Kin-related spatial structure in brown bears Ursus arctos

Kin-related social structure may influence reproductive success and survival and, hence, the dynamics of populations. It has been documented in many gregarious animal populations, but few solitary species. Using molecular methods and field data we tested: (1) whether kin-related spatial structure exists in the brown bear (Ursus arctos), which is a solitary carnivore, (2) whether home ranges of adult female kin overlap more than those of nonkin, and (3) whether multigenerational matrilinear assemblages, i.e., aggregated related females, are formed. Pairwise genetic relatedness between adult (5 years and older) female dyads declined significantly with geographic distance, whereas this was not the case for male–male dyads or opposite sex dyads. The amount of overlap of multiannual home ranges was positively associated with relatedness among adult females. This structure within matrilines is probably due to kin recognition. Plotting of multiannual home-range centers of adult females revealed formation of two types of matrilines, matrilinear assemblages exclusively using an area and dispersed matrilines spread over larger geographic areas. The variation in matrilinear structure might be due to differences in competitive abilities among females and habitat limitations. The influence of kin-related spatial structure on inclusive fitness needs to be clarified in solitary mammals.     

Why breed communally? Factors affecting fecundity in a communal breeding mammal: the banded mongoose (<Emphasis Type="Italic">Mungos mungo</Emphasis>)

Fecundity is an important component of fitness. In cooperatively breeding species, studies aimed at understanding the factors that affect fecundity have largely been restricted to species that exhibit high reproductive skew, where reproduction is monopolised by a few individuals. In such species, dominant suppression and inbreeding avoidance are the principal explanations for low fecundity in subordinate females. In this paper, we evaluate the relative effects of individual, social, and environmental factors on female fecundity in a low skew cooperative breeding mammal: the banded mongoose (Mungos mungo). Most females (80%) conceived in each breeding event, and most pregnant females (93%) carried their litter to term. The principal determinants of a females fecundity were intrinsic qualities, particularly age and body size. However, there was no evidence of dominant suppression of subordinate reproduction or inbreeding avoidance. Similarly, there was little indication that social or environmental factors influence fecundity. We suggest that in the banded mongoose, the apparent lack of costs to inbreeding, and the absence of dominant female suppression of reproduction in other females result in low reproductive skew. Indeed, in banded mongooses, like lions (Felis leo), multiple breeding may be a consequence of benefits to rearing young communally.Communicated by F. Trillmich     

Reproductive success in a low skew, communal breeding mammal: the banded mongoose, Mungos mungo

In most cooperatively breeding species, reproduction is monopolised by a subset of group members. However, in some species most or all individuals breed. The factors that affect reproductive success in such species are vital to understanding why multiple females breed. A key issue is whether or not the presence of other breeders is costly to an individual’s reproductive success. This study examines the factors that affect the post-parturition component of reproductive success in groups of communal-breeding banded mongoose (Mungos mungo), where up to ten females breed together. Per-litter reproductive success was low (only 18% of pups survived from birth to independence). Whilst singular breeding was wholly unsuccessful, there were costs associated with breeding in the presence of increasing numbers of other females and in large groups. Synchronisation of parturition increased litter success, probably because it minimises the opportunity for infanticide or decreases competitive asymmetry between pups born to different females. There was no evidence of inbreeding depression, and reproductive success was generally higher in litters where females only had access to related males within their group. I conclude that communal breeding in female banded mongooses represents a compromise between the benefits of group-living and communal pup care on the one hand, and competition between females to maximise their personal reproductive success on the other. Such conflicts are likely to occur in most communal breeding species. Whilst communal breeding systems are generally considered egalitarian, negative effects of co-breeders on individual reproductive success is still an issue.     

Age- and state-dependent reproductive effort in male mountain goats, Oreamnos americanus

In mammalian polygynous mating systems, male reproductive effort consists mainly of male–male competition and courting of females, which entail substantial somatic costs. Males are thus expected to adjust their reproductive effort according to their age and condition. In this study, we examined how activity budgets of male mountain goats (Oreamnos americanus), a polygynous ungulate, varied with age in a marked population over two periods: (1) summers 1995–2006 and (2) ruts 2004–2006. We then assessed if the proportions of time spent in male–male competition and courtship behaviors were influenced by age-specific body mass and social rank during the rut. Males spent most of their time foraging and resting during summer, and rested more and foraged less with increasing age. During the rut, pronounced shifts in activity budgets occurred as juveniles (1–2 years) increased time spent foraging, whereas adults (≥3 years) increased standing and time spent in social interactions at the expense of foraging. At old age, reproductive effort either stabilized or decreased slightly, providing weak support for the ‘mating strategy–effort’ hypothesis, predicting that courtship behaviors should peak in prime-aged males. Age-specific body mass did not affect time spent in male–male competition, but was positively related with time spent in courtship behaviors, providing support for the ‘individual quality’ hypothesis, predicting that males with more resources at the start of the rut should spend more time in mating-related activities. Age-specific social rank did not affect reproductive effort. Surviving to prime age while increasing mass each year should thus allow male ungulates to gain greater ability to court estrus females.     

Sex‐Biased Inbreeding Effects on Reproductive Success and Home Range Size of the Critically Endangered Black Rhinoceros

A central premise of conservation biology is that small populations suffer reduced viability through loss of genetic diversity and inbreeding. However, there is little evidence that variation in inbreeding impacts individual reproductive success within remnant populations of threatened taxa, largely due to problems associated with obtaining comprehensive pedigree information to estimate inbreeding. In the critically endangered black rhinoceros, a species that experienced severe demographic reductions, we used model selection to identify factors associated with variation in reproductive success (number of offspring). Factors examined as predictors of reproductive success were age, home range size, number of nearby mates, reserve location, and multilocus heterozygosity (a proxy for inbreeding). Multilocus heterozygosity predicted male reproductive success (p< 0.001, explained deviance >58%) and correlated with male home range size (p < 0.01, r² > 44%). Such effects were not apparent in females, where reproductive success was determined by age (p < 0.01, explained deviance 34%) as females raise calves alone and choose between, rather than compete for, mates. This first report of a 3‐way association between an individual male's heterozygosity, reproductive output, and territory size in a large vertebrate is consistent with an asymmetry in the level of intrasexual competition and highlights the relevance of sex‐biased inbreeding for the management of many conservation‐priority species. Our results contrast with the idea that wild populations of threatened taxa may possess some inherent difference from most nonthreatened populations that necessitates the use of detailed pedigrees to study inbreeding effects. Despite substantial variance in male reproductive success, the increased fitness of more heterozygous males limits the loss of heterozygosity. Understanding how individual differences in genetic diversity mediate the outcome of intrasexual competition will be essential for effective management, particularly in enclosed populations, where individuals have restricted choice about home range location and where the reproductive impact of translocated animals will depend upon the background distribution in individual heterozygosity. Efectos de la Endogamia Sesgada por el Sexo sobre el Éxito Reproductivo y el Rango del Tamaño de Hábitat del Rinoceronte Negro, Especie en Peligro Crítico     

There is a limbo under the moon: what social interactions tell us about the floaters’ underworld

The ultimate and proximate causes of natal dispersal have been extensively investigated, but the behaviour of dispersers in relation to social interactions has been largely neglected. Here, we investigated the social organisation of floating individuals during their dispersal by analysing the behaviour of 40 radio-tagged eagle owls Bubo bubo during the wandering and stop phases of dispersal. Unexpectedly, eagle owl floaters mixed with conspecifics independently of their sex, age, phase of dispersal, birthplace, health status and habitat features, showing an ‘underworld’ of interactions characterised by the absence of obvious social organisation or short-term strategies. Non-breeding owls were not transient floaters that occurred at numerous sites for short periods of time but rather had fairly stable home ranges: they attempted to settle as soon as possible within well-defined home ranges. The spatial distribution pattern of floaters and high rates of home range overlap support the prediction that floating individuals are not spatially segregated, challenging the expectation that dominance by size, age and/or health status may determine the exclusive use of some portions of the dispersal area. Finally, (1) the short distances among conspecifics and the extensive home range overlaps allowed us to discard the possibility that neighbouring floaters represent a real cost during dispersal and (2) floater interactions showed a lack of clear mechanisms for avoidance of kin competition among offspring or inbreeding.     

Philopatry and mother-daughter associations in bushy-tailed woodrats: space use and reproductive success

Social groups in mammals are often based on overlapping generations of philopatric females, yet few researchers have examined fitness consequences of associations between females and their daughters. We examined survival and reproductive consequences of sharing a natal site with one's mother or daughter in wild bushy-tailed woodrats (Neotoma cinerea). Bushy-tailed woodrats reside on discrete rocky outcrops, and females tend to spend their entire reproductive life on their natal outcrop. Mothers and daughters that co-existed on the natal outcrop were closely associated in space. Juvenile females were more likely to survive their first winter to become breeding yearlings at their natal site if their mother was also present at the natal site during the winter. Juveniles survived equally well with or without their mother where densities of adult females were experimentally reduced, suggesting that females enhance their daughter's survival by facilitating their access to limited resources. Yearling females exhibited no reproductive costs from association with their mothers at the natal site during the breeding season; in fact, yearlings that bred at the natal site simultaneously with their adult mothers tended (P = 0.07) to experience fewer reproductive failures than did those that bred in the absence of their mothers. Mothers did not experience any detectable costs when sharing a natal site with a daughter. We conclude that bushy-tailed woodrats exhibit stable, fitness-enhancing associations among mothers and their philopatric daughters.     

Scent-marking behavior in wild groups of common marmosets (Callithrix jacchus)

Scent-marking in mammals has been frequently related to within-group social and reproductive dominance and to defense of territory and resources. We studied the scent-marking behavior of five wild groups of common marmosets, Callithrix jacchus, during 5 months of the fruiting season in northeastern Brazil. Circumgenital marking was the most common type of marking. Marks were distributed throughout the home range and were deposited mainly during travel and intergroup encounters. Although marks were commonly deposited at gum trees, there was no evidence that the animals used scent marks to label fruiting trees or sleeping sites. Contrary to expectations, reproductively dominant females did not mark more than reproductively subordinate females. Moreover, during intergroup encounters, reproductively subordinate females displayed higher frequencies of scent-marking than the reproductively dominant females of their group. Our results suggest that scent-marking is not strictly tied to reproductive dominance or territorial (or resource) defense in common marmosets. Because marks provide information about individual identity and reproductive condition, scent marks could serve different functions when used by different individuals. The high frequency of marking by reproductively subordinate females during intergroup encounters suggests that scent-marking might be used to signal to individuals of neighboring groups. Our data highlight the importance of social and ecological variables in scent-marking behavior. Received: 26 October 1998 / Received in revised form: 20 May 1999 / Accepted: 30 May 1999     

Tug-of-war over reproduction in a cooperatively breeding cichlid

In group-living animals, dominants may suppress subordinate reproduction directly and indirectly, thereby skewing reproduction in their favour. In this study, we show experimentally that this ability (‘power’) is influenced by resource distribution and the body size difference between unrelated dominants and subordinates in the cichlid Neolamprologus pulcher. Reproduction was strongly skewed towards the dominant female, due to these females producing more and larger clutches and those clutches surviving egg eating better than those of subordinate females, but was not so when subordinates defended a patch. If breeding shelters were provided in two patches, subordinate females were more likely to exclusively defend a patch against the dominant female and breed, compared to when the same breeding resource was provided in one patch. Relatively large subordinate females were more likely to defend a patch and reproduce. Females also directly interfered with each other’s reproduction by eating the competitors’ eggs, at which dominants were more successful. Although dominant females benefited from subordinate females due to alloparental care and an increase in egg mass, they also showed costs due to reduced growth in the presence of subordinates. The results support the view that the dominant’s power to control subordinate reproduction determines reproductive partitioning, in agreement with the predictions from tug-of-war models of reproductive skew.     

Competition and group size in Thomas’s langurs (Presbytis thomasi): the folivore paradox revisited

Among primates, group size is highly variable. The standard ecological model assumes that better predation avoidance as group size increases favours living in larger groups, whereas increased travel costs and reduced net food intake due to within-group competition for resources set the upper limit. Folivorous primates, however, tend to defy this generalisation in that some live in small groups despite low costs of feeding competition. To resolve this ’folivore paradox’, it has been suggested that folivore group size is limited by social factors such as male harassment or infanticide, or that females can disperse more easily and thus maintain group size near optimum levels. In this paper, we examine the effects of group size on home range size, day-journey length, activity budget and diet in wild Thomas’s langurs (Presbytis thomasi), which live in one-male multi-female groups with a limited life cycle. We examined only data from the stable middle tenure phase when factors such as the strength of the breeding male or the way in which groups were formed did not influence ranging and activities. During this phase, group size affected day-journey length and home range size, and had a minor effect on diet, but did not influence time spent feeding or resting, allogrooming or birth rates. Hence the upper limit to group size during the middle tenure phase in Thomas’s langurs is not set by feeding competition. The folivore paradox is not due to frequent female dispersal in Thomas’s langurs. The timing of female dispersal is not as expected if it serves to keep group sizes near the ecological optimum, and groups seem to be below this optimum. Instead, female reproductive success is presumably maximised in small to mid-sized groups because larger groups show a clear trend to experience higher risk of take-over, often accompanied by infanticide. Because females can redistribute themselves among nearby groups when groups reorganise each time a new male starts up a new group, females can keep the group small. Thus, a social factor, risk of infanticide, seems to provide the selective advantage to small group size in Thomas’s langurs. Received: 29 July 1999 / Revised: 17 November 1999 / Accepted: 15 October 2000     

Delayed breeding in the cooperatively breeding Florida scrub-jay (Aphelocoma coerulescens): inhibition or the absence of stimulation?

To determine whether fundamental differences exist in the reproductive physiology of breeder and nonbreeder Florida scrub-jays (Aphelocoma coerulescens), we compared plasma levels of testosterone (T) and luteinizing hormone (LH) in males, and estradiol (E₂) and LH in females. Although male breeders had higher overall T and larger testes, nonbreeders’ T paralleled that of breeders, and their testes were more than an order of magnitude larger than regressed testes. Breeder and nonbreeder males had equivalent baseline LH, and equivalent changes in LH following a gonadotropin-releasing hormone (cGnRH-I) challenge. The T, LH and GnRH challenge data indicate that nonbreeder males have functional hypothalamo-pituitary-gonadal (HPG) axes. We found no hormonal evidence of inbreeding suppression in males: nonbreeders that did not live with their mothers and those that did had similar T. Male nonbreeders that were exposed to E₂-implanted females had higher T than did controls, suggesting that the lack of within-pair stimulation is a key factor in whether an individual delays breeding. Female nonbreeders had E₂ titres equal to or higher than breeders and neither basal LH nor LH following GnRH challenge differed by breeding status. Nonbreeders’ ovarian follicles were smaller than breeders’, but were larger than they would be during the non-breeding season. These data suggest that nonbreeders were primed for breeding and were simply waiting for an opportunity or a required stimulus. Female nonbreeders that lived in a territory with an unrelated male breeder had significantly higher E₂ than those that remained with their fathers. Similarly, nonbreeders that were captured away from their home territories had elevated E₂. However, nonbreeders that lived with their fathers had E₂ that was equivalent to breeding females, suggesting that inbreeding avoidance may not be the primary factor leading to delayed breeding in females. Received: 13 June 1995 /Accepted after revision: 27 April 1996     

Plasma levels of androgens and gonadal development of breeders and helpers in the bell miner (Manorina melanophrys)

Plasma concentration of androgens and gonad development were studied in wild cooperatively breeding bell miners (Manorina melanophrys) of different age, sex and social status. Plasma levels of androgens increased with age of birds. For sexually mature (SM) bell miners, androgen levels were higher in breeders than in nonbreeding (NB) helpers (P < 0.005), and they were marginally higher in females than in males (P < 0.10). Thus female breeders showed the highest plasma levels of androgens. This pattern may be explained by an asymmetry between the sexes in the levels of competition needed to reach and defend breeding status; the degree of intra-sexual competition seems to be higher among female bell miners than among males. In general, male breeders had larger gonads than male SMNB helpers; male breeder gonads produced spermatozoa whereas most helpers' gonads did not. However, old (i.e. 34 months) SMNB male helpers had large gonads producing spermatozoa in spite of having low plasma levels of androgens. We suggest that young male bell miners may be under a voluntary mechanism of reproductive suppression perhaps favoured by inbreeding avoidance; although some aggressive interactions observed between very young helpers (about 2 months) and adult members of the breeding unit suggests that a certain degree of imposition also occurs at young ages. Older male helpers may avoid aggression with the male breeder and imposed reproductive suppression, hence the low circulating levels of androgens, but they have the potential to sneak extra-pair copulations. Gonad size decreased during the non-breeding season in both sexes.     

Pup production,sex ratios,and survivorship in African wild dogs, <Emphasis Type="Italic">Lycaon pictus</Emphasis>

The local resource enhancement (LRE) model predicts that in cooperatively breeding species, sex ratios will be biased in favor of the more helpful sex. In this study, we assess the assumptions underlying the LRE model in a population of cooperatively breeding wild dogs (Lycaon pictus) in Northern Botswana monitored over a 15-year period. In this population, litter size and pup survival to 1 year are strongly affected by pack size and the breeding female’s age, but adult males have a stronger and more linear effect on females’ reproductive performance than do adult females. This asymmetry in the benefits derived from male and female helpers is reflected in male-biased sex ratios in litters at the time pups emerge from the den. Sex ratio biases are most pronounced in the litters of the youngest mothers who live in significantly smaller packs than older females. The presence of potential rivals for the dominant female’s position depresses pup production at the time of emergence, suggesting that competition among females for breeding positions may also contribute to the selective forces affecting birth sex ratios.     

Pregnancy failure and suppression by female-female interaction in enclosed populations of the red-backed vole,Clethrionomys rufocanus bedfordiae

Summary Experimental populations of the redbacked vole,Clethrionomys rufocanus bedfordiae (Thomas), were introduced into field enclosures and then subsequently checked for home range overlap and reproductive condition. The times to parturition of females that did not have exclusive home ranges were significantly delayed compared with those of females whose ranges did not overlap (Table 4). The delays in parturition were not significantly related to the age of the females or to the number and ages of males that overlapped with a female. Copulatory plugs observed in the females that did not have exclusive ranges indicated that they often failed to become pregnant, regardless of having copulated. After the acquisition of exclusive ranges, these same females came into estrus and became pregnant (Figs. 2, 3).     

The functional significance of infanticide in a population of California ground squirrels (Spermophilus beecheyi)

Naturally-occurring infanticide was observed in a population of California ground squirrels (Spermophilus beecheyi). In four seasons, 40 infanticides were observed. All victims were post-emergent pups. Of 37 killings in which the killer was sexed 36 were by females. All infanticidal females were mothers at the time they killed, but in no case was a mother seen to kill or harm her own young. The victim was cannibalized in 22 cases and taken immediately into the killer’s burrow in 16 others. In no case did killers gain access to the victimized mother’s burrow or territory and female pups were not killed preferentially over males. In light of evolutionary explanations, infanticide in this population may best fit the resource exploitation hypothesis, in which killers commit infanticide to gain a nutritional benefit. Resource competition is a possible auxiliary explanation, since any time a female kills unrelated young she is eliminating possible competitors to her own offspring. This behavior could confer a fitness advantage on killers or it could be an adaptively neutral, alternative feeding strategy. More data are necessary to distinguish between these hypotheses. Received: 26 January 1995/Accepted after revision: 9 September 1995     

Biparental investment and reproductive success in a subsocial desert beetle: the role of maternal effort

Parastizopus armaticeps is a nocturnal subsocial detritivorous desert tenebrionid that produces very few offspring per brood. The two environmental factors that constrain reproduction, rapid sand desiccation rate and food scarcity, are countered by biparental effort. Males dig and extend breeding burrows, maintaining their moisture level; females forage on the surface at night for high-quality detritus, the larval food. This was shown to be a scarce and unpredictable resource for which there is high competition. When food was supplemented in a field experiment, offspring number and survivorship doubled and burrow failure due to desiccation dropped from approximately half, the typical failure rate for unsupplemented burrows, to zero. Food supplementation did not, however, increase larval foodstore size and there was no difference in the size of the offspring produced. Supplemented females reallocated their time, foraging less and digging more with the male. This change in maternal behaviour patterns resulted in deeper burrows which remained moist longer, thus extending the larval production period. Female foraging efficiency, particularly food retrieval speed, determined how much time females could allocate to digging, consequently increasing the reproductive success of the pair. Burrow depth and sand moisture level at the burrow base were the major correlates of reproductive success, but the scarcity and unpredictability of high-quality food on the surface and the competition for this resource influenced the number of offspring indirectly through their effect on female behaviour. Received: 29 November 1996 / Accepted after revision: 7 December 1997     

Experimental evidence for helping and hindering by previous offspring in the cooperative-breeding Seychelles warbler Acrocephalus sechellensis

Prebreeding Seychelles warblers (Acrocephalus sechellensis) frequently act as helpers on their natal territory, aiding in territory defence, predator mobbing, nestbuilding, incubation (only females) and feeding dependent young of their parents. In some cases helpers could attain breeding status (e.g. by joint-nesting) in their natal group and become co-breeders. Comparisons of group size and reproductive success on a given quality territory suggest that the presence of alloparents (helpers and cobreeders) significantly affects the reproductive success of their parents. The influence of alloparents on reproductive success was examined by removing alloparents from breeding units and comparing the success of natural-sized and artificially reduced groups. Removal experiments, controlled for territory quality, group size and breeder age, showed that the presence of one alloparent significantly improved the reproductive success of its parents. Analysis strongly suggests that this was entirely due to helping behaviour (i.e. providing care to offspring of their parents), thereby improving the helper's inclusive fitness benefits from staying at home. However, these experiments showed also that the presence of two or more alloparents in medium-quality territories significantly decreased reproductive success, compared with groups with one alloparent. Several lines of evidence suggest that this may have been due to the joint-nesting and reproductive competition that could occur in breeding groups, or simply to resource depression when a large number of previous offspring remained on their natal territory.     

Natal philopatry in bannertailed kangaroo rats

Summary I describe prolonged retention of offspring in natal home ranges, or natal philopatry, in the bannertailed kangaroo rat Dipodomys spectabilis. Though weaning occurs at about one month of age, offspring shared natal burrows with their mothers for three to seven months, and 39% of surviving offspring remained within natal home ranges through reproductive maturity. Males as well as females were philopatric. Data on the availability of burrows and on patterns of resettlement suggest that natal philopatry in this species may be a means of providing juveniles with access to essential resources, in this case food caches and large complex burrow systems, that are not readily available outside natal home ranges.     

Determinants of male reproductive success in American black bears

We determined annual male reproductive success in black bears (Ursus americanus) using DNA and field data from two populations in New Mexico. We identified the likely father for 78 of 120 genotyped cubs born during 1994–2000 and calculated reproductive success for 102 males known or presumed present within designated mating regions. Age was a strong determinant of reproductive success. The observed peak in reproductive success occurred at roughly the same intermediate age (10 years) as milder peaks in body size characteristics (11–12 years) and frequency of bear-inflicted wounds (13 years), suggesting body size and fighting might be important for mating. Success was negatively associated with the distance between home range centers of males and mothers. Success of young males (<7 years old) was also negatively associated with mature male (≥7 years old) density, and increasing density shifted the peak age of reproduction higher. The dispersed distribution of females likely limited the capacity of large mature males to dominate reproduction; therefore, success was determined by a complex set of variables.     

Factors predicting male fertilization success in an external fertilizer

In postcopulatory sexual selection both sperm competition and cryptic female choice are considered to be important selective agents, but their relative importance for male fertilization success has received little attention. We tested whether sperm quality, male spawning coloration, male heterozygosity, and genetic overlap with the female explained a male’s fertilization success in controlled in vitro fertilization competition trials between equal numbers of sperm from pairs of male Arctic charr (Salvelinus alpinus), an external fertilizer. Offspring were genotyped to determine each males’ share of paternity. The velocity of a male’s sperm relative to the velocity of the competing male’s sperm was the best predictor of male fertilization success. Yet, sperm velocity was not related to spawning coloration or male heterozygosity. In fact, the most brightly colored male in a pair had the lowest fertilization probability. This could result from cryptic female choice for pale males, but might rather be a result of paler males producing more competitive sperm than more colored males. Furthermore, the more microsatellite alleles a male shared with the female relative to the competing male, the higher fertilization success he had. We argue that this latter may be an effect of assortative cryptic female choice, which might prevent hybridization with sympatric Arctic charr morphs or one form of kin selection.