Sociality,Bateman’s gradients,and the polygynandrous genetic mating system of round-tailed ground squirrels (<Emphasis Type="Italic">Xerospermophilus tereticaudus</Emphasis>)

Historically, most mammals have been classified as polygynous; although recent molecular evidence suggests that many mammals may be polygynandrous, particularly the ground-dwelling sciurids. We genotyped 351 round-tailed ground squirrels (Xerospermophilus tereticaudus) using seven microsatellite loci to determine paternity in 31 litters from 2004 to 2007. Polygyny was evident in all years except in 2007, when the population size was reduced. Multiple paternity occurred in the majority of litters (55%) with 2.5 ± 0.26 sires/litter (n = 31). Forty-nine percent of resident males (n = 114) sired offspring, and of males that sired offspring (n = 56) 27% sired young in multiple litters in a single breeding season. Litter size was positively correlated with the number of sires. Through an indirect analysis of paternity, we found 21 litters (68%) with an average relatedness of 0.5 or less. Males had a greater opportunity for sexual selection (I _s = 1.60) than females (I _s = 0.40); Bateman’s gradient was also greater in males (1.07 ± 0.04, n = 56) than females (0.82 ± 0.08, n = 31). The mating system in round-tailed ground squirrels defined through genetic analyses and Bateman’s gradients is polygynandrous compared to the previously suggested polygynous mating system as established by behavioral observations and fits within the predictions of the ground squirrel sociality models. Upon evaluating the predictions of the sociality models among sciurid species, we found a negative relationship between the level of sociality with litter size and the average percentage of multiple paternity within a litter. Thus, recent genetic information and reclassification of mating systems support the predictions of the ground-dwelling squirrel sociality models.     

Social organization and space-use in Gunnison's prairie dog

Summary Social organization of the Gunnison's prairie dog, Cynomys gunnisoni, was studied in two populations in south-central Colorado. Gunnison's prairie dogs live in complex, interactive societies fitting current definitions of highly social ground squirrels. Members of harems (coteries) cooperatively use and defend a common territory. Spatial overlap is extensive between the adult male(s) and adult females, and among adult females within the harem through the active season. Amicable behavioral interactions are frequent within the harem, whereas interactions between members of different harems are primarily agonistic and spatial overlap is minimal. Although their behavioral repertoire is more limited, social organization of the Gunnison's prairie dog most closely resembles that of the black-tailed prairie dog, C. ludovicianus. Although body size, age of first reproduction, and age of emigration differed between the two study populations (Rayor 1985a), a comparison of social traits did not reveal substantial differences.     

Complex social structure of southern flying squirrels is related to spatial proximity but not kinship

Social individuals have organized relationships that affect fitness and so a species' tendency to be social has important implications for its population ecology, gene flow, and its distribution in space and time. We quantitatively examined the social structure of southern flying squirrels (Glaucomys volans) and tested for a role of kinship and prior familiarity in predicting social structure. To quantify social structure, we monitored nest group composition of southern flying squirrels. All squirrels at the study site were marked with passive integrated transponder (PIT) tags and nest cavity entrances were monitored with automated PIT tag recorders for a period of 28 months. Squirrels were genotyped at eight microsatellite loci. Permutation tests of associations suggested that individuals nested with other specific individuals more often than expected by chance. The lagged association rate indicated that relationships were stable and persisted across seasons and years. Multiple summer nest associates came together in winter to form larger nest groups which were likely important for social thermoregulation. A measure of prior familiarity, but not kinship, was related to the proportion of time individuals nested together during winter. We suggest that the evolution of sociality in southern flying squirrels is driven largely by mutually beneficial behaviors related to social thermoregulation although other, as of yet, unidentified mechanisms are needed to explain sociality in the warm season. We hypothesize that minimum group size requirements associated with social thermoregulation could explain the absence of this species in patchy landscapes and aspects of range boundary dynamics near their northern range boundary.     

Sibling recognition in thirteen-lined ground squirrels: effects of genetic relatedness,rearing association,and olfaction

Summary I investigated sibling-sibling recognition in captive thirteen-lined ground squirrels (Spermophilus tridecemlineatus) by cross-fostering lab-born pups shortly after birth. When young reached about 45 days of age, I observed dyadic interactions in a test arena of pairs from four relatedness X rearing groups, and recorded the frequency of exploratory encounters between individuals. Sibs-reared together and nonsibs-reared together exhibited significantly fewer exploratory encounters than either sibs-reared apart or nonsibsreared apart. Young reared together were equally exploratory, regadless of relatedness; similarly, young reared apart, whether they were sibs or nonsibs, showed similar levels of exploration. Thus, the differential treatment of siblings in the lab appears to be based on rearing association and not genetic relatedness per se. I interpret this recognition based on association (rearing familiarity) in the context of the species' social organization and compare my results on S. tridecemlineatus with similar studies on S. beldingi, S. parryii, and S. richardsonii. I also used an olfactory impairment technique (zinc sulfate) and found that differential treatment in thirteen-lined ground squirrels was influenced by olfactory cues.     

Association patterns and kinship in female Indo-Pacific bottlenose dolphins (Tursiops aduncus) of southeastern Australia

Kinship has been shown to be an important correlate of group membership and associations among many female mammals. In this study, we investigate association patterns in female Indo-Pacific bottlenose dolphins (Tursiops aduncus) inhabiting an embayment in southeastern Australia. We combine the behavioral data with microsatellite DNA and mitochondrial DNA data to test the hypotheses that genetic relatedness and maternal kinship correlate with associations and social clusters. Mean association between females was not significantly different from a random mean, but the standard deviation was significantly higher than a random standard deviation, indicating the presence of nonrandom associates in the dataset. A neighbor-joining tree, based on the distance of associations between females, identified four main social clusters in the area. Mean genetic relatedness between pairs of frequent female associates was significantly higher than that between pairs of infrequent associates. There was also a significant correlation between mtDNA haplotype sharing and the degree of female association. However, the mean genetic relatedness of female pairs within and between social clusters and the proportion of female pairs with the same and different mtDNA haplotypes within and between clusters were not significantly different. This study demonstrates that kinship correlates with associations among female bottlenose dolphins, but that kinship relations are not necessarily a prerequisite for membership in social clusters. We hypothesize that different forces acting on female bottlenose dolphin sociality appear to promote the formation of flexible groups which include both kin and nonkin.     

Differential aggressiveness between fire salamander (<Emphasis Type="Italic">Salamandra infraimmaculata</Emphasis>) larvae covaries with their genetic similarity

Responding differentially to kin and non-kin is known to be adaptive in many species. One example is the inclusive fitness benefits of reducing aggression toward closer relatives. Little is known, however, about the ability of animals to assess differential degrees of genetic relatedness and to respond accordingly with differential levels of aggression. In the present study, we tested whether aggressiveness between body mass-matched pairs of fire salamander (Salamandra infraimmaculata) larvae covaried with the genetic similarity between them. We quantified aggressiveness at three levels of genetic similarity by selecting pairs within and across pools from recently genotyped populations. We also assessed aggression between pairs of siblings. Aggression and associated injuries decreased as genetic similarity increased across the groups. These findings suggest that cannibalistic salamanders can assess their degree of genetic relatedness to conspecifics and vary their behavioral responses depending on the degree of similarity between them along a genetic relatedness continuum.     

Effects of kinship on growth of a live-bearing reef fish

The influence of genetic relatedness on the individual performance (e.g. growth, development) of animals is often tied to agonistic or cooperative behaviors among conspecifics, and studies of the effects of kinship have produced mixed results. To explore genetic relatedness independent of these behaviors, we investigated the effects of kinship on the growth of the kelp perch Brachyistius frenatus, a live-bearing, planktivorous marine reef fish that is capable of only limited dispersal. Although juveniles occur in aggregations and compete for food resources, they do not exhibit overt aggressive or cooperative behavioral interactions. We hypothesized that under competition and in the absence of these behaviors, sibling and non-sibling groups of juvenile B. frenatus raised at the same densities in the field would not differ in average growth, but that siblings would exhibit lower variation in growth, simply due to genetic similarities in inherent growth rates. Pregnant, female kelp perch were collected and placed in cages until parturition was complete. Groups of young, recently born from the same mother or from different mothers, were then raised in the field for 9 wk. Our results revealed that average growth rates were similar between sibling and non-sibling treatments. While variation in growth increased initially in non-siblings, siblings showed little such variation. This divergence, however, was not consistent over the duration of the experiment, and variation in the growth of siblings ultimately converged with that of non-siblings. Effects of genetic relatedness would be most likely to manifest themselves early after birth, before environmental factors exert their influence, and this may explain the initial separation but eventual convergence in variation in growth between sibling and non-sibling treatments. For B. frenatus and other organisms that will encounter relatives and compete for resources without overt behavioral interactions, the degree of kinship may play a minor role in the demography of local populations. Received: 26 January 1998 / Accepted: 30 September 1998     

Behavioral responses of territorial red squirrels to natural and experimental variation in population density

The relative scarcity of studies at the intersection of behavioral and population ecology is surprising given the presumed importance of behavior in density-dependent population regulation. Here we tested whether North American red squirrels (Tamiasciurus hudsonicus) adjust their behavior in response to local population density and whether they use rates of territorial vocalizations in their local neighborhood to assess density. We examined these relationships using 18?years of live trapping and 20?years of behavioral data that were collected across natural variation in local population density. To disentangle the effects of population density on behavior from those due to changes in per capita food abundance or changes in the frequency of antagonistic interactions with neighbors, we also experimentally manipulated population density with long-term food supplementation as well as perceived population density with long-term playbacks of territorial vocalizations. The frequency with which squirrels emitted territorial vocalizations was positively associated with local population density. In contrast, antagonistic physical interactions observed between squirrels and territorial intrusions were rare and the frequency of intrusions was weakly and negatively, not positively, associated with population density. Squirrels experiencing naturally and experimentally high density conditions spent less time in the nest and feeding but more time being vigilant. Similar density-dependent changes in behavior were observed in response to our manipulations of perceived population density, indicating that vocalization rates and not physical interactions or food abundance were the mechanism by which squirrels assessed and responded behaviorally to changes in local density.     

Social attributes and associated performance measures in marmots: bigger male bullies and weakly affiliating females have higher annual reproductive success

Studying the structure of social interactions is fundamental in behavioral ecology as social behavior often influences fitness and thus natural selection. However, social structure is often complex, and determining the most appropriate measures of variation in social behavior among individuals can be difficult. Social network analysis generates numerous, but often correlated, measures of individual connectedness derived from a network of interactions. We used measures of individual connectedness in networks of affiliative and agonistic interactions in yellow-bellied marmots, Marmota flaviventris, to first determine how variance was structured among network measures. Principal component analysis reduced our set of network measures to four “social attributes” (unweighted connectedness, affiliation strength, victimization, and bullying), which revealed differences between patterns of affiliative and agonistic interactions. We then used these extracted social attributes to examine the relationship between an individual’s social attributes and several performance measures: annual reproductive success, parasite infection, and basal stress. In male marmots, bullying was positively associated with annual reproductive success, while in females, affiliation strength was negatively associated with annual reproductive success. No other social attributes were significantly associated with any performance measures. Our study highlights the utility of considering multiple dimensions when measuring the structure and functional consequences of social behavior.     

Macro-habitat patch structure,environmental harshness,and Marmota flaviventris

Summary Social behavior of 15 colonies of Yellow-bellied Marmots was studied at sites differing in both elevation (plant growing season length) and patch structure (density and spacing of suitable colony sites in large blocks of habitat) from 1976 through 1978. Colonies were typically composed of family groups. In low elevation colonies, offspring dispersed at the end of the juvenile year. In all high elevation areas, offspring dispersed as yearlings, and parent-offspring and sibling interactions during the juvenile year were highly amicable. In high elevation areas in continuous habitat, dispersal by yearlings occurred with no agonistic interactions. In high elevation areas with patchy habitat, however, brief but intense periods of extremely agonistic sibling and parent-offspring interactions occurred immediately prior to dispersal of yearlings.Social behavior and dispersal are uncorrelated with elevation, plant growing season length, or available foraging time. Social interaction among relatives in marmot colonies is amicable when other suitable colony sites are located nearby; in these areas, dispersal occurs without any antagonism. Agonistic behavior occurs only in areas with a patchy distribution of suitable colony sites. At these sites, dispersal occurs only after periods of extreme antagonism. Further, those offspring who do not disperse from the colony site are the ones who dominate and initiate agonistic interactions with their siblings. The large-scale structure of the habitat influences the propensity of the individual to disperse. When the animal is reluctant to disperse because of a combination of high transit difficulty and low desirability of a new site, angonistic social interactions with relatives force dispersal.     

Seasonal decline in intracolony genetic relatedness in eastern tent caterpillars: implications for social evolution

Summary Genetic relatedness in social insect colonies may vary spatially or temporally as a result of changes in colony membership due to immigration or to variation in patterns of maternity and paternity. We estimated relatedness for eastern tent caterpillars (Malacosoma americanum) in laboratory colonies derived from egg masses using multilocus genotypic data derived from electrophoresis. This estimate is compared with estimates obtained from colony samples taken in the field at four intervals spanning the larval developmental season. We found that average intracolony relatedness is close to 0.5 initially but declines through the developmental season due to colony merging, showing that caterpillars do not discriminate between siblings and nonsiblings in order to preserve colony family structure. Using the intracolony values together with relatedness values for higher levels of population structure, we estimated the effective mean number of simple families represented in single colonies through the season. The overall effective number of families per tent increased from one at the time of eclosion to 1.3 by the end of the season. Average intracolony relatedness remained relatively high despite the occurrence of colony merging, apparently as a result of the low density of tents on most trees, combined with high relatedness within the original colonies. Thus, high intracolony relatedness is maintained in M. americanum populations through the effects of adult dispersal, mating, and oviposition patterns, rather than through behavioral discrimination mechanisms of the larvae. These findings underscore the importance of considering the causes of temporal variation in genetic relatedness as well as the consequences for the indirect component of inclusive fitness. Correspondence to: J.T. Costa     

Interspecific associations of Cape ground squirrels with two mongoose species: benefit or cost?

Mixed-species associations have been described in many vertebrate species, but few behavioral studies have investigated associations between species from different mammalian orders. Cape ground squirrels (Xerus inauris) are highly social rodents that inhabit burrows with two species of mongoose, but the benefits of these interspecific associations to ground squirrels remain unresolved. We compared the behavior of squirrels while solitary, with conspecifics, and in the presence of suricates (Suricata suricatta) and yellow mongooses (Cynictis pencillatus). Squirrels spent less time alert and more time feeding when suricates were present, but increased vigilance in the presence of yellow mongooses. In a series of mobbing trials with a puff adder (Bitis arietans), a common predator of all three species, Cape ground squirrels were the most active in mobbing the snake. Our results suggest that Cape ground squirrels benefit from associating with suricates, but not necessarily with yellow mongooses. Both mongoose species benefit from the burrowing activities of the squirrels for thermoregulation and escape from predators, and a suite of other organisms may similarly benefit from the habitat modifications by Cape ground squirrels, suggesting they could be considered ecosystem engineers of the arid and semi-arid regions of southern Africa. Thus, the association between Cape ground squirrels and suricates appears mutually beneficial, whereas yellow mongooses may merely be commensals of the squirrels.     

Alarm calling behavior of the thirteen-lined ground squirrel,Spermophilus tridecemlineatus

Summary Alarm calling in a population of thirteenlined ground squirrels, Spermophilus tridecemlineatus, was studied over a three-year period. Data on ground squirrel reactions to human and canine approaches and to the approach or presence of avian predators were used to quantify alarm calling behavior.The results support the hypothesis that alarm calling in this species functions to warn genetic relatives. Human and canine approach-elicited calls were most frequently given by mothers and their recently emerged young; adult males and nonparous females rarely called. The onset of maternal calling coincided with the aboveground appearance of a mother's own litter, and both juvenile and maternal calling were terminated at approximately three weeks post-emergence. Alarm calls were rarely emitted during encounters with avian predators.Alarm calling behavior in S. tridecemlineatus thus appeared to be dependent upon the presence of newly emerged juvenile relatives. To investigate whether the population structure of S. tridecemlineatus was perhaps incompatible with the evolution of alarm calling directed toward adult relatives, the distance between the home ranges of adult relatives and the distance over which the alarm vocalization is audible to ground squirrels were measured. The results revealed that females were likely to have adult relatives relatives residing within audible range of the call.     

The association of female kin in the arctic ground squirrel Spermophilus parryii

Summary The association of kin in arctic ground squirrels (Spermophilus parryii) was studied near Haines Junction, Yukon Territory, during the summers of 1977, 1978, and 1979. Males dispersed in this population, thus only females were likely to live near relatives.Close female kin (sisters, mothers/daughters) had greater overlap of home ranges and interacted more amicably and less agonistically than did less closely related females. Closely related females clumped their young at emergence, whereas more distantly related females did not; there was little indication of clumping of pre-emerged young. The overlap of home ranges of distant relatives (known genetic relatives that had not associated in a natal burrow) was intermediate to that of close relatives and non-relatives. The types of interactions between distant relatives were more similar to those between non-relatives than between close relatives.I conclude that female arctic ground squirrels exhibit nepotism. Females may benefit from associations with relatives by sharing watching for predators once juveniles become conspicuous. Male arctic ground squirrels commit infanticide and several females may be more effective at protecting their young from infanticidal males than females living alone. I suggest that clumping of young by close relatives may provide a mechanism allowing distantly related females to identify each other.Address for offprint requests     

The genetic population structure of the gray mouse lemur (<Emphasis Type="Italic">Microcebus murinus), </Emphasis>a basal primate from Madagascar

The genetic structure of a population is closely connected to fundamental evolutionary processes and aspects of social behavior. Information on genetic structure is therefore instrumental for the interpretation of social behavior and evolutionary reconstructions of social systems. Gray mouse lemurs (Microcebus murinus) are basal primates endemic to Madagascar whose social organization is characterized by solitary foraging at night and communal resting during the day. Conflicting reports about population structure based on behavioral observations led us to examine the genetic structure of one population in detail in order to: (1) identify natural genetic units in this solitary primate, and (2) to test the assumption of current models of primate social evolution that solitary primates are organized in matrilines. DNA was extracted from tissue samples of 85 individuals from Kirindy forest to determine their variability at a 530 bp fragment of the mitochondrial D-loop and at six microsatellite loci. We found that this population was characterized by a great general diversity among mtDNA haplotypes, a pronounced sex difference in mtDNA haplotype diversity and spatial clustering of females with a particular haplotype, but low average relatedness among members of haplotype clusters. Specifically, we identified 13 different haplotypes, which were unevenly distributed among individuals. About 80% of all individuals, most of which were females or juvenile males, shared a single haplotype. Rare haplotypes were almost exclusively represented by single adult males, who apparently migrated into this population. One other haplotype was represented by a small group of females living at one edge of the study area. Microsatellite analysis revealed above-average relatedness among females with overlapping home ranges, as well as no signs of inbreeding, implying that male dispersal results in high levels of gene flow among matrilineal groups. We conclude that gray mouse lemur populations are hierarchically organized in small family units of closely related females that form stable sleeping groups, several of which are connected through a common mtDNA haplotype and form spatially distinct clusters. The presence of such matrilines supports a basic assumption of current models of primate social evolution.     

Seasonal nestmate recognition in the ant Formica exsecta

Under favorable conditions, the mound-building ant Formica exsecta may form polydomous colonies and can establish large nest aggregations. The lack of worker aggression towards nonnestmate conspecifics is a typical behavioral feature in such social organization, allowing for a free flux of individuals among nests. However, this mutual worker toleration may vary over the seasons and on spatial scales. We studied spatio-temporal variation of worker–worker aggression within and among nests of a polydomous F. exsecta population. In addition, we determined inter- and intracolony genetic relatedness by microsatellite DNA genotyping and assessed its effect on nestmate recognition. We found significant differences in the frequency of worker exchange among nests between spring, summer, and autumn. Moreover, we found significant seasonal variation in the level of aggression among workers of different nests. Aggression levels significantly correlated with spatial distance between nests in spring, but neither in summer nor in autumn. Multiple regression analysis revealed a stronger effect of spatial distances rather than genetic relatedness on aggressive behavior. Because nestmate discrimination disappeared over the season, the higher aggression in spring is most plausibly explained by cue intermixing during hibernation.     

Seasonal polydomy and unicoloniality in a polygynous population of the red wood ant Formica truncorum

Ant colonies may have a single or several reproductive queens (monogyny and polygyny, respectively). In polygynous colonies, colony reproduction may occur by budding, forming multinest, polydomous colonies. In most cases, budding leads to strong genetic structuring within populations, and positive relatedness among nestmates. However, in a few cases, polydomous populations may be unicolonial, with no structuring and intra-nest relatedness approaching zero. We investigated the spatial organisation and genetic structure of a polygynous, polydomous population of Formica truncorum in Finland. F. truncorum shifts nest sites between hibernation and the reproductive season, which raises the following question: are colonies maintained as genetic entities throughout the seasons, or is the population unicolonial throughout the season? Using nest-specific marking and five microsatellite loci, we found a high degree of mixing between individuals of the population, and no evidence for a biologically significant genetic structuring. The nestmate relatedness was also indistinguishable from zero. Taken together, the results show that the population is unicolonial. In addition, we found that the population has undergone a recent bottleneck, suggesting that the entire population may have been founded by a very limited number of females. The precise causes for unicoloniality in this species remain open, but we discuss the potential influence of intra-specific competition, disintegration of recognition cues and the particular hibernation habits of this species.     

Lack of detectable nepotism in multiple-queen colonies of the fire ant Solenopsis invicta (Hymenoptera: Formicidae)

Multiple-queen (polygyne) colonies of the introduced fire ant Solenopsis invicta present a paradox for kin selection theory. Egg-laying queens within these societies are, on average, unrelated to one another, and the numbers of queens per colony are high, so that workers appear to raise new sexuals that are no more closely related to them than are random individuals in the population. This paradox could be resolved if workers discriminate between related and unrelated nestmate sexuals in important fitness-related contexts. This study examines the possibility of such nepotism using methods that combine the following features: (1) multiple relevant behavioral assays, (2) colonies with an unmanipulated family structure, (3) multiple genetic markers with no known phenotypic effects, and (4) a statistical technique for distinguishing between nepotism and potentially confounding phenomena. We estimated relatedness between interactants in polygyne S. invicta colonies in two situations, workers tending egg-laying queens and workers feeding maturing winged queens. In neither case did we detect a significant positive value of relatedness that would implicate nepotism. We argue that the non-nepotistic strategies displayed by these ants reflect historical selection pressures experienced by native populations, in which nestmate queens are highly related to one another. The markedly different genetic structure in native populations may favor the operation of stronger higher-level selection that effectively opposes weaker individual-level selection for nepotistic interactions within nests. Received: 28 June 1996 / Accepted after revision: 6 October 1996     

Individual differences in the behavior of juvenile yellow-bellied marmots

Summary Yellow-bellied marmots express considerable individuality as measured by behavior in a maze, mirror-image stimulation (MIS), and social behavior in the field. Maze behavior discriminated between residents and dispersers; residents explored the maze more widely than did dispersers. Males could not be distinguished from females nor survivors from non-survivors by their maze behavior. A group of six yearling females was established to examine the relationship between individual behavioral phenotypes (as determined by MIS) and social behavior in the field. This experiment provided a situation in which social behavior was not influenced by age, sex, or reproduction (female yearlings are non-reproductive). The number of social interactions per individual ranged from 25 to 69. The number of observed interactions per individual differend significantly from the expected for greeting, allogrooming, total amicable, play, and total social interactions. Rankings of greeting, total amicable, and total interactions were directly correlated with rankings on the avoidance axis; play was inversely correlated with the approach axis. These results suggest that marmots have individual behavioral phenotypes that are expressed in their social interactions with their conspecifics.     

Chaotic genetic patchiness and high relatedness of a poecilogonous polychaete in a heterogeneous estuarine landscape

The genetic structure of benthic marine invertebrates is often described as “chaotic” when genetic structure cannot be explained and barriers to dispersal and gene flow cannot be identified. Here, chaotic patterns of genetic structure for the polychaete Pygospio elegans (Claparède) sampled at 16 locations from the heterogeneous Isefjord–Roskilde Fjord estuary complex in Denmark were found. There was no isolation by distance, and the geography of the estuary complex did not seem to pose a barrier to dispersal and gene flow in this species. We investigated whether characteristics of the environment could be related to the genetic structure and possibly restrict gene flow in this species. Additionally, since P. elegans is poecilogonous, producing larvae with different pelagic developmental periods, we investigated whether observed developmental modes in the samples might clarify the genetic patterns. None of the tested factors explained the population genetic structure. However, a high degree of relatedness among individuals in almost all samples was found. Samples with a larger percentage of young individuals had more related individuals, suggesting that different cohorts could be comprised of individuals with different degrees of relatedness. Relatedness within a site could be increased by limited larval dispersal, collective dispersal of related larvae, sweepstakes reproductive success, or asexual reproduction, but distinguishing between these requires further study. Using a “seascape genetics” approach allowed us to investigate some of the numerous potential factors that could influence population genetic structure in a poecilogonous species.