Sexual dimorphism in snakes: different reproductive roles favour different body plans

Because snakes have a highly simplified morphology, and many species have a wide (and broadly overlapping) range of adult body sizes within each sex, they offer an excellent opportunity to compare body composition of males and females. Evolutionary theory predicts that particular body components should be differentially enlarged in the two sexes. For example, we might expect the reproductive success of females to be enhanced by enlargement of organ systems involved in the processing and storage of energy (e.g. alimentary tract, liver, fat stores) whereas males would benefit from the enlargement of systems important for mate-searching, male–male combat and sperm competition (e.g. larger mass of skeletal muscles, tail, and kidneys). Dissection of 243 specimens of three snake species (117 Vipera aspis, 43 Elaphe longissima, 83 Coluber viridflavus) broadly supported these predictions. Strong sex differences were apparent in relative sizes (masses) of all the non-gonadal body components that we weighed. For example, males consistently had more musculature (relative to body length) than did conspecific females. Dimorphism in relative muscle mass is likely to be one of the most fundamental and widespread morphological differences between males and females in the Animal Kingdom.     

Related male Drosophila melanogaster reared together as larvae fight less and sire longer lived daughters

Competition over access to reproductive opportunities can lead males to harm females. However, recent work has shown that, in Drosophila melanogaster, male competition and male harm of females are both reduced under conditions simulating male-specific population viscosity (i.e., in groups where males are related and reared with each other as larvae). Here, we seek to replicate these findings and investigate whether male population viscosity can have repercussions for the fitness of offspring in the next generation. We show that groups of unrelated-unfamiliar (i.e., unrelated individuals raised apart) males fight more intensely than groups of related-familiar males (i.e., full siblings raised together as larvae), supporting previous findings, and that exposure to a female is required to trigger these differential patterns of male–male competition. Importantly, we show that differences in male–male competition can be associated with transgenerational effects: the daughters of females exposed to unrelated-unfamiliar males suffered higher mortality than the daughters of females exposed to related-familiar males. Collectively, these results suggest that population structure (i.e., variation in the relatedness and/or larval familiarity of local male groups) can modulate male–male competition with important transgenerational consequences.     

Sexual conflict over mating in Gnatocerus cornutus? Females prefer lovers not fighters

Female mate choice and male–male competition are the typical mechanisms of sexual selection. However, these two mechanisms do not always favour the same males. Furthermore, it has recently become clear that female choice can sometimes benefit males that reduce female fitness. So whether male–male competition and female choice favour the same or different males, and whether or not females benefit from mate choice, remain open questions. In the horned beetle, Gnatocerus cornutus, males have enlarged mandibles used to fight rivals, and larger mandibles provide a mating advantage when there is direct male–male competition for mates. However, it is not clear whether females prefer these highly competitive males. Here, we show that female choice targets male courtship rather than mandible size, and these two characters are not phenotypically or genetically correlated. Mating with attractive, highly courting males provided indirect benefits to females but only via the heritability of male attractiveness. However, mating with attractive males avoids the indirect costs to daughters that are generated by mating with competitive males. Our results suggest that male–male competition may constrain female mate choice, possibly reducing female fitness and generating sexual conflict over mating.     

Breeding dispersal and site-fidelity in three monogamous wader species in the Western Isles, U.K.

Adult and post-natal breeding-dispersal movements of Redshank Tringa totanus , Dunlin Calidris alpina and Ringed Plover Charadrius hiaticula were quantified in west Scotland. Data from 373 returning birds marked as breeding adults and 142 returning birds marked as chicks are presented. Unlike most previous studies, this study measured dispersal movements up to 40 km and attempted to overcome distance-related biases. For each species, adult males were significantly more nest site-faithful between years than were adult females. Likewise, first-time breeding males bred significantly closer to their natal site than did females. The settlement intensity per unit area of habitat showed marked differences between species, age classes and sex in the relative attractiveness of potential areas at different distances to settling birds. Adult dispersal was negatively and significantly related to breeding success in the previous year and positively related to capture on the nest in the previous year. Dispersal movements of adult Ringed Plover breeding on unstable cultivated habitats were significantly related to habitat quality (measured in terms of nest survival) and were always to habitat of better or equal quality.     

Density‐dependent demography and movements in a cyclic brown lemming population

Theoretical modeling predicts that both direct and delayed density‐dependence are key factors to generate population cycles. Deciphering density‐dependent processes that lead to variable population growth characterizing different phases of the cycles remains challenging. This is particularly the case for the period of prolonged low densities, which is inherently data deficient. However, demographic analyses based on long‐term capture–mark–recapture datasets can help resolve this question. We relied on a 16‐year (2004–2019) live‐trapping program to analyze the summer demography and movements of a cyclic brown lemming population in the Canadian Arctic. More specifically, we examined if inversely density‐dependent processes could explain why population growth can remain low during the prolonged low phase. We found that the proportion of females in the population was inversely density‐dependent with a strong male‐biased sex ratio at low densities but not at high densities. However, survival of adult females was higher than adult males, but both had lower survival at low densities than at high ones. Distances moved by both adult males and females were density‐dependent, and proportion of females in reproductive condition was weakly density‐dependent as it tended to increase at low density. Individual body condition, measured as monthly change in body mass, was not density‐dependent. Overall, the strong male‐biased sex ratio at very low densities suggests a loss of reproductive potential due to the rarity of females and appears to be the most susceptible demographic factor that could contribute to the prolonged low phase in cyclic brown lemmings. What leads to this sex‐bias in the first place is still unclear, potentially owing to our trapping period limited to the summer, but we suggest that it could be due to high predation rate on breeding females in winter.     

Balancing sexual selection through opposing mate choice and male competition

Male–male competition and female mate choice act contemporaneously in the cockroach Nauphoeta cinerea and the social pheromone of males influences the outcome of both forms of sexual selection. We therefore examined the joint and separate effects of male–male competition and female mate choice to determine if the selective optima for the pheromone were the same or different. Dominant males in a newly established hierarchy mated more frequently, but not exclusively. Manipulations of the multi-component social pheromone produced by males of N. cinerea showed that both long- and close-range attraction of females by males were influenced by the quantity and composition of the pheromone. The most attractive composition, however, differed from that which was most likely to confer high status to males. Since the outcome of male–male competition can conflict with mating preferences exhibited by females, there is balancing sexual selection on the social pheromone of N. cinerea. Such balancing selection might act to maintain genetic variation in sexually selected traits. We suggest that the different forms of sexual selection conflict in N. cinerea because females prefer a blend different to that which is most effective in male–male competition in order to avoid mating with overly aggressive males.     

Effects of dominance and female presence on secondary sexual characteristics in male tufted capuchin monkeys (Sapajus apella)

Alpha status may lead to physiological changes that enhance secondary sexual characteristics, which may serve as competitive signals to conspecific males, sexual signals to females, or possibly a combination of both. Here, we report measurements of secondary sexual characteristics in captive dominant and subordinate male tufted capuchin monkeys (Sapajus apella) with varying access to females. An adult male (who had previously been subordinate while housed with other males) was paired with an adult female. This male–female pair was introduced into a room that housed three other male–male pairs with stable hierarchy arrangements. We analyzed weight, body measurements, facial photographs, and hair cortisol before, during, and after introducing a female into the room. While there were no differences in weight or measurements between alphas and subordinates without physical access to the female prior to or during the female''s presence, we found that direct access to the female resulted in dramatic changes in facial appearance, body size, and testicular volume in the male who was paired with her. Overall, we found little evidence to suggest that alpha males advertise their status within all‐male groups via sexual secondary characteristics. However, direct physical access to females appears to trigger the development of such characteristics in alpha males. It remains of continued interest to identify the endocrine mechanisms responsible for the development, and possible loss, of secondary sexual characteristics.     

No evidence that relatedness or familiarity modulates male harm in Drosophila melanogaster flies from a wild population

Sexual selection frequently promotes the evolution of aggressive behaviors that help males compete against their rivals, but which may harm females and hamper their fitness. Kin selection theory predicts that optimal male–male competition levels can be reduced when competitors are more genetically related to each other than to the population average, contributing to resolve this sexual conflict. Work in Drosophila melanogaster has spearheaded empirical tests of this idea, but studies so far have been conducted in laboratory‐adapted populations in homogeneous rearing environments that may hamper kin recognition, and used highly skewed sex ratios that may fail to reflect average natural conditions. Here, we performed a fully factorial design with the aim of exploring how rearing environment (i.e., familiarity) and relatedness affect male–male aggression, male harassment, and overall male harm levels in flies from a wild population of Drosophila melanogaster, under more natural conditions. Namely, we (a) manipulated relatedness and familiarity so that larvae reared apart were raised in different environments, as is common in the wild, and (b) studied the effects of relatedness and familiarity under average levels of male–male competition in the field. We show that, contrary to previous findings, groups of unrelated‐unfamiliar males were as likely to fight with each other and harass females than related‐familiar males and that overall levels of male harm to females were similar across treatments. Our results suggest that the role of kin selection in modulating sexual conflict is yet unclear in Drosophila melanogaster, and call for further studies that focus on natural populations and realistic socio‐sexual and ecological environments.     

Dominance is not always an honest signal of male quality,but females may be able to detect the dishonesty

Females prefer dominant males as mating partners in numerous species. Male dominance rank is considered as an honest signal of male quality, because only healthy males in good condition are thought to be able to win fights with other males. Here, we tested whether activation of the immune system influences the success of males in male–male competition and mating in the field cricket, Gryllus integer. We activated the immune system of males with a nylon monofilament (to mimic a parasitoid larva), and arranged fights between male pairs to assess male dominance and associated mating success. Activation of the immune system with nylon monofilament substantially enhanced the fighting success of males during male–male competition but had no effect on mating success. However, sham-manipulation (a wound only) did not have any effect on fighting success although females mated more often with dominant males. Our study suggests that when male crickets meet an apparent survival threat they may behave more dominantly, probably owing to terminal investment. Male success during male–male competition is not always an honest signal of males’ quality, but females may be able to detect this dishonesty.     

Male Courtship Behavior and Weapon Trait as Indicators of Indirect Benefit in the Bean Bug,Riptortus pedestris

Females prefer male traits that are associated with direct and/or indirect benefits to themselves. Male–male competition also drives evolution of male traits that represent competitive ability. Because female choice and male–male competition rarely act independently, exploring how these two mechanisms interact is necessary for integrative understanding of the evolution of sexually selected traits. Here, we focused on direct and indirect benefits to females from male attractiveness, courtship, and weapon characters in the armed bug Riptortus pedestris. The males use their hind legs to fight other males over territory and perform courtship displays for successful copulation. Females of R. pedestris receive no direct benefit from mating with attractive males. On the other hand, we found that male attractiveness, courtship rate, and weapon size were significantly heritable and that male attractiveness had positive genetic covariances with both courtship rate and weapon traits. Thus, females obtain indirect benefits from mating with attractive males by producing sons with high courtship success rates and high competitive ability. Moreover, it is evident that courtship rate and hind leg length act as evaluative cues of female choice. Therefore, female mate choice and male–male competition may facilitate each other in R. pedestris. This is consistent with current basic concepts of sexual selection.     

Male competition reverses female preference for male chemical cues

Females must choose among potential mates with different phenotypes in a variety of social contexts. Many male traits are inherent and unchanging, but others are labile to social context. Competition, for example, can cause physiological changes that reflect recent wins and losses that fluctuate throughout time. We may expect females to respond differently to males depending on the outcome of their most recent fight. In Bolitotherus cornutus (forked fungus beetles), males compete for access to females, but copulation requires female cooperation. In this study, we use behavioral trials to determine whether females use chemical cues to differentiate between males and whether the outcome of recent male competition alters female preference. We measured female association time with chemical cues of two size‐matched males both before and after male–male competition. Females in our study preferred to associate with future losers before males interacted, but changed their preference for realized winners following male competitive interactions. Our study provides the first evidence of change in female preference based solely on the outcome of male–male competition.     

Mating Behavior of Daphnia: Impacts of Predation Risk,Food Quantity,and Reproductive Phase of Females

High predation risk and food depletion lead to sexual reproduction in cyclically parthenogenetic Daphnia. Mating, the core of sexual reproduction, also occurs under these conditions. Assessment of the environmental conditions and alteration of mating efforts may aid in determining the success of sexual reproduction. Here, we evaluated the impacts of predation risk, food quantity, and reproductive phase of females on the mating behavior of Daphnia obtusa males including contact frequency and duration using video analysis. Mating–related behavior involved male–female contact (mating) as well as male–male contact (fighting). Mating frequency increased while unnecessary fighting decreased in the presence of predation risk. In addition, low food concentration reduced fighting between males. Males attempted to attach to sexual females more than asexual females, and fighting occurred more frequently in the presence of sexual females. Duration of mating was relatively long; however, males separated shortly after contact in terms of fighting behavior. Thus, assessment of environmental factors and primary sexing of mates were performed before actual contact, possibly mechanically, and precise sex discrimination was conducted after contact. These results suggest that mating in Daphnia is not a random process but rather a balance between predation risk and energetic cost that results in changes in mating and fighting strategies.     

Impact of individual movement and changing resource availability on male–female encounter rates in an herbivorous insect

Species often confront changing resource distributions that result from natural and anthropogenic processes. For species that reproduce on or in close association with particular resources (e.g. host plants), changing resource distributions could affect the success of mate finding. We examine how mate-finding behaviours in an herbivorous insect mediate the impact of changing host plant spatial distribution. We tracked movements of 84 Melissa blue butterflies (Lycaeides melissa) in the Great Basin of western North America. Track data revealed sex differences in movement: males spent more time moving fast and females more time moving slowly; males moved more ballistically and females moved more diffusely. These differences vary quantitatively, but not qualitatively, between environments with contrasting resource distributions.From these data we created and parameterised a computer model of male–female encounters and used it to examine implications of changes to the patchiness and abundance of host plants. We use the cumulative encounter time between each simulated male–female pair as a proxy for mating success, thus allowing for the consideration of different female behaviours. The simulations suggest observed movement parameters exist in a trade-off between individuals maximising the number of potential mates they encounter and the probability that each encounter leads to mating success. Increasing host plant abundance decreases encounter rates thus encouraging males to be more diffuse to compensate. Changing the local resource density, i.e. increasing host plant patchiness, accentuated these trade-offs: by decreasing cumulative encounter time in resource rich environments and increasing it in resource sparse ones. Thus we see that both spatial resource geometry at multiple scales and plasticity in male movement strategies are important factors to consider when seeking to understand population reproductive behaviour, for example when assessing ecological impact of development, determining range boundaries and slowing invasions or outbreaks.     

Adaptive value of same-sex pairing in Laysan albatross

Same-sex pairing is widespread among animals but is difficult to explain in an evolutionary context because it does not result in reproduction, and thus same-sex behaviour often is viewed as maladaptive. Here, we compare survival, fecundity and transition probabilities of female Laysan albatross in different pair types, and we show how female–female pairing could be an adaptive alternative mating strategy, albeit one that resulted in lower fitness than male–female pairing. Females in same-sex pairs produced 80% fewer chicks, had lower survival and skipped breeding more often than those in male–female pairs. Females in same-sex pairs that raised a chick sometimes acquired a male mate in the following year, but females in failed same-sex pairs never did, suggesting that males exert sexual selection by assessing female quality and relegating low-quality females into same-sex pairs. Sexual selection by males in a monomorphic, non-ornamented species is rare and suggests that reconsideration is needed of the circumstances in which alternative reproductive behaviour evolves. Given the lack of males and obligate biparental care in this species, this research demonstrates how same-sex pairing was better than not breeding and highlights how it could be an adaptive strategy under certain demographic conditions.     

Breeding migrations by bighorn sheep males are driven by mating opportunities

In some species where male mating success largely depends on intrasexual competition, males can adopt migratory or resident strategies to seek breeding opportunities. The resulting mixture of resident and migrant tactics within a population can have important ecological, genetic, and evolutionary consequences for metapopulations. Bighorn sheep Ovis canadensis males establish a linear dominance hierarchy that influences their mating tactics. Some males perform breeding migrations during the pre‐rut and rut to seek mating opportunities, but little is known about these seasonal movements. We analyzed presence/absence data for 62 marked bighorn males during six mating seasons (20–32 males/year) in the Sheep River Provincial Park, Alberta, Canada, where hunting was not allowed. On average, about half of males left their natal population to rut elsewhere. The proportion of males leaving (yearly range 15%–69%) increased as the number of resident mature males increased and the populational sex ratio decreased, with fewer females during the pre‐rut. Among those leaving the park, 24% did so in October, while the trophy sheep hunting season was open. Detailed monitoring of breeding migrations in protected populations could inform management strategies to limit evolutionary impacts of hunting, which can alter size‐dependent mortality and create artificial pressures driving changes on heritable traits.     

Fairy wrasses perceive and respond to their deep red fluorescent coloration

Fluorescence enables the display of wavelengths that are absent in the natural environment, offering the potential to generate conspicuous colour contrasts. The marine fairy wrasse Cirrhilabrus solorensis displays prominent fluorescence in the deep red range (650–700 nm). This is remarkable because marine fishes are generally assumed to have poor sensitivity in this part of the visual spectrum. Here, we investigated whether C. solorensis males can perceive the fluorescence featured in this species by testing whether the presence or absence of red fluorescence affects male–male interactions under exclusive blue illumination. Given that males respond aggressively towards mirror-image stimuli, we quantified agonistic behaviour against mirrors covered with filters that did or did not absorb long (i.e. red) wavelengths. Males showed significantly fewer agonistic responses when their fluorescent signal was masked, independent of brightness differences. Our results unequivocally show that C. solorensis can see its deep red fluorescent coloration and that this pattern affects male–male interactions. This is the first study to demonstrate that deep red fluorescent body coloration can be perceived and has behavioural significance in a reef fish.     

Sex role similarity and sexual selection predict male and female song elaboration and dimorphism in fairy‐wrens

Historically, bird song complexity was thought to evolve primarily through sexual selection on males; yet, in many species, both sexes sing and selection pressure on both sexes may be broader. Previous research suggests competition for mates and resources during short, synchronous breeding seasons leads to more elaborate male songs at high, temperate latitudes. Furthermore, we expect male–female song structure and elaboration to be more similar at lower, tropical latitudes, where longer breeding seasons and year‐round territoriality yield similar social selection pressures in both sexes. However, studies seldom take both types of selective pressures and sexes into account. We examined song in both sexes in 15 populations of nine‐fairy‐wren species (Maluridae), a Southern Hemisphere clade with female song. We compared song elaboration (in both sexes) and sexual song dimorphism to latitude and life‐history variables tied to sexual and social selection pressures and sex roles. Our results suggest that song elaboration evolved in part due to sexual competition in males: male songs were longer than female songs in populations with low male survival and less male provisioning. Also, female songs evolved independently of male songs: female songs were slower paced than male songs, although only in less synchronously breeding populations. We also found male and female songs were more similar when parental care was more equal and when male survival was high, which provides strong evidence that sex role similarity correlates with male–female song similarity. Contrary to Northern Hemisphere latitudinal patterns, male and female songs were more similar at higher, temperate latitudes. These results suggest that selection on song can be sex specific, with male song elaboration favored in contexts with stronger sexual selection. At the same time, selection pressures associated with sex role similarity appear to favor sex role similarity in song structure.     

Sexual segregation of seasonal foraging habitats in a non-migratory marine mammal

Many animal species segregate by sex. Such segregation may be social in nature, or ecological, or both. Grey seals (Halichoerus grypus), like many large mammals, are sexually size dimorphic. In size dimorphic species, allometric differences in morphology, metabolic rate and reproductive costs are likely. Such differences may require the sexes to use different foraging strategies or different habitats. To investigate sexual segregation of habitat in grey seals, we used satellite tracks from 95 (male 46; female 49) adults breeding at Sable Island, Nova Scotia (44 degrees N, 60 degrees W) collected from 1995 to 2005. Location estimates were made from satellite fixes using a state-space movement model to estimate true locations and regularize them in time. Location estimates were used to calculate home range kernels of male and female habitat use each month. Month by sex kernel home ranges revealed striking differences and dynamics in habitat use between males and females on spatial scales broader than most terrestrial examples and at temporal and spatial resolutions rarely available for marine species. Differences were most pronounced just before (October-December) and immediately after breeding (February-March). During both periods, males primarily used areas along the continental shelf break, while females mainly used mid-shelf regions. Coupled with previously identified sex-specific seasonal patterns of energy storage, diving and diet, our findings suggest that males and females differ profoundly in their spatial foraging strategies. These differences may serve to maximize fitness by reducing intersexual competition during key foraging periods.     

Staying close to home: Ecological constraints on space use and range fidelity in a mountain ungulate

Understanding patterns of animal space use and range fidelity has important implications for species and habitat conservation. For species that live in highly seasonal environments, such as mountain goats (Oreamnos americanus), spatial use patterns are expected to vary in relation to seasonal changes in environmental conditions and sex‐ or age‐specific selection pressures. To address hypotheses about sex, age, and seasonality influence on space‐use ecology, we collected GPS location data from 263 radio‐collared mountain goats (males, n = 140; females, n = 123) in coastal Alaska during 2005–2016. Location data were analyzed to derive seasonal and sex‐specific fixed‐kernel home range estimates and to quantify the degree of seasonal range and utilization distribution overlap. Overall, we determined that home range size was smallest during winter, expanded coincident with the onset of green‐up and parturition, and was largest during summer. Home range size of males and females did not differ significantly during winter, but females had larger home ranges than males during summer, a relationship that was switched during the mating season. Pairwise comparisons involving individual females across subsequent years indicated home ranges were significantly smaller during years when they gave birth to offspring. Mountain goats exhibited a strong degree of range fidelity, and 99% (n = 138) of individual animals returned to their previous year''s seasonal range with an average annual Bhattacharyya''s affinity utilization distribution overlap index of 68%. Similarity of seasonal home range utilization distributions varied in relation to sex and season in some respects. Home range overlap was highest during the summer vegetation growing season, particularly among females. These findings advance our understanding about how environmental variation and sex‐ and age‐related reproductive constraints influence space use and range fidelity among alpine ungulates. Documentation of the high degree of range fidelity among mountain goats has important conservation implications in landscapes increasingly altered by anthropogenic activities.     

Estimation of breeding population size using DNA‐based pedigree reconstruction in brown bears

Robust estimates of demographic parameters are critical for effective wildlife conservation and management but are difficult to obtain for elusive species. We estimated the breeding and adult population sizes, as well as the minimum population size, in a high‐density brown bear population on the Shiretoko Peninsula, in Hokkaido, Japan, using DNA‐based pedigree reconstruction. A total of 1288 individuals, collected in and around the Shiretoko Peninsula between 1998 and 2020, were genotyped at 21 microsatellite loci. Among them, 499 individuals were identified by intensive genetic sampling conducted in two consecutive years (2019 and 2020) mainly by noninvasive methods (e.g., hair and fecal DNA). Among them, both parents were assigned for 330 bears, and either maternity or paternity was assigned to 47 and 76 individuals, respectively. The subsequent pedigree reconstruction indicated a range of breeding and adult (≥4 years old) population sizes: 128–173 for female breeders and 66–91 male breeders, and 155–200 for female adults and 84–109 male adults. The minimum population size was estimated to be 449 (252 females and 197 males) in 2019. Long‐term continuous genetic sampling prior to a short‐term intensive survey would enable parentage to be identified in a population with a high probability, thus enabling reliable estimates of breeding population size for elusive species.