Male reproductive success in free-ranging feral horses

In the social organization of feral horses, adult males compete to monopolize groups or bands of females, sometimes called harems. Alternative male strategies are to remain alone or with other bachelors or, less commonly, to accept subordinate status within a harem. The hypothesis that dominant harem stallion status confers a reproductive advantage was tested in free-ranging feral horses. The presence of foals in harems headed by vasectomized (VSX) versus intact stallions was used to assess the ability of these stallions to control reproduction in their harems. Of harems headed by VSX stallions, 17 and 33% contained foals during years 2 and 3 post-treatment, respectively. In contrast, 86 and 80% of harems headed by non-VSX stallions contained foals in those years. Acquisition of pregnant mares appeared more likely than sneak copulations by bachelor stallions to account for foals in harems with a single stallion. However, most foals were born into harems that included a subordinate stallion, an occurrence that was undoubtedly exacerbated by the extended breeding season resulting from the sterility of the harem stallion. Thus, in comparing alternative reproductive tactics, bachelors appeared less successful than subordinate stallions within a harem. However, the highest reproductive success was achieved by the harem stallion, further demonstrating that alternative tactics are not equally profitable. Received: 13 July 1999 / Received in revised form: 24 July 1999 / Accepted: 24 July 1999     

Harem choice and breeding experience of female southern elephant seals influence offspring survival

Female mammals can increase their lifetime fitness through modification of investment potential and by providing better rearing environments with improved breeding experience. We examined the relationships between reproductive fitness and the behavioural decisions that female southern elephant seals (Mirounga leonina) made during the breeding season. We examined whether mother age and breeding experience influenced reproductive success (measured as 1st-year survival probability), and whether there was a change in the choice of harem size with increasing age. Pups produced by young mothers had lower 1st-year survival probability than pups produced by older mothers. A significant increase in mean female mass with age required an analysis of both these effects on offspring survival. There was a significant positive effect of both female age and mass, and the interaction between the two, on 1st-year pup survival. The proportion of young mothers (<5 years old) decreased and the proportion of older mothers (>6 years old) increased with increasing harem size (harems surveyed from 1997 to 2001). Females chose larger harems in which to breed as they aged. Females demonstrated fidelity to breeding areas among successive breeding seasons, with older females displaying greater breeding-site fidelity than younger females. The mean number of previous breeding attempts per female within a harem (breeding experience) increased significantly with increasing harem size. Breeding females returned to breed later in the breeding season as they aged—we hypothesize that young, subordinate females gain a priority advantage by returning earlier. These results lend support to the hypothesis that there are fitness advantages, in terms of offspring survival, that are conferred to females that breed in successively larger harems with age. Potential mechanisms that select for females to improve their breeding conditions include improved mate selection and the avoidance of conspecific harassment in harems.Communicated by F. Trillmich     

Sexual selection favours small and symmetric males in the polygynous greater sac-winged bat Saccopteryx bilineata (Emballonuridae, Chiroptera)

We investigated how morphological traits of territorial males in the polygynous bat Saccopteryx bilineata were related to their reproductive success. Because of the frequency of aerial courtship displays and defence manoeuvres, and the high energetic costs of flight, we expected small and symmetric males to be better able to court females on the wing and to monopolize copulations with females in their harems. We predicted that small and symmetric males would sire more offspring within the colony and a larger portion of the young born within their harem than large or asymmetric males. We measured size and fluctuating asymmetry of 21 territorial males and analysed their reproductive success in 6 offspring cohorts (n=209 juveniles) using 11 microsatellite loci. As predicted, small and symmetric males had, on average, a higher reproductive success in the colony than large and asymmetric males. The percentage of young sired by males within their harem increased as males decreased in size, but was not influenced by fluctuating asymmetry. As fluctuating asymmetry of males correlated with their reproductive success within the colony but not within their harems, we infer that fluctuating asymmetry is probably related to female choice, whereas male size is probably important for harem defence on the wing.Communicated by G. Wilkinson     

Female aggression and male peace-keeping in a cichlid fish harem: conflict between and within the sexes in Lamprologus ocellatus

Summary Conflicts of interest within and between the sexes are important processes leading to variability in mating systems. The behavioral interactions mediating conflict are little documented. We studied pairs and harems of the snail-shell inhabiting cichlid fish Lamprologus ocellatus in the laboratory. Due to their larger size, males controlled the resource that limited breeding: snail shells. Males were able to choose among females ready to spawn. Females were only accepted if they produced a clutch within a few days of settling. When several females attempted to settle simultaneously the larger female settled first. Females were least aggressive when guarding eggs. Secondary females were more likely to settle when the primary female was guarding eggs. In established harems females continued to be aggressive against each other. The male intervened in about 80% of female aggressive interactions. Male intervention activity correlated with the frequency of aggression among the females in his harem. The male usually attacked the aggressor and chased her back to her own snail shell. When a male was removed from his harem, aggression between females increased immediately and usually the secondary female was expelled by the primary female within a few days. Time to harem break-up was shorter the more mobile the primary females' young were and did not correlate with the size difference between harem females. Male L. ocellatus interfere actively in female conflict and keep the harem together against female interests. Female conflict presumably relates to the cost of sharing male parental investment and to the potential of predation by another female's large juveniles on a female's own small juveniles.Correspondence to: F. Trillmich     

Parental care of nestlings by male red-winged blackbirds

Summary Nestling feeding by males is less common among birds with polygynous mating systems than in monogamous species, because of the pronounced trade-off between parental behavior and the attraction of additional mates. In this study, however, we found that male red-winged blackbirds (Agelaius phoeniceus) commonly assisted females in feeding nestlings in several Ontario marshes. Male parental care was additional to that provided by females, and it significantly enhanced the fledging success of nests (Table 2). Male redwings did not help to feed all nests on their territories: primary and secondary nests were much more likely to receive male parental care than tertiary and later nests. Contrary to expectation, male parental care was not restricted to the nests of primary females: a greater proportion of secondary than primary nests were assisted (Tables 4a and b). The presence of new females settling on the territory at the same time that a resident female had nestlings, resulted in males deferring parental care until a later brood. This suggests that males trade off the recruitment of females against parental care to an existing brood. Although the number of nestlings fledging from a male's territory was strongly influenced by the number of females in the harem, males could additionally increase their reproductive success by feeding nestlings in one or more nests on their territories (Fig. 2). The reproductive success of females was significantly enhanced by male assistance in feeding nestlings (Table 3). However, those females not receiving male assistance on territories of feeding males did not suffer a significantly reduced reproductive success in comparison to females on territories of non-feeding males (Table 2). Males varied considerably in the quality of brood care given. We therefore suggest that the quality of male parental care may be a factor considered by females in choosing a breeding situation.     

Mating system of a Neotropical roost-making bat: the white-throated, round-eared bat, Lophostoma silvicolum (Chiroptera: Phyllostomidae)

The vast majority of bats strongly depend on, but do not make, shelters or roosts. We investigated Lophostoma silvicolum, which roosts in active termite nests excavated by the bats themselves, to study the relationship between roost choice and mating systems. Due to the hardness of the termite nests, roost-making is probably costly in terms of time and energy for these bats. Video-observations and capture data showed that single males excavate nests. Only males in good physical condition attracted females to the resulting roosts. Almost all groups captured from excavated nests were single male-multifemale associations, suggesting a harem structure. Paternity assignments based on ten polymorphic microsatellites, revealed a high reproductive success of 46% by nest-holding males. We suggest that the mating system of L. silvicolum is based on a resource-defense polygyny. The temperatures in the excavated nests are warm and stable, and might provide a suitable shelter for reproductive females. Reproductive success achieved by harem males appears to justify the time and effort required to excavate the nests. Reproductive success may thus have selected on an external male phenotype, the excavated nests, and have contributed to the evolution of an otherwise rare behavior in bats.Communicated by G. Wilkinson     

Positive association between social and extra-pair mating in a polygynous songbird, the dickcissel (Spiza americana)

In polygynous species, males appear to gain additional offspring by pairing with multiple females simultaneously. However, this may not be true if some females copulate outside of the social pair bond. Polygynous males could experience lower paternity because of trade-offs among gaining multiple social mates, guarding fertility with these mates, and pursuing extra-pair matings. Alternatively, polygynous males could simultaneously gain extra social mates and have high paternity, either because of female preferences or because of male competitive attributes. We tested four predictions stemming from these hypotheses in a facultatively polygynous songbird, the dickcissel (Spiza americana). Unlike most previous studies, we found that males with higher social mating success (harem size) also tended to have higher within-pair paternity and that the number of extra-pair young a male sired increased significantly with his social mating success. Females that paired with mated males were not more likely to produce extra-pair young. In contrast, extra-pair paternity was significantly lower in the nests of females whose nesting activity overlapped that of another female on the same territory. This pattern of mating was robust to differences in breeding density. Indeed, breeding density had no effect on either extra-pair mating or on the association between polygyny and paternity. Finally, nest survival increased with harem size. This result, combined with the positive association between polygyny and paternity, contributed to significantly higher realized reproductive success by polygynous male dickcissels.     

Behavioral strategies of American kestrels during mate replacement

Summary To determine the influence of mate replacement on the behavior and reproductive success of wild American kestrels (Falco sparverius) we removed 4 female and 16 male members of breeding pairs during incubation in 1983 and 1984. Eight males and 1 female were replaced within a mean time of 43 h. Widowed females that received a replacement spent less time hunting, but incubated and performed aerial displays more frequently than females that did not receive a replacement. After replacement, widowed females continued to incubate the original clutch, yet copulated and performed other courtship behaviors with the incoming male. Overlapping of normal temporally separate behavioral cues may be a female strategy to gain assistance from a replacement mate. However, none of the original clutches was successfully hatched. Of 8 pairs with replacement males, 2 pairs abandoned their territory, two remained on territory but did not renest, and 4 renested within a mean interval of 18 days. The lone female replacement and her mate copulated and performed nest inspections but then abandoned the territory. Incubation behavior was similar between replacement and control pairs; however, replacement males fed more invertebrate prey to nestling and made many more nest visits. All 3 replacement nests that hatched young failed within 8 days.     

Kinship,reproductive strategies and social dynamics of yellow-bellied marmots

Summary Social behavior of yellow-bellied marmots was observed for three years in colonies where kinship was known and for one year in a high elevation colony where harems were contiguous. Social dynamics of yellow-bellied marmots is dependent on kinship, group composition, and age-sex classes. This pattern is a consequence of the reproductive strategies of males and females. Females behave cohesively toward sisters or daughters, but not with sons and agonistically toward other females. Males generally behave amicably toward females and agonistically toward males, including their sons. Thus, reproductive strategies limit nepotism. This behavior is consistent with a population process in which sons typically disperse as yearlings. At least some of the variation in the expected patterns of social behaviors is attributable to individual differences. Because male and female reproductive strategies differ, a marmot population consists of two social subsystems. The female unit is the closely-related kin group which may also be a burrow group. The male unit is a harem which usually consists of two or more competing female kin groups.     

Social organization and space-use in Gunnison's prairie dog

Summary Social organization of the Gunnison's prairie dog, Cynomys gunnisoni, was studied in two populations in south-central Colorado. Gunnison's prairie dogs live in complex, interactive societies fitting current definitions of highly social ground squirrels. Members of harems (coteries) cooperatively use and defend a common territory. Spatial overlap is extensive between the adult male(s) and adult females, and among adult females within the harem through the active season. Amicable behavioral interactions are frequent within the harem, whereas interactions between members of different harems are primarily agonistic and spatial overlap is minimal. Although their behavioral repertoire is more limited, social organization of the Gunnison's prairie dog most closely resembles that of the black-tailed prairie dog, C. ludovicianus. Although body size, age of first reproduction, and age of emigration differed between the two study populations (Rayor 1985a), a comparison of social traits did not reveal substantial differences.     

Variance in the male reproductive success of western gorillas: acquiring females is just the beginning

Variance in male reproductive success is expected to be high in sexually dimorphic mammals, even when it is modulated by the costs and benefits of group living. Here, we investigate the variance in reproductive success of male western gorillas (Gorilla gorilla), a highly dimorphic primate with long-term male–female associations, using 12.5 years of data collected at Mbeli Bai in northern Congo. Access to mates and offspring survival were both major sources of variance in male reproductive success. Males with larger harems had lower offspring mortality with no apparent reduction in female fertility or observed tenure length, so the size of harems did not seem to be limited by female feeding competition or by the risk of takeovers and infanticide by outsider males. The lower mortality in larger harems may reflect improved vigilance against predators, and females may cluster around males that enhance offspring survival. Thus, this study illustrates how a detailed analysis of the components of male reproductive success can shed light on the interrelated social and ecological aspects that affect it.     

Determinants of male mating success in the red bishop (Euplectes orix)

We studied sexual selection in the red bishop, Euplectes orix, a colonial, polygynous weaverbird widely distributed over sub-Saharan Africa. Male reproductive success measured in terms of the number of nests accepted by females and the number of eggs and nestlings in all the nests on a male's territory varied considerably. The standardized variance (variance/mean²) in male reproductive success ranged from 0.505 to 1.737 in different years, indicating a high potential for sexual selection in this species. An analysis of genetic parentage for 432 nestlings by non-radioactive, multilocus DNA fingerprinting confirmed that male reproductive success (number of young sired on the territory) in this species can be reliably estimated by the measures introduced above. In all 4 study years there was a strong positive correlation between male mating success and the total number of nests that males built in their territories. The number of nests built can be partitioned into the number of weeks a male held a territory and his nest-building performance. Both factors exert a significant positive effect on male mating success and in combination explained between 53.3 and 86.3% of the variation in male reproductive success. Male morphological characters were found to be of no importance. Males that established a territory in the following season built more nests and held their territories for longer than males that did not establish a territory in the following season, suggesting that these measures might be indicators of male condition and quality. Male nest-building performance (number of nests built per week) seems to be unrelated to male condition or quality. Received: 8 January 1999 / Received in revised form: 7 June 1999 / Accepted: 13 June 1999     

Reproductive partitioning in communally breeding guira cuckoos,<Emphasis Type="Italic"> Guira guira</Emphasis>

Guira cuckoos, Guira guira, exhibit a rare polygynandrous reproductive system with groups containing several male and female breeders, allowing for important tests of reproductive skew models. Female reproductive strategies involve leaving the group, varying clutch size, egg ejection and infanticide, among others. Here we examined the predictions of reproductive skew models relative to reproductive partitioning among females in groups. We used yolk protein electrophoresis to identify individual females eggs in joint nests. We found that reproductive partitioning favors early-laying females, which lay and incubate more eggs than females that begin laying later. Because the female that lays first tends to switch between repeated nesting bouts, and females do not always contribute eggs to each bout, female reproductive success tends to equalize within groups over time. The pattern of reproductive partitioning differs from that described for anis, another crotophagine joint-nester. We calculated reproductive skew indices for groups in 2 years, for both laying and incubation, as well as an overall population value. These were compared to random skew generated by simulations. Varying degrees of skew were found for different groups, and also across sequential nesting bouts of the same groups. Overall, however, skew did not deviate from random within the population. Nests that reached incubation tended to have lower skew values during the laying phase than nests terminated due to ejection of all eggs followed by desertion. Groups had higher reproductive skew indices in their first nesting bout of the season, and these nests frequently failed. These results illustrate the importance of social organization in determining not only individual, but group success in reproduction, and highlight the flexibility of vertebrate social behavior.Communicated by J. Dickinson     

Female reproductive strategies in bar-headed geese (Anser indicus): Why are geese monogamous?

Summary In a semi-captive flock of Anser indicus with a surplus of females, permanent harem groups consisting of one male with one to five females, lasting for up to several years, were regularly observed. Polygynous groups contain one paired female to which the male is most attentive and secondary females which follow the paired male and are tolerated by the pair. Average annual reproductive success was lowest in lone females (0.02 young fledged per year), higher in secondary females (0.23 young) and highest in paired females (0.56 young per year). Differences seemed due to different degrees of male assistance. Secondary females could not be shown to be competitors of paired females in annual reproductive success. Lone females became secondary females mainly after an age of 3 years, i.e. when their chances to pair had dropped significantly. Females were more likely to become secondary instead of paired females in years when the adult sex ratio was more heavily female biased. As sex ratios in wild geese are usually around 1:1 or even biased towards males, females will not usually need to resort to the suboptimal secondary-female strategy. Hence, geese usually live in monogamous pairs instead of harem groups.     

Are female boat-tailed grackle colonies neutral assemblages?

Female boat-tailed grackles (Quiscalus major) nest in colonies, and apparently settle independently of male activities. Associations between colony females may be mutually beneficial (cooperative hypothesis) or females may be penalized by associating (competitive hypothesis). Contrary to predictions based on either cooperative or competitive behavior, (1) reproductive success was not related to colony size nor to internest distance; (2) intracolony nesting synchronies were the same as those of the whole population, and within colonies, there was no relationship between the timing and spacing of nearest-neighbor nests; and (3) already established females were only occasionally aggressive toward females attempting to settle near them, and their aggressive response was independent of their stage of nesting. The results suggest that females act independently of each other and do not affect each other's fitness (neutral female hypothesis). Colonies may be neutral aggregations in sites secure from ground predators. Although females benefit by co-occupying predator-free sites, advantages and disadvantages of colomality do not appear to be related to intrasexual association.     

Alternative male mating tactics in a cichlid, Pelvicachromis pulcher: a comparison of reproductive effort and success

Pelvicachromis pulcher is a small African cichlid which breeds in holes. Males may either reproduce monogamously (pair males), polygynously (harem males), or be tolerated as helpers in a harem territory (satellite males). These helpers share in defence of the territory against conspecifics, heterospecific competitors and predators. There are two male colour morphs that are fixed for life and are apparently genetically determined. These differ in their potential mating strategy. Red morph males may become harem owners, while yellow morph males may become satellite males, and males of both morphs may alternatively pair up monogamously. We compared the reproductive effort and success of these three male reproductive strategies. Effort was measured as attack rates, time expenditure and the risk of being injured or killed when attacking competitors or predators of three sympatric fish species. Reproductive success was measured by observing how many eggs were fertilized by each male when this was possible, and by using genetic markers. The number of fry surviving to independence of parental care was used as a criterion of success. The reproductive success of harem males was 3.3 times higher than that of pair males and 7 times higher than that of the average satellite male. Dominant satellite males, however, were as successful as monogamous pair males, using the measure of fertilized eggs. To our knowledge, this has not been found previously in any fish species. Both harem and pair males had lower parental defence costs per sired offspring, however, than males using the alternative satellite tactic. Defence effort was significantly related to the risk of injury. Received: 17 January 1996 / Accepted after revision: 9 June 1997     

Pair and extra-pair mating success relative to male quality in red-winged blackbirds

We tested whether the reproductive success of male red-winged blackbirds (Agelaius phoeniceus) varied with male secondary sexual traits or with haematozoa prevalence, and whether these patterns were consistent with females preferring genetically superior males. We also determined whether the traits that correlated with male success on their own territories were also correlated with male success at siring young on other males territories. Our analysis included data from a 6-year study involving 617 nestlings for which paternity was determined by DNA profiling. Larger males sired more young on their territories, principally because they obtained larger harems. The success of larger males at acquiring more mates did not appear to be a consequence of larger males holding larger or better-quality territories. Older and longer-lived males sired more young by extra-pair fertilizations. Larger males sired the most offspring overall (on territory + off territory). Variation in epaulet size and color, responses to male and female models, nest defence and parasitism was not correlated with male success either on or off their territories or overall. Male success in a given year was significantly correlated with success the previous year, as expected if females were selecting genetically superior males. The male that was by far the most successful individual in this study was highly consistent from year to year. Because male body size is positively correlated with survival in this population (although not within the sample of males included in this study), female preference for larger males may have reflected a general preference for males with superior Survival ability. We propose that the direct advantage realized by older males in extra-pair matings might indicate that experience is important, such that experienced males are better at creating or exploiting the opportunities for extra-pair mating. This hypothesis is consistent with a pairwise analysis of cuckoldry that showed that cuckolders were most often older than the males they cuckolded.     

Resource quality or harem size: what influences male tenure at refuge sites in tree weta (Orthoptera: Anostostomatidae)?

Rather than seeking females directly, males in many animal species locate and defend sites that contain spatially limited resources essential for female survival and reproduction. In these cases, resident males successfully repelling conspecific rivals will mate with sexually receptive females that seek to use the resident’s resources. Theory predicts that if resources are limiting in nature, are costly to procure, and if residency at the resource site increases male reproductive success, each site should be monopolised by a single adult male. Moreover, if females aggregate at these sites, males should be sedentary and monopolise harems. This predicts that males should reside at resource sites longer than females and male tenure should be positively correlated with harem size. I address these hypotheses using a wild population of the Wellington tree weta, Hemideina crassidens, a sexually dimorphic insect in which males use mandibular weapons in fights to control resource sites (galleries in trees) required by females for diurnal refuge. In a longitudinal study using artificial galleries, I show that male tenure in a gallery is positively related to harem size and, contrary to prediction, not gallery size per se. Moreover, contrary to prediction, females are more sedentary than males: males appear to move frequently between galleries to search for novel mates. I show experimentally that galleries represent mating sites to males: males prefer galleries housing adult females whereas females prefer unoccupied galleries. Females likely avoid male-occupied galleries because they incur injuries when interacting with males.     

Aggression in female Red-Winged Blackbirds: A strategy to ensure male parental investment

Summary Female Red-winged Blackbirds (Agelaius phoeniceus) are often aggressive towards conspecific females during the breeding season. We hypothesize that the function of female-female aggression in this species is to guard the nonshareable portion of the male's parental investment.The investment-guarding hypothesis predicts that a female should be more aggressive toward another female evincing interest in mating with the territory-owning male than toward a female simply perching within the male's territory. Results of mount presentations to females with active nests confirmed this prediction. Nesting females attacked a stuffed conspecific female mounted in a precopulatory, soliciting posture significantly more often than a mount in a normal, perched posture.The male's nonshereable parental care consists of provisioning his young, and most of this care is invested in the brood of his primary (first-to-nest) female. It is therefore predicted that primary females should be more aggressive than secondary (later-nesting) females. Female mount presentations also confirmed this prediction. Primary females attacked the soliciting mount significantly more often than secondary females.     

Mate choice in the yellow-shouldered widowbird: correlates of male attractiveness

Recent investigations of male ornaments in sexual selection have used experimental manipulation of tail length in three widowbird species, but only for one of these have correlates of male reproductive success been reported. I examined correlates of male attractiveness to nesting females over two breeding seasons for the polygynous yellow-shouldered widowbird, Euplectes macrourus, in order to discover which cues females may be using to select mates. The black, long-tailed ( 10 cm) males defend large territories and build nest frames, or cock's nests, which females then line and use for nesting. I examined various aspects of male morphology, five behavioral displays, territory characteristics, and the number of cock's nests that males built. Few correlates of mating success were found. The best predictor was the number of cock's nests that a male builds, though one courtship display also correlated with male mating success in 1 year, as did average grass height. Tail length did not correlate with male mating success. A partial correlation analysis confirmed that cock's nests and, in 1 year, grass height, were the primary contributers to male success. Females may choose where to nest primarily on the availability of suitable nesting sites. Long tails may be used by females seeking extra-pair copulations or in male-male competition for territories.