Mating patterns and a test for female mate choice in Etheostoma spectabile (Pisces,Percidae)

Neither size nor breeding color correlated with spawning success of male orangethroat darters, Etheostoma spectabile (Pisces, Percidae), under natural field conditions. When females were presented experimentally with a simultaneous choice they spent no more time in proximity to large than small males, and were subsequently no more likely to spawn with large than with small males. Females also displayed no preference for bright versus dull males. Males and females did not differ significantly in size. Etheostoma spectabile may lack sexual size dimorphism as a result of the lack of female choice for size and the ineffectiveness of male attempts to monopolize females, or selection may be for increased size of females. Males are not dwarfs because of sperm competition. Contest competition among males appeared to be important in initiating spawnings but many males obtained spawnings by participating in ongoing spawning events. Etheostoma spectabile is an example of a sexually dimorphic species with no evident female preference for male size or color.     

Mating behavior of female rock shrimp Rhynchocinetes typus (Decapoda: Caridea)—indication for convenience polyandry and cryptic female choice

While studies on a wide diversity of organisms have demonstrated the importance of female behavior during matings, in crustacean studies, a strong bias towards male mating behavior prevails. Reproductively mature rock shrimp (Rhynchocinetes typus) exist as several ontogenetic stages that differ in their morphological and physiological capacities. In natural populations, the majority of males are in early ontogenetic stages (termed typus), many are in intermediate stages (intermedius), and few are in the terminal molt stage (robustus). Dominant robustus males, which have already demonstrated their biological fitness by surviving to this stage, have previously been shown to have a higher potential than subordinate typus males to defend receptive females against other males, and fertilize the entire clutch of a female. While females should thus show a preference for robustus males, they nevertheless frequently receive sperm from typus males. These observations suggested that females might have mechanisms to discriminate against sperm from subordinate males. In laboratory experiments, we observed that females avoided being seized by typus males for longer time periods in the absence of robustus males than in their presence. Following seizure, females that were initially held by typus males, required more time to initiate spawning than those held by robustus males. Many typus males transferred spermatophores to females before these started to spawn while robustus males waited until females began to spawn before they transferred spermatophores. Females manipulated spermatophores received from typus males for long time periods (minutes), but not those they received from robustus males. By accepting sperm from subordinate typus males, females may avoid further harassment (convenience polyandry), but they subsequently may discriminate against these subordinate males by delaying spawning and removing their sperm. These observations suggest that female behavior influences the outcome of matings, favoring fertilization of eggs by sperm from dominant males. Convenience polyandry and cryptic female choice may be common in other crustaceans as well.Electronic Supplementary Material Supplementary material is available in the online version of this article at .Communicated by T. Czeschlik     

Association patterns of spider monkeys: the influence of ecology and sex on social organization

Summary In this paper I consider how the costs and benefits of group living in spider monkeys (Ateles geoffroyi) vary between troop members. The results suggest that ecological factors set an upper limit to the number of spider monkeys that can associate and still efficiently exploit the available resources. In addition, the needs of the individual appears to influence the type and size of the subgroup it chooses. Adult males band together, travel over a large area, and are frequently sighted near the community's boundary. In contrast, adult females spend more time solitary than males and have association patterns that are strongly influenced by the presence of a dependent infant. Females with dependent infants tend to travel in small subgroups or alone, avoid the boundaries of the community's home range, and exhibit a restricted pattern of use of their range. The results suggest that males may be attempting to locate females with which they can breed, while mothers attempt to protect their infants by avoiding conspecifics and potentially dangerous situations near territorial boundaries.     

Fitness benefits of multiple mating versus female mate choice in the cellar spider (Pholcus phalangioides)

Recent studies have demonstrated that mating with multiple males can be beneficial for females and her offspring even if males contribute nothing but sperm. This was mainly established for species in which sperm from several males mix in the reproductive tract of the female, thus allowing sperm competition and/or female sperm choice. However, in species with last male sperm precedence, female re-mating decides against the previous male by strongly limiting his reproductive success. We tested the effect of female re-mating behaviour using the cellar spider Pholcus phalangioides, which shows strong last males sperm precedence and moderate levels of polyandry under natural situations. We predicted that females prevented from remating even though they are receptive would show reduced reproductive success compared to females that accept two copulations and females that reject a second male, since the latter two treatments were allowed to behave according to their decisions. However, if the number of matings per se had an effect on oviposition or on offspring performance, double-mated females should perform better compared to both treatments of once-mated females. We measured female fecundity and fertility over a period of 140 days, comparable to the species' natural reproductive peak season. Two thousand one hundred and fifty-two offspring from 67 first egg sacs were reared under two feeding levels. We registered development time and survival, and measured offspring adult size and mass. We found a positive effect of double mating, as in this treatment, oviposition probability was higher compared to the other treatments. Interestingly, adult female offspring of the DM treatment that were raised under low food level had a higher condition index compared to those from FS and RM, but development time, size and mass at adulthood were not affected by mating treatment. Female choice only seemed to affect hatching latency of the offspring. Overall, the main predictor of female reproductive output and success was female body size.     

Variation of life history traits and mating patterns in female langur monkeys (Semnopithecus entellus)

To determine nutritional and social influences on life history traits and mating behavior in female langur monkeys, new long-term data for a feral population living in multi-male groups under poor nutritional conditions are presented and compared with published long-term data for the same species living in one-male groups under excellent nutritional conditions. Under poor nutritional conditions, conceptions were confined to 5 months per year, coinciding with the period of the highest mating activity. Age at first parturition, gestation periods, lactation periods, and interbirth intervals were all significantly increased resulting in a more than 50% reduced reproductive rate under poor conditions. The pattern of sexual behavior during gestation was influenced by neither nutritional nor social conditions but the frequency was higher in multi-male groups. Furthermore, estrus length was significantly increased in multi-male groups where females tended to mate with as many resident males as available. Both phenomena, together with a high percentage of sexual behavior occurring outside the mid-cycle estrus, are interpreted in the light of paternity confusion in multi-male groups in a species where infanticide by males is regularly observed. The extreme variation in life history traits and mating patterns disclosed here in one species suggests that future cross-species comparisons should strictly select datasets based on local ecological and social conditions.     

Strategies of female mate choice: A theoretical analysis

Summary Females of many species face a difficult sampling problem while searching for a mate. How do they find the best male possible when the constraints of time, memory, and mobility prevent them from visiting all the males in the population?Five possible solutions are examined: random mating a fixed-threshold criterion, a fixed-threshold with last-chance option, an optimal one-step decision strategy, and the best-of-n-males strategy. Random mating is the worst strategy whenever the female gets more than one chance to mate. The two fixed-threshold strategies approach equal effectiveness as n increases but are always below the optimal one-step decision strategy. However, the best-of-n-males strategy always yields the highest expectation of fitness in a mate. The difference is especially great when n5.Plotting the average fitness of males chosen vs n, the number of males examined, yields a negatively accelerating curve. Since the cost of searching will be an increasing function of n, the two curves can be combined to yield an optimum n: the point where the difference between the curves is greatest.The one field study (Brown 1978) that addresses these problems in detail reveals that female mottled sculpins choose males on a relative, rather than absolute, basis, as theory suggests they should.     

Mate choice among sympatric fur seals: female preference for conphenotypic males

When closely related species breed in sympatry, and where hybrids have lower fitness, reinforcement theory predicts that selection should favour mechanisms that reduce the probability of interspecific matings. If this situation arises among species that exhibit resource defence polygyny where males and females of different species reside in the same territories, there may be some conflict between mate choice based on territory-holding ability (sexual selection) and mate choice for correct species. We investigated this in a population of fur seals where three species are sympatric and where some females breed in the territories of heterospecific males, and where interspecific matings and hybrid pups are observed. The territorial status of males and the birthing sites of females were determined during daily observations, as were the movements of males and females, the location of matings and mating partners. DNA extracted from skin samples was used to determine paternities using DNA fingerprinting and the mtDNA genotype of individuals. Individuals were also classed on the basis of species-typical phenotype. We found that extra-territory inseminations (ETIs) were significantly more prevalent (67%) when territorial males and resident females were of different phenotype than when of similar phenotype (27%), but mtDNA genotype had no effect on the rate of ETIs. ETIs were probably by males with the same phenotype, as pups born to these females in the following season had the same phenotype as their mothers, suggesting they were not hybrids. These results suggest that within the resource defence polygynous mating system of these sympatric fur seals, female mate choice is more influenced by male phenotype than genotype. Contrary to our predictions, our study indicates that potential conflict between mate choice based on sexual selection and species recognition is unlikely, because females have some capacity to discriminate between males both within and between species on phenotypic traits additional to those under sexual selection. Although at least 25% of the pups born in this study were hybrid, this study can only support reinforcement theory if hybrids have reduced fitness. The fitness of hybrids among the species studied is currently unknown. Received: 19 January 1998 / Accepted after revision: 12 September 1998     

Mate sampling behaviour of female pied flycatchers: evidence for active mate choice

Summary This paper presents major new evidence for active mate choice of female pied flycatchers, Ficedula hypoleuca. Fifteen color-ringed females were released into a study area containing 23 unmated males defending one nestbox each. Through intensive surveillance, the behavior of the females was observed during 2 consecutive days. Twenty-two of the males received a total of 131 female visits. Six of the females settled in the study area, and their premating period lasted 1.3–2.5 days. The females were seen searching for mates for at least 6–32 h and were seen visiting at least 1–9 different males. Hence, some of the females rejected males before mating. Nevertheless, the females settled close to the site of release (range: 16–243 m), suggesting that they mated with one of the first males encountered. Females visited males most frequently in the morning, and the diurnal distribution of visits was significantly correlated with male song activity. Offprint requests to: S. Dale     

Kipsigis women's preferences for wealthy men: evidence for female choice in mammals?

Summary In contrast to studies of birds, in mammals there is no clear evidence that polygyny evolved through female choice for males with high quality resources. Among the Kipsigis people of Kenya, polygyny may be a consequence of women's preferences for wealthy men, because strong correlations exist between land ownership and the number of a man's wives (Borgerhoff Mulder 1987 a), and the resources men hold are primary determinants of women's reproductive success (Borgerhoff Mulder 1987b). This paper has two aims: first, to test whether Kipsigis women prefer wealthy men by examining the sequence of marriages among a group of pioneers (Table 1) who established a settlement in the territory of their enemies (1930–1949); second, to determine whether women suffer reproductively as a result of polygynous marriage. Data show that Kipsigis women, or their parents on their behalf, preferentially chose men offering high quality breeding opportunities, with respect to the number of acres available on which to settle (Fig. 2) ; controlling for quality of breeding opportunity there is a preference for bachelors over monogamists over polygynists. Analyses of the full demographic sample show that there are reproductive costs associated with having a large number of cowives (Table 2), costs which women attempt to minimize through judicous marital choices. These results are discussed in relation to resource defence polgyny, female choice and, specifically, the polygyny threshold model.     

Condition-dependent control of paternity by female purple martins: implications for coloniality

Proposed causal links between extra-pair copulation (EPC) and colony formation in socially monogamous birds hinge on the question of which sex controls fertilizations. We examined in colonial purple martins Progne subis (1) whether EPCs were forced or accepted by females, and (2) the degree to which apparently receptive females were able to obtain EPCs against their mates’ paternity defenses. Paternity analyses of multilocus DNA fingerprinting confirmed previous findings of a marked relationship between age class and extra-pair fertilizations (EPFs), with young males losing paternity of 43% (n = 53) of their putative offspring compared to 4% (n = 85) by old males. All assignable extra-pair offspring were sired by old males, with one male obtaining most EPFs each year. Contrary to the hypothesis that EPCs are forced, EPF frequency within age class did not increase with seasonal increases in the number of males per fertile female. Whereas the male control hypothesis predicted that the male age class that mate-guarded more would be cuckolded less, the reverse was true: young males guarded significantly more intensely. The male age class difference in cuckoldry could not be explained by the possibility that young and inexperienced females (which are usually paired to young males) were more vulnerable to forced copulation because EPFs were unrelated to female age. These findings suggest that females (1) pair with old males and avoid EPCs, or (2) pursue a mixed mating strategy of pairing with young males and accepting EPCs from old males. The receptivity to EPCs by females paired to young males put them in conflict with their mates. Two factors determined the paternity achieved by young males: (1) the relative size of the male to the female, with young males achieving much higher paternity when they were larger than their mates, and (2) the intensity of mate-guarding. Both variables together explained 77% of the variance in paternity and are each aspects of male-female conflict. Given female receptivity to EPCs, mate-guarding can be viewed as male interference with female mating strategies. We conclude that EPCs are rarely or never forced, but the opportunity for females paired to young males to obtain EPCs is relative to the ability of their mates to prevent them from encountering other males. Evidence of mixed mating strategies by females, combined with other features of the martin mating system, is consistent with the female-driven “hidden lek hypothesis” of colony formation which predicts that males are drawn to colonies when females seek extra-pair copulations. Received: 23 March 1995/Accepted after revision: 14 January 1996     

Preference for male traits in female wolf spiders varies with the choice of available males,female age and reproductive state

Individual variation in female preference for male traits may influence mate choice, especially if benefits and costs of choosiness vary with the range of available males or reproductive timing. We examined variation in female preference for male leg tuft size in Schizocosa ocreata (Hentz) wolf spiders with video playback. Dichotomous (simultaneous) choice test experiments included all possible combinations of four stimuli (i.e., modified versions of the same video male stimulus): average tuft size (control), reduced (−25%), enlarged (+25%), and no tufts (removed). Females exhibited a directional preference for larger tuft size independent of the nature of the choice (except for reduced tufts vs no tufts where no difference was seen). Female preference in the short term (over a period of 4 days) was also highly repeatable for control vs reduced tufts, but not for control vs enlarged tufts. Responses of females in ‘no-choice’ presentations of a single (control) male stimulus varied with age post-maturity; females were less receptive in weeks 1 and 2, highly receptive at week 3, and less thereafter. Mated females were least receptive and most aggressive towards a male stimulus. Females offered choices repeatedly at different ages post-maturity consistently preferred the control male vs reduced tufts over all 3 weeks but varied in their preference for enlarged tufts vs control male. In the first 2 weeks, females preferred the enlarged tuft male stimulus, but showed no preference by the third week. Females tested in week 4 showed no preference in either choice. Results suggest that the potential interaction between female preference for male traits and female reproductive timing may be a critical consideration in mate choice.     

Mating with a kin decreases female remating interval: a possible example of inbreeding avoidance

Inbreeding depression is a relative decline in fitness in offspring of related parents. The magnitude of inbreeding costs varies among taxa and may increase under stressful conditions. Inbreeding tolerance is expected to be low and selection for inbreeding avoidance intense when both sexes invest substantially in shared offspring like in nuptial gift-giving butterflies. This is especially true for increasing mating rate for inbreeding avoidance as nuptial feeding decreases net costs of mating for females. We explored implications of inbreeding in the nuptial gift-giving green-veined white butterfly, Pieris napi. Compared to outbred ones, partially inbred (F = 0.25) eggs and neonate larvae had 25% lower hatching success and 30% lower survival until adult eclosion, respectively. Inbreeding was also associated with small size. Yet, the magnitude of inbreeding depression was independent of larval conditions. A lack of assortative mating and mating durations independent of mating type suggest that neither females nor males discriminate close relatives (r = 0.5) as mates. Indicative of a postcopulatory mechanism to avoid inbreeding, female remating intervals decreased following incestuous matings. Such a plastic response may affect the level of postcopulatory sexual selection as female remating interval (time between successive matings) is necessarily negatively correlated with mating rate (matings per unit time) and mating frequency (lifetime number of matings), and precopulatory mate choice appeared insignificant. Moreover, incest-induced shift in the phenotype towards the adaptive peak may contribute to the evolution of female mating rates, although alternative explanations for polyandry besides material benefits have rarely been invoked when nuptial feeding is involved.     

Understanding abundance patterns of a declining seabird: implications for monitoring.

The Kittlitz's Murrelet (Brachyramphus brevirostris) is a rare, non-colonial seabird often associated with tidewater glaciers and a recent candidate for listing under the Endangered Species Act. We estimated abundance of Kittlitz's Murrelets across space and time from at-sea surveys along the coast of Alaska (USA) and then used these data to develop spatial models to describe abundance patterns and identify environmental factors affecting abundance. Over a five-week period in the summer of 2005, we recorded 794 Kittlitz's Murrelets, 16 Marbled Murrelets (B. marmoratus), and 70 unidentified murrelets. The overall population estimate (N, mean +/- SE) during the peak period (3-9 July) was 1317 +/- 294 birds, decreasing to 68 +/- 37 by the last survey period (31 July-6 August). Density of Kittlitz's Murrelets was highest in pelagic waters of Taan Fjord (18.6 +/- 7.8 birds/km2, mean +/- SE) during 10-16 July. Spatial models identified consistent "hotspots" of Kittlitz's Murrelets, including several small areas where high densities of murrelets were found throughout the survey period. Of the explanatory variables that we evaluated, tidal current strength influenced murrelet abundance most consistently, with higher abundance associated with strong tidal currents. Simulations based on the empirically derived estimates of variation demonstrated that spatial variation strongly influenced power to detect trend, although power changed little across the threefold difference in the coefficient of variation on detection probability. We include recommendations for monitoring Kittlitz's Murrelets (or other marine species) when there is a high degree of uncertainty about factors affecting abundance, especially spatial variability.     

Mating in the red-sided garter snake,Thamnophis sirtalis parietalis: differential effects on male and female sexual behavior

Summary Female red-sided garter snakes, Thamnophis sirtalis parietalis, become unattractive to most males after mating in the field and in the laboratory. Male red-sided garter snakes vary in their latencies to court attractive females following copulation, with courtship resuming in minutes to hours. Unsuccessful males in mating balls disperse from mating pairs, but are not residually inhibited from courting attractive females. These patterns of behavior indicate that males have evolved mechanisms to maximize opportunities for copulation with several females, while females mate only once per season.     

Measuring mate choice using correlation: the effect of female sampling behaviour

For intersexual selection to occur, it is necessary that females choose between males. It is now well appreciated that constraints exist, which preclude females sampling all the available males in a population. These constraints are likely to have caused the evolution of sampling rules (such as the “best-of-n” rule) by which females sample males. Here we investigate the impact of female subsampling of the male population, not on the evolution of sampling behaviour, but on the population-level correlation between a male trait and currencies such as reproductive success. This study is important as it illustrates when population-level correlations can be safely used to infer the presence and strength of sexual selection in the field. We find that the correlation between a male trait and a mate choice variable rises steeply as the number of males sampled by each female increases, flattening above seven to ten males sampled. This shape is found to be remarkably robust, and little affected by, for example, the mate choice variable used, by noise in assessment, by sampling behaviour depending on female quality, or by population size. The only variable found to have a large impact is male clumping according to their “quality”. If females are sampling about four males, the maximum correlation that can be found at the population level is in the range 0.4–0.6, perhaps as little as 0.1 if males are strongly clumped. A recent review of the literature suggests that four is the average number of males that females sample. Thus, the absence of a strong correlation cannot by itself be used to infer that sexual selection is weak, as it may be due to females sampling few males. Received: 18 May 1998 / Accepted after revision: 18 July 1998     

Social and ecological influences on activity budgets of vervet monkeys,and their implications for group living

Summary Activity patterns were documented over a 20-month period in six groups of vervet monkeys (Cercopithecus aethiops) in Amboseli National Park, Kenya. Group size varied both among groups and within groups through time. The home ranges of two groups were located in Acacia xanthophloea habitat whereas those of four adjacent groups were located in A. tortilis habitat. Repeated measures analysis of variance, three-way analysis of variance, and meta-analysis of group size effects were carried out on feeding, moving, resting, scanning, allo-grooming, and other behavior. Time spent feeding varied seasonally, peaking during periods when A. tortilis seeds were eaten. However, time spent feeding did not vary within or among groups, or between habitats, males and females, or dominants and subordinates. Males spend more time scanning and less time allo-grooming than females, and high-ranking individuals of both sexes spent more time scanning than low-ranking individuals. Among females, subordinates spent more time moving, whereas among males, dominants spent more time moving. Groups living in A. xanthophloea habitat spent more time moving and allo-grooming than groups in A. tortilis habitat but this may have been confounded by the fact that these two groups were on average larger than the others. Significant differences in activity budgets among groups demonstrate that activity patterns observed in one group are not always readily generalized to other groups even when they come from the same population. Group size analyses examined potential costs and benefits of group living. Increased intragroup competition, measured by time spent moving and feeding, was only weakly evident in larger groups. Individuals in larger groups allo-groomed more than individuals in smaller groups. Self-grooming also increased in larger groups. Individuals in larger groups spent less time scanning than individuals in smaller groups, but there was more scanning per group in larger groups than in smaller groups. Correspondence to: L.A. Isbell     

Nectar secondary compounds affect self-pollen transfer: implications for female and male reproduction

Pollen movement within and among plants affects inbreeding, plant fitness, and the spatial scale of genetic differentiation. Although a number of studies have assessed how plant and floral traits influence pollen movement via changes in pollinator behavior, few have explored how nectar chemical composition affects pollen transfer. As many as 55% of plants produce secondary compounds in their nectar, which is surprising given that nectar is typically thought to attract pollinators. We tested the hypothesis that nectar with secondary compounds may benefit plants by encouraging pollinators to leave plants after visiting only a few flowers, thus reducing self-pollen transfer. We used Gelsemium sempervirens, a plant whose nectar contains the alkaloid gelsemine, which has been shown to be a deterrent to foraging bee pollinators. We found that high nectar alkaloids reduced the total and proportion of self-pollen received by one-half and one-third, respectively. However, nectar alkaloids did not affect female reproduction when we removed the potential for self-pollination (by emasculating all flowers on plants). We then tested the assumption that self-pollen in combination with outcrossed pollen depresses seed set. We found that plants were weakly self-compatible, but self-pollen with outcrossed pollen did not reduce seed set relative to solely outcrossed flowers. Finally, an exponential model of pollen carryover suggests that high nectar alkaloids could benefit plants via increased pollen export (an estimate of male function), but only when pollinators were efficient and abundant and plants had large floral displays. Results suggest that high nectar alkaloids may benefit plants via increased pollen export under a restricted set of ecological conditions, but in general, the costs of high nectar alkaloids in reducing pollination balanced or outweighed the benefits of reducing self-pollen transfer for estimates of female and male reproduction.     

A mechanism for female choice of large males in the treefrog Hyla chrysoscelis

Summary In this study, we combine both field and laboratory experiments to address the effects of female preference for certain call characteristics on a large-male mating advantage in the treefrog H. chrysoscelis. In laboratory-choice experiments, females always chose the call with the lower fundamental frequency when call rate and call intensity were held constant and the difference in frequency between the two calls was 15%. The lower frequency call was preferred by 8 out of 12 females when the difference in fundamental frequency was 7.4%. These results are consistent with field comparisons of the size of unmated males calling within 2 m of a mated male: Male body size was negatively correlated with fundamental frequency and the greater the size difference, the more likely that the larger male mated. In field choice experiments, females preferred males with higher call rates. Since size differences between males used in this experiment averaged only 2.3 mm, we would not expect the fundamental frequency of a male's call to be the best predictor of mating success. Laboratory results demonstrated that call rate could override female preference for the low frequency call over the high frequency call, while intensity could at least dilute this preference. However, individual males in the field varied both call rate and the call intensity as perceived by the female. We suggest that the interaction between call rate, male size and mating success should be studied further through the use of field-choice experiments.     

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a female's decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.     

Behavioural responses of Diana monkeys to male long-distance calls: changes in ranging,association patterns and activity

Although much is known about the relationship between vigilance, group size and predation risk, behavioural responses to predation risk and their resultant costs are less clear. We investigated the response of Diana monkeys to increased predation risk by looking at behavioural changes associated with male long-distance calls, which are reliably given to certain predators. After male long-distance calls, group spread and nearest-neighbour distance decreased whilst travel and association rates for the group increased. The average height and exposure level of individuals in the group did not change after calls. Individual Diana monkeys changed their behaviour and were more likely to be vigilant or travel and less likely to engage in social or resting behaviours after long-distance calls. In addition, movement rates increased with the number of species the Diana monkeys were associated with. Diana monkey long-distance calls facilitate the joining of groups of other species. Black and white colobus and lesser spot-nosed monkeys were more likely to be in an association following a long-distance call than before. Behavioural responses, such as increased travel or association rates, that reduce foraging efficiency are interpreted as evidence of a non-lethal impact of increased predation risk.