Variation in colony structure in the subterranean termite Reticulitermes flavipes

The genetic organization of colonies of the subterranean termite Reticulitermes flavipes in two subpopulations in Massachusetts was explored using five polymorphic allozymes and double-strand conformation polymorphism (DSCP) analysis of the mitochondrial control region. Empirically obtained estimates of worker relatedness and F-statistics were compared with values generated by computer simulations of breeding schemes to make inferences about colony organization. In one study site (G), worker genotypes indicated the presence of a mixture of colonies headed by monogamous outbred primary reproductives and colonies headed by inbreeding neotenic reproductives, both colony types having limited spatial ranges. A second site (S) was dominated by several large colonies with low relatedness among nestmates. Mixed DSCP haplotypes in three colonies indicated that nestmates had descended from two or three unrelated female reproductives. Computer simulations of breeding schemes suggested that positive colony inbreeding coefficients at site S resulted from either commingling of workers from different nests or different colonies. Such an exchange of workers between nests corresponds to the multiple-site nesting lifetype of many subterranean termites and resembles colony structure in polycalic Formica ants. Our study demonstrates considerable variation in R. flavipes colony structure over a small spatial scale, including colonies headed by monogamous outbred primary reproductives, colonies containing multiple inbred neotenic reproductives and large polydomous colonies containing the progeny of two or more unrelated queens, and suggests that the number of reproductives and nestmate relatedness change with colony age and size.     

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     

Relatedness,recognition errors,and colony fusion in the termite <Emphasis Type="Italic">Nasutitermes corniger</Emphasis>

Loss of aggression between social groups can have far-reaching effects on the structure of societies and populations. We tested whether variation in the genetic structure of colonies of the termite Nasutitermes corniger affects the probability of aggression toward non-nestmates and the ability of unrelated colonies to fuse. We determined the genotypes of workers and soldiers from 120 colonies at seven polymorphic microsatellite loci. Twenty-seven colonies contained offspring of multiple founding queens or kings, yielding an average within-colony relatedness of 0.33. Genotypes in the remaining 93 colonies were consistent with reproduction by a single queen and king or their progeny, with an average within-colony relatedness of 0.51. In standardized assays, the probability of aggression between workers and soldiers from different colonies was an increasing function of within-colony relatedness. The probability of aggression was not affected significantly by the degree of relatedness between colonies, which was near zero in all cases, or by whether the colonies were neighbors. To test whether these assays of aggression predict the potential for colony fusion in the field, we transplanted selected nests to new locations. Workers and soldiers from colonies that were mutually tolerant in laboratory assays joined their nests without fighting, but workers and soldiers that were mutually aggressive in the assays initiated massive battles. These results suggest that the presence of multiple unrelated queens or kings promotes recognition errors, which can lead to the formation of more complex colony structures.     

A study of the genetic relationships within and among wolf packs using DNA fingerprinting and mitochondrial DNA

Summary DNA fingerprinting and mitochondrial DNA analyses have not been used in combination to study relatedness in natural populations. We present an approach that involves defining the mean fingerprint similarities among individuals thought to be unrelated because they have different mtDNA genotypes. Two classes of related individuals are identified by their distance in standard errors above this mean value. The number of standard errors is determined by analysis of the association between fingerprint similarity and relatedness in a population with a known genealogy. We apply this approach to gray wolf packs from Minnesota, Alaska, and the Northwest Territories. Our results show that: (1) wolf packs consist primarily of individuals that are closely related genetically, but some packs contain unrelated, non-reproducing individuals; (2) dispersal among packs within the same area is common; and (3) short-range dispersal appears more common for female than male wolves. The first two of these genetically-based observations are consistent with behavioral data on pack structure and dispersal in wolves, while the apparent sex bias in dispersal was not expected.USA Offprint requests to: N. Lehman at the present address     

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     

Oligogyny by unrelated queens in the carpenter ant, Camponotus ligniperdus

Multilocus DNA fingerprinting and microsatellite analysis were used to determine the number of queens and their mating frequencies in colonies of the carpenter ant, Camponotus ligniperdus (Hymenoptera: Formicidae). Only 1 of 61 analyzed queens was found to be double-mated and the population-wide effective mating frequency was therefore 1.02. In the studied population, 8 of 21 mature field colonies (38%) contained worker, male, or virgin queen genotypes which were not compatible with presumed monogyny and therefore suggested oligogyny, i.e., the cooccurrence of several mutually intolerant queens within one colony. Estimated queen numbers in oligogynous colonies ranged between two and five. According to the results of the genetic analysis, most of the queens coexisting in oligogynous colonies were not closely related. Pleometrosis is very rare and queenless colonies adopt mated queens both in the laboratory and field. Therefore, the most plausible explanation for the origin of oligogynous colonies in C. ligniperdus is the adoption of unrelated queens by orphaned mature colonies. The coexistence of unrelated, but mutually intolerant queens in C. ligniperdus colonies demonstrates that oligogyny should be considered as a phenomenon distinct from polygyny. Received: 18 December 1997 / Accepted after revision: 20 June 1998     

RAPD markers suggest genotypic effects on forager specialization in a eusocial wasp

Genetic variability within insect societies may provide a mechanism for increasing behavioral diversity among workers, thereby augmenting colony efficiency or flexibility. In order to assess the possibility that division of labor has a genetic component in the eusocial wasp Polybia aequatorialis, I asked whether the genotypes of workers within colonies correlated with behavioral specialization. Workers specialized by foraging for one of the four materials (wood pulp, insect prey, nectar, or water) gathered by their colonies. I collected foragers on 2 days from each of three colonies and identified the material the foragers were carrying when collected. I produced random amplified polymorphic DNA (RAPD) markers from the genomic DNA of these foragers and estimated genotypic similarity of foragers based on sharing of variable RAPD marker bands. Contingency tests on 20 variable loci per colony showed statistically significant (P <0.05) biases in RAPD marker frequencies among forager types in the three colonies. Patterns of association of RAPD marker bands with specializations were constant in two colonies, but changed between collection days in one colony. RAPD marker biases suggest that division of labor among workers includes a genetic component in P. aequatorialis. Colony-level selection on variation in division of labor is a possible factor favoring the evolutionary maintenance of high genotypic variability (low relatedness) in epiponine wasp colonies and in other eusocial insects. Received: 18 July 1995/Accepted after revision: 1 October 1995     

Queen longevity,queen adoption,and posthumous indirect fitness in the facultatively polygynous ant Myrmica tahoensis

In many polygynous ant species, established colonies adopt new queens secondarily. Conflicts over queen adoption might arise between queens and workers of established colonies and the newly mated females seeking adoption into nests. Colony members are predicted to base adoption decisions on their relatednesses to other participants, on competition between queens for colony resources, and on the effects that adopted queens have on colony survivorship and productivity. To provide a better understanding of queen-adoption dynamics in a facultatively polygynous ant, colonies of Myrmica tahoensis were observed in the field for 4 consecutive years and analyzed genetically using highly polymorphic microsatellite DNA markers. The extreme rarity of newly founded colonies suggests that most newly mated queens that succeed do so by entering established nests. Queens are closely related on average (rˉ = 0.58), although a sizable minority of queen pairs (29%) are not close relatives. An experiment involving transfers of queens among nests showed that queens are often accepted by workers to which they are completely unrelated. Average queen numbers estimated from nest excavations (harmonic mean = 1.4) are broadly similar to effective queen numbers inferred from the genetic relatedness of colony members, suggesting that reproductive skew is low in this species. Queens appear to have reproductive lifespans of only 1 or 2 years. As a result, queens transmit a substantial fraction of their genes posthumously (through the reproduction of related nestmates), in comparison to direct and indirect reproduction while they are alive. Thus queens and other colony members should often accept new queens when doing so will increase colony survivorship, in some cases even when the adopted queens are not close relatives. Received: 20 February 1996/Accepted after revision: 25 May 1996     

Sociogenetic organization of the red ant Myrmica rubra

Knowledge of the sociogenetic organization determining the kin structure of social insect colonies is the basis for understanding the evolution of insect sociality. Kin structure is determined by the number and relatedness of queens and males reproducing in the colonies, and partitioning of reproduction among them. This study shows extreme flexibility in these traits in the facultatively polygynous red ant Myrmica rubra. Relatedness among worker nestmates varied from 0 to 0.82. The most important reason for this variation was the extensive variation in the queen number among populations. Most populations were moderately or highly polygynous resulting in low relatedness among worker nestmates, but effectively monogynous populations were also found. Polygynous populations also often tend to be polydomous, which is another reason for low relatedness. Coexisting queens were positively related in two populations out of five and relatedness was usually similar among workers in the same colonies. Due to the polydomous colony organization and short life span of queens, it was not possible to conclusively determine the importance of unequal reproduction among coexisting queens, but it did not seem to be important in determining the relatedness among worker nestmates. The estimates of the mating frequency by queens remained ambiguous, which may be due to variation among populations. In some populations relatedness among worker nestmates was high, suggesting monogyny and single mating by queens, but in single-queen laboratory nests relatedness among the worker offspring was lower, suggesting that multiple mating was common. The data on males were sparse, but indicated sperm precedence and no relatedness among males breeding in the same colony. A comparison of social organizations and habitat requirements of M. rubra and closely related M. ruginodis suggested that habitat longevity and patchiness may be important ecological factors promoting polygyny in Myrmica. Received: 15 May 1995/Accepted after revision: 17 October 1995     

Colony fusion causes within-colony variation in a parthenogenetic ant

The evolutionary stability of cooperation and altruism in colonies of social insects requires that nestmates be to some extent related. An efficient system of discrimination against non-nestmates protects the nest against unrelated conspecifics, which might exploit or parasitize the colony. The co-occurrence of unrelated individuals in mature colonies therefore is a rare event that deserves more attention. Here, we report on the relatively common incidence of colony fusion in the ant Platythyrea punctata. Workers of this ant can produce genetically identical female offspring from unfertilized eggs through thelytokous parthenogenesis. Consequently, the majority of colonies has a “clonal structure” and consists of individuals with identical multilocus genotypes. Nevertheless, field observations indicate that a surprisingly large percentage of colonies contain workers belonging to two or more different genetic lineages. Much of this genetic heterogeneity is incompatible with eventual recombination or mutation events, but instead appears to result from colony fusion or the adoption of unrelated individuals. Indeed, colonies of P. punctata from the Dominican Republic and Barbados readily merged in the laboratory and, after elimination of one of the two reproductive workers, formed stable, genetically heterogeneous colonies. We discuss the possible causes and benefits of colony fusion in natural populations.     

Colony life history and demography of a swarm-founding social wasp

Colonies of social insects are sometimes viewed as superorganisms. The birth, reproduction, and death of colonies can be studied with demographic measures analogous to those normally applied to individuals, but two additional questions arise. First, how do adaptive colony demographies arise from individual behaviors? Second, since these superorganisms are made up of genetically distinct individuals, do conflicts within the colony sometimes modify and upset optima for colonies? The interplay between individual and superindividual or colony interests appears to be particularly complex in neotropical, swarm-founding, epiponine wasps such as Parachartergus colobopterus. In a long-term study of this species, we censused 286 nests to study colony-level reproduction and survivorship and evaluated individual-level factors by assessing genetic relatedness and queen production. Colony survivorship followed a negative exponential curve very closely, indicating type II survivorship. This pattern is defined by constant mortality across ages and is more characteristic of birds and other vertebrates than of insects. Individual colonies are long-lived, lasting an average of 347 days, with a maximum of over 4.5 years. The low and constant levels of colony mortality arise in part from colony initiation by swarming, nesting on protected substrates, and an unusual expandable nest structure. The ability to requeen rapidly was also important; relatedness data suggest that colonies requeen on average once every 9–12 months. We studied whether colony optima with respect to the timing of reproduction could be upset by individual worker interests. In this species, colonies are normally polygynous but new queens are produced only after a colony reaches the monogynous state, a result which is in accord with the genetic interests of workers. Therefore colony worker interests might drive colonies to reproduce whenever queen number happens to cycled down to one rather than at the season that is otherwise optimal. However, we found reproduction to be heavily concentrated in the rainy season. The number of new colonies peaked in this season as did the percentages of males and queens. Relatedness among workers reached a seasonal low of 0.21–0.27, reflecting the higher numbers of laying queens. This seasonality was achieved in part by a modest degree of synchrony in the queen reduction cycle. Worker relatedness reached peaks of around 0.4 in the dry season, reflecting a decrease to a harmonic mean queen number of about 2.5. Thus, a significant number of colonies must be approaching monogyny entering the rainy season. Coupled with polygynous colonies rearing only males (split sex ratios), this makes it possible for a colony cycle driven by selfish worker interests to be consistent with concentrating colony reproduction during a favorable season.     

High degree of polyandry in Apis dorsata queens detected by DNA microsatellite variability

Workers of six colonies of the giant honeybee Apis dorsata from Sabah, Malaysia (five colonies) and Java (one colony) were genotyped using single locus DNA fingerprinting. The colonies from Sabah nested in colony aggregations of 5 and 28 nests respectively on two trees. Three DNA microsatellite loci (A14, A76, A88) with a total of 27 alleles provided sufficient genetic variability to classify the workers into distinct sub-families revealing the degree of polyandry of the queens. Queens mated on average with 30.17 ± 5.98 drones with a range from 19 to 53. The average effective number of matings per queen was 25.56 ± 11.63. In the total sample of 192 workers, 22 individuals were found that were not offspring of the colony's queen. Three of these were potentially drifted offspring workers from genotyped queens of colonies nesting on the same tree.     

Kin discriminators in the eusocial sweat bee <Emphasis Type="Italic">Lasioglossum malachurum</Emphasis>: the reliability of cuticular and Dufour’s gland odours

The ability to discriminate degrees of relatedness may be expected to evolve if it allows unreciprocated altruism to be preferentially directed towards kin (Hamilton in J Theor Biol 7:1–16, 1964). We explored the possibility of kin recognition in the primitively eusocial halictid bee Lasioglossum malachurum by investigating the reliability of worker odour cues that can be perceived by workers to act as indicators of either nest membership or kinship. Cuticular and Dufour’s gland compounds varied significantly among colonies of L. malachurum, providing the potential for nestmate discrimination. A significant, though weak, negative correlation between chemical distance and genetic relatedness (r = −0.055, p < 0.001) suggests a genetic component to variation in cuticular bouquet, but odour cues were not informative enough to discriminate between different degrees of relatedness within nests. This pattern of variation was similar for Dufour’s gland bouquets. The presence of unrelated individuals within nests that are not chemically different from their nestmates suggests that the discrimination system of L. malachurum is prone to acceptance errors. Compounds produced by colony members are likely combined to generate a gestalt colony chemical signature such that all nestmates have a similar smell. The correlation between odour cues and nest membership was greater for perceived compounds than for non-perceived compounds, suggesting that variability in perceived compounds is a result of positive selection for nestmate recognition despite potentially stabilising selection to reduce variability in odour differences and thereby to reduce costs derived from excessive intracolony nepotistic behaviour.     

Queen acceptance and the complexity of nestmate discrimination in the Argentine ant

In most social insect species, individuals recognize and behave aggressively towards non-nestmate conspecifics to maintain colony integrity. However, introduced populations of the invasive Argentine ant, Linepithema humile, exhibit pronounced variation in intraspecific aggression denoting diversity in nestmate recognition behavior, which possibly shapes their social structure and the varying levels of unicoloniality observed among these populations. One approach to better understand differential aggression behaviors towards conspecifics and recognition cue perception and response in L. humile is to examine variation in nestmate discrimination capability among genetically distinct colonies under different social contexts. Consequently, we investigated the dynamics of queen and worker recognition in southeastern US L. humile queenless and queenright colonies by measuring rates of non-nestmate worker and queen adoption and intercolony genetic similarity. Aggression levels between colony pairs differed and were associated with non-nestmate worker, but not queen adoption. Adoption of queens and workers was a function of host colony origin, while colony queen number affected adoption of queens, but not workers, with queens more readily accepted by queenless hosts. Fecundity of adopted non-nestmate queens was comparable to that of rejected non-nestmate and host colony queens, suggesting that queen fecundity did not affect adoption decisions. Genetic similarity between colonies ranged from 30 to 77% alleles shared, with more genetically similar colonies showing lower levels of intraspecific aggression. Non-nestmate queens and workers that were more genetically similar to host colony workers were more likely to be adopted. We provide the first evidence for the role of L. humile colony queen number on queen discrimination and suggest an effect of resident queens on worker conspecific acceptance thresholds. Our findings indicate a role for genetically based cues in L. humile nestmate recognition. However, subtle discrimination capability seems to be influenced by the social context, as demonstrated by more frequent recognition errors in queenless colonies.     

Transmission of a pathogen in Bombus terrestris,with a note on division of labour in social insects

Summary Parasites of social insect workers can be transmitted within the colony to other, related host individuals or, alternatively, to unrelated workers of other colonies. Division of labour affects the probability of transmission, as young individuals often work inside the nest whereas older ones often leave the nest to forage. Therefore, the relative probabilities of transmission within-vs. between-nests is also affected by the delay between host infection and the shedding of propagules, i.e. the latent period of the parasite strain. We therefore hypothesized that strains of the flagellate parasite Crithidia bombi (Trypanosomatidae, Zoomastigophorea) infecting workers of the bumble bee Bombus terrestris (Hymenoptera, Apidae) could differ in their delays and coexist in a population. This would be the case if strains that are shed after a short time delay were more efficiently transmitted to other colony members, whereas strains with long delays were more efficiently transmitted to non-related workers in the population. We tested this hypothesis by experimentally varying time delay and by allowing transmission to either sister workers from the same nest or unrelated workers from other nests. Transmission of C. bombi was measured as the number of parasitic cells shed by the exposed workers after a standard period. The results showed that relatedness as such had no effect, but that delay and nest identity were highly significant effects to explain variation in transmission success. There was a significant interaction between nest identity and delay, such that bees of some colonies acted as efficient transmitters for C. bombi under short delays and vice versa. We discuss how division of labour may affect parasitism in social insects and, vice versa, how division of labour may be under selection from the effects of parasitism, using available evidence from the literature. Correspondence to: P. Schmid-Hempel     

Estimation of kinship parameters: the island model with separate sexes

Summary Information from pedigrees provides the most accurate means of estimating relatedness within and between social groups, but the method has the disadvantage of requiring longterm breeding records. If a regular system of mating occurs, then kinship parameters may be estimated from recurrence relations which decribe the change in coancestry and inbreeding from generation to generation. Breeding records collected over a single generation are therefore sufficient. In this paper the island model is used for the derivation of general expression of the coancestry and inbreeding coefficients in a dioecious population with non-overlapping generations. An equation for the average coefficient of coancestry (f ^JJ ) within a juvenile cohort is first found, and the other coefficients are then easily obtained. Parameters required are the inbreeding effective numbers of male and female parents and the migration rates of the two sexes. — In species with overlapping generations, the average coefficient of coancestry of the juvenile cohort is not related simply by migration rates to the average coancestries of the adult classes. The difference arises because average coancestry between pairs of individuals chosen from different cohorts is not in general the same as that between pairs of the same cohort. The precise difference depends on the mating system and cohort structure of the social group. Suitable modification to the basic expression for f ^JJ is described. The use of the composite method is illustrated by reference to the social units of two large mammals: prides of lion and clans of impala.     

Mean colony relatedness is a poor predictor of colony structure and female philopatry in the communally breeding Bechstein's bat (Myotis bechsteinii)

In order to understand why animals are social and how group members interact with each other it is important to know their relatedness. However, few studies have investigated the genealogy in complete social groups of free-living animals with low reproductive skew. This holds particularly true for bats. Although almost all bat species are social, their sociobiology is not well understood. Because they are volant, nocturnal and have a rather cryptic life-style, bats are difficult to observe in the wild. Furthermore females are generally gregarious making genetic parent-offspring assignment a challenging task. We used genetic markers in combination with knowledge about age and colony membership of individually marked bats to construct pedigrees in completely sampled maternity colonies of Bechstein's bats (Myotis bechsteinii). Despite considerable fluctuations in population size, no immigration occurred over 5 years in four colonies living in close proximity. Additionally, confrontation tests showed that females of one maternity colony were able to detect and attempted to prevent the intrusion of foreign females into a roost they occupy. Although colonies were absolutely closed, and 75% of the colony members lived together with close relatives (rS=0.25), mean colony relatedness was nearly zero (0.02). Average relatedness therefore is a poor estimator for the potential of kin selection in Bechstein's bat colonies and may be misleading when attempting to understand the social structure of animals living in groups where many members breed. Based on our results we discuss the potential adaptive value of living in closed societies with low reproductive skew.     

Relatedness and avoidance of inbreeding in the lizard, Tiliqua rugosa

Two potential mechanisms for reducing the level of inbreeding, sex-biased dispersal and kin avoidance, were examined in the Australian sleepy lizard, Tiliqua rugosa. The home range centres, and the genotypes at four polymorphic microsatellite DNA loci were determined for adult lizards in a 70-ha study area near Mount Mary, South Australia. From estimates of genetic relatedness, females were as closely related to other females as they were to males, both within the whole study area, and within home ranges. Similarly, males were as closely related to other males as they were to females. This suggests that dispersal in the population is not sex-biased. Sleepy lizards form monogamous pairs during the spring. Partners were less closely related to each other than to other potential partners in the home range area. This suggests active choice of unrelated partners. The mechanism for recognising related from unrelated individuals is unknown, but the behaviour could reduce inbreeding. Received: 7 November 1998 / Accepted: 30 May 1999     

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.     

Kin associations and direct vs indirect fitness benefits in colonial cooperatively breeding sociable weavers Philetairus socius

Indirect fitness benefits are believed to be an important force behind the evolution of cooperative breeding. However, helpers may associate with their relatives as a result of delayed dispersal, hence, kin associations might be a consequence of demographic viscosity rather than active choice. In addition, recent studies showed that helpers may have access to reproduction therefore direct benefits might also play an important role. Here, we investigate the possible roles of direct genetic benefits and kin associations on helping behavior in the sociable weaver Philetairus socius, a colonial and cooperatively breeding passerine. We used a microsatellite-based genotyping method to describe the genetic structure within nests and colonies. Within a colony, we found considerable genetic structure between males but not females. Sociable weaver colonies have several nests that are simultaneously active, giving individuals a choice of associating with a range of first-order kin to unrelated individuals. Helpers were significantly more related to the young in the helped nests than in other nests of the colony, suggesting an active choice for associating with kin. The helpers were generally offspring or first-order relatives of one (50%) or both (43%) breeders, although more infrequently, seemingly unrelated individuals also helped (7%). We found no supporting evidence of extrapair parentage and hence no direct genetic gains from helping in our population. This strong reproductive skew is contrary to theoretical models predicting conflicts over reproduction in stepfamilies. We discuss whether female decisions and/or other direct benefits of remaining in kin associations or helping might explain the high skew observed.