Alternative reproductive tactics in male common shrews: relationships between mate-searching behaviour,sperm production,and reproductive success as revealed by DNA fingerprinting

The common shrew (Sorex araneus) is a solitary small mammal with a promiscuous mating system. Previous studies of this species suggest that females typically mate multiply, and that males may adopt alternative mate-searching tactics. We studied two generations of common shrews in a population near Oxford, England. Males were found to adopt two different mate-searching tactics. Those classed as type A occupied relatively small exclusive ranges during March, and made repeated long-distance movements to visit female ranges around the time of first oestrus in April. Males classed as type B established large overlapping ranges in areas of relatively high female density during March, and maintained these ranges throughout April. Type B males were larger than type A males at an early stage of sexual maturation, but there was no difference in the adult body size of the two types of male. Type A males had significantly higher epididymal sperm counts than type B males. Paternity analyses of litters born during the first year of the study reveal that the mean number of offspring fathered by type B males was greater than the mean number fathered by type A males. It is concluded that different mate-searching tactics may be conditional upon the timing of sexual maturation. Differences in sperm production are discussed in relation to sperm competition theory.     

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.     

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     

Egg mimicry and allopaternal care: two mate-attracting tactics by which nesting striped darter (Etheostoma virgatum) males enhance reproductive success

In a variety of fish species with paternal care of offspring, females prefer to spawn in nests that already contain eggs. This female preference has been hypothesized to explain egg thievery in male sticklebacks, allopaternal care of eggs in minnows, and the evolution of egg-mimicking body features in male cichlids and darters. Here we employ microsatellite-based parentage analyses to evaluate the reproductive success of striped darter (Etheostoma virgatum) males that appear to utilize two of these functionally related tactics to entice females to spawn in their nests. In an isolated population (Clear Creek, Ky.), we observed that breeding males develop conspicuous white spots on their pectoral fins. If these spots are egg mimics, as we suspect, then this represents the fourth independent evolutionary origin of egg mimicry documented to date in darters, the first based on pigmentation (as opposed to physical structures), and the first in which the egg mimics vary greatly in number among males. From direct counts of microsatellite genotypes in clutches of embryos, at least 3.8 females contributed to the progeny within a typical nest, and females tended to spawn preferentially with males that were larger and displayed more egg-mimic spots. In another population (Hurricane Creek, Tenn.) without egg mimics, the multi-locus genetic data document that allopaternal care is common, especially among the smallest males who sometimes tend nests containing their own as well as an earlier sire's offspring. Thus, these foster males had adopted egg-containing nests and then successfully spawned with subsequent females. Overall, the genetic data on paternity and maternity, in conjunction with field observations, suggest that egg mimicry and allopaternal care are two mate-attracting reproductive tactics employed by striped darter males to exploit female preferences for spawning in nests with 'eggs'. Electronic Publication     

Biased parental investment and reproductive success in Gabbra pastoralists

Demographic data from 848 Gabbra households are used to examine the relationships between herd size and reproductive success in relation to sex, in a traditional, pastoralist population. The number of camels in the household herd has a significant positive effect on the reproductive success of both men and women, although the effect of wealth is greater for men, as predicted from evolutionary theory. The greater the number of elder brothers a man has, the lower his reproductive success, as a result of a smaller initial herd and a later age at marriage. This is not true for women –number of elder sisters does not have a measurable effect on a woman’s fertility, although it does have a small, negative effect on the size of her dowry. These results are interpreted as competition between same-sex siblings for parental investment, in the form of their father’s herd, which is more intense between sons than daughters as parental investments are greatest in males. Received: 30 June 1995/Accepted after revision: 23 October 1995     

Differences in reproductive success between laboratory and wild-derived golden hamsters (Mesocricetus auratus) as a consequence of inbreeding

All laboratory golden hamsters (Mesocricetus auratus) originated from a sibling pairing back in 1930. Due to this extreme founder event, domestic golden hamsters are presumed to be one of the most bottlenecked animal populations. Nevertheless, domestic hamsters show no obvious signs of inbreeding depression in commonly used breeding stocks. To explore the existence of potentially masked inbreeding effects, we compared the reproductive success of laboratory (lab) and wild-derived (wild) golden hamsters. We allowed oestrus females to mate consecutively with lab and wild males. The resulting offspring was genotyped using microsatellites to assess paternity. Finally, we compared male reproductive success to genetic variability, sexual behaviour and different sperm characteristics. Both hamster strains exhibited the expected large difference in genetic diversity (H _wild =0.712±0.062 vs H _lab =0.007±0.007. The reproductive success of wild males dramatically exceeded that of lab males (87% of pups were sired by wild males). Sexual behaviour of wild and lab males only varied in the number of long intromissions (intromissions without ejaculation at the end of the mating). No significant differences were observed in relation to mounting, ejaculation and intromission. There were also no apparent differences in sperm motility, velocity and density or testis histology between wild and lab hamsters. We conclude that the reduced reproductive success of lab males represents a hidden inbreeding effect, although its precise physiological cause remains unclear. These results provide first evidence for a major fitness disadvantage in captive golden hamsters.     

Racketed tail of the male and female turquoise-browed motmot: male but not female tail length correlates with pairing success,performance, and reproductive success

Both males and females of many avian species maintain elaborate plumage traits, and elaborate monomorphic plumage may convey adaptive benefits to one or both sexes as inter- or intraspecific signals. Both sexes of the turquoise-browed motmot (Eumomota superciliosa) are elaborately plumed with long racket-tipped tail. I investigated whether the racketed tail functions as a sexually selected signal in one or both sexes by testing the predictions that males and/or females with the largest tails have: (1) greater pairing success, (2) greater reproductive performance (clutch-initiation date, clutch size, and hatching success), and (3) greater reproductive success. Yearling males with longer denuded rachises (wires) on the central tail feathers had greater pairing success. In addition, adult males with longer wires paired with females who laid larger clutches, had greater hatching success independent of clutch size, and fledged more young. There was no relationship between female tail plumage and pairing success, reproductive performance, or fledgling success. These results are consistent with the hypothesis that male tail plumage functions as a mate choice or status signal, but that the tail of the female does not function in a sexually selected context. I discuss alternative hypotheses for the evolutionary maintenance of the elaborate female tail plumage.     

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     

Intraspecific brood parasitism in the moorhen: parentage and parasite-host relationships determined by DNA fingerprinting

Parasitic female moorhens (Gallinula chloropus) lay from one to six eggs in the nests of conspecific neighbours. DNA fingerprinting was used to show that parasitic eggs could be correctly identified when they appeared in addition to or outside the host’s laying sequence. Moorhen hosts accept all parasitic eggs laid after the 2nd day of their laying period. To understand why moorhen hosts tolerate parasitic eggs, we tested two hypotheses. (1) The quasi-parasitism hypothesis: females lay their eggs in the evening when the host males are normally in attendance at the nest, so host males may allow parasitic females to lay in their nests in exchange for fertilizing their eggs. However, DNA fingerprinting showed that all the parasitic eggs were sired by the parasites’ mates. Parasitic moorhens frequently continue laying a clutch in their own nest, without a break in the laying sequence after a parasitic laying bout. The eggs laid by brood parasites in their own nests were also sired by their own mates. Therefore this hypothesis was rejected. (2) The kin selection hypothesis: if one or both members of the host pair are close relatives of the parasite, the costs of rearing parasitic chicks will be to some degree offset by inclusive fitness benefits. We examined the genetic relationships between parasites and their hosts using DNA fingerprinting and genealogical data. Natal philopatry by both sexes was relatively common in this population, and the probability that a neighbour of either sex was a first-order relative (parent-offspring) was calculated as 0.18. Although first-order relatives were not preferentially chosen as hosts over individuals that were not first-order relatives, even through random host selection there is almost a one-in-five chance that brood parasites in this population are closely related to their hosts. This may facilitate host tolerance of parasitic eggs. Other hypotheses are also discussed. Received: 3 February 1995/Accepted after revision: 27 August 1995     

Mating system and individual reproductive success of sympatric anadromous and resident brook charr, <Emphasis Type="Italic">Salvelinus fontinalis</Emphasis>, under natural conditions

Salmonids are known for the occurrence in sympatry of two life-history forms, one that undergoes migration to sea before returning to freshwater to reproduce (anadromous) and one that inhabits freshwater without a migration phase (resident). Whereas one breeding population is often suggested by population genetic studies, mating patterns have rarely been directly assessed, especially when both sexes are found within each life-history form. By using highly polymorphic microsatellite loci and parentage analysis in a natural population of sympatric anadromous and resident brook charr (Salvelinus fontinalis), we found that gene flow occurred between the two forms and was mediated by resident males mating with both resident and anadromous females. Determinants of reproductive success, estimated by the number of surviving juveniles (ages 1 and 2 years), differed between the sexes. No strong evidence of the influence of size on individual reproductive success was found for males, whereas larger females (and hence most likely to be anadromous) were more successful. The higher individual reproductive success of anadromous fish compared to residents was mainly explained by this higher reproductive success of anadromous females. We suggest that resident males adopt a “sneaking” reproductive tactic as a way of increasing their reproductive success by mating with females of all sizes in all habitats. The persistence of the resident tactic among females may be linked to their advantage in accessing spatially constrained spawning areas in small tributary streams unavailable to larger females.     

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     

Female control of reproductive skew in cooperatively breeding brown jays (<Emphasis Type="Italic">Cyanocorax morio</Emphasis>)

Brown jays (Cyanocorax morio) are long-lived, social corvids that live in large, stable, territorial groups (mean = 10 individuals). In this study, I determined the distribution of reproductive success within groups using multi-locus DNA fingerprinting. Breeding females produced virtually all (99%) of the young within their own nests. Reproduction within groups was highly skewed towards a single primary female, although long term data indicate that secondary females (female breeders that were usually younger and subordinate to the primary female) were sometimes successful. The high reproductive skew observed for females was associated with primary female aggression. Successful reproduction by secondary females may have been due to parental facilitation or the inability of primary females to completely suppress secondary females. Multiple paternity occurred in 31–43% of broods and extra-group paternity occurred in a minimum of 22% of broods. Patterns of paternity also varied between years, since females often switched or included new genetic mates. Although male consorts of nesting females fathered relatively few offspring (20%), they still had a higher chance of fathering offspring than any other single group male. Reproduction was less skewed for males than females as a result of female mating patterns. Female reproductive patterns are consistent with some of the predictions and assumptions from optimal skew models, while male reproductive patterns are not. The factors affecting skew in species with complex social systems such as incomplete control by breeders over subordinate reproduction, female control of paternity, and resource inheritance have not been well incorporated into reproductive skew models.Communicated by: J. Dickinson     

Female dispersal and reproductive success in wild western lowland gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>)

This paper presents data on the dispersal patterns and reproductive success of western lowland gorilla females from a long-term study at Mbeli Bai in the Nouabalé-Ndoki National Park, Republic of Congo. We find that female natal and secondary transfer is common. Female immigration rates are negatively related to group size, and emigration rates are positively related to group size, with the net result that larger groups are losing females and smaller groups are gaining females. Furthermore, females transferring between known groups show a preference for significantly smaller groups. However, there is no effect of group size on female reproductive success. Male protection and male quality are considered important in determining female transfer decisions. The case for infanticide is argued and females exhibit strategies that appear to minimise the probability of infanticide following the death of the silverback. Exclusively single-male groups and group formation through female acquisition by solitary males may bias female transfer to lone silverbacks and small groups. The effects of group size on female dispersal and reproductive success are not wholly consistent with an argument for increased foraging costs, and group size effects are more parsimoniously explained by demographic factors. Male protection from intra-group aggression is the most likely factor underlying grouping patterns across gorilla taxa, but differences in population structure and male reproductive strategies may account for inter-specific variation. We stress the need for intra-specific comparisons and more complete data sets on western lowland gorilla feeding behaviour.Communicated by S. Boinski     

Mating system of the cooperatively breeding noisy miner Manorina melanocephala,as revealed by DNA profiling

In this study we examined parentage within broods of the cooperatively breeding noisy miner using multi-locus DNA profiling. Previous studies of noisy miners described them as highly promiscuous, leading to the suggestion that promiscuous mating behaviour was a tactic used by females to recruit males as provisioners to their nests (Dow 1978). At our study site in south-east Queensland, Australia, we found that both multiple and extra-group paternity (i.e. a female mating with a male outside the group of male provisioners at her nest) were rare. In nests where multiple paternity was possible (i.e. clutch size > 1) 97% of 31 broods were sired by only a single male. Overall, 96.5% of all nestlings (n = 85) were the result of monogamous matings. Also, at the vast majority of nests, the male that sired the nestlings was also the main provisioner among all male nest attendants. Our results show that the mating system of the noisy miner can no longer be considered cooperative polyandry (Brown 1987) but is clearly genetic monogamy. We discuss the implications of this finding for understanding the complex social system of this species.     

Effects of paternal care on reproductive success in the polygynous spotless starling Sturnus unicolor

For males of socially polygynous avian species like the spotless starling, there may exist a trade-off between investing in paternal care and controlling several nests. To determine how the intensity of paternal care affects reproductive success per brood sired or expressed as the total number of young raised in all nests controlled by the same male, it is necessary to manipulate paternal care. Testosterone (T) has been shown to depress the tendency for males to care for their young, and induces them to acquire more mates. The effects of paternal care on reproductive success were studied by treating certain male starlings with exogenous T and others with the antiandrogen cyproterone acetate (CA), and comparing the parental behavior of T- and CA-males throughout the breeding season with that of controls. CA-males fed their chicks more during the first week after hatching than T-males, with controls feeding at intermediate rates, both on a per nest basis and as total effort for all nests controlled by the same male. Paternal feeding rates during the first week of chick life had a significant positive effect on the number of fledged young. The hormone treatment significantly affected the number of chicks raised per nest, CA-males having a higher breeding success per nest than T-males, and controls showing intermediate levels of success. There was no significant effect of treatment on total reproductive success attained by males throughout the season. In the polygonous spotless starling, the intensity of paternal care of young affects reproductive success per nest positively but not on a seasonal basis. Received: 6 February 1999 / Received in revised form: 30 June 1999 / Accepted: 11 July 1999     

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     

Cooperative breeding increases reproductive success in the social spider <Emphasis Type="Italic">Stegodyphus dumicola</Emphasis> (Araneae,Eresidae)

Sociality in some birds, mammals, and social insects was suggested to have evolved through the lengthening and extension of parental care behaviors to nondirect descendents. In these systems, group members care for young cooperatively and, thus, increase the reproductive success of the breeders and fitness of the young. Parental care behaviors, such as regurgitation feeding and matriphagy (consumption of the mother), occur in several subsocial and social spiders. However, it is not known whether females in a colony cooperate in caring for the young of other females and whether such cooperative care improves reproductive success. To answer this question, we created experimental colonies of the social spider Stegodyphus dumicola (Araneae, Eresidae), allowing only one female in a group to produce young, simulating reproductive skew occurring in nests in nature. In this paper, we show for the first time that females of S. dumicola cooperate in providing regurgitated food for young of other females and are even eaten by those young. Young raised by a group of females were larger and had greater survival than young raised only by their mother. Thus, fitness benefits from raising broods cooperatively may have favored the evolution of sociality in spiders.     

Determinants of male reproductive success in wild long-tailed macaques (Macaca fascicularis)—male monopolisation, female mate choice or post-copulatory mechanisms?

One of the basic principles of sexual selection is that male reproductive success should be skewed towards strong males in species with anisogamous sex. Studies on primate multi-male groups, however, suggest that other factors than male fighting ability might also affect male reproductive success. The proximate mechanisms leading to paternity in multi-male primate groups still remain largely unknown since in most primate studies mating rather than reproductive success is measured. Furthermore, little research focuses on a female’s fertile phase. The aim of this study was to investigate the relative importance of male monopolisation and female direct mate choice for paternity determination. We also investigated the extent to which paternity was decided post-copulatory, i.e. within the female reproductive tract. We used a combined approach of behavioural observations with faecal hormone and genetic analysis for assessment of female cycle stage and paternity, respectively. The study was carried out on a group of wild long-tailed macaques living around the Ketambe Research Station, Gunung Leuser National Park, Indonesia. Our results suggest that both male monopolisation and post-copulatory mechanisms are the main determinants of male reproductive success, whereas female direct mate choice and alternative male reproductive strategies appear to be of little importance in this respect. Female cooperation may, however, have facilitated male monopolisation. Since paternity was restricted to alpha and beta males even when females mated with several males during the fertile phase, it seems that not only male monopolisation but also post-copulatory mechanisms may operate in favour of high-ranking males in long-tailed macaques, thus reinforcing the reproductive skew in this species.     

Solitary nesting and reproductive success in the paper wasp Polistes aurifer

Female Polistes paper wasps are capable of independent nesting, yet many populations demonstrate a mixture of solitary and cooperative nest foundation. Previous studies of Polistes have found survival and/or productivity advantages of cooperative nest foundation compared to solitary nesting, and reproductive skew models have been designed to predict the dynamics of such flexible cooperation. In this paper, we examine the success of different nesting strategies in a previously unstudied population of Polistes aurifer in southern California. The colony cycle of this population is less synchronous than that of other temperate species, and the frequency of solitary nesting averages 86.2%. Our data suggest that this low rate of cooperative nest founding is adaptive, as demonstrated by the lack of survival or productivity advantages for cooperative foundress associations. Due to foundress turnover and nest foundation later in the season, many nests produce only one set of offspring. This results in a loss of the eusocial nature of some nests in the population. Data from a small sample of multifoundress nests show significant positive reproductive skew, despite concession model predictions that skew should be low in populations with low ecological constraints on independent nesting. This lack of support for the concessions skew model reflects a diminished incentive for cooperation.Communicated by L. Keller