Population genetic analysis of a recent range expansion: mechanisms regulating the poleward range limit in the volcano barnacle Tetraclita rubescens

As range shifts coincident with climate change have become increasingly well documented, efforts to describe the causes of range boundaries have increased. Three mechanisms—genetic impoverishment, migration load, or a physical barrier to dispersal—are well described theoretically, but the data needed to distinguish among them have rarely been collected. We describe the distribution, abundance, genetic variation, and environment of Tetraclita rubescens, an intertidal barnacle that expanded its northern range limit by several hundreds of kilometres from San Francisco, CA, USA, since the 1970s. We compare geographic variation in abundance with abiotic and biotic patterns, including sea surface temperatures and the distributions of 387 co‐occurring species, and describe genetic variation in cytochrome c oxidase subunit I, mitochondrial noncoding region, and nine microsatellite loci from 27 locations between Bahia Magdalena (California Baja Sur, Mexico) and Cape Mendocino (CA, USA). We find very high gene flow, high genetic diversity, and a gradient in physical environmental variation coincident with the range limit. We infer that the primary cause of the northern range boundary in T. rubescens is migration load arising from flow of maladapted alleles into peripheral locations and that environmental change, which could have reduced selection against genotypes immigrating into the newly colonized portion of the range, is the most likely cause of the observed range expansion. Because environmental change could similarly affect all taxa in a region whose distributional limits are established by migration load, these mechanisms may be common causes of range boundaries and largely synchronous multi‐species range expansions.     

Beyond habitat requirements: individual fine-scale site fidelity in a colony of the Galapagos sea lion (<Emphasis Type="Italic">Zalophus wollebaeki</Emphasis>) creates conditions for social structuring

Site fidelity has been widely discussed, but rarely been related explicitly to a species’ social context. This is surprising, as fine-scale site fidelity constitutes an important structural component in animal societies by setting limits to an individual’s social interaction space. The study of fine-scale site fidelity is complicated by the fact that it is inextricably linked to patterns of habitat use. We here document fine-scale site fidelity in the Galapagos sea lion (Zalophus wollebaeki) striving to disentangle these two aspects of spatial behaviour. Regardless of sex and age, all individuals used small, cohesive home ranges, which were stable in size across the reproductive and non-reproductive season. Home ranges showed a large individual component and did not primarily reflect age- or sex-specific habitat requirements. Site specificity could be illustrated up to a resolution of several metres. Long-term site fidelity was indicated by home range persistence over 3 years and the degree of site fidelity was unaffected by habitat, but showed seasonal differences: it was lower between reproductive and non-reproductive periods than between reproductive seasons. We further examined static and social interaction within mother–offspring pairs, which constitute a central social unit in most mammalian societies. Regardless of the occupied habitat type, adult females with offspring had smaller home range sizes than non-breeding females, demonstrating the importance of spatial predictability for mother–offspring pairs that recurrently have to reunite after females’ foraging sojourns. While social interaction with the mother dropped to naught in both sexes after weaning, analysis of static interaction suggested female-biased home range inheritance. Dispersal decisions were apparently not based on habitat quality, but determined by the offspring’s sex. We discuss the implication of observed fine-scale site fidelity patterns on habitat use, dispersal decisions and social structure in colonial breeding pinnipeds.     

Geographic range limits of species

Understanding the forms that the geographic range limits of species take, their causes and their consequences are key issues in ecology and evolutionary biology. They are also topics on which understanding is advancing rapidly. This themed issue of Proc. R. Soc. B focuses on the wide variety of current research perspectives on the nature and determinants of the limits to geographic ranges. The contributions address important themes, including the roles and influences of dispersal limitation, species interactions and physiological limitation, the broad patterns in the structure of geographic ranges, and the fundamental question of why at some point species no longer evolve the ability to overcome the factors constraining their distributions and thus fail to continue to spread. In this introduction, these contributions are placed in the wider context of these broad themes.     

Integrating individual behaviour and landscape genetics: the population structure of timber rattlesnake hibernacula

Individuals of many species show high levels of fidelity to natal populations, often due to reliance on patchily distributed habitat features. In many of these species, the negative impacts of inbreeding are mitigated through specialized behaviours such as seasonal mating dispersal. Quantifying population structure for species with these characteristics can potentially elucidate social and environmental factors that interact to affect mating behaviour and population connectivity. In the northern part of their range, timber rattlesnakes are communal hibernators with high natal philopatry. Individuals generally recruit to the same hibernaculum as their mother and remain faithful to that hibernaculum throughout their lives. We examined the genetic structure of Crotalus horridus hibernacula in the northeastern USA using microsatellite loci. Sampled hibernacula exhibited only modest levels of differentiation, indicating a significant level of gene flow among them. We found no significant correlation between genetic differentiation and geographical distance, but did find significant positive correlation between genetic differentiation and a cost-based distance metric adjusted to include the amount of potential basking habitat between hibernacula. Therefore, thermoregulation sites may increase gene flow by increasing the potential for contact among individuals from different populations. Parentage analyses confirmed high levels of philopatry of both sexes to their maternal hibernaculum; however, approximately one-third of paternity assignments involved individuals between hibernacula, confirming that gene flow among hibernacula occurs largely through seasonal male mating dispersal. Our results underscore the importance of integrating individual-level behaviours and landscape features with studies of fine-scale population genetics in species with high fidelity to patchily distributed habitats.     

Genetic structure of African buffalo herds based on variation at the mitochondrial D-loop and autosomal microsatellite loci: Evidence for male-biased gene flow

Sexual differences in herding behaviour ofAfrican buffalo (Syncerus caffer) werestudied by analysing at the herd levelmitochondrial D-loop hypervariable region I andfourteen autosomal microsatellites. Three herdsfrom Arusha National Park in Tanzania wereanalysed with mtDNA and five herds from KrugerNational Park in South Africa with mtDNA andmicrosatellites. Significant mtDNAdifferentiation was observed among herds inArusha NP (F_ST = 0.12, based on haplotypefrequencies). Assignment tests withmicrosatellite data from Kruger NP showed thatmost frequent migration between herds is bymales two years. This was confirmed bytests for herd differentiation and analyseswith Lynch and Ritland's relatedness estimator.Within a herd, males younger than two years andfemales showed a higher relationship through acommon father rather than a common mother,indicating that female herd members mate withonly a few dominant males. This in turnsuggests a female:male sex ratio larger than5:1. The migration rate per generation betweenherds was estimated to be 5–2% for femalesand close to 100% for males. Finally, theimplications for the management of buffalopopulations are being discussed.     

Post-breeding information gathering and breeding territory shifts in northern wheatears

1. Prospecting non-breeding individuals have been shown to collect information on breeding sites a year ahead of breeding, but whether experienced breeders prospect future breeding sites is less well-known. Using data on post-breeding movements and between-year site shifts from a long-term population study of the migratory northern wheatear (Oenanthe oenanthe L.), we investigated (1) whether breeding territory selection of experienced breeders was a two-step process, made partly in the post-breeding period and partly at the time of territory establishment in the subsequent year; (2) predictions of which factors and cues correlate to site shifts at these two selection periods; and (3) consequences of territory shifts. 2. Many wheatears stayed close to their breeding site during the post-breeding period, but about 20% shifted to new potential breeding sites. Males that shifted to a new post-breeding location were also more likely to shift territory between years. 3. Factors linked to site shifts differed in the two investigated episodes of site selection. Post-breeding site shifts were linked to an environmental predictor of individual fitness, which is also related to foraging conditions (i.e. wheatears moved from tall to short field layers). Post-breeding site shifts, however, were not more frequent among young and failed breeders which may benefit most from prospecting for alternative breeding sites. Instead, in line with predictions of improving breeding conditions between-year site shifts were more frequent among young (only males) and failed breeders, whereas the link to territory field layer height disappeared. 4. Young males occupied more attractive sites and were more likely to breed successfully in year t + 1, but this improvement could also be expected from random choice. 5. Our results suggest that even though site shifts are determined partly during the post-breeding period, post-breeding movements of experienced breeders may be costly, especially when the breeding site is also a good foraging and moulting site in the post-breeding season. Under such circumstances an individual's choice of breeding site may often be determined at arrival in the subsequent year.