A note on group sizes of oribi (Ourebia ourebi, Zimmermann, 1783) from two contrasting sites in Zambia, with and without predation

The oribi (Ourebia ourebi, Zimmermann, 1783) is a small species of antelope widely represented across open grasslands of sub‐Saharan Africa. Although largely territorial, differences in group size and mating systems (monogamy/polygamy), have been linked to habitat conditions and population density. At high population densities, additional males are recruited to assist in territorial defence. Other factors which might impact on group size, include predation threat, especially at low population densities. In this study (1995–98), we recorded group sizes of oribi across two contrasting areas (c. 300 km² each) of Zambia with (Kafue), and without (Bangweulu), large vertebrate predators. A total of 412 groups was recorded across the two sites, with mean sizes of 2.31 (n = 217) at Bangweulu and 2.33 (n = 195) at Kafue. The modal group size was two throughout (range 1–6). Groups were more variable in size and sex composition at Kafue than at Bangweulu, but there was no significant difference in estimates of population density between sites. Mean estimates were 2.02 and 1.90 km^?2 for Bangweulu and Kafue, respectively. Sex ratios (males : females) were biased towards females (1 : 1.72 and 1 : 3.55, respectively) at both sites. Adult males were particularly scarce at Kafue. However, values of density and group size, were both unexceptional for the species. Whilst a modal group size of two is consistent with a monogamous mating system, variations in group composition and size were more difficult to explain. Adult females appeared to retain or recruit additional animals to groups, irrespective of the needs of territorial defence. This suggests that vigilance is an important function of groups on large open plains, especially in the presence of predators. By extension, reversed sexual dimorphism in oribi could reflect increased selection for vigilance duties in females. Predators had no apparent effect on population density, but oribi groups were more variable in size and sex composition in their presence. Adult males may be particularly vulnerable to predation when defending territorial borders at low population densities. However, further work is needed to define group dynamics in this species and to identify causes of mortality amongst adult males.     

Evolution of ungulate mating systems: Integrating social and environmental factors

Ungulates exhibit diverse mating systems that range from monogamous pair territories to highly polygynous leks. We review mating systems and behaviors across ungulates and offer a new approach synthesizing how interacting factors may shape those mating systems. Variability exists in mating systems among and within species of ungulates and likely is affected by predation risk, availability of resources (food and mates), habitat structure, and sociality. Ungulate mating systems may be labile as a consequence of the varying strength of those interacting factors. In addition, degree of polygyny and sexual dimorphism in size are associated with the evolution of mating systems. Neither male–male combat nor paternal care, however, can completely explain differences in sexual size dimorphism for ungulates, a necessary component in understanding the development of some mating systems. Whatever the evolutionary pathway, sexual segregation limits paternal care allowing more intense male–male competition. Selection of habitat structure, because it modifies risk of predation, is a major determinant of sociality for ungulates. Likewise, ruggedness and steepness of terrain limit the types of mating systems that can occur because of limitations in group size and cohesiveness, as well as the ability of males to herd even small groups of females effectively. The quality and defensibility of resources affect mating systems, as does the defensibility of females. Population density of females also may be a critical determinant of the types of mating systems that develop. Size of groups likewise constrains the types of mating tactics that males can employ. Our aim was to use those relationships to create a broad conceptual model that predicts how various environmental and social factors interact to structure mating systems in ungulates. This model provides a useful framework for future tests of the roles of both ecological and social conditions in influencing the social systems of ungulates.     

The influence of nest predation on mating strategies under polygyny

The mating status of an advertising male is one important factorfor female assessment. In species where already-mated malestry to attract additional females at some distance from theirmate, mating status may be difficult to assess. Limitationsin the time available to breed and a decrease in reproductivesuccess over the breeding season could make the costs outweighthe benefits of a time-consuming mate assessment process, whichautomatically postpones the onset of breeding. Time and energyconstraints will then limit the behavioral options for femalesin search of breeding opportunities, and females would suffera cost when assessing male mating status. Female choice maythen be better understood by analyzing the behavioral optionsfor females; the success of a strategy in which costly assessmentsare avoided and females choose mates indiscriminately, disregardingmating status ("fast" strategy), could be compared to the successof a strategy in which females carefully try to assess matingstatus ("coy" strategy). Nest predation should be included insuch an analysis because it influences the amount of male assistancefemales of polygynous males receive, as nest predation for onlyone of a polygynous male's two females leads to more male assistancefor the other female. In this study, I compare the outcome ofa "fast" strategy with that of a "coy" strategy under differentrates of nest predation. Although high probabilities of becominga mate of a polygynous male associated with a fitness reductiondecrease the relative success of a "fast" strategy, simulationsshow that high rates of nest predation considerably lower thedifference in the outcome of the two strategies. Therefore,the cost of assessing mating status must be relatively smallto be economical.     

Dispersal Patterns in <Emphasis Type="Italic">Tarsius spectrum</Emphasis>

One of the most widely accepted explanations for the difference in the sex bias between mammals and birds is that male-biased dispersal in mammals is due to the preponderance of polygynous mating systems exhibited by this class, whereas birds are predominantly monogamous. Spectral tarsiers (Tarsius spectrum) are unusual in that they exhibit variation in its mating system. Although the majority of spectral tarsier groups are monogamous, ca. 15% are polygynous. If mating system influences dispersal, then I predicted that the polygynous groups would exhibit male biased dispersal whereas I predicted that the dispersal patterns of the monogamous groups would be analogous to that exhibited by birds, specifically female biased. Alternatively, I hypothesized that ecological variation may influence dispersal habits in this species. Specifically, I predicted that polygynous groups would exhibit greater habitat quality than monogamous groups. The 2 hypotheses are not mutually exclusive. On the basis of 14 individuals birdbanded between 1994 and 1999, I determined that individuals of both sexes were equally likely to disperse (males, n = 5; females, n = 9). Males dispersed twice as far as females did. The mean dispersal distance for males was 660 m, and for females it was 266 m. Females (77%) were more likely to form a territory adjacent to the parental territory than were males (20%). Individuals exhibited relatively high amounts of site fidelity (86%) that were related to physical characteristics of the sleeping site. Adults that dispersed a second time (n = 4) initially resided in trees that were shorter and had a smaller diameter-at-breast height than the trees of individuals that exhibited site fidelity. The results of my study partly support the parental mating system hypothesis and also support the habitat quality hypothesis.     

Nest and mate choice in the red bishop (Euplectes orix) : female settlement rules

We investigated female settlement in a colony of red bishops(Euplectes orix), a territorial and highly polygynous weaverbirdwidely distributed over sub-Saharan Africa. An earlier studyshowed that male reproductive success is mainly determinedby the number of nests a male builds in his territory, whichappeared to be a good indicator of male quality. Because malesprovide no parental care or food resources within the territory,females sharing a territory do not compete for material resourcesand might therefore be expected to settle preferentially interritories of males that build many nests to gain the possiblegenetic benefit of high-quality offspring. An analysis of femalesettlement, however, revealed that females did not show a preferencefor territories of males with many nests and that the distribution of female breeding attempts with regard to the number of vacantnests within a territory could be explained best by randomfemale settlement in 3 out of 4 years. Females settled moreoften than expected by chance (in 3 out of 4 years) in territoriesalready containing occupied nests, indicating that residentfemales did not discourage settlement of additional females.However, sharing a territory with other females might imposecosts in terms of an increased predation risk because nestsin territories that contained other nests with young sufferedfrom higher predation than nests in territories that did notcontain other nests with young. Females therefore might tradethe possible benefits of settling in territories of males withmany nests against the costs of sharing a territory with otherfemales. This might result in the mating pattern found withrandom female settlement and male reproductive success beingdirectly proportional to the number of nests built. We discuss possible implications of this mating pattern for sexual selectionon males.     

A role for female ornamentation in the facultatively polygynous mating system of collared flycatchers

In a polygynous mating system, females settling with alreadymated males often experience low mating success due to the reducedparental contribution of the male. However, there are numerousfactors that may still make it advantageous for some femalesto choose this mating status. Facultative polygyny is believedto be dominated by male advertisement and female choice. Althoughquality differences and competition among females are increasinglyrecognized as important determinants of polygynous settlementpatterns, the importance of signals of female quality in thismating system is largely unknown. Here we examined the relationshipof the white wing patch size (WPS) of female collared flycatchers,a phenotypically plastic and age-dependent ornament, with socialmating status, while controlling for settlement date and age.At the population level, monogamous, primary, and secondaryfemales did not differ in WPS. However, the primary female ofindividual males was more ornamented than the secondary female,and this difference declined with increasing distance betweenprimary and secondary nests. Secondary female ornamentationincreased, whereas that of the primary female did not changewith nest distance. These results suggest a subtle role forfemale ornamentation at polygynous mating. Future studies shouldtherefore take into account mating status when assessing thecosts and benefits of female signals. Moreover, patterns inquality-indicating female traits may contribute to the explanationof differences in reproductive success among females of differentmating status.     

Experimental confirmation of the polygyny threshold model for red-winged blackbirds

The polygyny threshold model assumes that polygynous mating is costly to females and proposes that females pay the cost of polygyny only when compensated by obtaining a superior territory or male. We present, to the authors' knowledge, the first experimental field test to demonstrate that females trade mating status against territory quality as proposed by this hypothesis. Previous work has shown that female red-winged blackbirds (Agelaius phoeniceus) in Ontario prefer settling with unmated males and that this preference is adaptive because polygynous mating status lowers female reproductive success. Other evidence suggests that nesting over water increases the reproductive success of female red-winged blackbirds. Here we describe an experiment in which females were given choices between two adjacent territories, one owned by an unmated male without any over-water nesting sites and the other by an already-mated male with over-water sites. Females overwhelmingly preferred the already-mated males, demonstrating that superior territory quality can reverse preferences based on mating status and supporting the polygyny threshold model as the explanation for polygyny in this population.     

Socio-genetic structure and mating system of a wild boar population

Wild boars Sus scrofa have a social organization based on female groups that can include several generations of adults and offspring, and are thus likely matrilineal. However, little is known about the degree of relatedness between animals living in such groups or occupying the same core area of spatial activity. Also, polygynous male mating combined with matrilineal female groups can have strong influences on the genetic structure of populations. We used microsatellite genotyping combined with behavioral data to investigate the fine-scale population genetic structure and the mating system of wild boars in a multi-year study at Châteauvillain-Arc-en-Barrois (France). According to spatial genetic autocorrelation, females in spatial proximity were significantly inter-related. However, we found that numerous males contributed to the next generation, even within the same social group. Based on our genetic data and behavioral observations, wild boars in this population appear to have a low level of polygyny associated with matrilineal female groups, and infrequent multiple paternity. Mortality due to hunting may facilitate the breakup of what historically has been a more predominantly polygynous mating system, and likely accelerates the turnover of adults within the matrilineal groups.     

South American sea lions in Peru have a lek‐like mating system

Five years of behavioral observations revealed significant effects of high air temperatures and breeding site topography on the mating system of South American sea lions in Peru. Unlike most polygynous mammals that defend females or fixed territories, male sea lions in Peru maintained positions along the shoreline where females passed each day to thermoregulate, and where most copulations occurred. Sex ratios (1 male per 17 females) and male mating success were extremely skewed (14% of males achieved 50% of the copulations, and 25% of them did not copulate at all). The mass daily movements of females toward the water and cool substrate of the shoreline, along with a highly skewed sex ratio, accentuated the difficulty for males to monopolize and restrict female movements. Females moved freely and chose their mates, unlike in temperate regions of their range where male South American sea lions control groups of females or access to tide pools. Our observations indicate that the South American sea lion in Peru has a lek‐like breeding system. This is a rare alternative to the common male strategies of defending females and resources, and is likely an evolutionary product of their highly skewed sex ratio, protracted breeding season, and the extreme subtropical climate where they breed.     

Is use of translocation for the conservation of subpopulations of oribi Ourebia ourebi (Zimmermann) effective? A case study

In South Africa, the oribi, Ourebia ourebi is an endangered small antelope that requires conservation management and intervention to prevent their extinction. Use of translocation for the conservation of subpopulations facing local extinction in the wild has been proposed. In this study, fifteen oribi from threatened populations were captured and released (November 2004) on a private game reserve in KwaZulu-Natal, South Africa. Radio telemetry was used to monitor the dispersal and survival of translocated oribi. Only one death occurred during that period. All other released oribi survived, and most remained in close proximity of the release site for more than a year postrelease. Reproduction also occurred. This suggests that, translocation could be a viable option for conserving wild populations of oribi, assuming that other factors, such as availability of suitable grassland habitat and that poaching has been curtailed, are met.     

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.     

Effects of maternal quality and mating status on female reproductive success in the polygynous spotless starling

Female mating status may affect reproductive success in avian polygynous mating systems through a combination of differences in female parental quality and status-dependent male assistance in parental care. Traditionally the literature has emphasized male assistance, neglecting evidence for consistency in female parental quality independent of mating status or repeatability in status. We studied the effects of male assistance on breeding success and its association with female mating status in a population of the polygynous spotless starling, Sturnus unicolor, during 3 years. Nestling provisioning by males improved the fledging success of late (mostly second) but not early (mostly first) broods. Reproductive success of females was affected mainly by female maternal quality: (1) primary and secondary females did not differ in output for early broods and in seasonal output despite a greater male assistance at primary nests; (2) monogamous females were more successful despite receiving no more help than other females; and (3) primary and secondary females had different clutch sizes before male assistance in parental care could operate. Female mating status was not significantly repeatable within seasons or between years. Females who changed status between years were as successful when rearing broods as secondary females as they were as primary or monogamous females. Breeding success (proportion of eggs resulting in fledglings) and clutch size were significantly repeatable between years for late but not for early clutches. Performance showed no strong association with female age or condition. Female breeding success seems to be weakly affected by male assistance in this sedentary, colonial species, and more dependent on inherent differences between females. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Intraspecific Variation in the Mating System of Spectral Tarsiers

Historically, a species’ social system was perceived to be a fixed attribute and deviations were usually treated as aberrant or maladaptive. Over the last few decades, socioecologists have started to recognize that variation in social systems is normal and adaptive. Here I explore how ecological variation affects the mating system of a nocturnal primate, the spectral tarsier, Tarsius spectrum. Several studies of the spectral tarsier have illustrated variation in this species’ mating system. Although most groups exhibit a monogamous mating system, a small proportion of the population consistently exhibits a polygynous mating system. I conducted this study at Tangkoko Nature Reserve in Sulawesi, Indonesia during 2007. I found that group size was highly variable, ranging from 2 individuals to as many as 8 individuals per sleeping site. Although most groups (21 of 26) were composed of a single adult male and a single adult female, ca. 19% of the groups contained 2 adult females. Three of the 5 groups with 2 adult females contained 2 young infants. As might be expected, polygynous groups were larger than were monogamous groups. The mean group size of monogamous groups was 2.9, whereas the mean group size of polygynous groups was 6.0. Polygynous groups were also more likely to use Ficus caulocarpa trees than were the monogamous groups. Polygynous groups consistently used more sleeping sites as well as larger diameter sleeping trees, than did monogamous groups. The large-diameter fig trees are ideal homes for the spectral tarsiers in that they offer multiple entrances and exits as well as protection from the elements. Polygynous and monogamous groups exhibited no differences in insect biomass available, home range size, or height of sleeping tree. These results support the hypothesis that ecological variation is an important determinant of mating system within spectral tarsiers.     

GENETIC VARIABILITY,QUEEN NUMBER,AND POLYANDRY IN SOCIAL HYMENOPTERA

Several hypotheses have been put forward to explain the adaptive significance of interspecific variation in mating frequencies by eusocial hymenopteran queens. Four of these hypotheses assert that polyandry is advantageous to queens because of the resultant increase in genetic variability within colonies (referred to as the “GV” hypotheses). Here we compare the frequency of polyandry between monogynous (single-queen) and polygynous (multiple-queen) ant species to test the hypotheses that (1) multiple mating functions primarily to increase intracolonial genetic variability, and (2) mating has costs (such as increased energetic losses or risk of predation or venereal disease). If one of the GV hypotheses is true and mating is costly, the frequency of polyandry should be lower in polygynous species (in which the presence of multiple queens results in low relatedness among workers) than in monogynous species. As predicted by the GV hypotheses, polyandry is less common among polygynous than among monogynous species. Furthermore, it seems that the causal relationship underlying this association is that the number of matings by queens depends on the number of queens present in the colony (rather than the number of queens being influenced by the number of matings), which also supports the GV hypotheses together with the assumption that mating has costs.     

Polygyny and the breeding success of the great reed warblerAcrocephalus arundinaceus

Settlement date, mating status, and breeding success of individually marked great reed warblers, Acrocephalus arundinaceus, were studied during the 1980–84 breeding seasons in Kahokugata, Ishikawa Prefecture, Japan. Twenty-five per cent of the territorial males were polygynous, of which the majority were bigamous. The settling periods of both sexes were long, extending for 65 days in males and 49 days in females. About 80% of males and females settled in the first half of the settling period, and the settlement date of 28–54% males overlapped with that of females. Many of the late settlers were bachelors and the males which mated earlier tended to be polygynous. The timing of a male's settlement is important in acquiring mates. Fifty-five per cent of eggs laid were lost before fledging, mainly due to predation. The mean number of fledglings was 3.19 per primary female, 2.41 per secondary female, and 2.80 per monogamous female. Comparison of the number of fledglings of females which mated during the same period showed that the presence of another female in the same territory did not adversely affect the breeding success of either of the polygynous females. Polygynous males have the advantage of decreasing the risk of breeding failure under high predation pressure.     

Effects of nestboxes on the breeding biology of Southern House Wrens Troglodytes aedon bonariae in the southern temperate zone

Nestboxes are known to increase clutch size, enhance breeding success and affect the social mating system of several cavity nesters. Although in recent years various cavity nesters have been studied in nestboxes in South America, the effects of boxes on the biology of the study species are unknown. We evaluated the effects of nestboxes on the breeding biology and social mating system of Southern House Wrens Troglodytes aedon bonariae by comparing birds breeding in nestboxes and tree cavities in two cattle ranches in Buenos Aires Province, Argentina. Southern House Wrens nesting in boxes had higher breeding success but, contrary to studies on the temperate zone, we did not find differences in clutch size between Wrens breeding in nestboxes and tree cavities. The main causes of nest failure in tree cavities were nest predation and flooding of the cavity (70 and 23% of the failures, respectively) while in nestboxes predation and desertion were the most important causes of failure (38 and 34% of the failures, respectively). The social mating system of Southern House Wrens is monogamy with biparental care, and neither was affected by the boxes. Males did not attract secondary females to additional nestboxes; however, nestboxes are safer breeding sites than tree cavities, and females seemed to prefer males with nestboxes on their territory. These results suggest that nest quality alone might be not enough for secondary females to accept polygyny.     

Female preference for large males in sailfin mollies,<Emphasis Type="Italic"> Poecilia latipinna</Emphasis>: the importance of predation pressure and reproductive status

We examined the effect of predation risk on female association patterns in the live-bearing sailfin molly (Poecilia latipinna). We tested two classes of females, with and without the risk of predation by a green sunfish (Lepomis cyanellus): (1) postpartum females (maintained with males until visibly gravid, then isolated and tested within 24–48 h of dropping a brood); and (2) non-postpartum females (different females, isolated from males for >50 days). When there was no apparent risk of predation, postpartum females showed a significant preference for large over small males, whereas non-postpartum females showed no size preference at all. When there was an apparent risk of predation, postpartum females maintained their preference for larger males and non-postpartum females continued to show no preference for large or small males. These results suggest that reproductive status (receptivity) plays a greater role in mate preference than predation risk. For postpartum females, the cost of not choosing a preferred mate may outweigh the potential cost of predation. Non-postpartum females either do not benefit from mating or are being indecisive about mating and thus are less likely to be choosy whether or not a predator is present.Communicated by I. Schlupp     

Infanticide as a male reproductive strategy has a nutritive risk effect in brown bears

Behavioural strategies to reduce predation risk can incur costs, which are often referred to as risk effects. A common strategy to avoid predation is spatio-temporal avoidance of predators, in which prey typically trade optimal resources for safety. Analogous with predator–prey theory, risk effects should also arise in species with sexually selected infanticide (SSI), in which females with dependent offspring avoid infanticidal males. SSI can be common in brown bear (Ursus arctos) populations and explains spatio-temporal segregation among reproductive classes. Here, we show that in a population with SSI, females with cubs-of-the-year had lower quality diets than conspecifics during the SSI high-risk period, the mating season. After the mating season, their diets were of similar quality to diets of their conspecifics. Our results suggest a nutritive risk effect of SSI, in which females with cubs-of-the-year alter their resource selection and trade optimal resources for offspring safety. Such risk effects can add to female costs of reproduction and may be widespread among species with SSI.     

Breeding Biology and Variable Mating System of a Population of Introduced Dunnocks (Prunella modularis) in New Zealand

Species with variable mating systems provide a unique opportunity to investigate whether females receive direct fitness benefits from additional male partners. The direct benefits provide an obvious explanation for why females would breed polyandrously, in a situation where males clearly do not attain their optimal reproductive success. Evidence for these direct benefits is, however, mixed. Here, we present a detailed study of the breeding biology of the dunnock, Prunella modularis, which inform an investigation into the effects of the social mating system on the reproductive success in a population of dunnocks in Southern New Zealand. We studied 80 different social groups over the course of three breeding seasons. Dunnocks in our population presented a variable mating system, with socially monogamous (45%), socially polyandrous (54%) and socially polygynandrous (1%) groups being observed in the same breeding season. We did not observe any polygynous social units in our study period although polygyny exists in the population. We found little difference in the numbers of eggs laid, and egg volume between monogamous and polyandrous nests. However, polyandrous groups had better hatching and fledging success than monogamous groups (composite d = 0.385, 95% CI: 0.307 to 0.463). Overall our results support the notion that polyandry is beneficial for females.     

Predation, sensitivity, and sex: why female black rhinoceroses outlive males

Among sexually dimorphic, polygynous mammals, adult femalestend to outlive males and respond more strongly to predatorsthan males. We asked whether a monomorphic, polygynous speciesvirtually immune to predation due to large size (black rhinoceros,Diceros bicornis) conforms to this pattern. Data on 193 interactionswith lions (Panthera leo) and spotted hyenas (Crocuta crocuta)in two nonhabituated populations in Namibia studied from 1991to 1993 revealed that: (1) females were more vigilant or aggressivethan males to either of the potential predators; and (2) whethersolitary or with calves, females attacked more often than males.Although solitary females tended to be more aggressive to lionsthan to hyenas, neither females with calves or males seemedto discriminate between the two carnivores. We also simulatedthe behavior of human predators (poachers) during 69 encounterswith rhinoceroses. While both sexes abandoned local sites becauseof our presence, females ran farther than males, covering upto 40 km in a day. These findings implicate a behavioral mechanismto explain why secondary sex ratios favor females - males aremore prone to human predation, a prediction consistent withdata from 12 populations throughout Africa. Black rhinocerosesappear to be an unanticipated exception to the well-establishedpattern of male-biased mortality in polygynous mammals; in theabsence of intense human predation (a recent event), male mortalityfails to exceed that of females, suggesting that intrasexualcompetition in a polygynous mammal: may not be the primary causeof unbalanced secondary sex ratios. Our results on the causesof sex differences in mortality and in responsiveness to differentpredators reinforce the relevance of behavioral ecology to conservation;such information is necessary for planning how best to minimizenegative human influences on the few remaining wild Africanrhinos.