Philopatric Males Research Articles

Bachelor groups form due to individual choices or environmental disrupters in African striped mice

In several mammal species, bachelor groups occur as a regular life history stage between dispersal and becoming the breeding male of a multifemale group. How such groups come into existence and how males that choose this strategy differ in life history traits from other males is rarely investigated. Males of the socially flexible African striped mouse, Rhabdomys pumilio , have been historically reported to adopt one of three alternative tactics: small group-living philopatric males, intermediate solitary roaming males or large group-living territorial males. Here, we describe for the first time, bachelors as a fourth male tactic. A long-term study on 13 generations of striped mice revealed two types of bachelor groups: (1) kin bachelor groups formed when all females disappeared, probably due to predation, and (2) nonkin bachelor groups formed by unrelated males that originated from different groups. Whereas kin bachelor groups occurred more frequently during the nonbreeding season, nonkin bachelor groups were most common in the breeding season and had a shorter tenure, since most of these males soon switched to the solitary roaming tactic or the territorial breeder tactic. Importantly, we found that nonkin bachelors were heavier and older than kin bachelors and that they did not differ from solitary roaming males in body mass and age. We therefore identified two different forms of bachelor groups, small kin bachelors whose female group members disappeared and larger nonkin bachelor groups showing traits of an alternative reproductive tactic. Both types of bachelor groups differed in multiple traits, demonstrating that it is important to follow the life histories of individuals to understand the mechanisms leading to alternative tactics. • We describe bachelor groups as a fourth male tactic in striped mice. • Kin bachelor groups form when all females disappear. • Nonkin bachelor groups are formed by unrelated males from different groups. • Both types of groups differ in body mass, age and seasonality of occurrence.

A review of philopatry and dispersal in felids living in an anthropised world

Abstract Typically, males of polygamous mammals are responsible for population connectivity and gene flow via dispersal, whereas females, showing stronger philopatry, strengthen local population stability and growth. These expectations can be disrupted by human disturbances; however, this possibility has been poorly examined in wide‐ranging mammals that are important targets for conservation. By reviewing philopatry and dispersal in felids, we aimed to evaluate: 1) whether the sex‐related patterns of philopatry and dispersal predicted for polygamous mammals are prevalent in felids, 2) possible major causes underlying each of these behaviours, and 3) if, and to what extent, anthropogenic disturbances can alter patterns of philopatry and dispersal in this animal lineage. We synthesised the available literature (n = 55 papers) comprising 12 species. Puma concolor was the most‐studied species, followed by other large species. Both philopatry and dispersal were heterogeneously defined, depending on the study aim and the method employed (telemetry, camera trapping or molecular tools). Most species followed the predicted philopatric and dispersal patterns, and most study areas (76%) were under some type of anthropogenic disturbance drivers. Philopatry was linked to females’ higher dependency on the quality and availability of resources, and to their social dynamics, higher reproductive success, inclusive fitness and demographic aspects of population. Dispersal was frequently linked to competition for mates and resources, and inbreeding avoidance. However, some plasticity was observed in both philopatry and dispersal, especially under the presence of anthropogenic drivers. For example, hunting can create open territories, increasing the number of philopatric females and opportunistic philopatric males. Habitat fragmentation can increase population isolation and male dispersal distance, and the presence of anthropogenic or natural barriers can result in unsuccessful dispersal attempts. We postulate that human activities affect long‐term population persistence in the Felidae, via disruption of sex‐related patterns of spatial dynamics.

Bonds of bros and brothers: Kinship and social bonding in postdispersal male macaques.

Group-living animals often maintain a few very close affiliative relationships-social bonds-that can buffer them against many of the inevitable costs of gregariousness. Kinship plays a central role in the development of such social bonds. The bulk of research on kin biases in sociality has focused on philopatric females, who typically live in deeply kin-structured systems, with matrilineal dominance rank inheritance and life-long familiarity between kin. Closely related males, in contrast, are usually not close in rank or familiar, which offers the opportunity to test the importance of kinship per se in the formation of social bonds. So far, however, kin biases in male social bonding have only been tested in philopatric males, where familiarity remains a confounding factor. Here, we studied bonds between male Assamese macaques, a species in which males disperse from their natal groups and in which male bonds are known to affect fitness. Combining extensive behavioural data on 43 adult males over a 10-year period with DNA microsatellite relatedness analyses, we find that postdispersal males form stronger relationships with the few close kin available in the group than with the average nonkin. However, males form the majority of their bonds with nonkin and may choose nonkin over available close kin to bond with. Our results show that kinship facilitates bond formation, but is not a prerequisite for it, which suggests that strong bonds are not restricted to kin in male mammals and that animals cooperate for both direct and indirect fitness benefits.

Dispersal and reproductive careers of male mountain gorillas in Bwindi Impenetrable National Park, Uganda

Dispersal is a key event in the life of an animal and it influences individual reproductive success. Male mountain gorillas exhibit both philopatry and dispersal, resulting in a mixed one-male and multimale social organization. However, little is known about the relationship between male dispersal or philopatry and reproductive careers in Bwindi mountain gorillas. Here we analyze data spanning from 1993 to 2017 on social groups in Bwindi Impenetrable National Park, Uganda to examine the proportion of males that disperse, age of dispersal, pathways to attaining alpha status, fate of dispersing males and philopatric males, and male tenure length as well as make comparisons of these variables to the Virunga mountain gorilla population. We report previously undocumented cases of dispersal by immature males and old males and we also observed the only known case of a fully mature male immigrating into a breeding group. We used genetic tracking of known individuals to estimate that a minimum of 25% of males that disperse to become solitary males eventually form new groups. No differences were found between the Bwindi and Virunga population in the age of male dispersal, the proportion of males that disperse, the age of alpha male acquisition, and dominance tenure length. The lack of differences may be due to small sample sizes or because the observed ecological variability does not lead to life history differences between the populations. Males in both populations follow variable strategies to attain alpha status leading to the variable one-male and multimale social organization, including dispersal to become solitary and eventually form a group, via group fissioning, usurping another alpha male, or inheriting the alpha position when a previous group leader dies.

Daily energy expenditure of males following alternative reproductive tactics: Solitary roamers spend more energy than group-living males

In many species, males follow alternative reproductive tactics (ARTs), where one tactic (called bourgeois) has much higher reproductive success than alternative tactics followed by males with lower competitive ability. The extent to which ARTs differ in energetic costs is unknown, but it is important to understand the fitness payoffs of ARTs. We studied male African striped mice (Rhabdomys pumilio) which follow one of three ARTs: heavy bourgeois males defend harems of females and have 10 times higher reproductive success than smaller roamers, which have ten times higher reproductive success than philopatric males, which remain in their natal group and are the smallest males. Bourgeois and philopatric males live in social groups that defend one territory, while roamers are solitary and roam over larger areas. We predicted that roamers will face higher energetic costs compared to group-living males because they do not gain thermoregulatory benefits of huddling in groups and might travel larger distances as they have larger home ranges. We measured daily energy expenditure (DEE) of 30 males, resting metabolic rate (RMR) of 79 males, travel distances and daily ranges of 31 males and changes in body mass of 51 males. Roamers had higher DEE and higher RMR than both types of group-living males. Philopatric males had shorter travel distances and smaller daily ranges than both roamers and bourgeois males, which did not differ from each other. This indicates that the higher DEE of roamers compared to bourgeois males cannot be explained by larger travel distances. Philopatrics gained body mass faster than bourgeois males and roamers, thereby increasing their competitive ability and thus the probability of later switching to a tactic of higher reproductive success. Our results suggest that roamers suffer energetic costs that might reduce their ability of gaining body mass and thus the likelihood of switching to the bourgeois tactic, indicating evolutionary trade-offs between investing energy into roaming versus gaining body mass.

The evolution of a rare mammalian trait \u2013 benefits and costs of male philopatry in proboscis bats

While inbreeding avoidance is widely accepted as the major driver of female natal dispersal, the evolution of male philopatry is still poorly understood and discussed to be driven by male mating strategy, mate competition among male kin and kin cooperation. During a twelve-year study, we gathered detailed genetic and observational data of individually marked proboscis bats to assess the degree of male philopatry as well as its costs and benefits to improve the understanding of its evolution. Our results reveal several patrilines with simultaneous presence of closely related males and a small proportion of unrelated immigrant males in their colonies. Philopatric males benefit from avoiding the costs of immigration into foreign colonies through significantly longer tenure, better integration (i.e. frequent nocturnal presence in the colonies) and consequently significantly higher reproductive success compared to immigrant males. Finally, we illustrate that despite a high proportion of philopatric males in the groups, the number of closely related competing males is low. Thus, the hypothesised costs of mate competition among male kin seem to be low in promiscuous mammalian societies with unrelated females and a small degree of male immigration and are readily outweighed by the benefits of staying in the natal group.

Sex-specific mortality costs of dispersal during the post-settlement stage promote male philopatry in a resident passerine

The costs and benefits of dispersal that select for sex-biased dispersal are still poorly understood. Many studies examine fitness consequences of dispersal after first breeding, while dispersal costs are most likely paid before first breeding during the movement, settlement, and post-settlement stages. We studied survival correlates of dispersal between flock settlement and first breeding during the first winter of juvenile willow tits (Poecile montanus), a small passerine that has female-biased natal dispersal, but shows no dispersal-associated survival differences after first breeding. This resident food-hoarding species winters in small stable non-kin territorial flocks. We collected capture-recapture data by following flocks from autumn to the following spring. We compared monthly survival and return rates of juveniles that were born and recruited within the study area (philopatric juveniles) and juveniles that originated from outside the study area (immigrant juveniles). Among males, survival was highest for philopatric juveniles whereas survival of females was higher among immigrant juveniles, providing one explanation for the female-biased natal dispersal observed in the species. Philopatric males may benefit from prior residency either through increased site familiarity and knowledge of winter food resources and/or by gaining higher social ranks during flock establishment. However, rank data provided little support for the latter hypothesis. Other mechanisms such as increased ability to find high-quality flocks and mates may be important for female survival. Our results provide further evidence that dispersal costs are paid mainly before first breeding and that sex-specific costs of dispersal play a role in the evolution of sex-biased dispersal. This paper shows that female-biased dispersal can be a consequence of sex-specific costs and benefits of dispersal during the post-settlement stage of the dispersal process which is a very poorly understood stage in dispersal theory. By examining correlates of dispersal before rather than after first breeding as it is usually done, our study aids in understanding the selection pressures modifying dispersal strategies. Our results have wide applicability because there are many resident taxa similar to the willow tit (our study species) that have a settlement stage and a prolonged non-reproductive phase before their first reproduction.

Sex Differences in the Use of Whinny Vocalizations in Spider Monkeys (Ateles geoffroyi)

In species in which males and females exhibit different association patterns, the use of vocalizations that regulate interindividual distance may differ between the sexes. Spider monkey social groups are characterized by high fission–fusion dynamics and sex differences in association patterns; female–female associations have been described as more passive than those between philopatric males. Individuals of both sexes produce whinny vocalizations, which may allow callers and receivers to mediate interindividual spacing based on existing social relationships. As such, we hypothesized individuals of each sex would use whinny vocalizations at different rates and in different contexts. To investigate sex differences in the rate of whinnying across behavioral contexts, we collected focal animal samples on Yucatan spider monkeys (Ateles geoffroyi yucatanensis) over 8 mo at Runaway Creek Nature Reserve, Belize. In addition, we recorded all changes in subgroup composition to investigate whether a female’s likelihood of calling was influenced by the number of conspecifics joining, or leaving their subgroup. We found that females called at higher rates than males in most behavioral contexts, especially foraging. The probability that females would call increased during subgroup fissions and fusions, and correlated positively with the number of individuals joining or leaving their subgroup. Male calling rates did not differ across contexts, and males generally called less than females. Our results suggest that whinnying by females may allow callers to mediate interindividual spacing in contexts where proximity risks increasing feeding competition. In species in which the sexes associate in qualitatively different ways, vocalizations may play a role in maintaining these differences.

Separation at weaning from the family is stressful for naturally group-living, but not solitary-living, male African striped mice Rhabdomys

Early separation from a family is stressful for young mammals, but might be more stressful for group-living than solitary species. Using juvenile males of three African striped mice Rhabdomys taxa that are either group (R. pumilio) or solitary (R. dilectus dilectus and R. d. chakae) living, we predicted greater separation anxiety in R. pumilio than R. dilectus because group-living could reduce anxiety in R. pumilio. Three brothers from each of 10 litters per taxon were randomly assigned soon after natural weaning (25 days) to one of three treatments for 10 days: (1) remained with the family (philopatric); (2) separated from the family by a wire mesh barrier (separated); and (3) isolated from the family (isolated). Males were individually tested in a four-arm maze to assess their anxiety responses and sampled for corticosterone concentrations 20 mins and 10 days later. Compared to R. dilectus males, R. pumilio males showed a greater treatment response to separation: philopatric males used the light arms of the maze less and had higher corticosterone concentrations compared to isolated males, which spent the most time in the light arms and had the lowest corticosterone concentrations overall; separated males showed an intermediate behavioural response, but had similar corticosterone concentrations to philopatric males. Thus, separation from a family group is more stressful in group-living Rhabdomys and this stress response dissipates with time. Philopatry and group-living may be more important for young R. pumilio, whereas dispersal at weaning is an important life history event for solitary R. dilectus.

Experimental increase of testosterone increases boldness and decreases anxiety in male African striped mouse helpers

Males of many species can adjust their behaviors to environmental conditions by changing reproductive tactics. Testosterone surges in adult breeding males typically inhibit the expression of paternal care while facilitating the expression of aggression during environmental changes. Similarly, in non-breeding philopatric males of cooperatively breeding species, up-regulation of testosterone may inhibit alloparental care while facilitating dispersal, i.e. males might become bolder and more explorative. We tested this hypothesis in philopatric male African striped mice, Rhabdomys pumilio. Striped mouse males can either remain in their natal groups providing alloparental care or they can disperse seeking mating opportunities. Compared to philopatric males, dispersed males typically show higher testosterone levels and lower corticosterone levels, and more aggression toward pups and same sex conspecifics. We experimentally increased the testosterone levels of the philopatric males kept in their family groups when pups were present. Testosterone-treated males did not differ significantly from control males in alloparental care and in aggression toward same-sex conspecifics. Compared to the control males, testosterone treated males were bolder, more active, and less anxious; they also showed lower corticosterone levels. The philopatric males were sensitive to our testosterone treatment for dispersal- and anxiety-like behavior but insensitive for social behaviors. Our results suggest a role of testosterone in dispersal.

Tradition over trend: Neighboring chimpanzee communities maintain differences in cultural behavior despite frequent immigration of adult females

The notion of animal culture has been well established mainly through research aiming at uncovering differences between populations. In chimpanzees (Pan troglodytes verus), cultural diversity has even been found in neighboring communities, where differences were observed despite frequent immigration of individuals. Female chimpanzees transfer at the onset of sexual maturity at an age, when the behavioral repertoire is fully formed. With immigrating females, behavioral variety enters the group. Little is known about the diversity and the longevity of cultural traits within a community. This study is building on previous findings of differences in hammer selection when nut cracking between neighboring communities despite similar ecological conditions. We now further investigated the diversity and maintenance of cultural traits within one chimpanzee community and were able to show high levels of uniformity in group-specific behavior. Fidelity to the behavior pattern did not vary between dispersing females and philopatric males. Furthermore, group-specific tool selection remained similar over a period of 25 years. Additionally, we present a study case on how one newly immigrant female progressively behaved more similar to her new group, suggesting that the high level of similarity in behavior is actively adopted by group members possibly even when originally expressing the behavior in another form. Taken together, our data support a cultural transmission process in adult chimpanzees, which leads to persisting cultural behavior of one community over time.

Endocrinology of sociality: Comparisons between sociable and solitary individuals within the same population of African striped mice

The social organization of species ranges from solitary-living to complex social groups. While the evolutionary reasons of group-living are well studied, the physiological mechanisms underlying alternative social systems are poorly understood. By studying group-living and solitary individuals of the same species, we can determine hormonal correlates of sociality without the problem of confounding phylogenetic factors. The African striped mouse (Rhabdomys pumilio) is a socially flexible species, which can be solitary or alternatively form complex family groups, depending on population density and the extent of reproductive competition. We predicted group-living striped mice to show signs of reproductive suppression and social stress, resulting in higher corticosterone but lower testosterone levels when compared to solitary-living individuals. To determine whether differences in social organization correlated with hormonal differences, we collected blood samples from free-living striped mice during four breeding seasons when we experimentally induced solitary-living in philopatric individuals by locally reducing population density. Striped mice that were group-living did not change their corticosterone or estosterone levels during the study, indicating that there was no temporal effect during the breeding season. Striped mice of both sexes had significantly lower corticosterone levels after switching from group- to solitary-living. Solitary males – but not solitary females – had higher testosterone levels than group-living conspecifics. Our results suggest that group-living results in physiological stress and can induce reproductive suppression, at least in philopatric males. The switch to solitary-living may thus be a tactic to avoid reproductive competition within groups, and is associated with decreased stress hormone levels and onset of independent reproduction.

Female‐biased dispersal in a bat with a female‐defence mating strategy

The ultimate causes for predominant male-biased dispersal (MBD) in mammals and female-biased dispersal (FBD) in birds are still subject to much debate. Studying exceptions to general patterns of dispersal, for example, FBD in mammals, provides a valuable opportunity to test the validity of proposed evolutionary pressures. We used long-term behavioural and genetic data on individually banded Proboscis bats (Rhynchonycteris naso) to show that this species is one of the rare mammalian exceptions with FBD. Our results suggest that all females disperse from their natal colonies prior to first reproduction and that a substantial proportion of males are philopatric and reproduce in their natal colonies, although male immigration has also been detected. The age of females at first conception falls below the tenure of males, suggesting that females disperse to avoid father-daughter inbreeding. Male philopatry in this species is intriguing because Proboscis bats do not share the usual mammalian correlates (i.e. resource-defence polygyny and/or kin cooperation) of male philopatry. They have a mating strategy based on female defence, where local mate competition between male kin is supposedly severe and should prevent the evolution of male philopatry. However, in contrast to immigrant males, philopatric males may profit from acquaintance with the natal foraging grounds and may be able to attain dominance easier and/or earlier in life. Our results on Proboscis bats lent additional support to the importance of inbreeding avoidance in shaping sex-biased dispersal patterns and suggest that resource defence by males or kin cooperation cannot fully explain the evolution of male philopatry in mammals.

Staying put or leaving home: endocrine, neuroendocrine and behavioral consequences in male African striped mice

Social flexibility occurs when individuals of both sexes can change their social and reproductive tactics, which in turn can influence the social system of an entire population. However, little is known regarding the extent to which individuals of socially flexible species vary in their social behavior and in the underlying physiological mechanisms that support different social tactics. The present study in African striped mice modeled in captivity three male tactics described from the field: (a) philopatric males remaining in the family; (b) solitary roamers; or (c) group-living breeding males. Sixteen pairs and their offspring were kept in captivity, while one male offspring from the family remained as singly housed after he reached 21days of age. Differences in behavior, morphology, hormone and neuropeptide levels were tested, and physiological measurements were correlated with behavioral measurements. In standardized arena experiments group-living males (philopatrics and breeders) were significantly more aggressive than singly housed males, in agreement with previous data suggesting that group-living, but not roaming males, are territorial. Philopatric males showed signs of reproductive suppression, small testes, lower testosterone and higher corticosterone levels than their singly housed brothers. Higher levels of arginine vasopressin (AVP) were measured in the PVN and BNST of singly housed males compared to group-living males. Based on these findings we hypothesize that roamers are physiologically primed, and capable, if the opportunity to mate arises, to release AVP, form social bonds and become territorial, thus quickly adopting the tactic as breeding male which would yield a higher reproductive success.

Differential investment into testes and sperm production in alternative male reproductive tactics of the African striped mouse (Rhabdomys pumilio)

Males that follow alternative reproductive tactics might differ in their investment into testis development and sperm production. The resource-allocation hypothesis predicts that males following a sneaker tactic should invest more into sperm production than dominant territorial males which should invest more into mate guarding. This hypothesis is supported by studies in species where individual males cannot switch between tactics (fixed tactics). Here we present the first data for a species where males can switch between tactics (plastic tactics). We studied African striped mice (Rhabdomys pumilio) in captivity, mimicking three tactics observed in the field: philopatric group-living males, singly-housed males representing roaming males, and group-living breeding males. We measured quantitative and qualitative reproductive traits, as well as serum and testis hormone concentrations. We found no support for the resource-allocation hypothesis, since breeding and singly-housed males invested similarly in testes and sperm. However, philopatric males had significantly smaller testes and epididymides, lower sperm counts, lower testosterone and higher corticosterone levels than males of the two other tactics. Philopatric males did not reach a larger body mass than singly-housed males with well developed reproductive traits, indicating that they did not trade investment in sperm production against growth. Interestingly, testis testosterone concentrations of philopatric males did not differ from those of other males. Our data suggest that philopatric males are reproductively suppressed by the breeding male, but might be ready to increase their serum testosterone levels when social and environmental conditions allow for this physiological switch accompanying the behavioral switch between tactics.

Philopatry and natal dispersal of Montagu's harriers (Circus pygargus) breeding in Spain: a review of existing data

Natal dispersal is an important component of bird ecology, plays a key role in many ecological and evolutionary processes, and has important conservation implications. Nevertheless, detailed knowledge on natal dispersal is still lacking in many bird species, especially raptors. We review and compile existing information from five tagging programmes of juvenile Montagu's harriers (Circus pygargus) in different Spanish regions, with PVC rings or wing tags, to provide an assessment of philopatry and natal dispersal of the species in Spain. Only 7% of all tagged harriers were observed as breeders in subsequent years. The percentage of philopatric (i.e. breeding within 10 km of the natal site) males and females was lower that 5%. Overall, there were no sexual differences in percentage of philopatric birds or dispersal distances, but we found study area differences. The low philopatry observed suggests a high capacity for natal dispersal in this species, for both sexes, and therefore high genetic mixing between populations. Differences in philopatry between study areas may be influenced by the different observation effort or detectability, or else reflect different philopatric strategies among populations. Finally, we found no significant differences in philopatry rate or dispersal distances related to tagging method, suggesting that tagging technique has a smaller effect than monitoring effort or observation ease on observation probability. Developing tagging programmes at a small scale and without procuring very large-scale and intensive subsequent monitoring is not worthwhile for evaluating philopatry and natal dispersal in this species.

Hormone levels of male African striped mice change as they switch between alternative reproductive tactics

Alternative reproductive tactics occur when individuals of the same species follow alternative ways to maximize reproductive success. Often younger and smaller males follow tactics that result in lower fitness than that of dominant larger males. The relative plasticity hypothesis predicts that hormone levels change as males change tactics, but direct tests of this hypothesis are missing. It has been demonstrated in a number of studies that males following different tactics also differ in hormone levels (unpaired data), but not that individual males change their hormone levels as they change tactic (paired data). We compared hormone levels in the same individuals before and after they changed their tactic, using field samples collected over a period of 6years. We studied male striped mice (Rhabdomys pumilio) following three alternative reproductive tactics: 1. alloparental philopatric males; 2. solitary roaming males, and 3. group-living dominant breeders. Testosterone levels increased and corticosterone levels decreased when philopatric males became roamers or breeders. The increase in testosterone levels tended to be higher in philopatric males that became roamers than in philopatric males that became breeders. Testosterone levels decreased when roamers became breeders. Prolactin levels increased when males of any other tactic became breeders. Thus, males significantly changed their hormone profiles as they changed tactics. These results are in agreement with the hypothesis that changes in hormone levels are associated with the switch from one alternative reproductive tactic to another.

A micro-geography of fear: learning to eavesdrop on alarm calls of neighbouring heterospecifics

Many vertebrates eavesdrop on alarm calls of other species, which is a remarkable ability, given geographical variation in community composition and call diversity within and among species. We used micro-geographical variation in community composition to test whether individuals recognize heterospecific alarm calls by: (i) responding to acoustic features shared among alarm calls; (ii) having innate responses to particular heterospecific calls; or (iii) learning specific alarm calls. We found that superb fairy-wrens (Malurus cyaneus) fled to cover to playback of noisy miner (Manorina melanocephala) aerial predator alarm calls only in locations where miners were present, suggesting that learning rather than acoustic structure determines response. Sites with and without miners were well within the dispersal distance of fairy-wrens, and philopatric males and dispersing females showed the same pattern, so that local genetic adaptation is extremely unlikely. Furthermore, where miners were present, fairy-wrens responded appropriately to different miner calls, implying eavesdropping on their signalling system rather than fleeing from miners themselves. Learned eavesdropping on alarm calls enables individuals to harvest ecologically relevant information from heterospecifics on an astonishingly fine spatial scale. Such phenotypic plasticity is valuable in a changing world, where individuals can be exposed to new species.

Relative fitness of alternative male reproductive tactics in a mammal varies between years

1. In many species, males can use different behavioural tactics to achieve fertilization, so-called alternative reproductive tactics (ARTs). Few field studies have measured fitness consequences of ARTs under varying environmental conditions. 2. Here, we describe fitness consequences of three phenotypically plastic ARTs in the African striped mouse (Rhabdomys pumilio) and show that relative fitness of ARTs differs between years. Each year represents a different generation. 3. For the generation living under high population density, tactics differed in relative fitness in accordance with the theory of conditional strategies, with highly successful territorial breeding males having 10 times higher success than solitary roaming males and 102 times higher success than adult natally philopatric males. 4. For the generation living under intermediate population density, the territorial breeding and roaming tactics yielded similar fitness, which would be in agreement with the theory of mixed strategies. No philopatric males occurred. 5. For the generation living under low population density, roaming was the only tactic used and some roamers had very high fitness. 6. The main prediction of status-dependent selection for conditional strategies is a correlation between fitness and status, often measured as body mass, but we did not find this correlation within tactics when more than one tactic was expressed in the population. 7. Female distribution seems to have an effect on which reproductive tactics male chose: female defence polygyny when females are clumped (interference competition), but a searching tactic when females are dispersed (scramble competition). In contrast to predictions arising from theory on scramble competition, male body mass was important in determining fitness only in the year when females were dispersed, but not in other years. 8. Our results indicate that the differentiation between conditional and mixed strategies is not an absolute one. In many other species, environmental conditions might fluctuate temporally and spatially so that the normally suboptimal tactic yields similar fitness to the (usually) dominant tactic or that only a single tactic prevails. 9. We suggest the term single strategy, independent of current fitness consequences, for systems where tactics are not genetically determined, in contrast to genetically determined alternative strategies.

No evidence for offspring sex-ratio adjustment to social or environmental conditions in cooperatively breeding purple-crowned fairy-wrens

When fitness returns or production costs vary between male and female offspring, selection is expected to favor females that adjust offspring sex ratio accordingly. However, to what extent vertebrates can do so is the subject of ongoing debate. Here, we explore primary sex ratios in 125 broods of cooperatively breeding purple-crowned fairy-wrens Malurus coronatus. We expected that females might adjust offspring sex ratio because this passerine species experiences considerable variation in social and environmental conditions. (1) However, although helpers substantially increase parental fitness, females (particularly in pairs and small groups) did not overproduce philopatric males (helper-repayment hypothesis). (2) Sex-ratio adjustment based on competition among individuals (helper-competition hypothesis) did not conceal helper-repayment effects or drive sex allocation on its own: while high-quality territories can accommodate more birds, brood sex ratios were independent of territory quality, alone or in interaction with group size. (3) Additionally, males are larger than females and are possibly more costly to produce (costly sex hypothesis), and (4) female offspring may benefit more from long-term effects of favorable conditions early in life (Trivers–Willard hypothesis). Nonetheless, large seasonal variation in food abundance was not associated with a consistent skew in primary sex ratios. Thus, overall, our results did not support the main hypotheses of adaptive sex-ratio adjustment in M. coronatus. We discuss that long-term differential costs and benefits may be insufficient to drive evolution of primary sex-ratio manipulation by M. coronatus females. More investigation is therefore needed to determine the general required sex differences in long-term fitness returns for mechanisms of primary sex-ratio manipulation to evolve.