Male Dominance Rank,Female Mate Choice and Male Mating and Reproductive Success in Captive Chimpanzees

Mating and consequently reproductive success in male vertebrates are predominantly determined by intermale competition and female mate choice. Their relative importance however, is still poorly understood. We investigated the interrelationship between male dominance rank — a formal indicator of male competitive ability — female mate choice, and male mating success in a multimale-multifemale group of captive chimpanzees. In addition, we examined the relationship between male dominance rank and reproductive success determined by genetic paternity analysis over a 13-yr period in the same captive population. We related the frequencies of sociosexual behaviors to the female anogenital swelling stage and female fertile phase as determined by urinary and fecal progestogen analysis. Rates of behaviors in both sexes increased with increasing intensity of female swelling, but they were not influenced by the timing of the fertile phase. Male mating success was clearly related to male dominance rank, with high-ranking males performing the overwhelming majority of copulations. This was mainly due to both rank-related rates of male soliciting behavior and intermale aggressiveness during the period of well-developed female anogenital swelling. Although females solicited copulations mainly from the high-ranking males and thus expressed a mate choice based on rank, their overall contribution in initiating copulations and thus influencing male mating success was low. The data on paternity from the population, which always contained 4 adult males, revealed that -males sired the majority (65%) of offspring. We conclude, that male dominance rank is an important determinant of male mating and reproductive success in captive (and presumably wild) chimpanzees and that female mate choice is of minor importance in modulating male reproductive outcome.     

The Effect of Female Quality on Male Ejaculatory Expenditure and Reproductive Success in a Praying Mantid

Strategic ejaculation is a behavioural strategy shown by many animals as a response to sperm competition and/or as a potential mechanism of cryptic male choice. Males invest more mating resources when the risk of sperm competition increases or they invest more in high quality females to maximize their reproductive output. We tested this hypothesis in the false garden mantid Pseudomantis albofimbriata, where females are capable of multiply mating and body condition is an indicator of potential reproductive fitness. We predicted male mantids would ejaculate strategically by allocating more sperm to high quality females. To determine if and how males alter their ejaculate in response to mate quality, we manipulated female food quantity so that females were either in good condition with many eggs (i.e. high quality) or poor condition with few eggs (i.e. low quality). Half of the females from each treatment were used in mating trials in which transferred sperm was counted before fertilisation occurred and the other half of females were used in mating trials where fertilisation occurred and ootheca mass and total eggs in the ootheca were recorded. Opposed to our predictions, the total number of sperm and the proportion of viable sperm transferred did not vary significantly between female treatments. Male reproductive success was entirely dependent on female quality/fecundity, rather than on the number of sperm transferred. These results suggest that female quality is not a major factor influencing postcopulatory male mating strategies in P. albofimbriata, and that sperm number has little effect on male reproductive success in a single mating scenario.     

Female Choice for Male Morphological Traits in House Sparrows,Passer domesticus

Female mate choice for male morphological traits was examined in captive house sparrows in New Mexico, USA. A study of house sparrows in Denmark indicated that male bib size is important in female choice (Moller 1988, 1989). To examine this issue further, and to compare the selective pressures on male morphological traits between the two populations, bibs of males were manipulated to one of four standard sizes and females were allowed to choose from four males, one of each bib size. The amount of time the female spent with each male was recorded for 40 trials. Males were then ranked in each trial, with a rank of 4 given to the male the female spent the most time with. The ranks a male obtained were then averaged to provide an indication of a male's attractiveness to females. Data from 23 different males indicated that bib size was not correlated with a male's average rank, even when the effects of other traits were controlled statistically. In addition, the size of the original (unaltered) bib was not correlated with male average rank. Bill depth was the only trait significantly correlated with a male's average rank. Differences in the populations studied or the methodologies used may have led to differences between the results obtained in this study and that of Moller (1988).     

Male Incubation and its Effect on Reproductive Success in the Black Swan,Cygnus atratus

Male incubation is very unusual among Anatidae. In the black swan, males spend more time incubating the clutch than females do. We investigated the incubation patterns of males and females in a breeding colony in Vienna over a one-year period. Breeding occurred year round, with a peak in winter. Inattentive periods during incubation were shorter in winter than at other times of the year, probably because the heat loss of eggs is then higher. We tested the importance of the costs of incubation by prolonging the breeding period experimentally. Eggs were exchanged for dummies, which were readily accepted and incubated for up to 342 days. The higher the incubation effort of females, measured as the product of their proportion of incubation and the total length of the incubation phase, the longer was the interval between the end of incubation and laying of a subsequent clutch. This suggests that the incubation effort of females is limiting the reproductive output of a pair. By taking charge of the greater part of incubation, males may increase the productivity of their female partners and thereby their own reproductive success. This amplification of female fecundity is a factor favouring a high share in male incubation which is discussed in polyandrous species (e.g. Jenni 1974) but generally overlooked in monogamous systems. The ultimate cause for male incubation in black swans may lie in the combination of a monogamous mating system and the capability of this species to breed whenever the environmental conditions turn favourable, at any time of the year.     

Costs and Benefits of Competitive Traits in Females: Aggression,Maternal Care and Reproductive Success

Recent research has shown that female expression of competitive traits can be advantageous, providing greater access to limited reproductive resources. In males increased competitive trait expression often comes at a cost, e.g. trading off with parental effort. However, it is currently unclear whether, and to what extent, females also face such tradeoffs, whether the costs associated with that tradeoff overwhelm the potential benefits of resource acquisition, and how environmental factors might alter those relationships. To address this gap, we examine the relationships between aggression, maternal effort, offspring quality and reproductive success in a common songbird, the dark-eyed junco (Junco hyemalis), over two breeding seasons. We found that compared to less aggressive females, more aggressive females spent less time brooding nestlings, but fed nestlings more frequently. In the year with better breeding conditions, more aggressive females produced smaller eggs and lighter hatchlings, but in the year with poorer breeding conditions they produced larger eggs and achieved greater nest success. There was no relationship between aggression and nestling mass after hatch day in either year. These findings suggest that though females appear to tradeoff competitive ability with some forms of maternal care, the costs may be less than previously thought. Further, the observed year effects suggest that costs and benefits vary according to environmental variables, which may help to account for variation in the level of trait expression.     

Predicting Current Reproductive Success of Goose Pairs Anser indicus from Male and Female Reproductive History

A multiple regression analysis of current reproductive success (goslings fledged) on three male and female life-history traits (age, previous breeding periods, previous successes) was carried out for 31 semicaptive pairs of barheaded geese Anser indicus. Of the female variables, age proved positively and the number of previous nesting periods negatively related to current success. The latter relationship suggests that incubation has costs in terms of future fecundity for the female. Of the male variables, age was negatively and the number of previous fledging successes positively related to current reproductive success. Attempts with additional data to explain these findings in the male either by a positive feedback of success on future success or by differences in male quality gave inconclusive results, as did earlier attempts to demonstrate reproductive costs in geese and swans. In monogamous species with long-term pair bonds, where male and female share most of their life history but specialize in different activities, reproduction may affect the sexes in different ways. If the correlations between life-history traits of mates are not accounted for, individual strategies or constraints may be obscured by opposing effects in the mate.     

Male House Finches with Elaborate Songs have Higher Reproductive Performance

The elaborate songs of male animals are thought to function in either territory defense (male–male communication) or mate attraction (male–female communication). In non‐territorial animals, male vocalizations are expected to function primarily in mate attraction, yet the reproductive consequences of male vocalizations in non‐territorial animals are poorly described. Here we explore the relationship between male song and male reproductive performance in a free‐living population of house finches, Carpodacus mexicanus, a non‐migratory, non‐territorial songbird. Based on recordings of 20 males, we analyzed three song features (song length, number of unique syllables per song, and song rate) and compared male song with two measures of within‐pair reproductive performance (nest initiation date and clutch size) and one measure of extra‐pair reproductive performance (whether males sired extra‐pair young). We demonstrate a positive association between male song and within‐pair reproductive performance; males that sang long songs initiated their first clutch significantly earlier and males that sang songs at a faster rate had larger clutches. Despite the fact that only one of our recorded males sired extra‐pair young in the nest of another male, this male's songs were the most elaborate for two of three song features measured, anecdotally suggesting that male song may play a role in both within‐pair and extra‐pair partner choice. These results suggest that male song is a sexually selected trait in non‐territorial house finches.     

Hatching asynchrony and maternal androgens in egg yolks of House Wrens

Synchronously and asynchronously hatching clutches of House Wrens Troglodytes aedon usually do not differ in reproductive success. Thus late-hatching nestlings in asynchronously hatching clutches somehow overcome any age- and size-related disadvantages of hatching after their nest-mates. One possible way for them to do this is for female House Wrens to add maternal androgens to the yolk of late-hatching eggs. We tested this hypothesis in a wild population of House Wrens that produces both asynchronously and synchronously hatching clutches. Yolks of eggs from both types of clutches were biopsied and the eggs returned to their nests to hatch. Radioimmunoassays revealed that total androgen levels in the yolk varied within and among clutches. However, total androgen levels in yolks did not vary predictably with egg position in either synchronously or asynchronously hatching clutches. Thus, deposition of androgens in yolk did not follow the expected pattern based on the potential for sibling competition in House Wrens.     

Male Reproductive Success in a Promiscuous Armoured Catfish Corydoras Aeneus (Callichthyidae)

Among a variety of fish mating systems, promiscuity with random-mating seems to be most prevalent. However, detailed studies of promiscuity have been rare due partly to the peculiar difficulty in examination of male mating and reproductive success in the random mating. Females of the armoured catfish Corydoras aeneus (no sexual dimorphism other than size of males > females) spawn 10–20 egg-clutches with multiple males at a time, but an entire egg clutch is inseminated by sperm of a single male. We studied mating system of this fish in aquarium. Males had neither mating territories nor monopolized females, never being aggressive against rival males. Evidence of female preference for certain male traits including size was not detected. Females mated a male in proportion to his relative courtship frequency among males. Courtship frequency was not related to male size, and male mating success was not different between small and large males. Clutch size and insemination rate were different neither between small and large males nor between frequently and less frequently courting males. Thus, the male reproductive success will not be related to the male size, but directly to courtship frequency, indicating the random mating in this fish. There seemed to be fecundity advantage with size in female, and the consequent sexual difference in energy allocation will be responsible to the sexual dimorphism. We also discuss the low male-GSI in this promiscuous fish in which sperm competition hardly occurred.     

Social Dominance among Male Meadow Voles is Inversely Related to Reproductive Success

Intrasexual selection can occur through direct aggressive interactions between males for access to females. We tested the relationship between social dominance and male reproductive success among meadow voles, Microtus pennsylvanicus. Dominance ranks of wild‐caught males were determined using neutral arena trials, with the winner of two of three trials considered dominant. These males were then released into field enclosures and allowed to visit females housed in nestboxes for 8 wk, and males’ home range sizes were determined using weekly grid trapping. Male reproductive success was determined using molecular paternity analysis (six microsatellite primers) for all pups born during the field experiment. Males with higher dominance ranks had larger home ranges. However, male dominance rank was not predictive of the number of total visits to females’ nestboxes or the number of visits to each male's most frequently visited nestbox. Males that made more visits to nestboxes sired more litters. Males that had higher dominance ranks sired fewer litters. These results suggest that there is a reproductive disadvantage to having higher dominance rank among male meadow voles.     

Female Preferences for Call Traits and Male Mating Success in the Neotropical Frog Physalaemus enesefae

Female preferences for male call traits may affect male mating success and the evolution of exaggerated secondary sexual traits. We used phonotaxis experiments to examine female preferences in the frog Physalaemus enesefae in relation to variation in male call duration, dominant frequency, intercall interval and amplitude (dB SPL). Females preferred long calls, low and average dominant frequency calls, short intercall intervals and more intense calls. We compared the patterns of female preferences with those of acoustic variation among males to test the prediction that properties with low within‐male variation are associated with stabilizing or weakly directional female preferences, whereas properties with high within‐male variation are associated with directional preferences. Females had weakly directional preferences for the dominant frequency of the call and strongly directional preferences for call duration and call rate. We also determined whether the temporal relationship between calls influenced preferences based on the dominant frequency of the call. Preferences for low‐frequency over high‐frequency calls disappeared when calls partially overlapped. Females preferred the leading call regardless of its dominant frequency. We also investigated mating patterns in the field. There was size‐assortative mating, as male and female body sizes snout‐vent length (SVL) were positively correlated. In addition, differences in the frequency distributions of body length (SVL) between mated and unmated males approached significance; lower SVL classes were underrepresented among mated males. These patterns may reflect female preferences for lower dominant frequency calls, as there is a negative correlation between male mass and the dominant frequency of the call.     

Increase in Male Reproductive Success and Female Reproductive Investment in Invasive Populations of the Harlequin Ladybird Harmonia axyridis

Reproductive strategy affects population dynamics and genetic parameters that can, in turn, affect evolutionary processes during the course of biological invasion. Life-history traits associated with reproductive strategy are therefore potentially good candidates for rapid evolutionary shifts during invasions. In a series of mating trials, we examined mixed groups of four males from invasive and native populations of the harlequin ladybird Harmonia axyridis mating freely during 48 hours with one female of either type. We recorded the identity of the first male to copulate and after the 48 h-period, we examined female fecundity and share of paternity, using molecular markers. We found that invasive populations have a different profile of male and female reproductive output. Males from invasive populations are more likely to mate first and gain a higher proportion of offspring with both invasive and native females. Females from invasive populations reproduce sooner, lay more eggs, and have offspring sired by a larger number of fathers than females from native populations. We found no evidence of direct inbreeding avoidance behaviour in both invasive and native females. This study highlights the importance of investigating evolutionary changes in reproductive strategy and associated traits during biological invasions.     

Correlated Responses in Male Reproductive Traits in Mice Selected for Litter Size and Body Weight

Correlated responses in male reproductive traits were determined at 4, 6 and 8 weeks of age in lines of mice selected for large litter size (L⁺), large 6-week body weight (W⁺), large litter size and small body weight (L⁺W^-) and small litter size and large body weight (L^-W⁺), and in an unselected control (K). Concentration of serum testosterone and weights of testes, seminal vesicles, epididymides and adrenal glands increased with age. Line differences in testosterone concentration were not detected. L⁺ and W⁺ males exhibited positive correlated responses in testes, epididymides and seminal vescile weights. Testis weight adjusted for body weight was significantly larger for L⁺ than controls and approached significance for W⁺. Realized genetic correlation betestis weight and litter size was 0.60 ± 0.04, and the realized partial genetic correlation holding body weight constant was 0.42. Therefore, pleiotropic loci, acting via the hypothalamic-pituitary axis, affect testis weight and litter size independently of body weight. Additionally, genes influencing overall growth have a pleiotropic effect on testis weight and litter size in mice; the realized genetic correlations of body weight with testis weight and with litter size were 0.60 ± 0.03 and 0.52 ± 0.10. Testis weight increased in both L⁺W^- and L^-W⁺ males. The positive correlated response in L⁺W^- may have resulted from changes in frequency of genes controlling reproductive processes; whereas, in L^-W⁺ it could have been the result of changes in the frequency of genes associated with body weight.     

Inferred Paternity and Male Reproductive Success in a Killer Whale (Orcinus orca) Population

We used data from 78 individuals at 26 microsatellite loci to infer parental and sibling relationships within a community of fish-eating ("resident") eastern North Pacific killer whales (Orcinus orca). Paternity analysis involving 15 mother/calf pairs and 8 potential fathers and whole-pedigree analysis of the entire sample produced consistent results. The variance in male reproductive success was greater than expected by chance and similar to that of other aquatic mammals. Although the number of confirmed paternities was small, reproductive success appeared to increase with male age and size. We found no evidence that males from outside this small population sired any of the sampled individuals. In contrast to previous results in a different population, many offspring were the result of matings within the same "pod" (long-term social group). Despite this pattern of breeding within social groups, we found no evidence of offspring produced by matings between close relatives, and the average internal relatedness of individuals was significantly less than expected if mating were random. The population's estimated effective size was <30 or about 1/3 of the current census size. Patterns of allele frequency variation were consistent with a population bottleneck.     

Environmental Effects on Male Reproductive Success and Parental Care in the Florida Flagfish Jordanella floridae

Models of optimal parental care predict that parental investment should depend on offspring value or the effect parental care has on offspring benefits. Few studies have examined the effect of external factors that influence offspring survival and the cost of care. In this study on the Florida flagfish (Jordanella floridae), a species with male parental care, we examined whether environmentally induced changes in care result from changes in egg requirements or in parental costs. We manipulated salinity and temperature, as these factors are known to affect the metabolic rate in both eggs and parents. We predicted that if the change in care behavior is determined by costs to the male then it should be paralleled by changes in non‐egg‐directed behavior. Conversely, if egg‐directed behavior changes independently of other behavior it would suggest that behavior is determined primarily by egg requirements. In addition we examined patterns of mating success under the assumption that if male care is affected by environmental factors then female preferences may change accordingly. Males decreased egg‐directed behavior (fanning and cleaning of eggs) at high salinity. Non‐egg‐directed behavior was unaffected by salinity. Temperature had no effects on behavior. Thus, we conclude that changes in egg demands are primarily responsible for the observed results. Successful males were bigger and more aggressive. This suggests that male dominance was an important determinant of male mating success. Unsuccessful males showed significantly more variation in number of red stripes with respect to salinity than successful males. Unsuccessful males may be less able to regulate color expression under varying environmental conditions, in which case color may be an indicator of male quality. We replicated the experiment early and late in the season. Males did not change their effort in care over the season. However, care (fanning) in the absence of eggs increased towards the end of the season. Since pre‐mating fanning was positively correlated to a male’s eventual mating success we conclude that males increased their effort to attract mates late in the season.     

Sexual Traits as Quality Indicators in Lekking Male Great Bustards

We studied the development of two sexual traits, whiskers and neck plumage, in relation to sexual selection in 41 free‐living great bustard, Otis tarda, males radio‐tracked at nine leks in central Spain in 1998–2001. During the pre‐breeding male–male competition period (Feb.) prior to female arrival, number and length of whiskers correlated with weight, but not with body size or age. Whiskers may thus have evolved as an intrasexual indicator of weight, which in the absence of other weapons in this species is decisive in male–male combats. Signalling through whiskers contributes to minimizing dangerous aggressive interactions in the lek. During the mating period (Apr.), both whisker and neck development were correlated with weight and age. Males reaching higher expression of both traits exhibited higher display intensity, a more prolonged display period through the mating season, and a higher estimated mating success. Moreover, interannual changes in a male’s expression of both traits were associated with changes in its display intensity and estimated mating success. Our results resolve earlier debates and contradictory results from previous authors, suggesting that these two secondary sexual traits, whiskers and neck, may function as reliable indicators of age and weight, the two main factors determining social rank of males in great bustard leks, during both rival assessment and mate choice. Their dual functions provide support for the pre‐existing trait and redundant signal hypotheses and suggest that multiple ornaments functioning as redundant signals might be more widespread than previously acknowledged.