Sex biases in kin shoaling and dispersal in a cichlid fish

Animal dispersal is associated with diverse costs and benefits that vary among individuals based on phenotype and ecological conditions. For example, females may disperse when males benefit more from defending territories in familiar environments. Similarly, size differences in dispersal propensity may occur when dispersal costs are size-dependent. When individuals do disperse, they may adopt behavioral strategies that minimize dispersal costs. Dispersing fish, for example, may travel within shoals to reduce predation risks. Further, kin shoaling may augment inclusive fitness by reducing predation of relatives. However, studies are lacking on the role of kin shoaling in dispersal. We explored how sex and size influence dispersal and kin shoaling in the cichlid Neolamprologus caudopunctatus. We microsatellite genotyped over 900 individuals from two populations separated by a potential dispersal barrier, and documented patterns of population structure, migration and within-shoal relatedness. Genetic differentiation across the barrier was greater for smaller than larger fish, suggesting larger fish had dispersed longer distances. Females exhibited weaker genetic differentiation and 11 times higher migration rates than males, indicating longer-distance female-biased dispersal. Small females frequently shoaled with siblings, possibly offsetting dispersal costs associated with higher predation risks. In contrast, small males appeared to avoid kin shoaling, possibly to avoid local resource competition. In summary, long-distance dispersal in N. caudopunctatus appears to be female-biased, and kin-based shoaling by small females may represent a behavioral adaptation that reduces dispersal costs. Our study appears to be the first to provide evidence that sex differences in dispersal influence sex differences in kin shoaling.     

Evidence for male-biased dispersal in lake malawi cichlids from microsatellites

This study addressed within-population dispersal patterns among the species-rich Lake Malawi cichlids, specifically among the rock-dwelling mbuna group. Relatedness values were calculated for 160 individuals belonging to two species from known locations in the field by screening six highly polymorphic microsatellite loci. The results from both species indicate that spatially adjacent females have higher average relatedness values than those separated by larger distances, but that this pattern is reversed in males. This therefore provides firm evidence for male-biased dispersal within the Malawian cichlid flock.     

Some evidence for different ecological pressures that constrain male and female body size

The dwarf morph of the Lake Tanganyika cichlid Telmatochromis temporalis uses empty snail shells as shelters and breeding sites in shell beds, in which many empty shells exist. Here, we assessed selection forces regulating body size in this fish. Field observations showed that large males tended to have a greater number of females in their territories, suggesting that sexual selection favours large males. Nonetheless, a transplant experiment suggested that male body size was limited by the ability to hide in empty shells from large piscivorous fish. In females, the number of ovarian eggs increased with body size, suggesting that fecundity selection favours large females. However, females are smaller than males. Females spawned eggs close to the apices inside the shells. The small space there would decrease the risk of egg predation by egg predators, and small body size of females may be a result of adaptation to spawn eggs in the small, safe spaces. This study provides support for the idea that male and female body sizes have been limited by different ecological pressures (predation on adult fish in males, predation on eggs in females), which has not been reported previously in any animal.     

Dispersal Patterns of Orang-utans (<Emphasis Type="BoldItalic">Pongo</Emphasis> spp.) in a Bornean Peat-swamp Forest

Knowledge regarding dispersal patterns in great apes may help in understanding the evolution of dispersal patterns and social grouping in early hominoids, as well as in our own species. However, the social structure and dispersal system of orang-utans (Pongo spp.) remains little understood despite past research. We addressed this question by conducting genetic analyses on a wild orang-utan (Pongo pygmaeus wurmbii) population from the Sabangau peat-swamp in Borneo. We estimated pairwise relatedness among 16 adult individuals using 19 polymorphic microsatellite markers. Mean relatedness among females was significantly higher than in males, irrespective of the relatedness estimator used, following the pattern predicted for male dispersal. Our results support field observations that average dispersal distance for females is less than for males, suggesting that female orang-utans are philopatric, whereas males disperse. This contrasts with previous findings from other sites where anthropogenic influences were present. Based on qualitative mitochondrial DNA analyses, it appears that unflanged adult males show some degree of site fidelity compared to flanged males. Thus, male orang-utans may disperse permanently from their natal range once they are fully flanged. Male-biased dispersal and female philopatry in orang-utans differ from those of extant African apes and are more similar to many Old World monkey species. Hence we hypothesize that this system may represent the ancestral state of early hominoids.     

Genetic relatedness in groups is sex-specific and declines with age of helpers in a cooperatively breeding cichlid

Kin selection can explain the evolution of cooperative breeding and the distribution of relatives within a population may influence the benefits of cooperative behaviour. We provide genetic data on relatedness in the cooperatively breeding cichlid Neolamprologus pulcher. Helper to breeder relatedness decreased steeply with increasing helper age, particularly to the breeding males. Helper to helper relatedness was age‐assortative and also declined with age. These patterns of relatedness could be attributed to territory take‐overs by outsiders when breeders had disappeared (more in breeding males), between‐group dispersal of helpers and reproductive parasitism. In six of 31 groups females inherited the breeding position of their mother or sister. These matrilines were more likely to occur in large groups. We conclude that the relative fitness benefits of helping gained through kin selection vs. those gained through direct selection depend on helper age and sex.     

Influence of brood size and offspring size on parental investment in a biparental cichlid fish,Neolamprologus moorii

Both males and females ofNeolamprologus moorii, a substrate-spawning cichlid fish in Lake Tanganyika, tend their eggs and fry. In this study, the influence of brood size and fry size on parental investment of this species was examined under natural conditions. Breeding parents intensively attacked fry-eating fishes that approached their broods. The attack rate by the parents against the approaching fry-eaters was positively correlated with brood size and negatively with fry size, but was much more strongly related to brood size. This suggests that the parental decision for brood defense appeared to be primarily determined by brood size. The reason for fry size being less important in the parental decision may be that even large fry have a low survival ability on their own under the high predation pressure in the study sites.     

A sex-specific trade-off between mating preferences for genetic compatibility and body size in a cichlid fish with mutual mate choice

Mating preferences for genetic compatibility strictly depend on the interplay of the genotypes of potential partners and are therein fundamentally different from directional preferences for ornamental secondary sexual traits. Thus, the most compatible partner is on average not the one with most pronounced ornaments and vice versa. Hence, mating preferences may often conflict. Here, we present a solution to this problem while investigating the interplay of mating preferences for relatedness (a compatibility criterion) and large body size (an ornamental or quality trait). In previous experiments, both sexes of Pelvicachromis taeniatus, a cichlid fish with mutual mate choice, showed preferences for kin and large partners when these criteria were tested separately. In the present study, test fish were given a conflicting choice between two potential mating partners differing in relatedness as well as in body size in such a way that preferences for both criteria could not simultaneously be satisfied. We show that a sex-specific trade-off occurs between mating preferences for body size and relatedness. For females, relatedness gained greater importance than body size, whereas the opposite was true for males. We discuss the potential role of the interplay between mating preferences for relatedness and body size for the evolution of inbreeding preference.     

Adjusting parental investment to changing environmental conditions: the effect of food ration on parental behaviour of the convict cichlid, Cichlasoma nigrofasciatum

Animals that care for their offspring may vary the amount of care provided for a particular brood in relation to environmental conditions. Food availablity is one factor that may affect the costs and benefits associated with parental investment. The convict cichlid, Cichlasoma nigrofasciatum, is a small, substrate-spawning cichlid from Central America. Both male and female provide parental care for eggs and fry. Paris were kept at one of three ration levels, high, medium or low. Time spent in parental fanning by females was positively related to ration. Males spent less time fanning than females and their parental behaviour varied considerably between individuals. Males on the high ration spent slightly more time fanning than those on the lower rations. The number of eggs produced per spawning and the post-spawning weight of both males and females were significantly and positively related to ration. Foraging frequency of both males and females was inversely related to ration. There was no significant effect of ration on the frequency of mouthing eggs and young or on intra-pair aggression. Eggs of low-ration fish hatched earlier than those of medium- and high-ration fish but there was no significant difference in the number of days that the young survived. These results suggest that the allocation of time and effort between parental and maintenance activities differs in relation to food supply. Parents may provide more care for the large brood produced when food is plentiful but place more emphasis on their own survival when food is short and broods are small.     

Socio-genetic correlates of unbiased sex dispersal in a population of black capuchin monkeys (<Emphasis Type="Italic">Sapajus nigritus</Emphasis>)

Most social mammal species exhibit male-biased dispersal. Sex bias in dispersal leads to a higher degree of relatedness among individuals of the philopatric sex, thus an atypical dispersal pattern might lead to deviations in the typical within-group kinship structure. Kinship, in turn, influences patterns of social interactions, as widely evident by kin-biased behaviors. We investigated the link between dispersal, relatedness structure, and sociopositive interactions established by adult females of black capuchin monkeys (Sapajus nigritus) living in a population that experiences female dispersal, an unusual pattern for capuchin monkeys. The study was conducted in Parque Estadual Carlos Botelho (PECB), within the Brazilian Atlantic Forest. We addressed dispersal and relatedness patterns by genotyping 20 adults of 3 groups across 9 microsatellite loci. We also sampled the monkeys’ behavior and compared spatial association frequencies and rates of grooming among same- and opposite-sex dyads. There was no difference between males and females in genetic parameters; both males and females show low coefficients of relatedness indicating that neither sex is consistently philopatric. The mean pairwise coefficient of relatedness for co-resident females was not higher than that for co-resident males. Compared to other populations of capuchin monkeys, female bond was weak, as evident by lower spatial association frequencies, reduced rates of grooming and lack of correlation between coefficients of relatedness and measures of dyadic sociopositive interactions. Our findings thus confirm that female dispersal is a habitual process in the capuchin population of PECB, and that, as expected, dispersal by females strongly influences the relatedness structure of the population as well as the affiliative relationships among female groupmates.     

Differences in Aggressiveness in the Midas Cichlid Fish (Cichlasoma citrinellum) in Relation to Sex,Reproductive State and the Individual

The Midas cichlid, Cichlasoma citrinellum, is monogamous and biparental. Because of competition for limited spawning sites and intense predation on their young, vigorous defense of their territory is essential. Although both sexes engage in defense, they differ in aggressiveness. The aggressive responses of both sexes were measured by counting the number of bites and bumps each fish directed toward its own mirror image. The size of the fish's genital papilla was also recorded to estimate its reproductive state. Compared with females, males had higher median mirror scores with greater variance. The scores of individual males were also more consistent through time than were those of females. Females close to spawning had the highest mirror scores, whereas male scores were highest early in the reproductive cycle. Selection has apparently favored aggressiveness in both sexes. We argue, however, that differences in aggression are the result of selection acting dissimilarly on the two sexes.     

Evidence for female-biased dispersal in the protandrous hermaphroditic Asian Seabass, Lates calcarifer

Movement of individuals influences individual reproductive success, fitness, genetic diversity and relationships among individuals within populations and gene exchange among populations. Competition between males or females for mating opportunities and/or local resources predicts a female bias in taxa with monogamous mating systems and a male-biased dispersal in polygynous species. In birds and mammals, the patterns of dispersal between sexes are well explored, while dispersal patterns in protandrous hermaphroditic fish species have not been studied. We collected 549 adult individuals of Asian seabass (Lates calcarifer) from four locations in the South China Sea. To assess the difference in patterns of dispersal between sexes, we genotyped all individuals with 18 microsatellites. Significant genetic differentiation was detected among and within sampling locations. The parameters of population structure (F(ST)), relatedness (r) and the mean assignment index (mAIC), in combination with data on tagging-recapture, supplied strong evidences for female-biased dispersal in the Asian seabass. This result contradicts our initial hypothesis of no sex difference in dispersal. We suggest that inbreeding avoidance of females, female mate choice under the condition of low mate competition among males, and male resource competition create a female-biased dispersal. The bigger body size of females may be a cause of the female-biased movement. Studies of dispersal using data from DNA markers and tagging-recapture in hermaphroditic fish species could enhance our understanding of patterns of dispersal in fish.     

Relatedness structure in Rhododendron metternichii var. hondoense revealed by microsatellite analysis

The relatedness structure of Rhododendron metternichii Sieb. et Zucc. var. hondoense Nakai was analysed in a 150 x 70-m quadrat in Hiroshima Prefecture, western Japan. The population of R. metternichii occurred as three subpopulations at the study site (A1-A3). Pairwise relatedness based on microsatellite genotypes at eight loci and Mantel tests revealed a hierarchical structure of relatedness within and among subpopulations: (i) relatedness between individuals within 10 m of one another was significantly positive; (ii) relatedness between individuals in the same subpopulation was significantly positive, but negative between individuals in distant subpopulations; and (iii) relatedness was not significantly different from zero among neighbouring subpopulations. In detail, however, relatedness within each subpopulation was significantly positive in subpopulation A1, relatively weak but significantly positive in subpopulation A2, and not significantly different from zero in subpopulation A3. Relatedness within each subpopulation was inversely related to correlations between interindividual distance and relatedness. The aggregation of related individuals at short distances from one another may lead to decreasing relatedness within subpopulations as a whole. Moreover, negative correlations between interindividual distance and relatedness corresponded to high flowering densities at less than 10-m distance, implying that high flowering densities reduce pollinator foraging distance and lead to stronger genetic structure within subpopulations. Small individuals (< 2.0 m in height) showed stronger genetic structure compared with that of large individuals (> or = 2.0 m in height). The different relatedness structure within and among subpopulations may be caused by various degrees of gene flow affected by distribution patterns of individuals and population density.     

Predation risk and alternative mating tactics in male Trinidadian guppies (Poecilia reticulata)

In the guppy (Poecilia reticulata), males have two alternative mating tactics. Individual males may either display to a receptive female prior to attempting to copulate with her or attempt to quickly sneakcopulate with a female without first displaying to her or without a prior receptive response from her. In this study, I experimentally investigated the effects of simulated local increases in the risk of predation (in the form of a cichlid fish predator model in situ) on the mating tactics used by free-ranging male guppies in two typical macrohabitats (riffle and pool) of a Trinidadian river. Focal male guppies displayed to females significantly less often on average, and conversely attempted sneak copulations more often, in the presence of the predator model than in its absence; this pattern was similar for both habitats. These fish therefore performed a lower proportion of sigmoid displays and increased their sneaky mating attempts when the apparent risk of predation had increased locally. This predator-mediated response is consistent with a trade-off between mating success and risk of mortality due to predation. The results are the first to confirm risk-sensitive mating behaviour in free-ranging male guppies within a population, and demonstrate the potential importance of predators in influencing the relative use of alternative mating tactics in this species on a microgeographical scale in the wild.     

Resources, not Kinship, Determine Social Patterning in the Territorial Gunnison's prairie Dog (Cynomys gunnisoni)

In this study, we describe patterns of relatedness in Gunnison's prairie dog ( Cynomys gunnisoni ) social groups. Kin selection is often cited as a mechanism for the evolution and maintenance of social groups, and Gunnison's prairie dog females are occasionally described as being strongly philopatric. Overall, randomization tests revealed that females within territorial groups were not more closely related to each other than expected at random. A similar pattern was found among males and between males and females, indicating that there was no sex-biased dispersal occurring in these populations. Ecological variables measured in this study, such as food abundance and food dispersion, were not correlated with relatedness. In addition, territory size and density/m² did not correlate significantly with relatedness. Although there was variability in the spatial overlap among individuals within groups, there was no indication that relatedness explained this variation. These results suggest that kin selection is not maintaining social groups in these populations, but that competition for access to resources required by both males and females may explain dispersal and social group patterns in these populations.     

Female‐biased dispersal alone can reduce the occurrence of inbreeding in black grouse (Tetrao tetrix)

Although inbreeding depression and mechanisms for kin recognition have been described in natural bird populations, inbreeding avoidance through mate choice has rarely been reported suggesting that sex‐biased dispersal is the main mechanism reducing the risks of inbreeding. However, a full understanding of the effect of dispersal on the occurrence of inbred matings requires estimating the inbreeding risks prior to dispersal. Combining pairwise relatedness measures and kinship assignments, we investigated in black grouse whether the observed occurrence of inbred matings was explained by active kin discrimination or by female‐biased dispersal. In this large continuous population, copulations between close relatives were rare. As female mate choice was random for relatedness, females with more relatives in the local flock tended to mate with genetically more similar males. To quantify the initial risks of inbreeding, we measured the relatedness to the males of females captured in their parental flock and virtually translocated female hatchlings in their parental and to more distant flocks. These tests indicated that dispersal decreased the likelihood of mating with relatives and that philopatric females had higher inbreeding risks than the actual breeding females. As females do not discriminate against relatives, the few inbred matings were probably due to the variance in female dispersal propensity and dispersal distance. Our results support the view that kin discrimination mate choice is of little value if dispersal effectively reduces the risks of inbreeding.     

Natal dispersal and parental escorting predict relatedness between mates in a passerine bird

Although relatedness between mates is of considerable evolutionary and ecological significance, the way in which the level of relatedness is determined by different behavioural processes remains largely unknown. We investigated the role of behaviour in predicting mate relatedness in great tits using genotypic markers and detailed observations. We studied how mate relatedness is influenced by natal dispersal, inbreeding/outbreeding avoidance after natal dispersal and a behaviour not previously considered that influences membership to social aggregations, namely family escorting behaviour by parents. Among locally born individuals, the level of mate relatedness decreased with natal dispersal distance for females, but not for males. In contrast, mate relatedness was negatively related to the extent of family movements for males, but not for females. However, family movements did not predict dispersal distance for either sex. Local recruits were more related to their mates than immigrants, but this was only significant for females. No evidence was found for inbreeding/outbreeding avoidance after dispersal. Our results suggest that, in highly mobile species, mating options are spatially and/or socially limited, and that parents influence mating options of their offspring before dispersal.     

Sex-related differences in the diet of the mink Mustela vison

The diets of free-ranging male and female mink were sampled by analysing faeces collected from radio-tagged individuals. Significant sex differences were apparent in the predation upon three of the five principal prey groups. The larger males preyed much more heavily upon lagomorphs, the largest prey taken, while females preyed more upon fish and crustaceans than did males. These differences were consistent in each season except the autumn (September to November), when males preyed more heavily upon fish and crustacea than did females. Due to their large size, adult lagomorphs are felt to be relatively unavailable to female mink. Male mink are apparently large enough to specialize on lagomorphs, and male mink niche breadth was consistently lower than that of females. Dietary overlap approximated to 40% in all seasons except summer (68%), when female predation upon lagomorphs reached a peak. This reduction in intraspecific feeding competition was felt to be a valuable side-effect of body-size dimorphism.     

Highly asymmetric fine-scale genetic structure between sexes of African striped mice and indication for condition dependent alternative male dispersal tactics

Sex-biased dispersal is observed in many taxa, but few studies have compared sex-biased dispersal among and within populations. We addressed the magnitude and habitat dependency of sex-biased dispersal in social African striped mice by separating group-related from population-related genetic variance to understand the contribution of each sex to deme structure. As dispersal over unoccupied habitat is likely to be more costly than dispersal within a population, we predicted that individuals leaving the natal population have a lower body condition, being inferior to heavier territorial individuals. Fine-scale genetic structure was detected in both sexes. Female relatedness decreased continuously from R = 0.21 at 25 m to zero at 500 m. Maximum male relatedness R = 0.05 was constant at distances between 25 and 75 m, becoming zero at 100 m. Genetic variance (F(ST) ) among seven locations was significantly higher in females than in males, while inbreeding estimates (F(IS) ) were significantly higher in males than in females. Assignment tests estimated significantly more migrants among males, while Bayesian clustering estimated only a single genetic unit cluster for males among the seven locations. The mean body mass of migrant males (44 g) was significantly lower than for males that remained resident and thus dispersed within their sub-population (48 g). Combined, the results showed habitat-independent male-biased dispersal and high female philopatry, and suggested that body condition was more important than kinship in male dispersal decisions. We suggest that locally inferior males are important for gene flow between sub-populations. Thus, males might follow alternative dispersal tactics.     

No evidence for female kin association,indications for extragroup paternity,and sex‐biased dispersal patterns in wild western gorillas

Characterizing animal dispersal patterns and the rational behind individuals’ transfer choices is a long‐standing question of interest in evolutionary biology. In wild western gorillas (Gorilla gorilla), a one‐male polygynous species, previous genetic findings suggested that, when dispersing, females might favor groups with female kin to promote cooperation, resulting in higher‐than‐expected within‐group female relatedness. The extent of male dispersal remains unclear with studies showing conflicting results. To investigate male and female dispersal patterns and extragroup paternity, we analyzed long‐term field observations, including female spatial proximity data, together with genetic data (10 autosomal microsatellites) on individuals from a unique set of four habituated western gorilla groups, and four additional extragroup males (49 individuals in total). The majority of offspring (25 of 27) were sired by the group male. For two offspring, evidence for extragroup paternity was found. Contrarily to previous findings, adult females were not significantly more related within groups than across groups. Consistently, adult female relatedness within groups did not correlate with their spatial proximity inferred from behavioral data. Adult females were similarly related to adult males from their group than from other groups. Using R _ST statistics, we found significant genetic structure and a pattern of isolation by distance, indicating limited dispersal in this species. Comparing relatedness among females and among males revealed that males disperse farer than females, as expected in a polygamous species. Our study on habituated western gorillas shed light on the dispersal dynamics and reproductive behavior of this polygynous species and challenge some of the previous results based on unhabituated groups.     

The role of sex ratios and resource availability on the pre-mating behavior of a monogamous fish

Convict cichlid fish (Archocentrus nigrofasciatus) are one of the few monogamous species where the female appears more colorful than the male. We examined whether this sexual dimorphism was reflected in a female-biased actual sex ratio (ASR) and whether convict cichlid females would exhibit behavioral traits typified in monogamous males when they possessed the extravagant colors and/or morphology. Our field observations revealed that females are marginally, but not significantly, more abundant than males but they did initiate more intersexual social interactions (i.e. approach behavior) than males. They also showed more intrasexual aggression. Furthermore, males more commonly chased females rather than vice versa. Our laboratory experiment also indicated that differences in intrasexual aggression were not related to differences in the ASR but did appear with the addition of a breeding site. Thus, while intrasexual competition was more frequently observed between females in the field it was probably related to the availability of breeding sites rather than our estimates of the ASR.