Contrasting genetic responses to population fragmentation in a coevolving fig and fig wasp across a mainland–island archipelago

Interacting species of pollinator–host systems, especially the obligate ones, are sensitive to habitat fragmentation, due to the nature of mutual dependence. Comparative studies of genetic structure can provide insights into how habitat fragmentation contributes to patterns of genetic divergence among populations of the interacting species. In this study, we used microsatellites to analyse genetic variation in Chinese populations of a typical mutualistic system – Ficus pumila and its obligate pollinator Wiebesia sp. 1 – in a naturally fragmented landscape. The plants and wasps showed discordant patterns of genetic variation and geographical divergence. There was no significant positive relationship in genetic diversity between the two species. Significant isolation‐by‐distance (IBD) patterns occurred across the populations of F. pumila and Wiebesia sp. 1 as whole, and IBD also occurred among island populations of the wasps, but not the plants. However, there was no significant positive relationship in genetic differentiation between them. The pollinator populations had significantly lower genetic variation in small habitat patches than in larger patches, and three island pollinator populations showed evidence of a recent bottleneck event. No effects of patch size or genetic bottlenecks were evident in the plant populations. Collectively, the results indicate that, in more fragmented habitats, the pollinators, but not the plants, have experienced reduced genetic variation. The contrasting patterns have multiple potential causes, including differences in longevity and hence number of generations experiencing fragmentation; different dispersal patterns, with the host's genes dispersed as seeds as well as a result of pollen dispersal via the pollinator; asymmetrical responses to fluctuations in partner populations; and co‐existence of a rare second pollinating wasp on some islands. These results indicate that strongly interdependent species may respond in markedly different ways to habitat fragmentation.     

Permeability of receptive fig fruits and its effects on the re‐emergence behaviour of pollinators

1. Figs and pollinating fig wasps provide a model system for studying mutualism. The permeability of the syconium changes during receptivity or between seasons, which may affect the behaviour of pollinators. Fig fruits are permeable during receptivity, and in some species, pollinators can enter and re‐emerge after oviposition/pollination. We studied the relationship between fig permeability and pollinator re‐emergence behaviour with a functional dioecious fig, Ficus hispida and the obligate pollinator Ceratosolen solmsi marchali. 2. The relationship reflects the interaction of figs and pollinators in the mutualism and also the conflicts of interests between the two partners: figs benefit from the enclosed fig fruits which have low permeability, but pollinators benefit from their re‐emergence behaviour, which requires high fig permeability. 3. The results showed that at the end of receptivity, the permeability of fig fruits lowered rapidly with changes to the ostiole structures, and re‐emergence rate was low, with more re‐emerging pollinators trapped in the ostiolar bracts. Our results also showed that in the rainy season, the length of receptivity was shorter and fig permeability was lower. The re‐emergence rates were also lower than those in the dry season. The results elucidated that figs' interests dominated in the conflicts between fig and pollinating wasp. 4. Based on a new criteria which employed the classification of pollinators found dead in the ostiolar bracts and which involved a survey of 6 monoecious and 12 dioecious fig species, we found that re‐emergence behaviour was prevalent among fig species, and was more prevalent in functional dioecious figs than monoecious ones.     

Pollinating fig wasps: genetic consequences of island recolonization

The levels of genetic diversity and gene flow may influence the long-term persistence of populations. Using microsatellite markers, we investigated genetic diversity and genetic differentiation in island (Krakatau archipelago, Indonesia) and mainland (Java and Sumatra, Indonesia) populations of Liporrhopalum tentacularis and Ceratosolen bisulcatus, the fig wasp pollinators of two dioecious Ficus (fig tree) species. Genetic diversity in Krakatau archipelago populations was similar to that found on the mainland. Population differentiation between mainland coastal sites and the Krakatau islands was weak in both wasp species, indicating that the intervening 40 km across open sea may not be a barrier for wasp gene flow (dispersal) and colonization of the islands. Surprisingly, mainland populations of the fig waSPS may be more genetically isolated than the islands, as gene flow between populations on the Javan mainland differed between the two wasp species. Contrasting growth forms and relative 'immunity' to the effects of deforestation in their host fig trees may account for these differences.     

Multiple evolutionary processes drive the patterns of genetic differentiation in a forest tree species complex

Forest trees frequently form species complexes, complicating taxonomic classification and gene pool management. This is certainly the case in Eucalyptus, and well exemplified by the Eucalyptus globulus complex. This ecologically and economically significant complex comprises four taxa (sspp. bicostata, globulus, maidenii, pseudoglobulus) that are geographically and morphologically distinct, but linked by extensive “intergrade” populations. To resolve their genetic affinities, nine microsatellites were used to genotype 1200 trees from throughout the natural range of the complex in Australia, representing 33 morphological core and intergrade populations. There was significant spatial genetic structure (F_ST = 0.10), but variation was continuous. High genetic diversity in southern ssp. maidenii indicates that this region is the center of origin. Genetic diversity decreases and population differentiation increases with distance from this area, suggesting that drift is a major evolutionary process. Many of the intergrade populations, along with other populations morphologically classified as ssp. pseudoglobulus or ssp. globulus, belong to a “cryptic genetic entity” that is genetically and geographically intermediate between core ssp. bicostata, ssp. maidenii, and ssp. globulus. Geography, rather than morphology, therefore, is the best predictor of overall genetic affinities within the complex and should be used to classify germplasm into management units for conservation and breeding purposes.     

Dating the fungus-growing termites' mutualism shows a mixture between ancient codiversification and recent symbiont dispersal across divergent hosts

The mutualistic symbiosis between fungus-growing termites and Termitomyces fungi originated in Africa and shows a moderate degree of interaction specificity. Here we estimate the age of the mutualism and test the hypothesis that the major splits have occurred simultaneously in the host and in the symbiont. We present a scenario where fungus-growing termites originated in the African rainforest just before the expansion of the savanna, about 31 Ma (19-49 Ma). Whereas rough age correspondence is observed for the four main clades of host and symbiont, the analysis reveals several recent events of host switching followed by dispersal of the symbiont throughout large areas and throughout different host genera. The most spectacular of these is a group of closely related fungi (the maximum age of which is estimated to be 2.4 Ma), shared between the divergent genera Microtermes, Ancistrotermes, Acanthotermes and Synacanthotermes (which diverged at least 16.7 Ma), and found throughout the African continent and on Madagascar. The lack of geographical differentiation of fungal symbionts shows that continuous exchange has occurred between regions and across host species.     

Analysis of microsatellite variation in the spider mite pest Tetranychus turkestani (Acari: Tetranychidae) reveals population genetic structure and raises questions about related ecological factors

The habitat fragmentation that characterizes agricultural systems made up of an array of sympatric crop and weed biotopes might be a major determinant of the population structure of plant‐feeding arthropods. Spider mites are interesting in this regard as several species are highly polyphagous, possibly generating homogenization by plant shifting, but also have the potential of forming host plant races which promotes specialization. This study analyses genetic diversity and structure in natural populations of a spider mite pest, Tetranychus turkestani, collected on both crops and weeds. Five microsatellite markers were used to genotype 283 individuals collected from 15 samples in four locations in southern France. The markers revealed considerable genetic variation, with an average heterozygosity of 0.68. Pairwise Φ_ST estimates calculated between localities showed differentiation in all comparisons. Although geographical distance appears to be a factor that influences T. turkestani population genetic structure at a regional scale, there was no clear evidence for differentiation between mites living on different host plants. A hierarchical analysis of the distribution of the genetic diversity within and between habitats showed that more than 97% of the observed genetic variation accounted for the differentiation between mites collected on the same host plant in a given locality. Significant heterozygote deficiency was found in 11 out of the 15 samples studied. Considering the biology of the mite, a Wahlund effect and inbreeding might explain such an excess of homozygosity. The data support the view that the plants in a given locality are colonized by mites that originate from diverse sources. They also support previous data suggesting that the demographic structure is made up of small demes of inbred individuals. The agricultural‐level implications of the data are discussed, notably the fact that mites may well be capable of moving between host plants, making weeds surrounding the crop fields potential reservoirs for the pest. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 69–78.     

How a haemosporidian parasite of bats gets around: the genetic structure of a parasite,vector and host compared

Parasite population structure is often thought to be largely shaped by that of its host. In the case of a parasite with a complex life cycle, two host species, each with their own patterns of demography and migration, spread the parasite. However, the population structure of the parasite is predicted to resemble only that of the most vagile host species. In this study, we tested this prediction in the context of a vector‐transmitted parasite. We sampled the haemosporidian parasite Polychromophilus melanipherus across its European range, together with its bat fly vector Nycteribia schmidlii and its host, the bent‐winged bat Miniopterus schreibersii. Based on microsatellite analyses, the wingless vector, and not the bat host, was identified as the least structured population and should therefore be considered the most vagile host. Genetic distance matrices were compared for all three species based on a mitochondrial DNA fragment. Both host and vector populations followed an isolation‐by‐distance pattern across the Mediterranean, but not the parasite. Mantel tests found no correlation between the parasite and either the host or vector populations. We therefore found no support for our hypothesis; the parasite population structure matched neither vector nor host. Instead, we propose a model where the parasite's gene flow is represented by the added effects of host and vector dispersal patterns.     

Gene dispersal inference across forest patches in an endangered medicinal tree: comparison of model‐based approaches

Propagule dispersal in plants is a fundamental mechanism for colonizing new sites and adapting to changing climates, as well as for maintaining genetic diversity. Contrasting past and current gene dispersal can provide useful insights to gauge the extent of recent human disturbances and guide management strategies. However, research on gene dispersal of plants is not yet exhaustive because evolutionary or environmental impacts are often species‐specific and most existing studies have focused on analysis of dispersal at a single site, which may not be helpful for landscape‐level inferences and management interventions. In the present study, we assessed whether current gene or propagule dispersal would be more restricted than past gene dispersal at multiple patches of the endangered medicinal tree, Prunus africana. We employed eight highly polymorphic microsatellite markers in conjunction with isolation‐by‐distance, spatial genetic structure (SGS), and parentage assignment models to estimate gene dispersal distance in a spatial extent of approximately 400 km². There was no significant difference between gene dispersal distances across the different models (Friedman chi‐squared = 7.286, d.f. = 5, P = 0.2002). Estimates of current gene dispersal distance were comparable to dispersal in the last few generations. However, gene dispersal distance was much shorter in smaller than bigger forest patches. Further, significant (P < 0.05) SGS was detected in most forest patches, with the extent of SGS among adults being stronger in the smaller than bigger patches. These results suggest the need for practicing enrichment plantings in most forest patches, particularly in the smaller ones, to assist gene exchange among individuals and patches. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 887–904.     

Major histocompatibility complex variation in insular populations of the Egyptian vulture: inferences about the roles of genetic drift and selection

Insular populations have attracted the attention of evolutionary biologists because of their morphological and ecological peculiarities with respect to their mainland counterparts. Founder effects and genetic drift are known to distribute neutral genetic variability in these demes. However, elucidating whether these evolutionary forces have also shaped adaptive variation is crucial to evaluate the real impact of reduced genetic variation in small populations. Genes of the major histocompatibility complex (MHC) are classical examples of evolutionarily relevant loci because of their well-known role in pathogen confrontation and clearance. In this study, we aim to disentangle the partial roles of genetic drift and natural selection in the spatial distribution of MHC variation in insular populations. To this end, we integrate the study of neutral (22 microsatellites and one mtDNA locus) and MHC class II variation in one mainland (Iberia) and two insular populations (Fuerteventura and Menorca) of the endangered Egyptian vulture (Neophron percnopterus). Overall, the distribution of the frequencies of individual MHC alleles (n=17 alleles from two class II B loci) does not significantly depart from neutral expectations, which indicates a prominent role for genetic drift over selection. However, our results point towards an interesting co-evolution of gene duplicates that maintains different pairs of divergent alleles in strong linkage disequilibrium on islands. We hypothesize that the co-evolution of genes may counteract the loss of genetic diversity in insular demes, maximize antigen recognition capabilities when gene diversity is reduced, and promote the co-segregation of the most efficient allele combinations to cope with local pathogen communities.     

Climate structures genetic variation across a species' elevation range: a test of range limits hypotheses

Gene flow may influence the formation of species range limits, and yet little is known about the patterns of gene flow with respect to environmental gradients or proximity to range limits. With rapid environmental change, it is especially important to understand patterns of gene flow to inform conservation efforts. Here we investigate the species range of the selfing, annual plant, Mimulus laciniatus, in the California Sierra Nevada. We assessed genetic variation, gene flow, and population abundance across the entire elevation‐based climate range. Contrary to expectations, within‐population plant density increased towards both climate limits. Mean genetic diversity of edge populations was equivalent to central populations; however, all edge populations exhibited less genetic diversity than neighbouring interior populations. Genetic differentiation was fairly consistent and moderate among all populations, and no directional signals of contemporary gene flow were detected between central and peripheral elevations. Elevation‐driven gene flow (isolation by environment), but not isolation by distance, was found across the species range. These findings were the same towards high‐ and low‐elevation range limits and were inconsistent with two common centre‐edge hypotheses invoked for the formation of species range limits: (i) decreasing habitat quality and population size; (ii) swamping gene flow from large, central populations. This pattern demonstrates that climate, but not centre‐edge dynamics, is an important range‐wide factor structuring M. laciniatus populations. To our knowledge, this is the first empirical study to relate environmental patterns of gene flow to range limits hypotheses. Similar investigations across a wide variety of taxa and life histories are needed.     

High gene flow on a continental scale in the polyandrous Kentish plover Charadrius alexandrinus

Gene flow promotes genetic homogeneity of species in time and space. Gene flow can be modulated by sex‐biased dispersal that links population genetics to mating systems. We investigated the phylogeography of the widely distributed Kentish plover Charadrius alexandrinus. This small shorebird has a large breeding range spanning from Western Europe to Japan and exhibits an unusually flexible mating system with high female breeding dispersal. We analysed genetic structure and gene flow using a 427‐bp fragment of the mitochondrial (mtDNA) control region, 21 autosomal microsatellite markers and a Z microsatellite marker in 397 unrelated individuals from 21 locations. We found no structure or isolation‐by‐distance over the continental range. However, island populations had low genetic diversity and were moderately differentiated from mainland locations. Genetic differentiation based on autosomal markers was positively correlated with distance between mainland and each island. Comparisons of uniparentally and biparentally inherited markers were consistent with female‐biased gene flow. Maternally inherited mtDNA was less structured, whereas the Z‐chromosomal marker was more structured than autosomal microsatellites. Adult males were more related than females within genetic clusters. Taken together, our results suggest a prominent role for polyandrous females in maintaining genetic homogeneity across large geographic distances.     

Fragmentation reduces regional‐scale spatial genetic structure in a wind‐pollinated tree because genetic barriers are removed

Gene flow strongly influences the regional genetic structuring of plant populations. Seed and pollen dispersal patterns can respond differently to the increased isolation resulting from habitat fragmentation, with unpredictable consequences for gene flow and population structuring. In a recently fragmented landscape we compared the pre‐ and post‐fragmentation genetic structure of populations of a tree species where pollen and seed dispersal respond differentially to forest fragmentation generated by flooding. Castanopsis sclerophylla is wind‐pollinated, with seeds that are dispersed by gravity and rodents. Using microsatellites, we found no significant difference in genetic diversity between pre‐ and post‐fragmentation cohorts. Significant genetic structure was observed in pre‐fragmentation cohorts, due to an unknown genetic barrier that had isolated one small population. Among post‐fragmentation cohorts this genetic barrier had disappeared and genetic structure was significantly weakened. The strengths of genetic structuring were at a similar level in both cohorts, suggesting that overall gene flow of C. sclerophylla has been unchanged by fragmentation at the regional scale. Fragmentation has blocked seed dispersal among habitats, but this appears to have been compensated for by enhanced pollen dispersal, as indicated by the disappearance of a genetic barrier, probably as a result of increased wind speeds and easier pollen movement over water. Extensive pollen flow can counteract some negative effects of fragmentation and assist the long‐term persistence of small remnant populations.     

Spatial scale affects landscape genetic analysis of a wetland grasshopper

Most landscape genetic studies assess the impact of landscape elements on species' dispersal and gene flow. Many of these studies perform their analysis on all possible population pairs in a study area and do not explicitly consider the effects of spatial scale and population network topology on their results. Here, we examined the effects of spatial scale and population network topology on the outcome of a landscape genetic analysis. Additionally, we tested whether the relevant spatial scale of landscape genetic analysis could be defined by population network topology or by isolation‐by‐distance (IBD) patterns. A data set of the wetland grasshopper Stethophyma grossum, collected in a fragmented agricultural landscape, was used to analyse population network topology, IBD patterns and dispersal habitats, using least‐cost transect analysis. Landscape genetic analyses neglecting spatial scale and population network topology resulted in models with low fits, with which a most likely dispersal habitat could not be identified. In contrast, analyses considering spatial scale and population network topology resulted in high model fits by restricting landscape genetic analysis to smaller scales (0–3 km) and neighbouring populations, as represented by a Gabriel graph. These models also successfully identified a likely dispersal habitat of S. grossum. The above results suggest that spatial scale and potentially population network topology should be more explicitly considered in future landscape genetic analyses.     

Prominent intraspecific genetic divergence within Anopheles gambiae sibling species triggered by habitat discontinuities across a riverine landscape

The Anopheles gambiae complex of mosquitoes includes malaria vectors at different stages of speciation, whose study enables a better understanding of how adaptation to divergent environmental conditions leads to evolution of reproductive isolation. We investigated the population genetic structure of closely related sympatric taxa that have recently been proposed as separate species (An. coluzzii and An. gambiae), sampled from diverse habitats along the Gambia river in West Africa. We characterized putatively neutral microsatellite loci as well as chromosomal inversion polymorphisms known to be associated with ecological adaptation. The results revealed strong ecologically associated population subdivisions within both species. Microsatellite loci on chromosome‐3L revealed clear differentiation between coastal and inland populations, which in An. coluzzii is reinforced by a unusual inversion polymorphism pattern, supporting the hypothesis of genetic divergence driven by adaptation to the coastal habitat. A strong reduction of gene flow was observed between An. gambiae populations west and east of an extensively rice‐cultivated region apparently colonized exclusively by An. coluzzii. Notably, this ‘intraspecific’ differentiation is higher than that observed between the two species and involves also the centromeric region of chromosome‐X which has previously been considered a marker of speciation within this complex, possibly suggesting that the two populations may be at an advanced stage of differentiation triggered by human‐made habitat fragmentation. These results confirm ongoing ecological speciation within these most important Afro‐tropical malaria vectors and raise new questions on the possible effect of this process in malaria transmission.     

Population genetic data suggest a role for mosquito‐mediated dispersal of West Nile virus across the western United States

After introduction, West Nile virus (WNV) spread rapidly across the western United States between the years 2001 and 2004. This westward movement is thought to have been mediated by random dispersive movements of resident birds. Little attention has been placed on the role of mosquito vectors in virus dispersal across North America. The mosquito vector largely responsible for WNV amplification and transmission of WNV in the western USA is Culex tarsalis. Here we present population genetic data that suggest a potential role for C. tarsalis in the dispersal of WNV across the western USA. Population genetic structure across the species range of C. tarsalis in the USA was characterized in 16 states using 12 microsatellite loci. structure and geneland analyses indicated the presence of three broad population clusters. Barriers to gene flow were resolved near the Sonoran desert in southern Arizona and between the eastern Rocky Mountains and High Plains plateau. Small genetic distances among populations within clusters indicated that gene flow was not obstructed over large portions of the West Coast and within the Great Plains region. Overall, gene flow in C. tarsalis appears to be extensive, potentially mediated by movement of mosquitoes among neighbouring populations and hindered in geographically limited parts of its range. The pattern of genetic clustering in C. tarsalis is congruent with the pattern of invasion of WNV across the western United States, raising the possibility that movement of this important vector may be involved in viral dispersal.     

Population genetic structure and its implications for adaptive variation in memory and the hippocampus on a continental scale in food-caching black-capped chickadees

Food-caching birds rely on stored food to survive the winter, and spatial memory has been shown to be critical in successful cache recovery. Both spatial memory and the hippocampus, an area of the brain involved in spatial memory, exhibit significant geographic variation linked to climate-based environmental harshness and the potential reliance on food caches for survival. Such geographic variation has been suggested to have a heritable basis associated with differential selection. Here, we ask whether population genetic differentiation and potential isolation among multiple populations of food-caching black-capped chickadees is associated with differences in memory and hippocampal morphology by exploring population genetic structure within and among groups of populations that are divergent to different degrees in hippocampal morphology. Using mitochondrial DNA and 583 AFLP loci, we found that population divergence in hippocampal morphology is not significantly associated with neutral genetic divergence or geographic distance, but instead is significantly associated with differences in winter climate. These results are consistent with variation in a history of natural selection on memory and hippocampal morphology that creates and maintains differences in these traits regardless of population genetic structure and likely associated gene flow.     

Pollen flow in fragmented landscapes maintains genetic diversity following stand-replacing disturbance in a neotropical pioneer tree,Vochysia ferruginea Mart

In forests with gap disturbance regimes, pioneer tree regeneration is typically abundant following stand-replacing disturbances, whether natural or anthropogenic. Differences in pioneer tree density linked to disturbance regime can influence pollinator behaviour and impact on mating patterns and genetic diversity of pioneer populations. Such mating pattern shifts can manifest as higher selfing rates and lower pollen diversity in old growth forest populations. In secondary forest, where more closely related pollen donors occur, an increase in biparental inbreeding is a potential problem. Here, we investigate the consequences of secondary forest colonisation on the mating patterns and genetic diversity of open-pollinated progeny arrays for the long-lived, self-compatible pioneer tree, Vochysia ferruginea, at two Costa Rican sites. Five microsatellite loci were screened across adult and seed cohorts from old growth forest with lower density, secondary forest with higher density, and isolated individual trees in pasture. Progeny from both old growth and secondary forest contexts were predominantly outcrossed (t_m=1.00) and experienced low levels of biparental inbreeding (t_m−t_s=0.00–0.04). In contrast to predictions, our results indicated that the mating patterns of V. ferruginea are relatively robust to density differences between old growth and secondary forest stands. In addition, we observed that pollen-mediated gene flow possibly maintained the genetic diversity of open-pollinated progeny arrays in stands of secondary forest adults. As part of a natural resource management strategy, we suggest that primary forest remnants should be prioritised for conservation to promote restoration of genetic diversity during forest regeneration.     

Gene flow in the green mirid,Creontiades dilutus (Hemiptera: Miridae), across arid and agricultural environments with different host plant species

Creontiades dilutus (Stål), the green mirid, is a polyphagous herbivorous insect endemic to Australia. Although common in the arid interior of Australia and found on several native host plants that are spatially and temporally ephemeral, green mirids also reach pest levels on several crops in eastern Australia. These host‐associated dynamics, distributed across a large geographic area, raise questions as to whether (1) seasonal fluctuations in population size result in genetic bottlenecks and drift, (2) arid and agricultural populations are genetically isolated, and (3) the use of different host plants results in genetic differentiation. We sequenced a mitochondrial COI fragment from individuals collected over 24 years and screened microsatellite variation from 32 populations across two seasons. The predominance of a single COI haplotype and negative Tajima D in samples from 2006/2007 fit with a population expansion model. In the older collections (1983 and 1993), a different haplotype is most prevalent, consistent with successive population contractions and expansions. Microsatellite data indicates recent migration between inland sites and coastal crops and admixture in several populations. Altogether, the data suggest that long‐distance dispersal occurs between arid and agricultural regions, and this, together with fluctuations in population size, leads to temporally dynamic patterns of genetic differentiation. Host‐associated differentiation is evident between mirids sampled from plants in the genus Cullen (Fabaceae), the primary host, and alternative host plant species growing nearby in arid regions. Our results highlight the importance of jointly assessing natural and agricultural environments in understanding the ecology of pest insects.     

Clinal colour variation within a panmictic population of tree squirrels,Tamiasciurus douglasii (Rodentia: Sciuridae), across an ecological gradient

Local adaptation occurs when a population in a heterogeneous environment experiences divergent ecological selection but only if selection is stronger than the homogenizing effects of gene flow. The forest environments of Oregon vary along a physical and biotic gradient from a wet, closed‐canopy forest near the coast to a drier open‐canopy forest eastward across the Cascade Mountains. The present study explores patterns of local adaptation in Douglas squirrels (Tamiasciurus douglasii) in relation to these transitions in forest structure and ecology. We test for the presence of morphological clines in relation to gene flow and, more specifically, whether any such character clines correspond with environmental clines. We sampled animals at six locations (10 specimens each) and evaluated environmental parameters across a 240‐km west‐to‐east transect. Population structure analysis of 18 microsatellite loci indicates a single, panmictic squirrel population across the entire transect. Coalescent‐based estimates show bidirectional gene flow at similar west–east intensities between squirrels in coastal and interior forests. Of the four skull traits examined, none shows a significant clinal transition. By contrast, ventral fur colour shows a strong clinal transition, from deep‐orange in coastal forest to whitish–yellow in the interior forest. This pattern of phenotypic divergence coincides with the gradient in tree‐canopy cover. Ventral fur colour of T. douglasii exemplifies a gradation of continuous phenotypic variation maintained despite ongoing gene flow in a panmictic population. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 536–546.     

Population genetic connectivity of an endemic New Zealand passerine after large‐scale local extirpations: a model of re‐colonization potential

Little is known about how a 70% loss of native forests has affected the genetic connectivity of remnant bird populations in New Zealand. We use the common and widely distributed New Zealand Bellbird Anthornis melanura as an indicator species of population connectivity for well‐flighted birds. Using eight microsatellite loci, we identified five main genetic populations in the North Island, South Island, sub‐Antarctic Auckland Islands and two small remnant island populations adjacent to a large region of avian extirpations in northern North Island. Only one remnant island population, on a 30‐year‐old conservation reserve at Tiritiri Matangi, displayed a clear signature of recent genetic bottleneck. The 7% migration rate at Tiritiri Matangi indicates that bottlenecks can be maintained despite habitat rehabilitation, possibly through behavioural barriers to gene flow. Adjacent to the same extirpation zone, Bellbirds on the Poor Knights Islands were found to have low genetic diversity and low re‐colonization potential. Two gaps concordant with deforestation patterns separated the Kapiti Coast of southern North Island from populations to both the north and the south. In summary, we identified linked avian habitats, as well as isolated and inbred populations and suggest that Bellbirds are good re‐colonizers. We emphasize the importance of genetic studies that assess animal dispersal among newly rehabilitated habitat patches.