Agonistic dominance in male baboons: An alternative view

Data on baboons have generated both the concepts normally associated with male dominance hierarchies among primates and the tests of their significance. The priority-of-access model has been used to predict the relationship between dominance rank and resource acquisition. While the correlation between these two factors, or between rank and measures of reproductive success, has varied among different primate species, most recent baboon field-workers have interpreted their results to be consistent with the model. Based on 1200 hr of observation of a troop of savannah baboons near Gilgil, Kenya, this paper presents data on male agonistic interactions and on male acquisition of resources. Predictions of the priority-of-access model are tested and an inverse relationship is found between agonistic dominance rank and acquisition of two limited resources, estrous females and meat. The importance of the residency status of males is explored and an alternative hypothesis is presented to account for the anomalous pattern in the data. The relationship of male reproductive success and dominance rank is evaluated in light of the data on these baboons and the “residency” hypothesis.     

Mate selection,consortship formation,and reproductive tactics in adult female savanna baboons

Mate selection is a proximate mechanism influencing the probability that two individuals will engage in reproductive activity. The degree to which mate selection by female savanna baboons (Papio cynocephalus anubis) affects consortship formation and the characteristics of males that they prefer to mate with were analyzed based upon a 19-month study in Kenya. Sexual solicitations by females had a positive impact on the probability that males would establish consort relationships. The most preferred mating partners were young, high ranking, newcomer males, but these same males did not have the highest mating success. Friendships between male and female baboons almost always developed subsequent to conception cycles. Female savanna baboon reproductive tactics appear to be based upon maximizing the probability that males will provide care for offspring.     

Determinants of male mating success in a large group of Barbary macaques (Macaca sylvanus) at Affenberg Salem

Male mating activities in relation to the likelihood of ovulation and conception were studied in a large group of semifree-ranging Barbary macaques (Macaca sylvanus) during two successive mating seasons. In both mating seasons, adult males attained a significantly higher mating success than subadult males, and they monopolized high ranking females more effectively than low ranking females during the period when conception was most probable. Also, in both mating seasons male rank was significantly correlated with male mating success if all sexually mature males were included. Nevertheless, mating success was not a linear function of age or rank. In both mating seasons mating success of 5-year-old males was much higher than that of dominant but peripheralized 6- and 7-year-old males. Moreover, a significant correlation between rank and measures of mating success among adult males was found in the second but not in the first mating season. The results indicate that mating and, most probably, reproductive success of male Barbary macaques is dependent on the male's social position in the group, which is defined not only by the outcome of dyadic agonistic encounters but also by the ability to get a central position in the group, and on the stability of rank relations.     

Consortship and Mating Success in Chacma Baboons (Papio cynocephalus ursinus)

Chacma baboons (Papio cynocephalus ursinus) show a lower consortship take‐over rate and longer consortship duration than the other savannah baboons ( Bulger 1993 ). It has been argued that researchers have focused on atypically small troops with few adult males, resulting in low competition for access to oestrous females. Consortship data from two mountain baboon troops containing seven and four males, respectively, were analysed to determine whether the troop with the greater number of males showed a weaker correlation between mating success and rank due to an expected higher consortship take‐over rate. No consort take‐overs were observed in either study troop and mating success in both troops was correlated strongly with male rank. The distribution of days spent in consortship amongst the males could be explained by the priority‐of‐access‐model. The degree of cycle overlap determined the number of males observed consorting oestrous females, whereas the number of males did not influence the relationship between rank and consorting activity.     

Reproductive strategies of rhesus macaques

Reproductive strategies incorporate a multitude of mechanisms that have evolved to promote the reproductive success of individuals. Evolutionary perspectives tend to emphasize the advantages of male-male competition and female choice as mediators of differential reproduction. Male rhesus macaques have not been observed to fight for access to sexually receptive females, although they suffer more wounds during the mating season. An increased likelihood of attacks appears to coincide with male troop entry. Males who spend more time in consort and mate with more females tend to sire more offspring. Genetic analysis of paternity has pinpointed age and endurance rivalry, rather than agonistic competition, as key variables associated with variation in progeny production. Female rhesus macaques often copulate with multiple males during their ovulatory period, and tend to conceive on the first cycle of the mating season. Female reproductive success is more likely to be a function of offspring survivorship than the identity of particular male partners. The role of female choice as a direct mediator of male reproductive success is unresolved, but female mate selection seems to indirectly affect male reproductive success because female preference for mating with novel males seems to foster male dispersal. Evaluating whether mating preferences for particular male phenotypes affectsfemale reproductive success is a task for the future. A common denominator to the reproductive strategies of both female and male rhesus macaques is that feeding patterns affect body condition which influences reproductive output and regulates relative reproductive success.     

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.     

BODY SIZE,SPERM COMPETITION,AND DETERMINANTS OF REPRODUCTIVE SUCCESS IN MALE SAVANNA BABOONS

One component of sexual selection is sperm competition. It has been reasoned that the intensity of sperm competition may be reflected in the relative testicular sizes of animals. Among males residing in multimale breeding systems, testicular size is relatively larger than among males residing in unimale mating systems. Information on whether differences in testicular size within a species can account for differences in male reproductive success is unavailable for natural populations of primates. A population of six troops of savanna baboons in Kenya was surveyed for morphometric analysis, and one of these troops was the subject of extensive behavioral observations afterwards. Testicular weights could not be obtained, but measurements of linear dimensions were transformed into volumetric estimates. Male weight accounted for 30% of the variance in testicular volume. Neither body size nor testicular volume was associated with differences in male reproductive activity. The outcome of fights over access to females could not be related to male body size, and ejaculatory patterns of males were independent of testicle size. Both sperm competition and aggressive competition intensified during the four-day optimum conception period, but fights over access to consort females were infrequent. Among savanna baboons, the probability of an ejaculation resulting in a conception is fairly low, which may account for the infrequency of injurious fights. Although testicle size influences sperm production, it does not influence either the timing of mating or the fertilizing capacity of spermatozoa, and both of these factors probably account for a substantial fraction of the variance in male baboon paternity. Sperm competition is an adjunct to agonistic competition as a mechanism affecting male baboon reproductive success. It is concluded that male reproductive success in baboons is affected more by social factors than by morphological traits associated with size.     

Intrasexual selection and male mating strategies in baboons and macaques

If baboon and macaque mating systems constitute a form of female defense polygyny, male mating strategies should be intrasexually selected and should vary in predictable ways with female defensibility, and demographic factors which affect the numbers of competing males per estrous female in populations. Substantial behavioral evidence exists for intrasexual selection of male mating strategies in baboons and macaques. Limited evidence also offers tentative support for theorybased predictions about the relationship between male mating strategy and female defensibility. Although male dominance rank generally predicts mating success, there are a number of factors which tend to increase the success of subordinate males above that expected from a simple dominance-based model of priority of access to mates.     

Dominance and reproductive success in primates

Theoretical models of the relationship between competition and differential reproduction in primates share the premise that agonistic dominance hierarchies determine differential access to those limited resources which are essential to reproduction. In particular, the priority-of-access-to-estrous-female model is based on the postulate that high rank in males enhances reproductive success. Tests for a correlation between rank and reproductive success in males have produced mixed results. Problems in measuring male reproductive success and in conceptualizing dominance systems are implicated in the diversity of conclusions reached. Other attributes which affect reproductive success have been proposed, and alternative models of mating systems, based on the concepts of reproductive strategies, social skills, and life histories have also been developed. Studies of differential reproduction in female primates are fewer, but have the advantage over male studies of directly measuring infant production and offspring survival. Research on the relationship between rank and reproductive success in females has shown that under conditions of low resource availability, subordinate females sometimes suffer reduced fecundity and infant survival as a result of restricted access to food and water. Under conditions of social stress, low-ranking females may suffer from disruption of their reproductive cycles, resulting in lowered fecundity. Dominant females may also incur reproductive costs as well as benefits for their social position. Better insights into the relationship between dominance and reproduction are potentially offered by improved genetic paternity measures, new models of social and demographic processes, and the recent availability of life history data from field studies.     

Do Female Mandrills Prefer Brightly Colored Males?

Although secondary sexual adornments are widespread in male primates, few studies have examined female choice for these characters. Mandrills (Mandrillus sphinx) present an extreme example of sexual dimorphism, with males exhibiting an array of striking adornments. The most dominant adult male in a group exhibits the brightest and most extensive red coloration, while the other males are less brightly colored. I examined whether female mandrills prefer brightly colored males using data on periovulatory sexual behavior during the 1996 mating season for all males 8 years old (n = 5) and all parous females (n = 9) in a semifree-ranging colony at CIRMF, Gabon. Brightness of male coloration is significantly positively correlated with time spent within 2 m of females, female responsibility for proximity, number of sexual presentations received, % approaches accepted by females, and % inspections with which females cooperated. Females also groomed only the brightest male. Behaviors indicating female preference are not correlated significantly with male dominance rank, and partial correlations confirm that the influence of male color on female behavior is stronger than that of male rank. With the influence of male dominance rank controlled, correlation coefficients between female behaviors and male mating success are high and positive. In further support of the hypothesis that females show mate choice for brightly colored males, independent of dominance rank, I report an unusual case wherein the alpha male fell in rank without loss of coloration. He experienced no significant change in female responsibility for proximity, sexual presentations received, or female reaction to approaches or inspections, though he was no longer observed to mate. Accordingly, female mandrills attend to differences in male secondary sexual characters and favor brightly colored males. As brightly colored males are also dominant this reinforces the influence of male-male competition on male reproductive success and may explain the very high reproductive skew in mandrill males and their extraordinary appearance.     

Male Mate Choice in Lemur catta

Though females are generally more selective in mate choice, males may also derive reproductive benefits from exercising mate selectivity if one or more factors limit male reproductive success and females differ in reproductive potential. I used male mating effort as a proxy for male mate choice in ring-tailed lemurs (Lemur catta). I calculated mating effort as the rate of male-male agonism during each female's estrous period 30 min before and 30 min after the first and last mountings with intromission. I collected data on 1 free-ranging Lemur catta troop during 2 consecutive breeding seasons on St. Catherines Island, USA. In both yrs, male mating effort differed significantly among troop females once I adjusted male-male agonistic rates to reflect agonistic intensity, and I corrected for the number of observed mates per female (2000: χ² = 27.43, df = 3, p < 0.0001; 2001: χ² = 21.10, df = 3, p < 0.001). Results strongly suggest male mate choice. Contrary to expectation, males did not expend the greatest mating effort for females with the highest dominance status nor the highest reproductive success. Males preferred females that either: (1) belonged to the age class in which fecundity and infant survival is the highest at this site (4–9 yrs), or 2) were older females (≥10 yrs) with high reproductive success. Female reproductive potential appears to be an important variable determining male mating effort in Lemur catta.     

Promiscuous Mating among Blue Monkeys in the Kakamega Forest,Kenya

13 females and 12 of 19 males present in a group of blue monkeys (Cercopithecus mitis stuhlmanni) mated promiscuously during an intense bout of sexual activity. Several aspects of their behavior are considered as mechanisms for determining mating patterns. Estrus synchrony coincided with a multi-male influx, allowing promiscuity. The length of stay and rank of an incoming male were correlated with his number of copulations in the group, while the original resident and highest ranking male was never seen to mate. Thus, although agonistic behavior, especially among males, affected mating success, it did not guarantee copulations. Adjustments of position relative to other individuals also affected who mated with whom. These and earlier observations suggest that there may be more than one way to be successful in mate competition, and lead us to question whether the kinds of behavior we observed have merely a reproductive function.     

Trade-off between pre and post-copulatory traits depends on locomotor activity in male Tribolium castaneum beetles

Locomotor performance is an indicator of dynamic exercise; thus, it is a central trait in many animal behaviours. Although higher locomotor endurance may increase male reproductive success (e.g., in mate searching and male–male contests), investment in other male reproductive traits (e.g., male attractiveness and sperm competition) may be decreased through energy consumption due to higher activity levels. Here, I investigated male attractiveness, mating success, and paternity success using males of the red flour beetle Tribolium castaneum selected for higher (H) and lower (L) locomotor endurance. Although there was no difference in male attractiveness between the selection regimes, H males had significantly higher mating success than L males. Conversely, L males had significantly higher paternity success than H males. Therefore, there was a trade-off between mating success and paternity success among the selection regimes, suggesting that locomotor endurance affects male reproduction in T. castaneum, and individual variation of locomotor endurance may be maintained within a population.     

Multiple Mating Increases Fecundity, Fertility and Relative Clutch Mass in the Female Leopard Gecko (Eublepharis macularius)

Females across many taxa may mate with several males or mate more than once with the same male within one reproductive event. Although many researchers have discussed the effects of multiple mating on reproductive success of females, few studies have attempted to disentangle whether the reproductive success of females differs with respect to whether females mate with multiple males or mate more than once with one male. In this study, we hypothesized that female leopard geckos (Eublepharis macularius) increase aspects of their reproductive success, such as fecundity, fertility and relative clutch mass, by mating more than once within one reproductive event, either by mating repeatedly with the same male or multiply mating with different males. We controlled for the potentially confounding variables of mating frequency and mate number by allowing females to mate once with one male, twice with the same male, or twice with two different males. We found that females that mated with more than one male laid more clutches, exhibited increased egg fertility and invested more in clutches relative to females that mated only once with one male, whereas females that mated twice to the same male were intermediate for these variables. Thus, reproductive success is higher among female leopard geckos that mated with more than one male compared to female leopard geckos that mated only once.     

Dominance rank reversals and rank instability among male Lemur catta: the effects of female behavior and ejaculation

In this study, dominance rank instability among male Lemur catta during mating was investigated. Also, data on agonism and sexual behavior across five consecutive mating seasons in a population of L. catta on St. Catherines Island, USA, were collected. Instances of male rank instability were categorized into three types. Type 1 consisted of a temporary switch in the dominance ranks of two males, which lasted for a period of minutes or hours. Type 2 dyadic male agonistic interactions showed highly variable outcomes for a period of time during which wins and losses were neither predictable nor consistent. Type 3 interactions consisted of a single agonistic win by a lower-ranked male over a more dominant male. More Type 2 interactions (indicating greater dominance instability) occurred when males had not spent the previous mating season in the same group, but this trend was not statistically significant. The majority of periods of male rank instability were preceded by female proceptivity or receptivity directed to a lower-ranked male. As such, exhibition of female mate choice for a lower-ranking male appeared to incite male-male competition. Following receipt of female proceptivity or receptivity, males who were lower-ranking took significantly longer to achieve their first agonistic win over a more dominant male than did males who were higher-ranked. Ejaculation frequently preceded loss of dominance. In conclusion, temporary rank reversals and overall dominance rank instability commonly occur among male L. catta in mating contexts, and these temporary increases in dominance status appear to positively affect male mating success.     

Emergence and mating behavior of the oriental fruit moth Cydia molesta (Lepidoptera: Tortricidae) and its potential for reproduction

The Oriental fruit moth, Cydia molesta (Busck, 1916) (Lepidoptera: Tortricidae), is a key pest of fruit and is widely distributed around the world. There are important connections between its behavior and biology and its management in agriculture, but few studies have investigated the associations between adult behaviors and oviposition. In this study, adult emergence, mating, and reproduction were investigated under laboratory and field conditions. The ratio of females to males at eclosion was approximately 1:1. When one virgin female had access to one virgin male, 66% and 34% of the couples copulated just once and twice, respectively; and the infertility rate of eggs (21.39 ± 1.25%) did not vary daily. Males, given access to one new female daily, could copulate multiple times, whereas females seldom mated more than once, indicating a male-biased operational sex ratio, but mating status of the male parent had no effect on progeny egg reproduction. Also, the number of eggs that hatched by all female partners of a male was inversely proportional to copulation duration for the female laying the eggs for total female reproductive success; and the number of eggs laid by all female partners of a male was proportional to their number of matings for total male reproductive success. However, the total number of eggs that hatched did not significantly differ for eggs laid by a female given new virgin males daily for mating (17.75 ± 4.28) versus eggs laid by virgin females (19.17 ± 7.51) presented daily with a male that re-mated daily with the series of females. Therefore, our results showed that females engaged in mate choice and males engaged in mate competition, affecting egg production, a factor that may be used to enhance mating disruption technology against Cydia molesta.     

Male age affects female mating preference but not fitness in the monandrous moth Dendrolimus punctatus Walker (Lepidoptera: Lasiocampidae)

It is widely accepted that male age can influence female mating preference and subsequent fitness consequences in many polyandrous species, yet this is seldom investigated in monandrous species. In the present study, we use the monandrous pine moth Dendrolimus punctatus to examine the effects of male age on female mating preference and future reproductive potential. In multiple male trials, when permitted free mating from an aggregation consisting of virgin males aged 0 (young), 2 (middle-aged) and 4 (old) days, virgin females preferentially mate with young and middle-aged males, although mating latency and mating duration are independent of male age. In single male trials, when virgin females are randomly assigned single virgin males of known age, a negative correlation is found between mating success and male age in this species. However, we find that male age also has no effect on mating latency and mating duration. Further fitness analysis reveals that females do not receive benefits in terms of oviposition period, total egg production, average daily egg production, percentage of egg hatching, longevity, expected reproduction and relative expected reproduction from mating with young and middle-aged males compared with mating with old males. The results of the present study are the first demonstrate that females mated preferentially with younger males but gain no apparent fitness benefits in a monandrous moth species.     

A review of the potential of surface flow constructed wetlands to enhance macroinvertebrate diversity in agricultural landscapes with particular reference to Integrated Constructed Wetlands (ICWs)

Discriminating female mate preferences enhance the variance in reproductive success among males of a population and create a potential for sexual selection, which can account for trait evolution and diversification. Fish color patterns are among the prime targets of mate choice-driven sexual selection. Populations of the cichlid Tropheus from Lake Tanganyika display remarkable geographic color pattern variation, but the role of female choice in their rapid and rich phenotypic diversification is unclear. Males and females establish a pair bond prior to spawning monogamously, but as brood care is strictly maternal, female investment in reproduction is high and the operational sex ratio is male-biased. Therefore, variance in male reproductive success can accrue if individual males succeed repeatedly in securing a mate. To test this prediction in the red colored Tropheus moorii “Chimba”, four pairs of males were presented to a series of females and female mate preferences were inferred from pairwise interactions. There was a significant difference in mating success between the males of each pair (P < 0.001 over all trials), as—with one exception—females shared preferences for the same males. Male courtship activity was strongly correlated with female choice. Our experiment suggests that female choice contributes to the variance in male reproductive success in the tested population.     

Courtship song, male agonistic encounters, and female mate choice in the house cricket,Acheta domesticus (Orthoptera: Gryllidae)

Whether female crickets choose among males based on characteristics of the courtship song is uncertain, but in many species, males not producing courtship song do not mate. In the house cricket,Acheta domesticus, we examined whether a female chose or rejected a male based on his size, latency to chirp, latency to produce courtship song, or rate of the high-frequency pulse of courtship song (“court rate”). We confirmed that females mated only with males that produced courtship song, but we found no evidence that the other factors we measured affected a female’s decision to mate. In addition, we investigated whether the outcome of male agonistic encounters affected the subsequent production of courtship song. In one experiment, we observed courtship and mating behavior when a single female was placed with a pair of males following a 10-min interaction period between the two males. Winners of male agonistic encounters had higher mating success. However, winners and losers of agonistic encounters were not different in their likelihood or latency to produce courtship song or in the number of times they were disrupted by the other male in the pair. In a second experiment, we allowed two males to interact for a 10-min period, but following this interaction period, we placed a female with each male separately and observed courtship and mating behavior. The mating success of winners and losers was not different under these circumstances, and we found no differences between winners and losers in any subsequent courtship or mating behavior examined. We conclude that winning agonistic encounters influences a male’s mating success in ways other than his production of courtship song and this effect is lost when winning and losing males are separated and each is given an opportunity to mate.     

Queuing and queue-jumping: long-term patterns of reproductive skew in male savannah baboons, Papio cynocephalus

In many animals, variance in male mating success is strongly correlated with male dominance rank or some other measure of fighting ability. Studies in primates, however, have varied greatly in whether they detect a relationship between male dominance rank and mating success. This variability has led to debate about the nature of the relation between rank and mating success in male primates. We contribute to the resolution of this debate by presenting an analysis of the relationship between dominance rank and male mating success over 32 group-years in a population of wild savannah baboons. When data were pooled over the entire period, higher-ranking males had greater access to fertile females. However, when we examined successive 6-month blocks, we found variance in the extent to which rank predicted mating success. In some periods, the dominance hierarchy functioned as a queue in which males waited for mating opportunities, so that rank predicted mating success. In other periods, the queuing system broke down, and rank failed to predict mating success when many adult males were in the group, when males in the group differed greatly in age, and when the highest-ranking male maintained his rank for only short periods. The variance within this single population is similar to the variance observed between populations of baboons and between species of primates. Our long-term results provide strong support for the proposition that this variance is not an artefact of methodological differences between short-term studies, but is due to true variance in the extent to which high-ranking males are able to monopolize access to females. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.