Female House Mice do not Differ in Their Exploratory Behaviour from Males

Dispersal is an important ecological trait affecting genetic variation and dynamics of populations. Hence, the exploratory behaviour prior to actual dispersal may be crucial for potentially dispersing individuals. In mammals, females are traditionally seen as the more philopatric sex and dispersal as male‐biased behaviour, and so behavioural strategies related to the exploration of novel resources should be differentially expressed in males and females. In addition, due to sexual selection exploratory strategies may be expected to vary according to females’ reproductive phase. We employed a standard open‐field test as an approximation of the first phase of dispersal, using adult house mice representing two subspecies, M. m. musculus and M. m. domesticus. We tested the prediction that exploration of neutral area varies in females during different phases of the oestrus cycle and is different between both sexes and subspecies. We expected to find higher exploration in males, as the more dispersing sex and less pronounced subspecies‐specific differences in females than in males. We found no significant effect of the oestrous phase on any of the parameters of the exploratory behaviour measured. Sexual dimorphism was found only in latency to enter the arena in M. m. domesticus where females hesitated longer to enter a new area than males. Significant subspecies‐specific differences were found in three of four tested exploration parameters, so we conclude that females of both subspecies follow similar strategies to those displayed by males. Musculus mice show shorter latency to enter a new area, but once inside, domesticus mice explore the arena significantly longer, with less frequent retreats to a shelter. Our results thus highlight that the role of female dispersal in interdemic gene flow should not be neglected.     

Sex Differences in Discrimination of Shoal Size in the Guppy (Poecilia reticulata)

For a diversity of species, differences in sexual and parental roles, along with differences in body morphology, often result in males and females having different diets, distinct predators and even different patterns of habitat use. As a consequence, the two sexes often face different environmental challenges and selection may favour the evolution of sex differences in cognition. We tested this prediction in the guppy (Poecilia reticulata). Under perceived hazard, individual guppies join the larger available social group, a behaviour that is thought to minimise predation risk. In this species, females are more frequently exposed to predation and more averse to predation risk; we therefore expected greater accuracy in shoal size discrimination in females. We compared the accuracy of male and female guppies in discriminating shoals of 4 and 6 conspecifics, which represents the upper limit of discrimination for this species. Overall, we found no sex differences in the accuracy of discriminating the two shoals. However, while females showed this ability at the beginning of the test, males began to select the larger group only after several minutes. In three control experiments, we found indications that this sex difference cannot be accounted for by differences in motivation or antipredator strategies between the two sexes, suggesting female guppies are more efficient at rapidly estimating shoal size.     

The behavioural response of Great Tits to novel environment and handling is affected by the DRD4 gene

Studies have provided inconsistent evidence regarding the effects on behaviour of nucleotide polymorphism at the DRD4 gene. For instance, the DRD4 gene affects exploratory behaviour in some bird species and populations but not in others. Our earlier experiment with a wild Great Tit Parus major population revealed an important impact of the DRD4 gene on behavioural variation during the breeding season. The aim of this study was to clarify these results by measuring individual behavioural differences (exploration and anxiety) in Great Tits under controlled conditions before the beginning of egg‐laying in order to exclude the potential impacts of breeding (egg‐laying, motivation to feed nestlings, the potential effect of a partner) and the natural environment (e.g. the presence of predators). Birds were caught from their natural habitat and their behaviour was tested in a trial room the following morning. Individuals with the CT heterozygous genotype (SNP830) performed more actively in different contexts (visiting a novel object in the trial room and vocal behaviour during handling) compared with homozygotes, and the relationships were consistent in males and females. In light of the results from our previous study (heterozygous males, but not females, had a shorter latency to feed nestlings than CC homozygotes), we conclude that the same DRD4 gene variants affect behavioural responses in a controlled environment and wild contexts in the same directions, but that these relationships are not sex‐specific under standardized conditions.     

Fitness Costs of Sexual Harassment–The Price of Persuasion

Females are often subjected to unwanted mating advances from males. Such advances can be costly to both parties. The short‐term costs of harassment to females have been widely explored in the literature; however, few studies have measured the direct fitness costs. Moreover, male costs are seldom considered. Conventional wisdom would lead us to hypothesise that sexual harassment is costly; thus, when males and females are housed together, harassment should reduce foraging, growth and reproductive output and may disrupt social interactions. This study quantified harassment costs in both sexes by observing behavioural responses and long‐term effects of unsolicited mating in a controlled setting. Sexually mature guppies were subjected to two housing treatments: equal sex ratios or single‐sex groups. The effects of male harassment on males and females were assessed by measuring behaviour, growth rate and the number of offspring produced over a period of 6 mo. Contrary to our expectations, our results indicated no significant differences in foraging and growth rates between mixed‐ and single‐sex shoals for either sex. Moreover, there was no significant difference in fry production between mixed‐ and all‐female shoals. Large males showed higher mortality when housed with females. Both sexes showed a reduction in shoaling when in mixed‐sex groups. Thus, there appear to be few direct costs of harassment for females in natural, mixed‐sex shoals, but males appear to bear significant harassment costs. The study provides insights into reproductive behaviour and life‐history traits.     

Zebra finches are bolder in an asocial, rather than social, context

Despite an expanding interest in animal personalities, the influence of social interactions and sex differences on individual differences in behaviour remains poorly understood. Using the social zebra finch (Taeniopygia guttata), we tested for behavioural differences in exploration of a novel environment and objects, between individuals of both sexes in relation to a social context; the presence of three male companions, three female companions or no companion birds. We predicted that the presence of conspecific companions should result in focal birds reacting to novelty by exploring more extensively because the companion birds contribute to anti-predator vigilance behaviour and because social isolation often causes behavioural inhibition in social species. We found that exploratory behaviour of focal individuals was significantly reduced in the presence of conspecific companions, irrespective of the companion's sex. Moreover, we found a weak trend towards females being more exploratory than males, irrespective of the social context. These results demonstrate the importance of considering the social context in animal personality studies and of exploring sex differences in personalities.     

The effect of sex and shoal size on shoaling behaviour in Danio rerio

Male and female zebra fish Danio rerio were given choices of shoals that differed in sex and size. Male zebra fish preferred to associate with female shoals over male shoals, but had no preference when given a choice between a mixed‐sex shoal and either a male or female shoal. Female zebra fish showed no significant preference when given a choice between male and females shoals, nor between mixed‐sex shoals and either male or female shoals. When given choices between shoals of differing size, females preferred to associate with the larger shoal, whether or not they were composed of males or females. Males, however, had no preference for larger shoals over smaller shoals, whether or not they were composed of males or females. These results showed that male zebra fish were capable of distinguishing between males and females solely on the basis of visual cues. Furthermore, these results demonstrated a significant difference between the shoaling choices of male and female zebra fish, which may indicate a difference in the function of shoaling for the two sexes.     

Do Female Zebra Finches,Taeniopygia guttata,Choose Their Mates Based on Their ‘Personality’?

A major challenge in behavioural and evolutionary ecology is to understand the evolution and maintenance of consistent behavioural differences among individuals within populations, often referred to as animal ‘personalities’. Here, we present evidence suggesting that sexual selection may act on such personality differences in zebra finches (Taeniopygia guttata), as females seem to choose males on the basis of their exploratory behaviour per se, while taking into account their own personality. After observing a pair of males, whose apparent levels of exploration were experimentally manipulated, females that exhibited low‐exploratory tendencies showed no preference during mate choice for males that had appeared to be either ‘exploratory’ or ‘unexploratory’. In contrast, intermediate and highly exploratory females preferred apparently exploratory males over apparently unexploratory ones. Our results suggest that behavioural or genetic compatibility for personality traits might be important for mate choice, at least for exploratory individuals.     

Choice of Female Groups by Male Mosquitofish (Gambusia holbrooki)

Even though females are usually more selective in choosing their mates, males are also capable of exercising mate choice. Despite the large body of evidence on the individual features preferred in sexual selection, little attention has been devoted to the first stage of male–female interaction. As poeciliid fish are known to be social, in the wild, initially mate choice may concern a preliminary selection among shoals. Only after this primary choice, males may subsequently direct their attention to a specific mate. We observed spontaneous preference of male mosquitofish (Gambusia holbrooki) when choosing between groups differing in size and sex ratio. In partial agreement with our predictions, males preferred to join a group of females rather than an isolated one (expt 1) and the larger group when two female groups were presented (expt 2). An all‐female group was preferred to a mixed‐sex group (expt 3), whereas no preference was observed when the two mixed‐sex groups differed in the number of males (expt 4) or in the size of the males (expt 5). These results suggest that male mosquitofish are capable of discriminating among different quantities of individuals within a group and use such information to select among groups in order to optimize the likelihood of successful matings.     

Environmental habituation and sexual composition affect juveniles' shoaling activity in a cichlid fish (Pelvicachromis taeniatus)

Group living is widespread in animals, and many fishes form shoals. Examining within-group interactions in fishes may contribute to the general understanding of dynamic social structures in animals. The sex ratio of a group has been shown to influence grouping decisions of fishes and can be expected to affect behaviour at group level. Behavioural experiments usually involve relatively short acclimatisation times, although the establishment of environmental habituation in fishes is understudied. This study tests whether the sex ratio and long-term habituation to experimental conditions influence general shoal performance (activity parameters, density) and responses of shoals to an acoustic-mechanical disturbance cue in juveniles of the cichlid fish Pelvicachromis taeniatus via individual tracking. The disturbance consisted of a defined hit against the experimental tank, which caused sudden noise and water movement. We found that a higher proportion of females increases shoal activity (swimming speed and distance covered), suggesting that female P. taeniatus are more active than males. Furthermore, shoal activity declined when shoals habituated to the experimental settings and with the time that the shoals were grouped together, which may reflect intensified group member familiarity. Moreover, behavioural changes after disturbance were weaker when individuals were kept with their group longer and more familiar to the experimental conditions. For prey species, lower activity might be beneficial under natural conditions due to lower conspicuousness of the group. We did not find any significant effects of the investigated factors on shoal density (mean interindividual distance) and speed synchronisation. The results indicate that sexual composition, familiarity between shoal members and habituation to the experimental environment affect shoal performance in a cichlid fish.     

Combining noninvasive genetics and a new mammalian sex‐linked marker provides new tools to investigate population size,structure and individual behaviour: An application to bats

Monitoring wild populations is crucial for their effective management. Noninvasive genetic methods provide robust data from individual free‐ranging animals, which can be used in capture–mark–recapture (CMR) models to estimate demographic parameters without capturing or disturbing them. However, sex‐ and status‐specific behaviour, which may lead to differences in detection probabilities, is rarely considered in monitoring. Here, we investigated population size, sex ratio, sex‐ and status‐related behaviour in 19 Rhinolophus hipposideros maternity colonies (Northern France) with a noninvasive genetic CMR approach (using faeces) combined with parentage assignments. The use of the DDX3X/Y‐Mam sexual marker designed in this study, which shows inter‐ and intrachromosomal length polymorphism across placental mammals, together with eight polymorphic microsatellite markers, produced high‐quality genetic data with limited genotyping errors and allowed us to reliably distinguish different categories of individuals (males, reproductive and nonreproductive females) and to estimate population sizes. We showed that visual counts represent well‐adult female numbers and that population composition in maternity colonies changes dynamically during the summer. Before parturition, colonies mainly harbour pregnant and nonpregnant females with a few visiting males, whereas after parturition, colonies are mainly composed of mothers and their offspring with a few visiting nonmothers and males. Our approach gives deeper insight into sex‐ and status‐specific behaviour, a prerequisite for understanding population dynamics and developing effective monitoring and management strategies. Provided sufficient samples can be obtained, this approach can be readily applied to a wide range of species.     

Successful despite poor flight performance: range expansion is associated with enhanced exploratory behaviour and fast development

Anthropogenic interference forces species to respond to changing environmental conditions. One possible response is dispersal and concomitant range shifts, allowing individuals to escape unfavourable conditions or to track the shifting climate niche. Range expansions depend on both dispersal capacity and the ability to establish populations beyond the former range. We here compare well‐established core populations with recently established edge populations in the currently northward expanding butterfly Lycaena tityrus. Edge populations were characterized by shorter development times and smaller size, a higher sensitivity to high temperature and an enhanced exploratory behaviour. The differences between core and edge populations found suggest adaptation to local climates and an enhanced dispersal ability in edge populations. In particular, enhanced exploratory behaviour may be advantageous in all steps of the dispersal process and may have facilitated the current range expansion. This study describes differences associated with a current range expansion, knowledge which might be useful for a better understanding of species responses to environmental change. We further report on variation between males and females in morphology and flight behaviour, with males showing a longer flight endurance and more pronounced exploratory behaviour than females.     

A further cost for the sicker sex? Evidence for male‐biased parasite‐induced vulnerability to predation

Males are typically the sicker sex. Data from multiple taxa indicate that they are more likely to be infected with parasites, and are less “tolerant,” or less able to mitigate the fitness costs of a given infection, than females. One cost of infection for many animals is an increased probability of being captured by a predator. A clear, hitherto untested, prediction is therefore that this parasite‐induced vulnerability to predation is more pronounced among males than females. We tested this prediction in the sexually size dimorphic guppy, Poecilia reticulata, in which females are typically larger than males. We either sham or experimentally infected guppies with Gyrodactylus turnbulli, elicited their escape response using an established protocol and measured the distance they covered during 60 ms. To discriminate between the effects of body size and those of other inherent sex differences, we size‐matched fish across treatment groups. Infection with G. turnbulli reduced the distance covered during the escape response of small adults by 20.1%, whereas that of large fish was unaffected. This result implies that parasite‐induced vulnerability to predation is male‐biased in the wild: although there was no difference in escape response between our experimentally size‐matched groups of males and females, males are significantly smaller across natural guppy populations. These results are consistent with Bateman's principle for immunity: Natural selection for larger body sizes and longevity in females seems to have resulted in the evolution of increased infection tolerance. We discuss the potential implications of sex‐ and size‐biased parasite‐induced vulnerability to predation for the evolutionary ecology of this host–parasite interaction in natural communities.     

Discrimination reversal learning reveals greater female behavioural flexibility in guppies

Behavioural flexibility allows an animal to adapt its behaviour in response to changes in the environment. Research conducted in primates, rodents and domestic fowl suggests greater behavioural persistence and reduced behavioural flexibility in males. We investigated sex differences in behavioural flexibility in fish by comparing male and female guppies (Poecilia reticulata) in a reversal learning task. Fish were first trained on a colour discrimination, which was learned equally rapidly by males and females. However, once the reward contingency was reversed, females were better at inhibiting the previous response and reached criterion twice as fast as males. When reward reversing was repeated, males gradually reduced the number of errors, and the two sexes had a comparable performance after four reversals. We suggest that sex differences in behavioural flexibility in guppies can be explained in terms of the different roles that males and females play in reproduction.     

Phenotype Matching and Early Social Conditions Affect Shoaling and Exploration Decisions

Early social conditions are vital for the establishment of future social interactions. Less, however, is known about how differences in early social conditions contribute to the process of individual recognition and subsequently in the decision of associating and exploring behaviours. In this study, we address this gap in the Trinidadian guppy Poecilia reticulata and test the prediction that fish would show a higher tendency to recognize and associate with individuals of similar phenotype. This prediction was tested by comparing the likelihood of association and latency to explore a novel area in males and females when in the presence of individuals that were familiar (reared together but from different populations), from the same population of origin (from the same population but deprived of interaction with each other), or were unfamiliar (different population and have never interacted). Both early social conditions and population of origin (phenotype matching) affected the tendency to shoal and explore. Females, but not males, exhibited identical preference to associate either with unfamiliar females as with familiar females from a different population. Importantly, female preference did not occur with unfamiliar individuals from a different population. In contrast with our prediction, tendency to shoal did not predict exploration propensity. Males always start exploration before the group whenever unfamiliar males composed the group. Females on the other hand only moved to the novel area after seeing the group doing so, revealing sexual dimorphism in exploratory behaviour. Our results provide evidence for a familiarity and phenotypic matching recognition mechanism at the population level and also highlight the importance of accounting for differences between sexes when investigating the effects of early social conditions.     

Intersexual segregation in foraging microhabitat use by Magellanic Woodpeckers (Campephilus magellanicus): Seasonal and habitat effects at the world's southernmost forests

Animals facing seasonal food shortage and habitat degradation may adjust their foraging behaviour to reduce intraspecific competition. In the harsh environment of the world's southernmost forests in the Magellanic sub‐Antarctic ecoregion in Chile, we studied intersexual foraging differences in the largest South American woodpecker species, the Magellanic Woodpecker (Campephilus magellanicus). We assessed whether niche overlap between males and females decrease when food resources are less abundant or accessible, that is, during winter and in secondary forests, compared to summer and in old‐growth forests, respectively. We analysed 421 foraging microhabitat observations from six males and six females during 2011 and 2012. As predicted, the amount of niche overlap between males and females decreased during winter, when provisioning is more difficult. During winter, males and females (i) used trees with different diameter at breast height (DBH); (ii) fed in trunk sections with different diameters; and (iii) fed at different heights on tree trunks or branches. Vertical niche partitioning between sexes was found in both old‐growth and secondary forests. Such a niche partitioning during winter may be a seasonal strategy to avoid competition between sexes when prey resources are less abundant or accessible. Our results suggest that the conservation of this forest specialist, dimorphic and charismatic woodpecker species requires considering differences in habitat use between males and females.     

Density‐dependent sex‐biased development of macroptery in a water strider

In wing‐polymorphic insects, wing morphs differ not only in dispersal capability but also in life history traits because of trade‐offs between flight capability and reproduction. When the fitness benefits and costs of producing wings differ between males and females, sex‐specific trade‐offs can result in sex differences in the frequency of long‐winged individuals. Furthermore, the social environment during development affects sex differences in wing development, but few empirical tests of this phenomenon have been performed to date. Here, I used the wing‐dimorphic water strider Tenagogerris euphrosyne to test how rearing density and sex ratio affect the sex‐specific development of long‐winged dispersing morphs (i.e., sex‐specific macroptery). I also used a full‐sib, split‐family breeding design to assess genetic effects on density‐dependent, sex‐specific macroptery. I reared water strider nymphs at either high or low densities and measured their wing development. I found that long‐winged morphs developed more frequently in males than in females when individuals were reared in a high‐density environment. However, the frequency of long‐winged morphs was not biased according to sex when individuals were reared in a low‐density environment. In addition, full‐sib males and females showed similar macroptery incidence rates at low nymphal density, whereas the macroptery incidence rates differed between full‐sib males and females at high nymphal density. Thus complex gene‐by‐environment‐by‐sex interactions may explain the density‐specific levels of sex bias in macroptery, although this interpretation should be treated with some caution. Overall, my study provides empirical evidence for density‐specific, sex‐biased wing development. My findings suggest that social factors as well as abiotic factors can be important in determining sex‐biased wing development in insects.     

Parting ways: parasite release in nature leads to sex‐specific evolution of defence

We evaluated the extent to which males and females evolve along similar or different trajectories in response to the same environmental shift. Specifically, we used replicate experimental introductions in nature to consider how release from a key parasite (Gyrodactylus) generates similar or different defence evolution in male vs. female guppies (Poecilia reticulata). After 4–8 generations of evolution, guppies were collected from the ancestral (parasite still present) and derived (parasite now absent) populations and bred for two generations in the laboratory to control for nongenetic effects. These F2 guppies were then individually infected with Gyrodactylus, and infection dynamics were monitored on each fish. We found that parasite release in nature led to sex‐specific evolutionary responses: males did not show much evolution of resistance, whereas females showed the evolution of increased resistance. Given that male guppies in the ancestral population had greater resistance to Gyrodactylus than did females, evolution in the derived populations led to reduction of sexual dimorphism in resistance. We argue that previous selection for high resistance in males constrained (relative to females) further evolution of the trait. We advocate more experiments considering sex‐specific evolutionary responses to environmental change.     

Sex differences in nutrient intake can reduce the potential for sexual conflict over fitness maximization by female and male crickets

As females and males have different roles in reproduction, they are expected to require different nutrients for the expression of reproductive traits. However, due to their shared genome, both sexes may be constrained in the regulation of nutrient intake that maximizes sex‐specific fitness. Here, we used the Geometric Framework for nutrition to examine the effect of macronutrient and micronutrient intakes on lifespan, fecundity and cuticular hydrocarbons (CHCs) that signal mate quality to prospective mates in female field crickets, Teleogryllus oceanicus. In addition, we contrasted nutritional effects on life‐history traits between males and females to determine how sex differences influence nutrient regulation. We found that carbohydrate intake maximized female lifespan and protein intake influenced CHC expression, while early life fecundity (cumulative fecundity at day 21) and lifetime fecundity were dependent on both macronutrient and micronutrient intakes. Fecundity required different nutrient blends to those required to optimize sperm viability in males, generating the potential for sexual conflict over macronutrient intake. The regulation of protein (P) and carbohydrate (C) intakes by virgin and mated females initially matched that of males, but females adjusted their intake to a higher P:C ratio, 1P:2C, that maximized fecundity as they aged. This suggests that a sex‐specific, age‐dependent change in intake target for sexually mature females, regardless of their mating status, adjusts protein consumption in preparation for oviposition. Sex differences in the regulation of nutrient intake to optimize critical reproductive traits in female and male T. oceanicus provide an example of how sexual conflict over nutrition can shape differences in foraging between the sexes.     

Does selection on horn length of males and females differ in protected and hunted populations of a weakly dimorphic ungulate?

Weaponry in ungulates may be costly to grow and maintain, and different selective pressures in males and females may lead to sex‐biased natural survival. Sexual differences in the relationship between weapon growth and survival may increase under anthropogenic selection through culling, for example because of trophy hunting. Selection on weaponry growth under different scenarios has been largely investigated in males of highly dimorphic ungulates, for which survival costs (either natural or hunting related) are thought to be greatest. Little is known, however, about the survival costs of weaponry in males and females of weakly dimorphic species. We collected information on horn length and age at death/shooting of 407 chamois Rupicapra rupicapra in a protected population and in two hunted populations with different hunting regimes, to explore sexual differences in the selection on early horn growth under contrasting selective pressures. We also investigated the variation of horn growth and body mass in yearling males (n = 688) and females (n = 539) culled in one of the hunted populations over 14 years. The relationship between horn growth and survival showed remarkable sexual differences under different evolutionary scenarios. Within the protected population, under natural selection, we found no significant trade‐off in either males or females. Under anthropogenic pressure, selection on early horn growth of culled individuals showed diametrically opposed sex‐biased patterns, depending on the culling regime and hunters’ preferences. Despite the selective bias between males and females in one of the hunted populations, we did not detect significant sex‐specific differences in the long‐term pattern of early growth. The relationship between early horn growth and natural survival in either sex might suggest stabilizing selection on horn size in chamois. Selection through culling can be strongly sex‐biased also in weakly dimorphic species, depending on hunters’ preferences and hunting regulations, and long‐term data are needed to reveal potential undesirable evolutionary consequences.