Intragroup Relatedness Research Articles

Age- and sex-dependent variation in relatedness corresponds to reproductive skew, territory inheritance, and workload in cooperatively breeding cichlids.

Kin selection plays a major role in the evolution of cooperative systems. However, many social species exhibit complex within‐group relatedness structures, where kin selection alone cannot explain the occurrence of cooperative behavior. Understanding such social structures is crucial to elucidate the evolution and maintenance of multi‐layered cooperative societies. In lamprologine cichlids, intragroup relatedness seems to correlate positively with reproductive skew, suggesting that in this clade dominants tend to provide reproductive concessions to unrelated subordinates to secure their participation in brood care. We investigate how patterns of within‐group relatedness covary with direct and indirect fitness benefits of cooperation in a highly social vertebrate, the cooperatively breeding, polygynous lamprologine cichlid Neolamprologus savoryi. Behavioral and genetic data from 43 groups containing 578 individuals show that groups are socially and genetically structured into subgroups. About 17% of group members were unrelated immigrants, and average relatedness between breeders and brood care helpers declined with helper age due to group membership dynamics. Hence the relative importance of direct and indirect fitness benefits of cooperation depends on helper age. Our findings highlight how both direct and indirect fitness benefits of cooperation and group membership can select for cooperative behavior in societies comprising complex social and relatedness structures.

Eusociality outcompetes egalitarian and solitary strategies when resources are limited and reproduction is costly.

Explaining the evolution and maintenance of animal groups remains a challenge. Surprisingly, fundamental ecological factors, such as resource variance and competition for limited resources, tend to be ignored in models of cooperation. We use a mathematical model previously developed to quantify the influence of different group sizes on resource use efficiency in egalitarian groups and extend its scope to groups with severe reproductive skew (eusocial groups). Accounting for resource limitation, the model allows calculation of optimal group sizes (highest resource use efficiency) and equilibrium population sizes in egalitarian as well as eusocial groups for a broad spectrum of environmental conditions (variance of resource supply). We show that, in contrast to egalitarian groups, eusocial groups may not only reduce variance in resource supply for survival, thus reducing the risk of starvation, they may also increase variance in resource supply for reproduction. The latter effect allows reproduction even in situations when resources are scarce. These two facets of eusocial groups, resource sharing for survival and resource pooling for reproduction, constitute two beneficial mechanisms of group formation. In a majority of environmental situations, these two benefits of eusociality increase resource use efficiency and lead to supersaturation—a strong increase in carrying capacity. The increase in resource use efficiency provides indirect benefits to group members even for low intra‐group relatedness and may represent one potential explanation for the evolution and especially the maintenance of eusociality and cooperative breeding.

Reproductive promiscuity in the variegated fairy-wren: an alternative reproductive strategy in the absence of helpers?

Cooperative breeding, in which auxiliary group members help rear related, but nondescendent young, is often explained by kin selection. Reproductive monogamy is predicted in cooperatively breeding systems, as monogamy increases intragroup relatedness and maximizes auxiliary inclusive fitness. While monogamy is observed across many systems, including eusocial insects and cooperatively breeding mammals, some cooperatively breeding birds exhibit high rates of extrapair paternity. Here we quantify paternity and examine the role of auxiliaries on extrapair paternity in the highly cooperative variegated fairy-wren, Malurus lamberti, a species with both male and female auxiliaries. Extrapair paternity occurred in 55.4% of nests, and 39.8% of offspring were the result of extrapair matings. The presence of both male and female auxiliaries had a positive relationship with the percentage of within-pair young sired by dominant males, however, the presence of male auxiliaries had a stronger impact than the presence of females. The number of extrapair young sired by dominant males decreased as the number of male auxiliaries increased. The total number of young sired by dominant males, however, was not predicted by group size or relatedness to their social partner, nor did group composition or relatedness to the breeding pair predict the reproductive success of subordinate males. We hypothesize that breeders use alternative reproductive strategies in the presence or absence of auxiliaries. Males and females may seek extrapair reproductive opportunities when no help is available in their group and nest survival is expected to be low. When help is available, breeders may reduce extrapair paternity, either to increase intragroup relatedness or because confidence in nest survival is high. Our data suggest that group composition is important in understanding extrapair paternity rates in cooperatively breeding birds and that variation in extrapair paternity rates may be the result of flexible breeding strategies when auxiliary presence and identity varies.

Direct benefits and evolutionary transitions to complex societies.

The selective forces that drive the evolution of cooperation have been intensely debated. Evolutionary transitions to cooperative breeding, a complex form of cooperation, have been hypothesized to be linked to low degrees of promiscuity, which increases intragroup relatedness and the indirect (that is, kin selected) benefits of helping. However, ecological factors also promote cooperative breeding, and may be more important than relatedness in some contexts. Identifying the key evolutionary drivers of cooperative breeding therefore requires an integrated assessment of these hypotheses. Here we show, using a phylogenetic framework that explicitly evaluates mating behaviours and ecological factors, that evolutionary transitions to cooperative breeding in cichlid fishes were not associated with social monogamy. Instead, group living, biparental care and diet type directly favoured the evolution of cooperative breeding. Our results suggest that cichlid fishes exhibit an alternative path to the evolution of complex societies compared to other previously studied vertebrates, and these transitions are driven primarily by direct fitness benefits.

Higher reproductive success for chimeras than solitary individuals in the kelp Lessonia spicata but no benefit for individual genotypes

Chimerism is a peculiar, yet widespread, type of group living in which genetically heterogeneous entities are created through fusion between conspecifics. Here we tested whether chimerism provides direct benefits to the kelp Lessonia spicata, by analyzing its consequences on reproductive investment and success, at both the genotype and thallus levels. In addition, we quantified the frequency of chimerism in two natural populations, tested if group members were close kin, and evaluated the effects of relatedness and the number of genotypes per thallus on reproduction. Chimeric thalli were frequent (>60 %) in natural populations of L. spicata. In most cases, average intragroup relatedness was not significantly different from the background population. Reproductive investment was not significantly affected by the type of thallus (chimeric versus non-chimeric), by the number of genotypes per thallus or the average relatedness within thallus. Chimerism did not result in net benefits or costs in terms of genotypic reproductive success or probability of reproducing at the genotypic level. Yet, at the thallus level, chimerism increased reproductive success and the probability of reproducing, since more than one genotype reproduced in chimeric thallus. At the population level, chimerism affects L. spicata reproductive success by allowing the coexistence of a higher density of potential reproducers and mates compared to a scenario with only non-chimeric thallus. Chimerism may then have an important effect on the effective population size and possibly in reducing selfing rates.

Intragroup genetic relatedness in two howler monkey species (Alouatta pigra and A. palliata): Implications for understanding social systems and dispersal.

The degree of genetic relatedness among group members is influenced by dispersal, group formation and composition, mating systems, and other socioecological factors. Making inferences about differences between species in their socio-genetic structure is difficult because studies rarely compare multiple species. In this study, we use multilocus microsatellite genotype data to analyze intragroup genetic relatedness in two howler monkey species (Alouatta palliata and A. pigra). We test the prediction that their patterns of intragroup genetic relatedness will be distinct based on expectations derived from their distinct social systems. Alouatta palliata is expected to have low levels of intragroup relatedness, given that both males and females are reported to disperse from their natal groups, and to join groups with no close kin. Levels of relatedness among A. pigra group members are expected to be variable according to the history of group formation, with new groups formed by unrelated individuals and well-established groups having close kin due to female nepotism and sometimes by takeovers by coalitions of related males. Our results indicate that in both species, most groups contain closely related same-sex and/or inter-sex dyads. This suggests that philopatry in A. palliata may be more common than reported or that individuals are using alternative strategies to reside with close kin. We found greater variation among groups in female-female relatedness in A. palliata than in A. pigra, implying that these species have distinct socio-genetic structures. Further studies including both long-term observational and genetic data are necessary to understand the mechanisms that determine the degree of variation in intragroup genetic relatedness within and among populations for both species. Ecological and demographic data are also necessary to determine the importance of other factors, especially habitat loss and fragmentation, in determining the degree of relatedness in howler monkey groups.

Intra-group relatedness affects parental and helper investment rules in offspring care

In any system where multiple individuals jointly contribute to rearing offspring, conflict is expected to arise over the relative contributions of each carer. Existing theoretical work on the conflict over care has: (a) rarely considered the influence of tactical investment during offspring production on later contributions to offspring rearing; (b) concentrated mainly on biparental care, rather than cooperatively caring groups comprising both parents and helpers; and (c) typically ignored relatedness between carers as a potential influence on investment behavior. We use a game-theoretical approach to explore the effects of female production tactics and differing group relatedness structures on the expected rearing investment contributed by breeding females, breeding males, and helpers in cooperative groups. Our results suggest that the breeding female should pay higher costs overall when helpful helpers are present, as she produces additional offspring to take advantage of the available care. We find that helpers related to offspring through the breeding female rather than the breeding male should contribute less to care, and decrease their contribution as group size increases, because the female refrains from producing additional offspring to exploit them. Finally, within-group variation in helper relatedness also affects individual helper investment rules by inflating the differences between the contributions to care of dissimilar helpers. Our findings underline the importance of considering maternal investment decisions during offspring production to understand investment across the entire breeding attempt, and provide empirically testable predictions concerning the interplay between maternal, paternal and helper investment and how these are modified by different relatedness structures.

Kin association during brood care in a facultatively social bird: active discrimination or by‐product of partner choice and demography?

Intra-group relatedness does not necessarily imply kin selection, a leading explanation for social evolution. An overlooked mechanism for generating population genetic structure is variation in longevity and fecundity, referred to as individual quality, affecting kin structure and the potential for cooperation. Individual quality also affects choosiness in partner choice, a key process explaining cooperation through direct fitness benefits. Reproductive skew theory predicts that relatedness decreases with increasing group size, but this relationship could also arise because of quality-dependent demography and partner choice, without active kin association. We addressed whether brood-rearing eider (Somateria mollissima) females preferentially associated with kin using a 6-year data set with individuals genotyped at 19 microsatellite loci and tested whether relatedness decreased with increasing female group size. We also determined the relationship between local relatedness and indices of female age and body condition. We further examined whether the level of female intracoalition relatedness differed from background relatedness in any year. As predicted, median female intra-group relatedness decreased with increasing female group size. However, the proportion of related individuals increased with advancing female age, and older females prefer smaller brood-rearing coalitions, potentially producing a negative relationship between group size and relatedness. There were considerable annual fluctuations in the level of relatedness between coalition-forming females, and in 1year this level exceeded that expected by random association. Thus, both passive and active mechanisms govern kin associations in brood-rearing eiders. Eiders apparently can discriminate between kin, but the benefits of doing so may vary over time.

Linkage disequilibrium in French wild cherry germplasm and worldwide sweet cherry germplasm

A basic knowledge on linkage disequilibrium (LD) is necessary in order to determine resolution of association studies. We investigated the extent and patterns of LD in a self-incompatible species (Prunus avium L.), in 3 groups (wild cherry, sweet cherry landraces and sweet cherry modern varieties), using a set of 35 microsatellite markers and the gametophytic self-incompatibility locus. Since population structure might create spurious LD, we thus used the information provided by a structure analysis published in a previous study to perform the LD analysis. In the current study, we detected a greater LD extent in sweet cherry than in wild cherry, which is plausibly due to the bottleneck associated with domestication and breeding. Higher LD values in sweet cherry sub-groups may be explained by smaller sample sizes. We also showed that the remaining structure in the groups of sweet cherry, in particular landraces, is responsible for a part of the LD extent. Intra-group relatedness may also account for extensive LD in two sub-groups. These results demonstrate, if ever necessary, the importance of controlling the genetic structure and relatedness when estimating LD. Moreover, LD decays very rapidly with genetic linkage distance in both wild and sweet cherries, which seems promising for future association studies.

Nestmate recognition and intraspecific chemical and genetic variation in Temnothorax ants

The evolutionary stability of social cooperation requires altruistic acts to be directed mainly towards related individuals. To maintain significant intragroup relatedness, social insects have evolved sophisticated recognition systems. We compared nestmate recognition abilities of three native European species of the ant genus Temnothorax in relation to variation in cuticular chemicals and intraspecific genetic diversity. Colony take-over and fusions are common in two sibling species T. nylanderi and T. crassispinus, but rare in T. unifasciatus, suggesting relatively inefficient nestmate recognition in the former species. Temnothorax nylanderi and T. crassispinus discriminated less effectively between nestmates and non-nestmates than T. unifasciatus. While chemical profiles of different colonies strongly overlapped in T. crassispinus and T. nylanderi, 97% of all T. unifasciatus samples were, in a discriminant analysis, correctly assigned to their respective colony. Chemical distances between colonies correlated with aggression levels in T. unifasciatus, but not in the other two species. Finally, preliminary genetic results indicated that haplotype divergence within T. nylanderi and T. crassispinus is remarkably low, whereas T. unifasciatus and other congeners are genetically more variable. In theory, loss of genetic variation, including depletion at loci involved in the production of cuticular hydrocarbons, could decrease colony recognition efficiency. We suggest that this might be the case in T. nylanderi and T. crassispinus, leading to colony fusions in the field. Despite their different life histories, these native species show similarities to invasive ants where genetic bottlenecks arising from founder effects are thought to result in a loss of colony boundaries.

Maintaining low intragroup relatedness: evolutionary stability of nonkin social groups

Communal sociality, involving cooperation among unrelated group members, occurs in all of the six families of bees. However, most communal species also occupy nests that persist over many generations. Why, then, do females not remain in their natal nests with their relatives? We examined how sex-biased dispersal contributes to maintaining low intragroup relatedness by documenting dispersal using microsatellite data in the communal halictine bee, Lasioglossum (Chilalictus) hemichalceum. The composition and genotypes of brood from nests excavated late in the reproductive season (late-season) were compared with those of adults from nests excavated from the same aggregation in early winter (post-season). Females left their natal nests if nests were large and if their brothers were present, demonstrating female-biased dispersal to avoid inbreeding in a social insect for the first time. Thus, the occurrence of within-nest mating in communal species, combined with strong selection against inbreeding in the Hymenoptera, appears to select for female-biased dispersal. This in turn results in a population-wide decrease in intracolony relatedness, thereby maintaining cooperation among nonkin in communal Hymenoptera.

“If This Is Week Three, We Must Be Doing ‘Feelings’”: An Essay on the Importance of Client-Paced Group Work

ABSTRACT This essay elaborates a primary assumption about client-paced group work: that if a treatment group operates as a microcosm of the relationship responsibilities in the real world, then the ownership of the treatment process through client-paced assimilation of treatment goals can be seen as a more appropriate reflection of and role model for healthy relationships. Whether a treatment group follows an open or closed format, the notion that all clients are expected to accomplish specified goals in a given amount of time may provoke client, therapist, and contextual resistance. This essay uses the McGill Domestic Violence Clinic's treatment group for men who batter as a model to illuminate the concepts of client responsibility and strong intra-group relatedness through client-paced treatment and emergent theme discussion.

When to parasitize? A dynamic optimization model of reproductive strategies in a cooperative breeder

We consider a cooperatively breeding group and find the optimal pattern of reproductive parasitism by a subordinate helper as a function of its body size, and hence the share of reproduction obtained by the subordinate. We develop the model for the social system of the cooperatively breeding cichlid fish Neolamprologus pulcher but the general framework is also applicable to other cooperative systems. In addition to behaving cooperatively by sharing tasks, sexually mature male cichlid helpers may directly parasitize the reproduction of dominant breeders in the group. We investigate the relative influence of life history and behavioural variables including growth, parasitism capacity, future reproductive fitness benefits and costs, relatedness and expulsion risk on the optimal reproductive strategy of subordinates. In a detailed analysis of the parameter space we show that a male helper should base its decision to parasitize primarily on an increase in expulsion risk resulting from reproductive parasitism (punishment), intra-group relatedness and the parasitism capacity. If expulsion risk is high then helpers should not parasitize reproduction at medium body size but should parasitize either when small or large.

The price of eggs: increased investment in egg production reduces the offspring rearing capacity of parents

Understanding the selective pressures shaping the number of offspring per breeding event is a key area in the study of life-history strategies. However, in species with parental care, costs incurred in offspring production, rather than rearing, have been largely ignored in both theoretical and empirical studies until relatively recently. Furthermore, the few experimental studies that have manipulated production costs have not yet teased apart effects that operate via the parental phenotype from effects on the quality of the resulting young. To examine whether increased egg production influences parental brood rearing capacity independently of effects operating via egg quality, we experimentally increased egg production in gulls and then examined their capacity to rear a control clutch. We found that the capacity of parents to rear the control brood was substantially reduced solely as a consequence of having themselves produced one extra egg. The paradox that, in many species, parents apparently aim for fewer young per breeding event than the experimentally and theoretically demonstrated optimum, has partly arisen from the failure to take into account the constraints imposed by production costs.

Vibrio pectenicida sp. nov., a pathogen of scallop (Pecten maximus) larvae.

Five strains were isolated from moribund scallop (Pecten maximus) larvae over 5 years (1990-1995) during outbreaks of disease in a hatchery (Argenton, Brittany, France). Their pathogenic activity on scallop larvae was previously demonstrated by experimental exposure. The phenotypic and genotypic features of the strains were identical. The G + C content of the strains was in the range 39-41 mol%. DNA-DNA hybridization showed a minimum of 73% intragroup relatedness. Phylogenetic analysis of small-subunit rRNA sequences confirmed that these strains should be affiliated within the family Vibrionaceae and that they are closely related to Vibrio tapetis and Vibrio splendidus. Phenotypic and genotypic analyses revealed that the isolates were distinct from these two vibrios and so constitute a new species in the genus Vibrio. They utilized only a limited number of organic substrates as sole carbon sources, including betaine and rhamnose, but did not utilize glucose and fructose. In addition, their responses were negative for indole, acetoin, decarboxylase and dihydrolase production. The name Vibrio pectenicida is proposed for the new species; strain A365 is the type strain (= CIP 105190T).

Infant-nonmother interactions of free-ranging mantled howlers (Alouatta palliata) in Costa Rica

Proximate and ultimate explanations of interactions between infants and nonmothers vary depending upon the relatedness of the interactors. We investigated interactions of infants and nonmothers from a 22-month continuous study and from the long-term monitoring of the mantled howler population of La Pacifica, Guanacaste Province, Costa Rica. Relatedness is low or absent in these mantled howler groups. Juvenile females appeared to practice care skills with older infants, but as most first infants died, they failed to benefit. Infant positive interactions with adults occurred with the mother and probable father. Other adult females behaved aggressively toward the youngest infants. Mothers were retentive of infants and responded negatively to these interactions, suggesting that they perceived them as threatening. Interactions with infants appear to reflect competition in groups of unrelated adults. A review of other populations of Alouatta palliata and other species of howlers indicate variability in social group size and suggest variability in intragroup relatedness. We suggest that further study will confirm that social behavior (including interactions with infants) will vary by resource availability (group size) and associated demographic patterns (male and female migration) that affect relatedness in howler social groups.

Reproductive Bribing And Policing As Evolutionary Mechanisms For The Suppression Of Within‐Group Selfishness

We show that a new, simple, and robust general mechanism for the social suppression of within-group selfishness follows from Hamilton's rule applied in a multilevel selection approach to asymmetrical, two-person groups: If it pays a group member to behave selfishly (i.e., increase its share of the group's reproduction, at the expense of group productivity), then its partner will virtually always be favored to provide a reproductive "bribe" sufficient to remove the incentive for the selfish behavior. The magnitude of the bribe will vary directly with the number of offspring (or other close kin) potentially gained by the selfish individual and inversely with both the relatedness r between the interactants and the loss in group productivity because of selfishness. This bribe principle greatly extends the scope for cooperation within groups. Reproductive bribing is more likely to be favored over social policing for dominants rather than subordinates and as intragroup relatedness increases. Finally, analysis of the difference between the group optimum for an individual's behavior and the individual's inclusive fitness optimum reveals a paradoxical feedback loop by which bribing and policing, while nullifying particular selfish acts, automatically widen the separation of individual and group optima for other behaviors (i.e., resolution of one conflict intensifies others).

Paenibacillus apiarius sp. nov.

The name "Bacillus apiarius" Katznelson 1955 was not included on the Approved Lists of Bacterial Names and thus lost standing in bacterial nomenclature. The genetic homogeneity of "B. apiarius" strains was assessed by determining their G+C contents by the buoyant density method and by measuring the levels of DNA relatedness by spectrophotometric reassociation procedures. The G+C contents of the 15 strains examined, ranged from 52 to 54 mol%. DNA reassociation revealed the presence of two clusters, each with high levels of intragroup relatedness (60 to 100%). One cluster consisted of six strains highly related to Bacillus thiaminolyticus, and the other consisted of nine strains related to the designated type strain of "B. apiarius." The strains in the second cluster were not closely related genetically to the type strains of organisms frequently associated with honey bees (namely, Paenibacillus alvei, Paenibacillus larvae, Bacillus laterosporus, and Paenibacillus pulvifaciens). The "B. apiarius" strains in the second cluster were also phenotypically homogeneous and distinguishable from the previously described species. Comparative analyses of the 16S rRNA gene DNA sequence showed that the proper phylogenetic position of the second cluster was in the genus Paenibacillus. These findings justify the proposal of a new species with the name Paenibacillus apiarius. The type strain is NRRL NRS-1438.

Vibrio tapetis sp. nov., the Causative Agent of the Brown Ring Disease Affecting Cultured Clams

A taxonomic characterization was carried out on strains of the bacteria that cause the brown ring disease of clams. On the basis of their phenotypic and genotypic characteristics, these strains can be considered to constitute a new taxonomic unit, distinct from other Vibrio species. The guanine-plus-cytosine content of the strains ranged between 42.9 and 45.5 mol% (43.2 mol% for the proposed type strain). DNA-DNA hybridization studies showed 100% intragroup relatedness, but levels of genetic relatedness to the reference strains of different Vibrio species tested ranged between 15 and 58%. The strains have all the properties characteristic of the genus Vibrio and can be clearly differentiated from other species of this genus by their growth at 4°C and their negative responses for growth at 30°C and in 6% NaCl, arginine dehydrolase, lysine decarboxylase, ornithine decarboxylase, and Voges-Proskauer reaction. The name Vibrio tapetis is proposed for the new species; strain B1090 (CECT 4600) is the type strain.

DNA relatedness among Bacillus thuringiensis serovars.

The genetic relationships of Bacillus cereus and of the Bacillus thuringiensis serovars were assessed from measurements of DNA reassociation. A study of 8 to 10 strains each of 13 of the most commonly encountered serovars revealed that the levels of intragroup DNA relatedness for most serovars ranged from 90 to 100%. In contrast, B. thuringiensis serovars canadensis and kenyae consisted of two DNA relatedness groups, each of which exhibited levels of intragroup relatedness of 80% or higher and levels of intergroup relatedness of 60 to 70%. Analyses of DNA relatedness performed with all of the serovars revealed that the taxa were segregated into 11 phena differentiated from each other at about the 65% level; within each phenon the level of relatedness was 80% or higher. Three phena contained strains belonging to more than one serovar; B. thuringiensis serovars alesti and dendrolimus clustered in phenon 1, serovars aizawai, kurstaki, galleriae, and morrisoni clustered in phenon 7, and serovar darmstadiensis and some strains of serovar kenyae clustered in phenon 11. The levels of DNA relatedness between B. cereus and B. thuringiensis strains ranged between 65 and 70%. My results suggest that many of the B. thuringiensis serovars are genetically distinct but closely related.