Genetic structure in a solitary rodent (Ctenomys talarum): implications for kinship and dispersal

The genetic structure of a population provides critical insights into patterns of kinship and dispersal. Although genetic evidence of kin structure has been obtained for multiple species of social vertebrates, this aspect of population biology has received considerably less attention among solitary taxa in which spatial and social relationships are unlikely to be influenced by kin selection. Nevertheless, significant kin structure may occur in solitary species, particularly if ecological or life history traits limit individual vagility. To explore relationships between genetic structure, kinship, and dispersal in a solitary vertebrate, we compared patterns of genetic variation in two demographically distinct populations of the talar tuco-tuco (Ctenomys talarum), a solitary species of subterranean rodent from Buenos Aires Province, Argentina. Based on previous field studies of C. talarum at Mar de Cobo (MC) and Necochea (NC), we predicted that natal dispersal in these populations is male biased, with dispersal distances for males and females being greater at NC. Analyses of 12 microsatellite loci revealed that in both populations, kin structure was more apparent among females than among males. Between populations, kinship and genetic substructure were more pronounced at MC. Thus, our findings were consistent with predicted patterns of dispersal for these animals. Collectively, these results indicate that populations of this solitary species are characterized by significant kin structure, suggesting that, even in the absence of sociality and kin selection, the spatial distributions and movements of individuals may significantly impact patterns of genetic diversity among conspecifics.     

From population structure to individual behaviour: genetic analysis of social structure in the European wild rabbit (Oryctolagus cuniculus)

The European wild rabbit (Oryctolagus cuniculus) lives in stable, territorial breeding groups, with male-biased natal dispersal, female philopatry, and a polygynous mating system. It was introduced into Britain in the 11 th century and kept in captive warrens as a food and fur resource. Populations expanded in the wild in the 18th century. Microsatellite markers were employed to examine the genetic structure of wild rabbit populations on three spatial scales: macrogeographic structure of 17 populations in East Anglia, microgeographic structure of a tagged population in the grounds of the University of East Anglia over four consecutive years, and pairwise kinships and individual movement within this tagged population. Populations across East Anglia were found to be genetically distinct, and heterozygote deficits were observed at all loci indicating sub-division within sampled populations. Analysis of the tagged population confirmed that rabbit populations are genetically sub-divided among social groups. Studying this population over four consecutive years revealed that as the population size increased, the number of social groups increased. Analysis of individual pairwise relatedness of females indicated that individuals did not necessarily group with kin, and behavioural data indicate that an optimum group size may exist which maximizes reproductive success.     

A molecular analysis of African lion (Panthera leo) mating structure and extra‐group paternity in Etosha National Park

The recent incorporation of molecular methods into analyses of social and mating systems has provided evidence that mating patterns often differ from those predicted by group social organization. Based on field studies and paternity analyses at a limited number of sites, African lions are predicted to exhibit a strict within‐pride mating system. Extra‐group paternity has not been previously reported in African lions; however, observations of extra‐group associations among lions inhabiting Etosha National Park in Namibia suggest deviation from the predicted within‐pride mating pattern. We analysed variation in 14 microsatellite loci in a population of 164 African lions in Etosha National Park. Genetic analysis was coupled with demographic and observational data to examine pride structure, relatedness and extra‐group paternity (EGP). EGP was found to occur in 57% of prides where paternity was analysed (n = 7), and the overall rate of EGP in this population was 41% (n = 34). Group sex ratio had a significant effect on the occurrence of EGP (P < 0.05), indicating that variation in pride‐level social structure may explain intergroup variation in EGP. Prides with a lower male‐to‐female ratio were significantly more likely to experience EGP in this population. The results of this study challenge the current models of African lion mating systems and provide evidence that social structure may not reflect breeding structure in some social mammals.     

Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and a late Pleistocene reduction in genetic diversity

Lions were the most widespread carnivores in the late Pleistocene, ranging from southern Africa to the southern USA, but little is known about the evolutionary relationships among these Pleistocene populations or the dynamics that led to their extinction. Using ancient DNA techniques, we obtained mitochondrial sequences from 52 individuals sampled across the present and former range of lions. Phylogenetic analysis revealed three distinct clusters: (i) modern lions, Panthera leo ; (ii) extinct Pleistocene cave lions, which formed a homogeneous population extending from Europe across Beringia (Siberia, Alaska and western Canada); and (iii) extinct American lions, which formed a separate population south of the Pleistocene ice sheets. The American lion appears to have become genetically isolated around 340 000 years ago, despite the apparent lack of significant barriers to gene flow with Beringian populations through much of the late Pleistocene. We found potential evidence of a severe population bottleneck in the cave lion during the previous interstadial, sometime after 48 000 years, adding to evidence from bison, mammoths, horses and brown bears that megafaunal populations underwent major genetic alterations throughout the last interstadial, potentially presaging the processes involved in the subsequent end-Pleistocene mass extinctions.     

The effect of female polyandry and sperm precedence on the evolution of sexual difference in dispersal timing

Predispersal copulation and unpredictable environment facilitate the evolution of female-biased dispersal in species, where females are functionally monandrous. Females should migrate and reproduce over different habitats to spread their risks due to environmental fluctuation. On the other hand, males do not have to disperse because their risks are spread by their mating partners who produce their offspring in different habitats. However, when females are functionally polyandrous, it is expected that they will not contribute to spreading the male's risk extensively. Therefore, by simulation with the individual based model, the present study evaluated how female polyandry influences the sexual difference in dispersal timing. This model revealed that when females are polyandrous, the timing of female remating and sperm priority patterns have an important influence on the evolution of sex-biased dispersal. Particularly when female remating is not synchronized with dispersal or when last-male sperm precedence does not exist, female-biased dispersal is evolved.     

Lion density and population structure in the Selous Game Reserve: evaluation of hunting quotas and offtake

In 1992, tourist hunting in the Selous Game Reserve generated 1.28 million dollars for the Tanzanian government, of which 0·96 million dollars were returned to wildlife conservation. Lions ( Panthera leo ) are one of three critical species for tourist hunting, consistently generating 12%–13% of hunting revenue from 1988 to 1992. Because of their ecological and economic importance (and intrinsic value), it is important that lion quotas be set so that offtake is sustainable. The population density of lions in Selous ranges from 0·08 to 0·13 adults km⁻², comparable to unhunted ecosystems. The adult sex ratio (36–41% male) and the ratio of cubs to adults (29% cubs) are similar to those of unhunted populations. The ratio of lions to hyaenas is lower in heavily hunted areas (0·17 lions/hyena) than in unhunted areas (0·43 lions/hyena). Hunting levels between 1989 and 1994 took 2·7–4·3% of adult males annually, which is sustainable. The current quota is 10–16% of the adult male population, which exceeds natural mortality rates for male lions. To remain stable if the quota was filled, the population would have to compensate via increased fecundity, increased juvenile survival, or an altered sex‐ratio. Compensation occurs in Selous by producing (or raising) more male than female cubs (66–81% of juveniles are male). Only 28% of the Selous quota was filled in 1992. The percentage of quota filled (both in Selous and nationwide) has dropped since 1988 as quotas have increased. The current intensity of lion hunting in Selous is sustainable, but the quota cannot be filled sustainably.     

Development and characterization of novel microsatellite markers in the Asiatic lion (Panthera leo persica)

All cat species, except domestic cats, are either threatened or endangered. The Asiatic lion (Panthera leo persica), a critically endangered species, is driven to a single habitat zone in Gir forests in India for its survival. The present status of this big cat compels us to acquire deeper knowledge of its genetic variability and phylogenetic relationships. Here we report the isolation of highly polymorphic microsatellite loci from a partial genomic library of the Asiatic lion. These loci were characterized and have shown high levels of variation ranging from six to 11 alleles per locus in the population studied.     

Pollen flow in the wildservice tree, Sorbus torminalis (L.) Crantz. IV. Whole interindividual variance of male fecundity estimated jointly with the dispersal kernel

Interindividual variance of male reproductive success (MRS) contributes to genetic drift, which in turn interacts with selection and migration to determine the short-term response of populations to rapid changes in their environment. Individual relative MRS can be estimated through paternity analysis and can be further dissected into fecundity and spatial components. Existing methods to achieve this decomposition either rely on the strong assumption of a random distribution of pollen donors (TwoGener) or estimate only the part of the variance of male fecundity that is explained by few covariates. We developed here a method to estimate jointly the whole variance of male fecundity and the pollen dispersal curve from the genotypic information of sampled seeds and their putative fathers and geographical information of all individuals in the study area. We modelled the relative individual fecundities as a log-normally distributed random effect. We used a Bayesian approach, well suited to the hierarchical nature of the model, to estimate these fecundities. When applied to Sorbus torminalis , the estimated variance of male fecundity corresponded to an effective density of trees 13 times lower than the observed density ( d_obs / d_ep ~  13). This value is between the value ( ~ 2) estimated with a classical mating model including three covariates (neighbourhood density, diameter, flowering intensity) that affect fecundity and the value ( ~ 30) estimated with TwoGener. The estimated dispersal kernel was close to previous results. This approach allows fine monitoring of ongoing genetic drift in natural populations, and quantitative dissection of the processes contributing to drift, including human actions.     

Sleeping site ecology in a rain-forest dwelling nocturnal lemur (Lepilemur mustelinus): implications for sociality and conservation

Suitable sleeping sites as potentially restricted resources are suggested to shape sociality in primates. We investigated sleeping site ecology of a rain-forest dwelling sportive lemur in eastern Madagascar for the first time. Using radiotelemetry, we characterized the type, quality and usage of sleeping sites as well as social sleeping habits of 11 focal individuals of the weasel sportive lemur (Lepilemur mustelinus) during the dry and the onset of the rainy season. Morphometric measurements provided additional information. The sexes showed an unusual sexual dimorphism for primates. Males and females did not differ in body length, but females surpassed males in body mass suggesting female dominance. Both sexes used dense vegetation and holes in hollow trees high above the ground as shelters for sleeping during the day. No sex difference in the quality of tree holes was found, but focal individuals used tree holes more often than open sleeping sites in dense vegetation. Both sexes showed high sleeping site fidelity limited to two to six different sites that they used primarily solitarily. The results imply that suitable sleeping sites are limited and survival of this species will strongly depend on the availability of mature rain forests with suitable hollow trees. Furthermore, these findings provide evidence of a solitary sleeping and ranging system in this rain-forest dwelling sportive lemur with suitable sleeping sites as defendable resources.     

Gene flow and immigration: genetic diversity and population structure of lions (Panthera leo) in Hwange National Park,Zimbabwe

The genetic diversity and population structure of a population of African lions in Hwange National Park, Zimbabwe, was studied using 17 microsatellite loci. Spatial genetic analysis using Bayesian methods suggested a weak genetic structure within the population and high levels of gene flow across the study area. We were able to identify a few individuals with aberrant or admixed ancestry, which we interpreted as either immigrants or as descendants thereof. This, together with relatively high genetic diversity, suggests that immigrants from beyond the study area have influenced the genetic structure within the park. We suggest that the levels of genetic diversity and the observed weak structure are indicative of the large and viable Okavango-Hwange population of which our study population is a part. Genetic patterns can also be attributed to still existing high levels of habitat connectivity between protected areas. Given expected increases in human populations and anthropogenic impacts, efforts to identify and maintain existing movement corridors between regional lion populations will be important in retaining the high genetic diversity status of this population. Our results show that understanding existing levels of genetic diversity and genetic connectivity has implications, not only for this lion population, but also for managing large wild populations of carnivores.     

Polymorphic microsatellites for the study of Aconitum napellus L. (Ranunculaceae), a rare species in France

Aconitum napellus is a perennial European Ranunculaceae that is currently decreasing in France. Six polymorphic microsatellite markers were characterized to study the polymorphism among individuals sampled from six populations in France. The total number of alleles per locus varied from two to four. Mean observed heterozygosities at each microsatellite locus ranged from 0.05 to 0.19. This set of microsatellites offers an efficient tool for more detailed investigations on population genetic structure of a rare species that can reproduce both sexually and by vegetative spread.     

Mate discrimination in Littorina littorea (L.) and L. saxatilis (Olivi) (Mollusca:Prosobranchia)

The ability of males of Littorina littorea and L. saxatilis to discriminate between mates of different sex, species and size was examined. In partner choice experiments males of L. littorea had the possibility to initiate a copulation with either a female or a male. The males did not show a preference for either sex. There was therefore no evidence that they could determine the sex of a conspecific prior to copulation. The duration of intrasexual copulation was considerably shorter than for intersexual copulation, both in the field and in laboratory experiments. For the two species, intersexual copulations were far more frequent than intrasexual ones. This can partly be explained by the difference in copulation time.Few interspecific copulating pairs were found on the shore. This may reflect a low interspecific encounter rate rather than a mechanism of species recognition. On all of these occasions, however, the active male was of L. saxatilis. It is argued that selection against precopulatory species and sex recognition is a more likely explanation than an hypothesis that states that the required mutations for precopulatory mate identification has not yet occurred. L. littorea males copulated longer with large than with small females. Copulation time was short with parasitized females, which are sterile or of low fecundity. The allocation of mating effort by males is discussed.     

Processing power limits social group size: computational evidence for the cognitive costs of sociality

Sociality is primarily a coordination problem. However, the social (or communication) complexity hypothesis suggests that the kinds of information that can be acquired and processed may limit the size and/or complexity of social groups that a species can maintain. We use an agent-based model to test the hypothesis that the complexity of information processed influences the computational demands involved. We show that successive increases in the kinds of information processed allow organisms to break through the glass ceilings that otherwise limit the size of social groups: larger groups can only be achieved at the cost of more sophisticated kinds of information processing that are disadvantageous when optimal group size is small. These results simultaneously support both the social brain and the social complexity hypotheses.     

Genetic diversity,evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa

Aim In recent decades there has been a marked decline in the numbers of African lions (Panthera leo), especially in West Africa where the species is regionally endangered. Based on the climatological history of western Africa, we hypothesize that West and Central African lions have a unique evolutionary history, which is reflected by their genetic makeup. Location Sub‐Saharan Africa and India, with special focus on West and Central Africa. Method In this study 126 samples, throughout the lion’s complete geographic range, were subjected to phylogenetic analyses. DNA sequences of a mitochondrial region, containing cytochrome b, tRNAPro, tRNAThr and the left part of the control region, were analysed. Results Bayesian, maximum likelihood and maximum parsimony analyses consistently showed a distinction between lions from West and Central Africa and lions from southern and East Africa. West and Central African lions are more closely related to Asiatic lions than to the southern and East African lions. This can be explained by a Pleistocene extinction and subsequent recolonization of West Africa from refugia in the Middle East. This is further supported by the fact that the West and Central African clade shows relatively little genetic diversity and is therefore thought to be an evolutionarily young clade. Main conclusions The taxonomic division between an African and an Asian subspecies does not fully reflect the overall genetic diversity within lions. In order to conserve genetic diversity within the species, genetically distinct lineages should be prioritized. Understanding the geographic pattern of genetic diversity is key to developing conservation strategies, both for in situ management and for breeding of captive stocks.     

Genetic evidence for female-biased dispersal in the bullfrog, Rana catesbeiana (Ranidae)

Sex-biased dispersal is an important but unexplored area of amphibian ecology. We predicted female-biased dispersal in the bullfrog (Rana catesbeiana) based on aspects of their mating system and tested this prediction using data from seven polymorphic DNA microsatellite loci. Allelic (F-statistics) and genotypic [assignment index, (AIc)] frequencies from nine Ontario populations support our prediction, although significant sex differences in inbreeding and variance of AIc were not detected. The diversity of mating systems found in amphibians represents an important avenue for investigating the relationship between reproductive systems, dispersal and phylogeny.