Effects of male sexual harassment on female time budgets,feeding behavior,and metabolic rates in a tropical livebearing fish (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>)

Sexual harassment by males has the potential to affect almost any aspect of female behavior and life history. Using Atlantic mollies (Poecilia mexicana) as a model—a species in which males do not court but almost constantly try to forcefully mate with females—we asked whether and how male harassment influences (a) females’ time budgets and (b) feeding rates (e.g., through frequent flight from male approaches), and (c) whether metabolic rates are increased as a response to stress. Field observations in a natural P. mexicana population revealed that males (average feeding rate 15%) spent far less time feeding than females (60%), and clearly traded off frequent pursuit of females with foraging. Most importantly, females’ feeding times were dramatically reduced when being pursued by a harassing male. Also in standardized lab experiments, females spent significantly less time feeding when accompanied by a male as compared to being in the presence of another female. This effect was also observed when partner fish (male or female) were presented only visually, but could not interact physically with the focal female. It seems, therefore, that females increase vigilance when a harassing male is around, which keeps them from feeding even before males actually approach them. Based on the latter result, we asked whether a stress-induced increase in metabolic rates would be discernible. We measured oxygen consumption and gill ventilation frequencies (opercular rates) of females in different social contexts (alone, with another female, or a male). The predicted, strong body mass dependency of both physiological parameters was uncovered, but no evidence for an effect of social context was detected. We argue that male harassment represents such a constant (but non-lethal) stressor for poeciliid females that their metabolic stress responses have adapted to this through habituation.     

Age- and state-dependent reproductive effort in male mountain goats, Oreamnos americanus

In mammalian polygynous mating systems, male reproductive effort consists mainly of male–male competition and courting of females, which entail substantial somatic costs. Males are thus expected to adjust their reproductive effort according to their age and condition. In this study, we examined how activity budgets of male mountain goats (Oreamnos americanus), a polygynous ungulate, varied with age in a marked population over two periods: (1) summers 1995–2006 and (2) ruts 2004–2006. We then assessed if the proportions of time spent in male–male competition and courtship behaviors were influenced by age-specific body mass and social rank during the rut. Males spent most of their time foraging and resting during summer, and rested more and foraged less with increasing age. During the rut, pronounced shifts in activity budgets occurred as juveniles (1–2 years) increased time spent foraging, whereas adults (≥3 years) increased standing and time spent in social interactions at the expense of foraging. At old age, reproductive effort either stabilized or decreased slightly, providing weak support for the ‘mating strategy–effort’ hypothesis, predicting that courtship behaviors should peak in prime-aged males. Age-specific body mass did not affect time spent in male–male competition, but was positively related with time spent in courtship behaviors, providing support for the ‘individual quality’ hypothesis, predicting that males with more resources at the start of the rut should spend more time in mating-related activities. Age-specific social rank did not affect reproductive effort. Surviving to prime age while increasing mass each year should thus allow male ungulates to gain greater ability to court estrus females.     

Males are faster foragers than females: intersexual differences of foraging behaviour in the Apennine chamois

Availability of food resources and individual characteristics can influence foraging behaviour, which can differ between males and females, leading to different patterns of food/habitat selection. In dimorphic species, females are usually more selective in food choice, show greater bite rates and spend more time foraging than males. We evaluated sexual differences in foraging behaviour in Apennine chamois Rupicapra pyrenaica ornata, during the warm season, before the rut. Both sexes selected nutritious vegetation patches and spent a comparable amount of time feeding. However, males had a significantly greater feeding intensity (bite rate) and a lower search effort for feeding (step rate), as well as they spent more time lying down than females. Females selected foraging sites closer to refuge areas than males. In chamois, sexual size dimorphism is seasonal, being negligible in winter–spring, but increasing to 30–40 % in autumn. Our results suggest that males enhance their energy and mass gain by increasing their food intake rate during the warm season, to face the costs of the mating season (November). Conversely, females seem to prioritize a fine-scale selection of vegetation and the protection of offspring. A great food intake rate of males in the warm season could have developed as a behavioural adaptation leading herbivores to the evolutionary transition from year-round monomorphism to permanent dimorphism, through seasonal dimorphism.     

Vocalizations and courtship intensity correlate with mounting success in marginated tortoises<Emphasis Type="Italic"> Testudo marginata</Emphasis>

The courtship and mounting behaviour of tortoises is elaborate, and based on a multiple signalling system involving visual, olfactory and acoustic signals. Vocalizations related to mounting seem to be particularly significant because tortoises vocalize mainly at this time. Vocalizations and courtship behaviour may be costly for males, and if these costs increase differentially for different males, then the potential exists for vocalizations and displays to reveal male individual quality. In this correlative study, we analysed relationships between male mounting success and morphological and behavioural traits, particularly acoustic signals, exhibited by male marginated tortoises (Testudo marginata) during courtship, in a group of 94 individuals breeding in semi-natural enclosures. For each male, we estimated general body condition, courtship intensity and mounting success; calls of mounting males were recorded and four sonagraphic features were measured. Calls differed significantly among males, and two features varied according to body condition. Male mounting success significantly increased according to the male/female size-ratio, suggesting the existence of a size-based assortative mating. Mounting success was also highly correlated with courtship intensity, measured as number of bites and rams given to females before mounting, and with number of calls emitted during mounting. Finally, mounting success was negatively related to call duration. To our knowledge, this is the first study in which features of tortoise vocalizations are shown to convey reliable information about male quality in socio-sexual contexts.Communicated by T. Czeschlik     

Changes in the activity budgets of yellow baboons (Papio cynocephalus) during sexual consortships

Summary The energetic cost of guarding estrous females has been assumed to be a potentially major factor in limiting the duration of male consortships in a number of polygamous species. This issue has been examined in the present study by assessing changes in the usual activity budgets of 13 male and 20 female yellow baboons which occurred during their consortships.The percent of time adult males spent feeding was strongly related to their agonistic rank: Higher-ranking males fed proportionately more than lower-ranking males while not in consort, and they showed a relatively greater decrease in feeding time during consortships.The proportion of time females spent feeding was related to their agonistic rank and (to a lesser degree) to their parity. Higher-ranking females fed more than lower-ranking females outside of consortships, and showed a relatively greater decrease in time spent feeding while in consort. Nulliparous females tended to feed more than parous females while not in consort; during consortships, the feeding time of nulliparous females decreased, while that of parous females increased slightly.The activity of male and female consort partners was most coordinated during the fertile cycle stage of the female partner, and least coordinated once the female had passed the fertile stage.Although reductions in feeding time during consortships were generally much more pronounced in males than in females, neither males nor females appeared to attain the undisturbed optimal feeding level for their sex while in consort.     

Influence of male harassment and female competition on female feeding behaviour in a sexual–asexual mating complex of mollies (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, <Emphasis Type="Italic">P. formosa</Emphasis>)

Differential interests between the sexes regarding the number of copulations can result in sexual harassment. Hence, females may have less time available for foraging. Male sexual harassment often leads to fitness reduction in females. We used the mating complex of the bisexual fish Poecilia mexicana and the co-occurring all-female Poecilia formosa to study sexual harassment and its incurred cost on female feeding efficiency. P. formosa is a sperm-dependent parthenogen that requires mating with host males to induce embryogenesis, but the male genes are not used. We therefore predicted P. mexicana males to prefer conspecific females. Hence, costs of male sexual harassment should not occur in unisexuals. While P. formosa are at a disadvantage compared to P. mexicana females due to male mate choice (leading to sperm limitation), this could be traded-off by suffering less from sexual harassment. In our experiment, we found males to direct significantly more pre-copulatory mating behaviour towards conspecific females, whereas actual mating attempts did not differ between species. Contrary to our prediction, both types of females started feeding later and spent less time feeding in the presence of a male partner compared to the time spent feeding with another female, suggesting that females of both species suffer from male harassment. The focal females' feeding time declined with increasing body size of the female competitor, and the same pattern was found when a male was present. We discuss that—besides sexual harassment—other factors such as food competition and female mate choice may affect female feeding efficiency.     

Energy expenditure and food consumption of foraging Imperial cormorants in Patagonia, Argentina

Energy management during the breeding season is crucial for central place foragers since parents need to feed themselves and their offspring while being spatially and temporally constrained. In this work, we used overall dynamic body acceleration as a measure of activity and also to allude to the foraging energy expenditure of breeding Imperial cormorants Phalacrocorax atriceps. We also analyzed how changes in the time or energy allocated to different activities affected the foraging trip energy expenditure and estimated the daily food requirements of the species. Birds spent 42 % of the total energy flying to and from the feeding areas and 16 % floating at sea. The level of activity underwater was almost 1.5 times higher for females than for males. The most expensive diving phase in terms of rate of energy expenditure was descending though the water column. The total foraging trip energy expenditure was particularly sensitive to variation in the amount of time spent flying. During the breeding season, adult cormorants breeding along the Patagonian coast would consume approximately 10,000 tons of food.     

The effect of reproductive condition on the foraging behavior of female hoary bats,Lasiurus cinereus

Summary Female mammals experience larg changes in time and energy budgets associated with reproduction and these may influence the foraging strategies of individuals. I studied the changes in foraging behavior associated with reproduction in female hoary bats, Lasiurus cinereus. As lactation progressed, individuals departed to forage earlier in the evening and spent more time foraging per night and less time roosting with their young. Foraging time increased by at least 73% between early lactation and fledging and then declined as the young became independent. Females with two young foraged for longer than did those with one and females with pre- and postfledging young foraged in different habitats. The changes in foraging time suggest that foraging activity of female L. cinereus is constrained and individuals act as time minimizers, adjusting their foraging behavior to meet current energy demand. Predation risk is unlikely to constrain the behavior of these bats. However, maximizing energy intake throughout lactation may not be the optimal strategy because storing excess energy increases flight cost and may reduce foraging efficiency. The need to keep newborn young warm may also influence foraging time. Such constraints, causing changes in foraging activity, may alter the availability of habitats and prey and must be considered when modelling foraging strategies. In addition, changes in flight time may significantly alter the energy budgets of bats in different stages of reproduction.     

Evidence for division of labor in the social caterpillar Eucheira socialis (Lepidoptera: Pieridae)

The caterpillars of Eucheira socialis westwoodi cooperatively spin and maintain a hollow silken nest and an elaborate network of silken foraging trails on their host plant, madrone (Arbutus spp.: Ericaceae). Nests typically contain several hundred larvae. Two populations are known to harbor a sex ratio distorter. The primary sex ratio in these two populations for four generations has been exceedingly male biased (64–79% male). Lepidoptera larvae are easily sexed using external morphology, allowing us to uniquely mark male and female larvae and to assemble larval groups of particular sex ratios. We report here the results of experiments on sex-specific larval behavior and physiology and the effect of colony sex ratio on individual behavior. We found that male larvae spent more time spinning silk on the nest and less time feeding than female larvae. Males were the first to emerge from the nest and the first to venture out along trails to feed. Male-biased nests had a significantly greater amount of silk deposited on their surfaces than female-biased nests. In the field, male-biased nests produced heavier male and female pupae than female-biased nests. Male and female larvae in 75% male nests became active earlier than males and females in other sex ratio treatments. Received: 11 September 1998 / Received in revised form: 24 February 1999 / Accepted: 27 March 1999     

The special role of male Cebus monkeys in predation avoidance and its effect on group composition

Summary Vigilance behavior, predator detection abilities, and responses to real and model predators were studied in two species of capuchin monkeys (Cebus albifrons and C. apella) in a Peruvian lowland rain forest. Adult males were more vigilant than adult females in both species, mainly because the males spent less time feeding and foraging and partly because they were at the periphery more often than the females. The increased vigilance of adult males is reflected in their superior performance in the detection of (model) predators. Adult and subadult males were also far more likely to approach and mob real and model predators. Adults that were outside the center of the group increased foraging activities but cut back an feeding, much of which was done in exposed tree crowns. Current theory suggests that primate groups are multi-male when a single male is unable to defend sexual access to the group of females. In these small capuchin groups, which are multimale, the enhanced safety of females and young provided by extra adult males furnishes a more plausible explanation. A comparison of the two capuchins with the ecologically similar Southeast Asian Macaca fascicularis suggests that the high predation risk outside the group may also have caused the unusual male career profile in capuchins, which have a long tenure of dominants and a very long potential lifespan. Further predictions of this hypothesis are developed.     

Effects of sex-associated competitive asymmetries on foraging group structure and despotic distribution in Andean condors

Phenotype-limited interference models assume competitive asymmetries among conspecifics and unequal sharing of resources. Their main prediction is a correlation between dominance status and patch quality: dominant individuals should preferentially exploit better-quality habitats. We tested assumptions and predictions of the phenotype-limited interference model in Andean condors (Vultur gryphus), a New World vulture with strong sexual size dimorphism (males are 30–40% heavier than females). We recorded searching birds in habitats differing in quality: mountains and plains. We also observed scavenging behaviour at 20 sheep carcasses, and videotaped 5 of them. Intraspecific hierarchy at carcasses was based on size: males dominated females and, within each sex, older birds dominated younger ones. Adult males and juvenile females occupied extreme positions in the feeding hierarchy. Aggression was directed at those individuals belonging to lower hierarchical levels. In high-quality areas (mountains), more condors arrived at carcasses. Juvenile females were more often observed searching in low-quality areas (plains), far from breeding areas and main roost sites. GLM analyses of individual behaviour showed that the hierarchy did not influence time of arrival, but low-ranking individuals spent more time at carcasses, especially if the number of condors at arrival was high. Additionally, low-ranking condors spent less time feeding at carcasses when individuals of higher hierarchical levels were present. On the other hand, the number of condors present had a positive effect on feeding rates of dominant individuals, probably because of a reduction in individual vigilance. These results support most of the assumptions and predictions of the phenotype-limited distribution model, although a spatial truncated distribution between phenotypes was not observed. Asymmetric feeding pay-off, unequal parental roles and sexual selection constraints could favour sexual divergence in body size in Andean condors. Received: 6 April 1998 / Accepted after revision: 11 July 1998     

Food or sex—males and females in a sex role reversed pipefish have different interests

In a sex role reversed pipefish, Syngnathus typhle, we found that basic life history allocations were directly influenced by sexual selection. We investigated time allocation to foraging and mating, respectively, in a choice experiment, giving males and females, of small or large body size, a choice between food and a potential partner. We found that males were more interested in foraging than mating, i.e., were more frequently observed in front of the food than in front of the partner, whereas females were more interested in the potential partner. This reflects sexual selection operating differently on the two sexes, as males and females are relatively similar in other life history traits, such as growth, mortality, age of maturity, dispersal, and parental expenditure. Moreover, large individuals allocated more time to mating activities, small to feeding. Individuals more interested in mating compared to food were subsequently more critical when given a choice between a large (high-quality) and a small (low-quality) partner, whereas individuals more interested in food were not selective. These findings are consistent with our predictions: sex-role reversed males can be relatively sure of achieving one or more matings, and should allocate more time to feeding and, hence, to parental investment, growth and/or future reproduction. Females, on the other hand, have more uncertain mating prospects and should allocate time to imminent reproductive activities, thereby foregoing other life history traits such as growth and future egg production. By this, they also sacrifice future fecundity and attractiveness.     

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     

Timing of the evening emergence from day roosts of the grey-headed flying fox, Pteropus poliocephalus: the effects of predation risk, foraging needs, and social context

This study addresses the functional question of how variation in foraging strategy, predation risk, and social context influence the timing of the evening emergence from day roosts of the grey-headed flying fox, Pteropus poliocephalus. The onset of evening emergence was expected to vary according to the relative costs and benefits of emerging early and should, therefore, reflect an optimal trade-off between predation risks and foraging needs. The onset of the colony-wide emergence was closely correlated with the time of sunset and cloud cover. However, as expected, the onset of the colony-wide emergence was delayed when a diurnal avian predator was present, whereas the onset was advanced during lactation when presumably energetic demands are higher. The trade-off between predation risks and foraging needs was further reflected in the emergence times of individual bats: adult females emerged earlier when they had higher foraging needs as indicated by their body condition; young emerged later when they were smaller and likely to be more at risk from predation due to their less developed flying skills. However, the emergence time of adult males depended on their social status: smaller bachelor males emerged from the colony earlier than larger harem-holding males who guard their harems until the last female had left. Thus, whereas the colony-wide emergence time reflected the outcome of a trade-off between predation risks and general foraging needs, on an individual level, the outcome of this trade-off depended on sex, age, body condition, and structural size and was modified by social context.     

Effects of male removal on female foraging behavior in the Eurasian treecreeper

In old, spruce-dominated forests of central Finland, Eurasian treecreepers Certhia familiaris divide their territories spatially during the breeding season. Females forage primarily on the upper parts of the tree trunks, while males use the lower parts of the tree trunks. In this study we removed males from eight territories in the early nestling period to see if the mate's absence would change the foraging patterns of the resident female. Widowed females foraged at lower heights, thus behaving more like paired males. These females also spent less time on each tree and on each foraging bout than did paired females. We conclude that male removal facilitated the change in a female's foraging niche and foraging time at the trees. Females may re-optimize their foraging site selection owing to the absence of dominant males and a consequent need to increase their parental care. Behavioral plasticity may be the mechanism of niche partitioning between the sexes in this species. Received: 28 June 1996 / Accepted after revision: 27 March 1997     

Bite rates in Rocky Mountain bighorn sheep (<Emphasis Type="Italic">Ovis canadensis</Emphasis>): effects of season,age, sex and reproductive status

We investigated the effects of vegetation biomass, crude protein content of consumed forage, age, sex and reproductive status on bite rates in Rocky Mountain bighorn sheep. We expected higher bite rates and vigilance in lactating females with young and higher bite rates in young growing individuals, than in non-reproducing females or rams. Lactating ewes had higher bite rates than yeld ewes and than subadult or adult rams. Subadult rams had higher bite rates than adult rams or yeld ewes. On recently burned grassland in spring, however, rams had a higher bite rate than adult females, while the contrary was true on control plots and on the burned plots in autumn. Bite rates declined for both ewes and rams from April to September and varied from year to year. While rams of different ages had significantly different bite rates, there was no effect of age on bite rates for ewes. There was no correlation between bite rates and available total biomass or biomass of live vegetation, or the numbers of steps taken while foraging for either ewes or rams. Adult rams had a lower vigilance rate than adult ewes, and vigilance decreased with increasing bite rates for all sheep. Bite rates in bighorn sheep vary greatly according to age, season and vegetation structure. An increase in bite rates during the forage growing season may compensate for higher energy demands during lactation and growth. There is a potential trade-off between foraging and vigilance as vigilance decreased with increasing bite rates.     

The reproductive behavior and energetics of male gray seals (Halichoerus grypus) breeding on a land-fast ice substrate

The reproductive behavior of male gray seals (Halichoerus grypus) breeding on land-fast ice at Amet Island, Nova Scotia, was studied. Data on energy expenditure (rate of mass loss over time) were collected. The average time budget of males at Amet Island was comparable to that of land-breeding males. The behavior of males showed seasonal changes, with a decrease in the proportion of time spent in the water and an increase in agonistic behavior during the peak mating period. The estimated amount of body mass lost over the season ranged between 25.6 and 77.1 kg, and the estimated percent of initial body mass lost ranged between 7.7 and 26.5% (n=10). The maximum number of observed copulations for an individual male was nine. Only 15 out of 42 males observed during 1992 and 1993 were seen copulating. The number of observed copulations per male was strongly correlated with success in remaining close to, or attending, females (r=0.91, P<0.001, n=42). The mean duration of attendance was 4.5 ± 5.54 days (n=42). Large size was not an important factor in determining attendance success, but reproductive effort (the estimated proportion of body mass lost over the season) and success in agonistic interactions with other males were both correlated with male success.     

Threat-sensitive foraging by larval threespine sticklebacks (Gasterosteus aculeatus)

Summary The threat-sensitive predator avoidance hypothesis predicts that prey can assess the relative threat posed by a predator and adjust their behaviour to reflect the magnitude of the threat. We tested the ability of larval threespine sticklebacks to adjust their foraging in the presence of predators by exposing them to conspecific predators of various sizes and recording their foraging and predator avoidance behaviours. Larvae (<30 days post-hatch) displayed predator escape behaviours only towards attacking predators. At 3 weeks post-hatch larvae approached the predator after fleeing, a behaviour which may be the precursor to predator inspection. Larvae reduced foraging and spent less time in the proximity of large and medium-sized predators compared to small predators. The reduction in foraging was negatively correlated to the predator/larva size ratio, indicating that larvae increased their foraging as they increased in size relative to the predator. We conclude that larval sticklebacks can assess the threat of predation early in their ontogeny and adjust their behaviour accordingly.Correspondence to: J.A. Brown     

Impacts of flight distance on sex ratio and resource allocation to offspring in the leafcutter bee, Megachile rotundata

Fisher's theoretical prediction of equal investment in each sex for a panmictic population (The genetical theory of natural selection. Clarendon, Oxford, 1930) can be altered by a number of factors. For example, the sex ratio theory predicts variation in equal investment in each sex when the maternal fitness gains from increased investment differ between sexes. Changing sex allocation because of changing payoffs may result from different ecological situations, such as foraging conditions. We investigated the impact of foraging travel cost on relative investment in sons vs daughters. Field studies were carried out with the central-place-foraging leafcutter bee Megachile rotundata (Fabricius), which has smaller males than females. Therefore, less investment is required to produce a viable son compared with a daughter. We found that with increased flight distance to resources, females produced a greater proportion of sons. Females also invested fewer resources in individual sons and daughters and produced fewer offspring with increased flight distance.     

Male mating competitiveness and age-dependent relationship between testosterone and social rank in bighorn sheep

In males, the acquisition and development of behavioral and morphological secondary sexual traits typically depends on testosterone and correlates with mating success. Testosterone level could affect competition for mates and thus be a target of sexual selection. We sought to relate testosterone levels to male mating competitiveness, by teasing apart the relationships between testosterone, behavior, and growth before the mating period. We monitored 24 adult bighorn rams (Ovis canadensis) at Ram Mountain, Alberta, from 2008 to 2011. Using linear mixed models, we tested the relationships between testosterone metabolites in feces, social rank, and both growth and size of two sexually selected traits: horns and body mass. The correlation between testosterone and social rank varied with age. Testosterone and rank were weakly and negatively correlated for young rams, positively correlated for prime-aged rams, and negatively correlated for older rams. Although testosterone had an increasingly positive effect on total horn length until 8 years of age, we could not detect any effects on annual growth rate of horns or body mass. Testosterone may be related to male’s ability to compete for mates through its relationship with behaviors determining social rank, rather than by influencing the development of morphological traits. Differences in testosterone levels among competitors may be a proximate cause of variance in fitness.