Trade-offs of predation and foraging explain sexual segregation in African buffalo

1. Many studies have investigated why males and females segregate spatially in sexually dimorphic species. These studies have focused primarily on temperate zone ungulates in areas lacking intact predator communities, and few have directly assessed predation rates in different social environments. 2. Data on the movement, social affiliation, mortality and foraging of radio-collared African buffalo (Syncerus caffer) were collected from 2001-06 in the Kruger National Park, South Africa. 3. The vast majority of mortality events were due to lion (Panthera leo) predation, and the mortality hazard associated with being an adult male buffalo in a male-only 'bachelor' group was almost four times higher than for adult females in mixed herds. The mortality rates of adult males and females within mixed herds were not statistically different. Mortality sites of male and female buffalo were in areas of low visibility similar to those used by bachelor groups, while mixed herds tended to use more open habitats. 4. Males in bachelor groups ate similar or higher quality food (as indexed by percentage faecal nitrogen), and moved almost a third less distance per day compared with mixed herds. As a result, males in bachelor groups gained more body condition than did males in breeding herds. 5. Recent comparative analyses suggest the activity-budget hypothesis as a common underlying cause of social segregation. However, our intensive study, in an area with an intact predator community showed that male and female buffalo segregated by habitat and supported the predation-risk hypothesis. Male African buffalo appear to trade increased predation risk for additional energy gains in bachelor groups, which presumably leads to increased reproductive success.     

Extra-pair paternity in relation to male age in Bullock's orioles

Single-locus minisatellite DNA profiling was used to assign paternity in a population of Bullock's orioles, Icterus galbula bullockii, and to determine the contribution of age to a male's success in obtaining extra-pair paternity. There was a very low rate of intraspecific brood parasitism (2/202 = 1.0% of chicks). Older adult males lost less within-pair paternity and gained more extra-pair fertilizations than did yearling subadult males. This resulted in adult males benefiting from an annual reproductive success more than double that of subadult males. Behavioural observations, used to determine the role of female choice in extra-pair copulations (EPCs), indicated that females actively participate in EPCs and that they prefer to obtain them from older males. While it was possible that females obtained EPCs as an insurance against the possible infertility of their social mate, the results of this study fit best with the hypothesis that females were attempting to obtain better-quality genes for their offspring by obtaining EPCs with older, better-quality males.     

Social segregation in male,but not female yearling rhesus macaques (Macaca mulatta)

Males and females of many species sex‐segregate, ranging from complete separation of habitats to social segregation within the same space, sometimes varying across seasons and lifespan development. Mechanisms for such segregation are not well understood, though some have suggested that sex differences in preferred juvenile behaviors lead to greater behavioral compatibility within than between sexes. This within‐sex behavioral compatibility may be the source of sex‐segregation. As juvenile behavioral sex differences are well‐documented in rhesus monkeys, we examined sex‐segregation patterns of yearling rhesus monkeys engaged in three different types of behavior: rough play, parallel play, and grooming. We observed male and female rhesus yearlings from five stable long‐term age‐graded social groups of 67–183 animals. Behavioral observations were designed to collect equal numbers of rough play, grooming, and parallel play bouts. In addition, sex composition and proximity to adults was recorded for each bout. Across all behaviors, more all‐male groups and fewer mixed sex‐groups were observed than expected by chance. All‐female groups occurred at the level expected by chance. Thus, males sex‐segregated regardless of type of behavior, while females did not sex‐segregate. Female groups were observed in proximity to adults more often than expected by chance. These results suggest that behavioral compatibility may produce sex‐segregation in male yearling rhesus monkeys, possibly preparing males and females for different social roles and segregation as adults. Am. J. Primatol. 72:87–92, 2010. © 2009 Wiley‐Liss, Inc.     

Natural and sexual selection on a plumage signal of status and on morphology in house sparrows,Passer domesticus

Temporal patterns of natural and sexual selection on male badge size and body traits were studied in a population of house sparrows, Passer domesticus. Badge size was a heritable trait as revealed by a significant father-son regression. Survival during autumn dispersal and winter was not related to badge size or body traits in yearling male house sparrows. Badges that signal dominance status were affected positively by directional selection for mating. Adult male house sparrows suffered an opposing selection pressure on badge size during autumn. Contrary to males, female house sparrows did not experience significant directional or stabilizing selection on any body trait. Directional sexual selection on male badge size due to female choice moves male sparrows away from their survival optimum. Opposing directional natural selection on badge size due to autumn mortality caused by predation maintains a stable badge size.     

Male aggressiveness in a polygynous ungulate varies with social and ecological context

Aggression is a social behaviour which can be affected by numerous factors. The quality and quantity of food resources may play an important role in the aggressiveness of territorial ungulates as the defence of these resources influences female choice and mating opportunities. However, the relationship between food resources and aggression remains poorly understood. We assessed the ecological and social factors that influence aggression in Lama guanicoe, a territorial ungulate exhibiting resource‐defence polygyny, during three periods (group‐formation, mating and post‐mating) in the reproductive seasons of 2014 and 2016. We recorded 460 focal observations of territorial (family groups, solitary) and non‐territorial (mixed and bachelor groups) males. We performed analyses at the population level (including all focal observations) and at the group level (each social unit separately), to test whether the factors that influence aggression differ at these different scales. We also identified proxies of vegetation quality as potential predictors of aggression. At the population level, we found that the presence of aggressive behaviour peaked during the mating season and that post‐mating aggression may have been driven by inter‐annual environmental variations. For family groups and solitary males, variables reflecting high vegetation quality/quantity were predictors of aggressive behaviour, reflecting the resource‐defence strategy of this species. Conversely, for mixed‐group males, aggression may be more associated with social instability and group size, although this hypothesis has yet to be tested. Our research reinforces the idea that aggression can occur in multiple contexts depending on male status (e.g. territorial or non‐territorial) and contributes to our understanding of how ecological (i.e. availability of food resources) and social factors influence aggression in a territorial ungulate.     

Male-male competition and mating success in the orb-web spider,Nephila clavata,with reference to temporal factors

Seasonal occurrence patterns of adults of both sexes, intensity of male-male interactions, and mating success in the spider,Nephila clavata, were examined in the field. Adult males began to attend female webs about 2 weeks before female maturation. Large adult males were abundant in the early breeding season, but small males increased later in the season. From the distribution of males among female webs and size relationship of males within a web, male-male interactions seemed to be more intense when most females were still subadult. This was verified by a field experiment in which males were artificially introduced to female webs that were attended by other males. It was found that the probability of introduced males remaining on subadult female webs was lower than that on adult webs. As mating occurred mostly in the period shortly after the female final molt and first male sperm precedence was known in all spiders reported so far, intense male-male competition on subadult female webs seemed to be reasonable. Male longevity had an important influence on the mating success of males with just-molted females. Mating success was also affected by the relative body size of males present in a given period. Since larger males occupied the position closest to females within a web and stayed there longer, relative body size appeared to influence mating success through male-male competition. Female body size at maturation declined with time; hence, males that attained sexual maturity earlier had the advantage of mating with larger and more fecund females. Therefore, early maturation as well as larger size seem to be two important trairs influencing the reproductive success of males.     

Hunting differentially affects mixed-sex and bachelor-herds in a gregarious ungulate, the impala (Aepyceros melampus: Bovidae)

We investigated herd-sizes and herd-compositions of Impala ( Aepyceros melampus ) inside a protected area [Lake Mburo National Park (LMNP) in western Uganda] and the unprotected adjacent ranchland [the Ankole Ranching Scheme (ARS)]. Impala experience intense hunting and poaching in the study area, and poaching is especially strong on the ARS. We found evidence for changes in overall group-sizes in both mixed-sex and pure bachelor herds between areas in and outside LMNP. Mixed-sex herds strongly decreased in size outside the National Park, but bachelor herds even slightly increased in size. While the group-composition of mixed-sex herds was very similar in areas in and outside LMNP, bachelor herds comprised more yearlings and subadult males on the ARS. Our study suggests that effects of hunting and other human nuisance may differ between herd types: mixed herds probably decrease in size because females are more strongly hunted. Around LMNP, impala are usually hunted using nets and spears, thereby increasing the hunters' chance of being injured. Poachers therefore prefer hornless females (and their calves), as it is less dangerous to handle net-caught females than males. As a result, males are less hunted, but increased vigilance and, therefore, reduced aggression among the members of a bachelor herd, may account for the observed increase in herd sizes and changes in group-compositions.     

Female transfer between one-male groups of proboscis monkey (<Emphasis Type="Italic">Nasalis larvatus</Emphasis>)

Successful or unsuccessful female transfers were observed seven times during a 32-month field study of proboscis monkeys (Nasalis larvatus) inhabiting a riverine forest along a tributary of the Kinabatangan River, Sabah, Malaysia. In all cases, the females voluntarily left their own groups and immediately joined with another one. When adult females tried to shift to other groups, adult males called them back to their own groups, but appeared to be indifferent to subadult females. When the adult females returned, the males never attacked the females physically, but instead often emitted herding sounds to them. One subadult female was repelled by a resident adult female. When one adult female transferred into a new one-male group, she left her behind son in an all-male group. The number of females often fluctuated in most study groups, with this fluctuation being more prominent among subadult females than adult females. It is likely that female transfer in proboscis monkeys is not a rare occurrence and that it is especially common among sub-adult females.     

Alternative strategies in the acquisition of home ranges by male pine martens in a high-density population

Two strategies of home range acquisition by male subadult pine martens (Martes martes) were described from a high-density population inhabiting Bia?owie?a National Park. Four mother?Coffspring pairs were identified by genetic parentage assignments. Four subadult males showed two different strategies of home range acquisition: dispersal and sedentary. The dispersing males used an area 4?C10 times larger than in sedentary subadult males. A sedentary subadult male used his natal area with his mother, and in the following mating season, this male left this area and established a home range that overlapped greatly with another unrelated female near the natal range. A similar high overlap between another subadult male and an unrelated adult female persisted for 3?years. After the death of this female, the male extended his range to overlap slightly with two to four other females. The sedentary strategy adopted by some subadult males may explain the great variation in spacing patterns of solitary mammals.     

Demographic and reproductive patterns in moustached tamarin monkeys (Saguinus mystax): Implications for reconstructing platyrrhine mating systems

In this paper we address a series of questions concerning reproductive opportunities, kinship, dispersal, and mating patterns in free-ranging moustached tamarin monkeys (Saguinus mystax). Between 1980 and 1990 information on group size, composition, and migration patterns was collected on marked groups of moustached tamarins inhabiting Padre Isla, an island in the Amazon Basin of northeastern Peru. In 1990, 86% of 114 animals residing in 16 social groups were trapped, examined, and released. Mean group size was 7.0, including 2.2 adult males and 2.0 adult females. None of these groups was characterized by a single adult male-female pair. In groups with more than one adult female, only the oldest female produced offspring. An examination of dispersal patterns indicates that transfers between groups were common and fell into several categories, including immigration of individual males and females, simultaneous transfer of pairs of subadult and/or adult males (sometimes relatives) into the same social groups, and group fissioning in which males and females of the splinter group join another small social group. We have no unambiguous cases of 2 adult/subadult females migrating together into the same social group. All 6 groups for which reproductive data were available were characterized by either a polyandrous or polygynous (polygyandrous) mating pattern. The results of this study indicate that moustached tamarins reside in small multimale multifemale groups that are likely to contain both related and unrelated adult group members. Kinship and social ties among males appear to be stronger and more longlasting than kinship and social ties among females. We contend that the modal mating system of moustached and many other tamarins is not monogamous, and offer the possibility that cooperative infant care and mating system flexibility in callitrichines evolved from a polygynous mating pattern. © 1993 Wiley-Liss, Inc.     

Social group dynamics and composition of managed wild vicuñas (<Emphasis Type="Italic">Vicugna vicugna vicugna</Emphasis>) in Jujuy,Argentina

The vicuña is a high Andean wild camelid that lives year-long in groups. We analyzed the social organization of 98 marked vicuñas after capture and live shearing, focusing on group composition, spatial patterns, territoriality and habitat use. The social units analyzed were families, bachelor groups and solo animals. Location fixes of 54 males and 36 females were registered over a 2-year period, and home range was estimated. All females belonged to family groups, and 8–12 month yearlings changed their home ranges prior to first reproduction at 2 years. Female yearlings dispersed to a new family group, while male yearlings dispersed to a bachelor group. Solitary males appeared during the interval between leaving bachelor or family groups, and vice versa. Our study showed that vicuña polygyny was based on female grazing area requirements. Females that stayed with the same males were defined as loyal, and had a mean 18.9 ha home range. Females that changed both to a different family and male; and grazed over the same home range size, were considered disloyal. Family groups selected the habitat that had a low stratum and the highest coverage of palatable plants, while bachelors used their habitat randomly.     

Choice of Female Groups by Male Mosquitofish (Gambusia holbrooki)

Even though females are usually more selective in choosing their mates, males are also capable of exercising mate choice. Despite the large body of evidence on the individual features preferred in sexual selection, little attention has been devoted to the first stage of male–female interaction. As poeciliid fish are known to be social, in the wild, initially mate choice may concern a preliminary selection among shoals. Only after this primary choice, males may subsequently direct their attention to a specific mate. We observed spontaneous preference of male mosquitofish (Gambusia holbrooki) when choosing between groups differing in size and sex ratio. In partial agreement with our predictions, males preferred to join a group of females rather than an isolated one (expt 1) and the larger group when two female groups were presented (expt 2). An all‐female group was preferred to a mixed‐sex group (expt 3), whereas no preference was observed when the two mixed‐sex groups differed in the number of males (expt 4) or in the size of the males (expt 5). These results suggest that male mosquitofish are capable of discriminating among different quantities of individuals within a group and use such information to select among groups in order to optimize the likelihood of successful matings.     

A preliminary behavioral comparison of two captive all-male gorilla groups

Gorillas live in polygamous harem groups, generally composed of one male, several adult females, and their offspring. With an equal numbers of male and female gorillas born in captivity, however, housing gorillas in social breeding units inevitably means that some males will not have access to female social partners. Thus, the future of the captive gorilla population depends on the collective ability of zoos to house equal numbers of males and females. This study examined the behavioral profiles of two all-male groups of captive lowland gorillas (Gorilla gorilla gorilla) to provide information on this type of housing situation. One group consisted of three subadult individuals, while the other consisted of two subadults and a silverback. Data were collected during two 6-month intervals, for a total of 284 hr. The behavioral profiles of the animals were stable over the course of the study but proximity patterns changed. Differences in feeding, solitary play, and object-directed behavior were found between groups, while no significant differences were observed in affiliative or agonistic social behavior. At both institutions, group cohesion appeared to be high, particularly between subadults; these individuals spent approximately 10% of their time engaging in social behavior and 25-50% of their time in close proximity (within 5 m). However, the Zoo Atlanta males spent significantly more time within 1 m and 5 m of each other than the Santa Barbara males, which may reflect a higher level of cohesiveness among members of the Zoo Atlanta group. The behavioral profiles of the animals in this study were similar to those found in bachelor groups of wild mountain gorillas. One notable exception was the absence of homosexual behavior between the silverback and subadults in Santa Barbara and the low frequency of this behavior between subadults in both groups. Although more longitudinal data are needed, these data suggest that all-male groups can be a feasible housing strategy for males at certain periods of their life span. Zoo Biol 20:27-40, 2001. Copyright 2001 Wiley-Liss, Inc.     

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.     

Adoption of a Wild orphaned ringtailed lemur infant by natal group members: Adaptive explanations

In December 1992 an infant ringtailed lemur, approximately 7 weeks of age, was orphaned in one of the regularly-censused social groups at the Beza-Mahafaly Reserve, southwestern Madagascar. The infant was initially adopted by a subadult (2 yr-old) male from the group. Continuous-time focal animal data were collected for a 12-hr period, from the time that the infant was retrieved by the young male, in order to document the adoption process. Ten members of the infant's social group (total group number=18) engaged in infant care behaviors over the 12-hr period. The subadult male spent the most time engaged in infant care, and he and one adult female exhibited the highest frequency of caregiving behaviors over the 12-hr period (p<0.001). Four adult males also initially cared for the infant. The orphan was one of only six infants in the reserve population to survive that year. She was censused two years later as an adolescent member of her natal group. Adaptive explanations for this adoption vary depending upon the care-giver. For the subadult male and adult female caregivers, kin selection can be suggested, as the infant was related to all females and immature animals in the group. Adult males may have exhibited caregiving behaviors as a strategy related to affiliation with adult females which could lead to potential mating and reproductive success.     

王朗国家级自然保护区岩羊集群结构及季节变化

岩羊(Pseudois nayaur)是青藏高原常见的食草动物,但对边缘分布区的种群了解较少.2018年6月到2019年8月,我们利用红外相机在四川王朗国家级自然保护区对岩羊的集群结构、特征及其季节变化进行了描述与分析.结果显示:调查共记录到岩羊1 921群次,共计6 623只次.按照性别和年龄组,可将岩羊集群划分为混...     

The influence of increasing density on body weight, kid production, home range and winter grouping in roe deer (Capreolus capreolus)

In this paper, we present an analysis of the consequences of increasing density, over a period of nine years (from 1980 to 1988), on the dynamics, and the social and spatial organization of a forest roe deer population. Hunting of this population ceased in 1979, after which time there was a significant increase in population density, with three distinct periods easily identified: 1980–1983 (PI), immediately following cessation of hunting, characterized by a relatively low density (d = 5–7 animals/100 ha), 1984–1985 (P2), a period of rapid population growth, and 1986–88 (P3), a period of high density (d = 25 animals/100 ha). During PI, the population was irregularly distributed across the study site but, as density increased, distribution became more uniform, and eventually covered the whole of the available area. Home-range structure and shape remained unchanged from PI to P3 but, by the end of the study, average range size was 30% lower for adult males only, and the period prior to subadult males, but not females, establishing a permanent home range had increased from c. 18 month to c. 30 months. Winter group size increased overall from PI to P3 with, for the first time, observations of groups of five or more animals and a reduction in the frequency of observations of solitary females, with does more commonly observed in pairs or small groups; the proportion of solitary males, however, did not change between the two periods. The mean number of kids per female declined significantly from PI to P3 and body weights recorded for juveniles of both sexes and for adult males were also significantly lower during the period of relatively high density (P3). However, for adult females, absolutely no body weight change was observed. This divergence between the sexes of response to increasing density is discussed.     

Male Competition,Mating Success and Female Choice in a Seasonally Breeding Primate (Erythrocebus patas)

Mammals with restricted breeding seasons often show brief but intense bouts of male competition for mates and male reproductive success has been attributed to male competitive abilities, with the most aggressively successful males able to control access to fertile females, or with females choosing to mate with such males. We studied male competition, mating behaviour and female mate selection in patas monkeys, a primate with a restricted breeding period. We observed two habituated patas groups in Laikipia District, Kenya, during Jun.-Aug. 1983. During the study, one group had a single resident male while the other group had multiple adult males. Within the multimale group, experienced adult males were no more successful than the subadult male. The sole resident male had a significantly higher rate of fights won (p < 0.02) although he did not differ from the multimale group males in rate of aggression or initiation of fights. We found no significant differences in either mating success or female preference based on males' experience or residency. The rates at which males copulated with and were solicited by females were not significantly correlated. We found no evidence of stable dominance ranks among males in the multimale group and aggressive success was not significantly correlated with copulation rate for males in the multimale group. Subadult males were responsible for the majority of copulations observed during the final third of the breeding season. Our observations of this patas population showed a fluid number of males in groups, with the same groups able to shift rapidly from single to multimale structure. This fluidity may result from the large fluctuations in numbers of breeding-age males and females observed over 4 yr of studying this population.     

Constant Proportionality in the Female Segment of a Roosevelt Elk Population

Abstract: Incomplete population counts indicate change in population sizes when constant proportionality holds, a condition that is rarely met. However, researchers have not explored whether constant proportionality holds for a segment of a population. I examined whether the female segment (juv, subadult M, subadult and ad F) of a Roosevelt elk (Cervus elaphus roosevelti) population displayed constant proportionality. When most food is in particular habitats, females of polygynous species should use that habitat frequently, even when food is limited, because they are more familiar with food distribution and abundance than males. I obtained counts of elk and tallies of naturally marked animals from vehicle surveys of a population inhabiting a landscape where forage was in meadows that were interspersed in closed-canopied forest. I conducted population surveys in January or February and estimated population size with Bowden's mark-resight estimator. Population size estimates declined from 130 in 1997 to 37 in 2006. The proportion of the population counted during surveys was inversely related to population size estimates. Estimated population sizes were inversely related to male (r² = 0.56) but not female sighting probabilities (r² = 0.004), which were ≥0.9. Constant proportionality in counts held for only the female segment of the population. Counts of the female segment of the population can inform managers about changes in this segment of the population over time.     

Sex-specific plasticity of growth and maturation size in a spider: implications for sexual size dimorphism

Sex-specific plasticity in body size has been recently proposed to cause intraspecific patterns of variation in sexual size dimorphism (SSD). We reared juvenile male and female Mediterranean tarantulas (Lycosa tarantula) under two feeding regimes and monitored their growth until maturation. Selection gradients calculated across studies show how maturation size is under net stabilizing selection in females and under directional selection in males. This pattern was used to predict that body size should be more canalized in females than in males. As expected, feeding affected male but not female maturation size. The sex-specific response of maturation size was related to a dramatic divergence between subadult male and female growth pathways. These results demonstrate the existence of sex-specific canalization and resource allocation to maturation size in this species, which causes variation in SSD depending on developmental conditions consistent with the differential-plasticity hypothesis explaining Rensch's Rule.