Testosterone predicts future dominance rank and mating activity among male chacma baboons

Despite the many benefits that testosterone has on male reproduction, sustaining high levels of testosterone for long periods can be costly. The challenge hypothesis predicts that males will show temporarily sustained elevations of testosterone at critical periods, counterbalanced by decreased levels during noncritical periods. We investigated male testosterone measures extracted from fecal samples in a group of chacma baboons (Papio hamadryas ursinus) living in the Okavango Delta, Botswana. Because rank serves as a proxy for competition for mates, we examined how male testosterone was related to dominance rank, age, aggression, and mating activity. Males showed an elevation in testosterone at maturity; young adult males had the highest testosterone levels followed by a steady decline with age. Among dispersing males, testosterone was temporarily elevated in the month following dispersal. After controlling for age, testosterone and rank were unrelated, but testosterone and changes in rank were positively correlated, such that males rising in rank had higher testosterone than males falling in rank. Thus, for males in this group, testosterone was predictive of a male's rank trajectory, or future rank. Similarly, male testosterone levels predicted future, rather than current, mating activity. Finally, male testosterone and aggression rates were unrelated during stable periods in the dominance hierarchy but positively related during unstable periods when high ranks were being contested. In general, our results support the challenge hypothesis with males exhibiting elevated testosterone in association with the acquisition of high rank (ensuring access to mates), rather than with mating itself.     

The costs of male parental care and its evolution in a neotropical frog

Summary Parental care is practiced exclusively by males of the Puerto Rican frog, Eleutherodactylus coqui. Males brood clutches of direct-developing eggs in non-aquatic nest sites and defend eggs against cannibalistic nest intruders. Here, I report on energetic and mating costs incurred by males that provide parental care, and suggest how these proximate costs affect male fitness and the evolution of male parental care in this species. Energetic costs are small for brooding males in comparison to non-brooding, calling males. Brooding males had a higher frequency of empty stomachs and lost small, but significant, fractions of their initial body mass during parental care. Abdominal fat bodies of brooding males during the middle third of parental care were significantly smaller than those of calling males; those of males brooding eggs in earlier or later stages were not different. The mating cost of parental care is greater. Most brooding males cease calling during parental care. However, gravid females are available (i.e., known to mate) on most nights during the principal breeding season; hence non-calling males miss potential opportunities to mate. A mating cost was estimated by calculating nightly mating probabilities for calling males in a plot where nightly calling male densities and daily oviposition schedules were known. On average, a male exhibiting normal calling behavior would be expected to obtain a new mate once every 35.7 days. Hence a brooding male that ceased calling for a 20-day parental care period would miss, on average, 0.56 additional mates. Males that were more successful than average in attracting mates could miss up to 1.63 matings. A marginal value model (Fig. 1) is used to analyze the net effect on male fitness of parental care benefits and costs in E. coqui (Fig. 3). The model indicates that males garner the highest reproductive success by providing care from oviposition through hatching. There is no stage during the pre-hatching period at which a desertion strategy would yield higher reproductive success. In fact, the model suggests that males should provide full parental care even in the face of much higher mating costs than currently obtain in the system.     

Male care and mating effort among Hadza foragers

Paternal care figures prominently in many scenarios of human evolution. Recently, however, such scenarios have been challenged on two scores. First, the level of male contribution may be insignificant. Second, male care may be provided as a form of mating effort, rather than parenting effort. In theory, since men can enhance their Darwinian fitness both by providing care to their own offspring if this raises offspring fitness and by pursuing additional mates if this leads to additional offspring, men should respond to payoffs from both mating and parenting effort. If men respond to payoffs from parenting effort, paternity ought to make a difference. And if men respond to payoffs from mating effort, mating opportunities ought to make a difference. I analyzed the impact of these two factors on variation in male care among the Hadza, a foraging society in Tanzania. Two predictions were tested: (1) biological children will receive more care than stepchildren, and (2) men will provide less care to their biological children as their mating opportunities increase. Both predictions were supported. These results suggest men provide care, in part, as parenting effort, and that they trade off parenting effort for mating effort when they have greater mating opportunities. Received: 21 January 1998 / Received in revised form: 24 January 1999 / Accepted: 1 February 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     

Correlates of male mating success in great bustard leks: the effects of age, weight, and display effort

We examined how mating success varied in relation to age, weight, body size, and display behavior among great bustard Otis tarda males. The estimated mating success was strongly skewed, with 45% of adult males being involved in copulation attempts and only 9.7% actually seen copulating successfully. Unlike most birds, body size continued increasing in great bustards several years after reaching sexual maturity. Age, weight, and display effort were all significant and independent predictors of male mating success. The higher display effort involved performing longer full-display bouts. Older males could detach from the male flock earlier in the season as well as on each day and spend longer seasonal and daily periods displaying as solitary birds, which contributed to increase their mating success. In contrast, males weighing more did not invest more in display, which suggests that they could be recognized as dominants by other males and selected by females through assessment of their plumage sexual traits. In contrast to most other bird species, the system described for great bustards resembles that found in some lek-mating ungulates, where social rank is a complex trait determined by both age and mass, and as in these mammals, it suggests that sexual selection continues to favor a high male weight in this extremely sexually dimorphic species.     

Female mallard mating preferences for multiple male ornaments

Mating preferences were studied in captive wild-stock female mallards. Independent observers scored ornament brightness for the bill and eight plumage areas of male mallards on a scale of 1 to 6. Ornament scores were generally positively correlated with each other as indicated by univariate correlations and principal component analysis. Males with higher bill scores were significantly more likely to obtain pairings. None of eight individual plumage ornaments were significant predictors of pairing success, but males with higher average plumage scores had significantly higher pairing success. Males that completed the molt first were also significantly more likely to be paired. Thus females may have paid some attention to plumage brightness. Stepwise multiple regression produced a model that included only bill and molt, both of which contributed a similar degree (r ² = 0.39); plumage ornaments and size measurements did not enter the model. These results are interpreted in light of hypotheses for the evolution of multiple ornaments, and phylogenetic plumage patterns in dabbling ducks. Received: 18 December 1995 / Accepted after revision: 22 September 1996     

The effect of female mating status on male offspring traits

In haplodiploid insects, males develop from unfertilized eggs; consequently, unmated females can reproduce. In a patchy, highly structured population, where brothers compete for mates and the reproductive return through sons is lower, females should minimize the number of male offspring. Consequently, unmated females are likely to have a reduced fitness compared to mated females. Here, we tested the oviposition behaviour of the haplodiploid beetle Coccotrypes dactyliperda. In this species, the unmated female can mate with her son to produce daughters. We predicted that unmated females could increase their fitness by (1) producing only few and small sons sufficient for mother–son mating and (2) dispersing to a patch occupied by conspecific females in order to increase their or their sons’ chance of mating. We demonstrate that (1) unmated females are common (23 % of all females), (2) they oviposit more frequently than mated females in occupied patches, (3) unmated females oviposit more eggs than mated females—this is in spite of the trade-offs, evident in this study, between the number of sons and the number of the mother’s future offspring after mating, (4) unmated females have a higher proportion of dispersing sons, and (5) sons of unmated females are smaller than sons of mated females. We conclude that the incidence of unmated females in the structured populations of C. dactyliperda is explained by plasticity in their oviposition behaviour. We discuss conditions where a high incidence of unmated females can persist as a successful strategy in structured populations.     

Age-related changes in reproductive effort of male bison

Summary The behavior of male bison during the rut was studied, to test the prediction that reproductive effort should increase with age. Because bull bison do not show parental care, the major component of reproductive effort is competition with other bulls to obtain copulations with estrous females. Data were collected on activity budgets during the pre-rut and rut seasons, on interactions with other bulls, and on proximity to tended females during the rut. Participation in the rut started at ages 5–6. Older bulls showed greater percent time active and less percent time eating than younger bulls. The older bulls also had higher counts of dominance and fight-related interactions and lower counts for behavior associated with submission than did the younger bulls. Risk taken seemed to increase with age since bulls aged 6 to 12 participated more frequently in dangerous activities associated with fighting. Within the 6 to 12 year old group, activity patterns changed little with age. However, from the interaction information, peak ages of 8 to 11 years appeared. For the activities: back up, chase, clash, head nod, jump away, move away, run away, run toward, push heads, dominant, and risk, 11 and 12 year olds more closely resembled younger bulls. This suggests a reproductive effort curve that is flat at zero until about age 6, increases to age 8, flattens out again to around age 10, and then decreases somewhat. Reproductive effort in male bison thus generally conforms with life-history theory predictions.     

Nectar dehydration by male carpenter bees as preparation for mating flights

Summary Females of all social and many solitary bees dehydrate nectar before storing it or adding it to larval provisions. Nectar dehydration by males has rarely been documented. We report on the neotropical facultatively social carpenter bee Xylocopa nigrocincta, in which the nest constant males are fed nectar by their female nestmates. Males dehydrate the nectar at the nest entrance before leaving the nest for mating territories. We show that males thereby minimize their water load, resulting in an improvement of their energy budget during hovering flights in their territories. Males can prolong the duration of territorial flights if they cany highly concentrated nectar. We assume that nectar dehydration as a pre-mating behavior is not correlated with the social organization in Xylocopa species. However, the behavior is particularly weil-developed in X. nigrocincta, where during the mating period males remain integrated in the nest society and are fed by their mothers and sisters.     

Comparative costs of promiscuity and monogramy: A test of reproductive effort theory

Summary Sharp-tailed sparrows Ammospiza caudacuta and seaside sparrows A. maritima co-occupy salt marshes in northeastern North America and are ecologically similar. Female caudacuta care for their young unaided, in comparison to martima's dual parental behavior. The annual reproductive effort of female caudacuta is higher than that of female maritima. Under reproductive effort theory, as both species' reproductive output is the same, female caudacuta should have a lower probability of survival than female maritima. However, the annual survival of both groups was the same. We believe that these results are possible only in highly productive habitats, where the relative cost differences of derived and ancestral mating systems may not be as great as in less productive environments. Lack of penalty in terms of lifetime fitness may have set the stage for the evolution of promiscuity in the sharp-tailed sparrow.In spite of their emancipation from parental duties, male caudacuta survive no better than females. We conclude that activities not directly related to care of young, such as patrolling large home ranges, may entail as much risk or energy expenditure as parental duties do.     

Costing reproduction: effects of mating opportunity on mating success in male Bicyclus anynana butterflies

It is frequently assumed that males have an almost unlimited reproductive capacity, while access to receptive females is typically limiting. Consequently, sexual selection is expected to favor vigorous courtship behavior in males. If such behavior is associated with non-trivial costs, ample current mating opportunities should be accompanied by a reduction in future mating vigor. To test this hypothesis, three treatments differing in sex ratio were established using the tropical butterfly Bicyclus anynana: 50 males each were housed either with 0, 25, or 50 females. Mating trials involving the competition among males from each treatment for a single virgin female were carried out on days 3, 9, and 15 after allocation to treatments. While there was no difference on day 3, prior mating opportunity clearly reduced mating success on days 9 and 15, being lowest if identical numbers of males and females were housed together. This finding suggests accumulating costs associated with high courtship and/or mating activity. Further, older males were more active and initiated copulation earlier than the younger ones, consistent with the residual reproductive value hypothesis. We found no evidence for a survival or fertility cost of mating.     

Preparation and activation: determinants of age at reproductive maturity in male baboons

Age at maturity is a particularly important life history trait, but maturational data are rare for males in natural populations of mammals. Here we provide information on three maturational milestones and their social and demographic correlates among 43 wild male baboons, Papio cynocephalus, in a natural population in Amboseli National Park, Kenya. We examined (1) age at testicular enlargement, which signals puberty and the onset of subadulthood, (2) age at attainment of adult dominance rank, which we consider to be the beginning of adulthood, and (3) age at first sexual consortship, which is the best measure available for age at first reproduction in male baboons. Testicular enlargement (median age = 5.69 years) occurred earlier among sons of high ranking mothers, and was not influenced by rainfall or seasonality. Attainment of adult dominance rank (median age = 7.41 years) was also accelerated among sons of high-ranking mothers, and among males whose mothers had died while the males were juveniles. First sexual consortship (median age = 7.92 years) was not influenced directly by maternal characteristics, but attainment of adult dominance rank always preceded first consortship. The lag time between attainment of adult rank and first consortship (median = 2.5 months; range = 5–526 days), was predicted by the number of sexually cycling females in the group when the male attained rank, and by how high ranking the male became in his first months as an adult. We suggest that the age at which a male baboon is ready to begin reproducing is influenced by a relatively stable maternal characteristic that exerts its influence early in development, but the timing with which this potential is realized depends on activation by more proximate, often stochastic triggers such as female availability. This two-level organization of influences is likely to contribute to the variance both in age at first reproduction and in lifetime fitness. Differences in the relative magnitude of the two levels will lead to both intra- and interspecific variability in the opportunity for maternal selection and sexual selection.     

A radiotracer technique for the determination of male mating success in natural populations

Summary Males of a natural population of Antechinus stuartii (Marsupialia) were injected at the beginning of their short, synchronous breeding period with one of twelve gamma-emitting nuclides which are individually recognizable by their unique spectial properties. This label passed to the females during ejaculation and was identified when the females were captured and counted in a whole-body counter. This technique established mating success of males and provided an estimate of reproductive success.     

Foliar damage to parental plants interacts to influence mating success of Ipomoea purpurea

Studies of how herbivory affects plant fitness often determine whether damage to one parent alters reproductive output (i.e., seed set or paternity) but ignore the possibility that the outcome may be different if both parents were damaged (i.e., the presence of maternal x paternal damage interactions). Using inbred lines of the common morning glory, Ipomoea purpurea, I conducted a series of greenhouse experiments to test whether foliar damage from a generalist insect herbivore, Trichoplusia ni, alters male and female fitness components when neither parent, one parent, or both I. purpurea parents had been damaged. In a single-donor experiment, flowers on both damaged and undamaged maternal plants received pollen from either damaged or undamaged paternal plants. I. purpurea flowers were more likely to be aborted when they received pollen from damaged paternal plants, or when maternal plants were both damaged and grown under low-resource conditions. Foliar damaged plants also produced less seed and pollen than undamaged plants, although seed mass and pollen viability were not affected by damage. In a multiple-donor experiment, flowers on damaged and undamaged maternal plants simultaneously received pollen from damaged and undamaged paternal plants, and F1 seeds were analyzed for paternity. Damaged paternal plants had reduced siring success compared to undamaged paternal plants, and this discrepancy was most pronounced when competition occurred on damaged maternal plants. Thus, damaged maternal plants were more "selective" than undamaged maternal plants. Although previous studies have demonstrated that herbivory can alter fruit and seed production and paternity patterns, this is the first study to show that the magnitude of herbivore damage experienced by both parents can interact to influence maternal and paternal mating success.     

Oviposition site guarding by male walnut flies and its possible consequences for mating success

Field studies showed that male Rhagoletis juglandis and R. boycei flies guard egg-laying punctures (and the eggs within) on host walnut (Juglans major) fruit and defend those sites from conspecific and heterospecific males. In field experiments with artificially punctured fruit,as well as field observations on unmanipulated fruit, males were consistently more likely to be sighted and stayed longer on damaged fruit than on undamaged fruit. On artificially punctured fruit, they consistently spent more time in the vicinity of a puncture than expected by chance alone. Males together on damaged fruit were more likely to engage in contests over those fruit than males together on undamaged fruit. Copulations were consistently more frequent for either species on damaged than undamaged fruit, both in observations of unmanipulated fruit and in artificial puncture experiments. Analyses which controlled for the longer male residence time on damaged fruit suggested strongly that copulations were consistently achieved at higher per capita rates on damaged than on undamaged fruit, indicating that puncture-guarding functions to increase access to females. An exception to the pattern in male mating success was noted at a site where both species used host fruit on the same trees. In this case, R. juglandis males were only slightly more common on punctured fruit than on control fruit and male success in copulation did not differ significantly between the two types of fruit. This anomalous result was apparently due to an almost absolute advantage enjoyed by R. boycei males in on-fruit contests with R. juglandis males. A likely basis for improvements in mating success associated with puncture guarding was a propensity for females to deposit eggs into existing punctures. Both in observations of unmanipulated fruit and in artificial puncture experiments, females consistently attempted oviposition more often in damaged than undamaged fruit. In artificial puncture experiments, both species at both sites deposited most clutches in damaged fruit. Mating generally took place as females initiated oviposition. The possible functions of puncture use by females as well as alternative functions of puncture guarding by males are discussed.     

Evolution of male mating behavior: male spadefoot toads preferentially associate with conspecific males

The evolution of male breeding aggregations is difficult to explain because males may reduce their reproductive success by associating with their closest competitors. We examined aggregative behavior by male New Mexico spadefoot toads, Spea multiplicata, which form breeding choruses in rain-filled pools. We specifically asked whether males are attracted to conspecific calls and, if so, whether they preferentially associate with those male calls that are also attractive to females. Field observations revealed that males showed significant clustering with conspecifics within breeding ponds, whereas laboratory phonotaxis experiments revealed that males preferentially associated with conspecific male calls. Moreover, when males were presented with conspecific calls that differed in call rate, smaller males associated with the stimulus preferred by females (average call rate). Thus, males appear to evaluate the attractiveness of competitors using the same trait employed by females to assess potential mates, and males adjust their positions relative to competitors depending on their size. We discuss these results in the light of several current hypotheses on the adaptive significance of male breeding aggregations. Received: 20 December 1999 / Accepted: 18 March 2000     

The effects of mating system on male mate choice in a coral reef fish

Summary We show how mate limitation appears to be critical in determining whether or not males exercise mate choice among available females. Thalassoma bifasciatum is a Caribbean reef fish with two distinct mating patterns: group-spawning and pair-spawning. In both mating systems, female fecundity is variable and size dependent, and female availability is high. However, sperm competition among group-spawning males apparently limits the number of effective matings in which a male may engage, whereas territorial pair-spawning males have little or no such limitation. Group-spawning males should be discriminating in their choice of mates and our data confirm this: there is strong evidence for assortative mating in group-spawns, with more large males joining in mating groups around large females. In contrast, pair-spawning males show no indication of mate preferences, and spawn with all females who arrive at their territories.