize them in time. Location estimates were used to calculate home range kernels of male and female habitat use each month. Month by sex kernel home ranges revealed striking differences and dynamics in habitat use between males and females on spatial scales broader than most terrestrial examples and at temporal and spatial resolutions rarely available for marine species. Differences were most pronounced just before (October-December) and immediately after breeding (February-March). During both periods, males primarily used areas along the continental shelf break, while females mainly used mid-shelf regions. Coupled with previously identified sex-specific seasonal patterns of energy storage, diving and diet, our findings suggest that males and females differ profoundly in their spatial foraging strategies. These differences may serve to maximize fitness by reducing intersexual competition during key foraging periods.     

Non‐sex‐biased Dominance in a Sexually Monomorphic Electric Fish: Fight Structure and Submissive Electric Signalling

Males and females commonly compete for limited resources. When interaction costs are similar for both sexes and there are no sexual differences in resource value estimation, a non‐sex‐biased dominance is expected. Moreover, only non‐sex‐biased assessment of contenders fighting ability (Resource Holding Potential, RHP) should influence contest decisions. To test these predictions, we evaluated non‐breeding agonistic intra‐ and intersexual dyadic interactions in the weakly electric fish, Gymnotus omarorum. During the non‐breeding season, resource value is not expected to depend on individuals’ reproductive status and should thus be equal for males and females. In addition, as G. omarorum presents no sexual differences in body size, interaction costs can be considered symmetric between sexes. We confirmed that body size differences, but not individuals’ gender, is the best predictor of dominance. We correlated RHP asymmetries with contest duration and evidenced that body size but not sex influences assessment in intrasexual and intersexual encounters. All dyads tested engaged in agonistic interactions (N = 33) in which a clear dominant emerged. The analysis of conflict phases evidenced the submissive role of electric displays. Electric organ discharge (EOD) interruptions appear early in the contest as an electric hiding attempt, whereas chirps are post‐resolution signals of subordinate status. Interestingly, the decision of interrupting the EOD was also influenced by RHP asymmetries, whereas chirping activity was influenced by the intensity of the attacks received. Our results confirm that body size is the best RHP proxy in non‐breeding intra‐ and intersexual contests of this monomorphic species and demonstrated a sequential pattern of submissive signalling by means of two different electric displays.     

The way in which testosterone controls the social and sexual behavior of the red deer stag (Cervus elaphus)

For 10 months of the year red deer stags live together in social groups. These groups then break up, and the animals exhibit a complex pattern of sexual behavior during a short rutting season in September and October. We have utilized this clear separation between social and sexual behavior to try and establish experimentally in free-ranging wild animals the way in which testosterone influences the two types of activity.     

Selectivity of wolf predation on red deer in the Bieszczady Mountains, Poland

A pattern of wolfCanis lupus Linnaeus, 1758 predation on red deerCervus elaphus Linnaeus, 1758 was studied in Bieszczady Mountains in 1991–2002. In total 324 remains of red deer > 4 months old, killed by wolves throughout the year, were found. The sex, age and bone marrow fat content of wolf kills were compared with the same characteristics within the free living red deer population. The overall contribution of calves killed by wolves (24%) in October-May was higher than in the population (17%), and decreased from autumn to spring. Adult males were more vulnerable to wolf predation than adult females: stags constituted 62% and hinds 38% of adult red deer killed by wolves, whereas in the population, the percentages were 37 and 63%, respectively. Stags killed by wolves were younger ([`(x)] = 4.1\bar x = 4.1 years old) than hinds ([`(x)] = 8.9\bar x = 8.9 years old). Wolves killed more > 8 years old hinds and < 5 years old stags than available in the population. In wolf kills, the average fat content in femur marrow was higher among hinds (84.9%) than stags (69.3%) and calves (66.1%). Only 8% of hinds had < 70% femur marrow fat content, whereas 40% of calves and 38% of stags had marrow fat values below that level. Marrow fat content showed seasonal variation and was the lowest in March among all sex-age classes. The monthly share of stags in all kills, and hinds in hind-calf part of the sample was negatively correlated with their monthly average bone marrow fat content, and monthly share of calves was positively correlated with monthly average bone marrow fat content of adults. The segregation of social units (hind-calf and stag groups), except during the rutting season, and the low fat reserves of males from midwinter until spring contribute to the high overall incidence of calves and adult males and the relatively low incidence of adult females among wolf kills.     

A New Gene Mapping Resource: Interspecies Hybrids between Pere David''s Deer (Elaphurus Davidianus) and Red Deer (Cervus Elaphus)

Three male F(1) hybrids between Pere David's deer and red deer were mated to red deer to produce 143 backcross calves. The pedigrees are a rare example of a fertile between evolutionarily divergent species. We examined the use of these families for genetic mapping of evolutionarily conserved (Type I) loci by testing for genetic linkage between five species-specific protein variants and 12 conserved DNA probes. Two probes were homologous, and the remainder syntenic, to the protein coding loci in cattle or humans. Using six restriction enzymes, each DNA probe detected one or more restriction fragments specific to Pere David's deer. Linkage analyses among the species-specific variants placed the loci into four linkage groups within which linkage between adjacent loci and gene order was supported by a LOD > 3. The linkage groups were (HPX, HBB)-FSHB- ACP2, LDHA-CD5-IGF2, BMP3- (GC, ALB)-(KIT, PDGFRA) and LDLR-C3-FGF1. Southern and protein analysis of LDHA and ALB provided identical segregation data. These linkage groups were consistent with the cattle gene map and provide new information for comparing the gene maps of ruminants, humans and mice. The deer hybrids are an important new resource that can contribute to the comparative analysis of the mammalian genome.     

Spatial and temporal components of the mating systems of kudu bulls and red deer stags

Observations were made on greater kudu (Tragelaphus strepsiceros) in the Kruger National Park in South Africa. Data on movement patterns, courtship, dominance interactions and proportion of time spent in various activities by males during the breeding season are presented. Comparative data on the apportionment of time by red deer (Cervus elaphus) setages were obtained on the Isle of Rhum, Scotland. While spatial components of male reproductive behaviour were similar in the two species, time-investments differed markedly. Kudu bulls showed little change in time spent feeding and moving during the breeding season, did not herd females and tolerated the presence of younger males. In contrast, red deer stags almost ceased feeding and greatly increased their time spent moving and in sexual or dominance-related activities. Thus the spatial and temporal components of male mating strategies can vary independently and require separate evolutionary explanaions. Comparative data for other ungulates are sparse, but suggest that time-investments are influenced by the duration of the breeding season and the breeding sex ratio. However, kudu bulls incur high mortality costs despite their low time-investments in reproductive activities.     

麋鹿幼仔的活动同步性与同性聚群倾向

哺乳动物幼体从出生到性成熟这段时间存在生理和行为上的巨大变化.麋鹿幼仔出生1周内,与成鹿和其它仔鹿呈隔离状态,且藏卧于隐蔽处,母鹿哺乳是引起幼仔活动的主要因素.