A multi‐basin approach determines variability in host fish suitability for unionids in tributaries of the Laurentian Great Lakes

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Umbonal musculature and relationships of the Late Triassic filibranch unionoid bivalves

Exceptionally well‐preserved Late Triassic unionoids from Silesia, Poland, show prominently ornamented juvenile shells and umbonal muscle attachments that are similar to Margaritifera, which are anatomically the least derived among extant unionoids. Their phosphatized (originally chitinous and impregnated with calcium phosphate) gill supports lacked transverse connections, and occasionally some of them were separated from others, being thus at the filibranch grade, like their trigonioid ancestors. Several separate small foot elevator attachments in these unionoids indicate Trigonodidae adaptation to marine or brackish water, in which the original trigonioid strong single attachment was already split into two in the Early Triassic. The ribbing of juvenile shells suggests a change to deeper infaunal life for juvenile stages, and generally less efficient near‐surface locomotion, with a wedge‐like foot, in adults. Much later the unionoids became eulamellibranchial, which promoted the brooding of the fish that their larvae parasitize. To accomodate the classification of the order under this scenario of evolutionary changes, a new suborder Silesunionina is proposed for its filibranch members. It includes the Silesunionidae fam. nov. , with the location of umbonal muscles similar to that in the extant underived unionoids, and the Unionellidae fam. nov. , with umbonal muscles attached to the external wall of the umbonal cavity. The early Late Triassic (Carnian) Silesunio parvus gen. et sp. nov. and latest Triassic (Rhaetian) Tihkia(?) silesiaca sp. nov. are proposed. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 863–883.     

Impact of a mouth parasite in a marine fish differs between geographical areas

Considerable variation exists in parasite virulence and host tolerance which may have a genetic and/or environmental basis. In this article, we study the effects of a striking, mouth‐dwelling, blood‐feeding isopod parasite (Ceratothoa italica) on the life history and physiological condition of two Mediterranean populations of the coastal fish, Lithognathus mormyrus. The growth and hepatosomatic index (HSI) of fish in a heavily human‐exploited population were severely impacted by this parasite, whereas C. italica showed negligible virulence in fish close to a marine protected area. In particular, for HSI, the parasite load explained 34.4% of the variation in HSI in the exploited population, whereas there was no significant relationship (0.3%) between parasite load and HSI for fish in the marine protected area. Both host and parasite populations were not differentiated for neutral genetic variation and were likely to exchange migrants. We discuss the role of local genetic adaptation and phenotypic plasticity, and how deteriorated environmental conditions with significant fishing pressure can exacerbate the effects of parasitism. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 842–852.     

Is life history a barrier to dispersal? Contrasting patterns of genetic differentiation along an oceanographically complex coast

Extreme variation in early life‐history strategies is considered a moderately good predictor of genetic subdivision and hence dispersal for a range of marine species. In reality, however, a good deal of population differentiation must reflect historical effects, more subtle variation in life histories, and, particularly, the interaction of larvae with oceanographic processes. Using a combination of allozyme and microsatellite markers, we show that the large‐scale genetic structure of populations of three species (direct and planktonically developing cushion stars and a planktonic developing sea anemone that is also asexually viviparous) varies consistently, in line with the predicted capacity for dispersal within three geographic regions. We detected high levels of genetic subdivision for the direct developing cushion star (F_ST = 0.6), low levels for the planktonically developing cushion star (F_ST = 0.009), and intermediate levels for the sexual/asexual sea anmone (F_ST = 0.19). These patterns are exhibited despite the highly variable patterns of current movement and the presence of biogeographic barriers. Our results suggest that, although there is large scale genetic differentiation for two species, patterns of population connectivity are remarkably consistent within major regions and do not reflect variation in major oceanographic processes or genetic discontinuity coincident with biogeographic boundaries. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 106–116.     

Comparative population genetic structures of the fruit fly Urophora cardui and its primary parasitoid Eurytoma robusta

The interaction between two species may depend on geographic scale and this in turn can affect co‐evolution among them. The present study comparatively examines population genetic structures of the tephritid gall fly Urophora cardui and its primary ectoparasitoid Eurytoma robusta for inference of relative dispersal patterns and host‐parasitoid specificity. Genetic differentiation patterns indicated two levels of hierarchical structure in both species: locally similar distance‐dependencies but globally differences. Locally, both species showed isolation by distance and a high correlation between host and parasitoid F_ST for the same population‐pairs was found. At the local level, E. robusta populations were most structured. These findings suggest that locally E. robusta is tracking behind its host, U. cardui, and that colonisation of new patches by both species underlie a stepping‐stone dispersal process. The investigation as a whole showed that U. cardui populations were hierarchically structured across a genetic‐geographical cline. There was no sign of a comparable cline in E. robusta where populations globally became independent of one another and of the host. The different degree of hierarchical genetic structure of the two species suggests that dispersal processes or interactions differ relative to geographical scale and population history.     

Isolation and high genetic diversity in dwarf mountain toads (Capensibufo) from South Africa

Traditional models of amphibian dispersal and gene flow point to low dispersal and high philopatry. In recent years, this traditional view has been challenged and it appears that no general model holds across taxa. Conservation of amphibians cannot be addressed on an over‐arching scale, but must come on a case‐by‐case basis, especially for range‐restricted species where information on gene flow and migration must be incorporated into conservation efforts. The only two members of the genus Capensibufo Grandison, 1980 (Anura: Bufonidae) are range restricted small bufonids, with distributions limited to montane areas in South Africa. Using a Bayesian analysis of two mitochondrial markers (16S and ND2), we examined the genetic patterns in Capensibufo rosei and Capensibufo tradouwi in order to understand both taxonomic and geographic boundaries. These species were not monophyletic, and demonstrate no clear taxonomic boundaries. Instead, the genus is extremely diverse genetically, with distinct lineages confined to isolated mountains that represent geographic boundaries. In addition, bioclimatic modelling using MAXENT and scenarios of climatic conditions at both the present and last glacial maximum suggest multiple bioclimatic and physical barriers to gene flow at present and in the past. We conclude that members of the genus have very low vagility, that current taxonomic boundaries are inadequate, and that strong geographic structuring has undoubtedly contributed to genetic diversity at the species level, rather than the population level. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 822–834.     

Historical demography of freshwater mussels (Bivalvia: Unionidae): genetic evidence for population expansion and contraction during the late Pleistocene and Holocene

Genetic variation was examined in two endangered mussel species, Epioblasma brevidens and Epioblasma capsaeformis, and in a non‐listed species, Lampsilis fasciola, in the Clinch River, Tennessee, USA, by screening mitochondrial DNA (mtDNA) sequences and nuclear DNA microsatellites. Patterns of mtDNA polymorphism exhibited different trends in long‐term population sizes for each species during the late Pleistocene and Holocene (～20 000 ya to present); namely, E. brevidens has declined over time, E. capsaeformis has remained demographically stable, and L. fasciola has expanded. However, analyses using microsatellites did not exhibit similar trends, perhaps because homoplasy had eliminated long‐term population signatures for the loci examined. For both marker types, long‐term effective population size (N_e) was low in E. brevidens, intermediate in E. capsaeformis, and high in L. fasciola. Moderately diverged mtDNA lineages, perhaps indicative of secondary contact, were observed in E. brevidens and E. capsaeformis. Perhaps the most surprising result of this study was the high level of genetic variation observed at both mtDNA and microsatellite DNA markers for L. fasciola, variation seemingly contrary to the relatively small demes that currently reside in the Clinch River. However, the data are consistent with known demographic and life‐history traits of these three mussel species and their fish hosts, namely that they each use hosts with different dispersal capabilities, ranging from low, moderate, and high, respectively. The low divergence of mtDNA sequence variation reported in this and other recent mussel studies indicates that considerable extant population genetic variation probably originated during the late Pleistocene and Holocene. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 376–397.     

Population structure and dispersal in a patchy landscape: nuclear and mitochondrial markers reveal area effects in the spider Theridion californicum (Araneae: Theridiidae)

Changes in land use have a major effect on patterns of biodiversity. However, few studies have examined the demographic and genetic shifts associated with a return to semi‐natural habitat following extended periods of human disturbance. Here we examine patterns of population structure in a spider restricted to the Pacific coastal strip of North America that exhibits an exuberant colour polymorphism. We use mitochondrial DNA and AFLP markers to examine genetic structure and estimate gene flow. The results show contrasting, gender‐specific patterns between these markers that suggest limited dispersal, combined with area effects most likely caused by expansion from refugial habitat patches following land‐management changes in a region of the San Francisco East Bay. Colour‐morph frequencies are not correlated with this complex genetic structure. Thus, unlike the classical area effects that were based on colour morphs, we demonstrate in T. californicum signals of historical contingency at neutral loci but not at the Colour locus, where traces of past events have been obliterated by balancing selection. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 600–620.     

Fine‐scale patterns of genetic divergence within and between morphologically variable subspecies of the sea urchin Heliocidaris erythrogramma (Echinometridae)

The spatial scale over which genetic divergences occur between populations and the extent that they are paralleled by morphological differences can vary greatly among marine species. In the present study, we use a hierarchical spatial design to investigate genetic structure in Heliocidaris erythrogramma occurring on near shore limestone reefs in Western Australia. These reefs are inhabited by two distinct subspecies: the thick‐spined Heliocidaris erythrogramma armigera and the thin‐spined Heliocidaris erythrogramma erythrogramma, each of which also have distinct colour patterns. In addition to pronounced morphological variation, H. erythrogramma exhibits a relatively short (3–4 days) planktonic phase before settlement and metamorphosis, which limits their capacity for dispersal. We used microsatellite markers to determine whether patterns of genetic structure were influenced more by morphological or life history limitations to dispersal. Both individual and population‐level analyses found significant genetic differentiation between subspecies, which was independent of geographical distance. Genetic diversity was considerably lower within H. e. erythrogramma than within H. e. armigera and genetic divergence was four‐fold greater between subspecies than among populations within subspecies. This pattern was consistent even at fine spatial scales (< 5 km). We did detect some evidence of gene flow between the subspecies; however, it appears to be highly restricted. Within subspecies, genetic structure was more clearly driven by dispersal capacity, although weak patterns of isolation‐by‐distance suggest that there may be other factors limiting gene exchange between populations. Our results show that spatial patterns of genetic structure in Western Australian H. erythrogramma is influenced by a range of factors but is primarily correlated with the distribution of morphologically distinct subspecies. This suggests the presence of reproductive barriers to gene exchange between them and demonstrates that morphological variation can be a good predictor of genetic divergence. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 578–592.     

Extensive local‐scale gene flow and long‐term population stability in the intertidal mollusc Katharina tunicata (Mollusca: Polyplacophora)

The dispersal capabilities of intertidal organisms may represent a key factor to their survival in the face of global warming, as species that cannot adapt to the various effects of climate change will have to migrate to track suitable habitat. Although species with pelagic larval phases might be expected to have a greater capacity for dispersal than those with benthic larvae, interspecies comparisons have shown that this is not always the case. Consequently, population genetic approaches are being increasingly used to gain insights into dispersal through studying patterns of gene flow. In the present study, we used nuclear single‐nucleotide polymorphisms (SNPs) and mitochondrial DNA (mtDNA) sequencing to elucidate fine‐scale patterns of genetic variation between populations of the Black Katy Chiton, Katharina tunicata, separated by 15–150 km in south‐west Vancouver Island. Both the nuclear and mitochondrial data sets revealed no genetic differentiation between the populations studied, and an isolation‐with‐migration analysis indicated extensive local‐scale gene flow, suggesting an absence of barriers to dispersal. Population demographic analysis also revealed long‐term population stability through previous periods of climate change associated with the Pleistocene glaciations. Together, the findings of the present study suggest that this high potential for dispersal may allow K. tunicata to respond to current global warming by tracking suitable habitat, consistent with its long‐term demographic stability through previous changes in the Earth's climate. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 589–597.     

Patterns of gene flow and source‐sink dynamics in high altitude populations of the common toad Bufo bufo (Anura: Bufonidae)

Understanding patterns of genetic structure is fundamental for developing successful management programmes for deme‐structured organisms, such as amphibians. We used five microsatellite loci and DNA sequences of the mitochondrial control region to assess the relative influences of landscape (geographic distance, altitude and rivers as corridors for dispersal) and historical factors on patterns of gene flow in populations of the toad Bufo bufo in Central Spain. We sampled 175 individuals from eight populations distributed along two major river drainages and used maximum‐likelihood and Bayesian approaches to infer patterns of gene flow and population structure. The mitochondrial DNA data show closely‐related haplotypes distributed across the Iberian Peninsula with no geographic structuring, suggesting recent differentiation of haplotypes and extensive gene flow between populations. On the other hand, microsatellites provide finer resolution, showing that high altitude populations (> 2000 m) exchange lower numbers of migrants with other populations. The results of Bayesian estimates for recent migration rates in high altitude populations suggest source‐sink dynamics between ponds that are consistent with independent data from monitoring over the past 20 years. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 824–839.     

The radiation of Satyrini butterflies (Nymphalidae: Satyrinae): a challenge for phylogenetic methods

We have inferred the most comprehensive phylogenetic hypothesis to date of butterflies in the tribe Satyrini. In order to obtain a hypothesis of relationships, we used maximum parsimony and model‐based methods with 4435 bp of DNA sequences from mitochondrial and nuclear genes for 179 taxa (130 genera and eight out‐groups). We estimated dates of origin and diversification for major clades, and performed a biogeographic analysis using a dispersal–vicariance framework, in order to infer a scenario of the biogeographical history of the group. We found long‐branch taxa that affected the accuracy of all three methods. Moreover, different methods produced incongruent phylogenies. We found that Satyrini appeared around 42 Mya in either the Neotropical or the Eastern Palaearctic, Oriental, and/or Indo‐Australian regions, and underwent a quick radiation between 32 and 24 Mya, during which time most of its component subtribes originated. Several factors might have been important for the diversification of Satyrini: the ability to feed on grasses; early habitat shift into open, non‐forest habitats; and geographic bridges, which permitted dispersal over marine barriers, enabling the geographic expansions of ancestors to new environments that provided opportunities for geographic differentiation, and diversification.

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Seed dispersal and population structure in Vriesea gigantea,a bromeliad from the Brazilian Atlantic Rainforest

Seed dispersal, population structure and the mating system of plant species can have great consequences on the genetic structure of populations. Vriesea gigantea is a bromeliad from southeastern Brazil which is self‐compatible and pollinator dependent for fruit set. Its populations are fertile in terms of the production of flowers, pollen, fruits and seeds. To assess the importance of seed supply for gene flow, colonization and distribution of adult individuals, the seed dispersal and population structure of V. gigantea were studied. Seeds are dispersed over short distances; most seeds land close to the mother plant. This pattern coincides with the reported aggregate distribution of bromeliad seedlings. Population structure results showed high seedling recruitment, because 51.3% developed into adults, although few juveniles reached this stage. This result is different from that for other bromeliad species from different habitat conditions. Seed dispersal and population structural patterns are consistent with previous molecular studies, revealing that V. gigantea populations are genetically structured, with low gene flow and a moderate outcrossing rate. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 317–325.     

The interplay of dispersal limitation,rivers, and historical events shapes the genetic structure of an Amazonian frog

Disentangling the impact of landscape features such as rivers and historical events on dispersal is a challenging but necessary task to gain a comprehensive picture of the evolution of diverse biota such as that found in Amazonia. Adenomera andreae, a small, territorial, terrestrial frog species of the Amazonian forest represents a good model for such studies. We combined cytochrome b sequences with 12 microsatellites to investigate the genetic structure at two contrasted spatial scales in French Guiana: along a ～6‐km transect, to evaluate dispersal ability, and between paired bank populations along a ～65‐km stretch of the Approuague river, to test the effect of rivers as barriers to dispersal. We observed significant spatial genetic structure between individuals at a remarkably small geographical scale, and conclude that the species has a restricted dispersal ability that is probably tied to its life‐history traits. Mitochondrial and microsatellite data also indicate a high level of differentiation among populations on opposite banks of the river, and, in some cases, among populations on the same riverbank. These results suggest that the observed population structure in A. andreae is the result of restricted dispersal abilities combined with the action of rivers and Quaternary population isolation. Given that Amazonia hosts a great portion of anurans, as well as other small vertebrates, that display life‐history traits comparable with A. andreae, we argue that our analyses provide new insights into the complex interactions among evolutionary processes shaping Amazonian biodiversity. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 356–373.     

Positive effects of flower abundance and synchronous flowering on pollination success,and pollinia dispersal in rewardless Changnienia amoena (Orchidaceae)

Pollination success and pollen dispersal in natural populations depend on the spatial‐temporal variation of flower abundance. For plants that lack rewards for pollinators, pollination success is predicted to be negatively related to flower density and flowering synchrony. We investigated the relationships between pollination success and flower abundance and flowering synchrony, and estimated pollinia dispersal distance in a rewardless species, Changnienia amoena (Orchidaceae). The results obtained in the present study revealed that male pollination success was negatively influenced by population size but was positively affected by population density, whereas female pollination success was independent of both population size and density. Phenotypic analysis suggested that highly synchronous flowering was advantageous through total pollination success, which is in contrast to previous studies. These results indicate that pollination facilitation rather than competition for pollinator visits occurs in this rewardless plant. The median distance of pollinia dispersal was 11.5 m (mean distance = 17.5 m), which is comparable to that of other rewardless plants but longer than for rewarding plants. However, pollen transfer occured mainly within populations; pollen import was a rare event. Restricted gene flow by pollinia and seeds probably explains the previous population genetic reporting a high degree of genetic differentiation between populations. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 477–488.     

Effects of dams and their environmental impacts on the genetic diversity and connectivity of freshwater mussel populations in Poyang Lake Basin,China

1. Habitat fragmentation is one of the main threats to biodiversity. Barriers to dispersal caused by anthropogenic habitat alteration may affect phylogeographic patterns in freshwater mussels. Knowledge of the phylogenetic and phylogeographic patterns of unionoids is vital to inform protection of their biodiversity.
2. Here, we assessed influences of dams and their environmental effects on the genetic diversity and population connectivity of a broadly distributed freshwater mussel, Nodularia douglasiae, in Poyang Lake Basin.
3. The results showed high genetic diversity in areas without dams and low genetic diversity in areas with dams. High genetic differentiation and low gene flow were found among the 11 populations. Genetic variation was significantly correlated with dissolved oxygen levels.
4. The observation of low genetic diversity in populations separated by dams indicated that those populations were subjected to genetic erosion and demographic decline because they are disconnected from other populations with higher diversity. High genetic differentiation and low gene flow among the 11 populations could be correlated with anthropogenic habitat alteration.
5. These results indicated that anthropogenic habitat alterations have led to the decline in freshwater mussel diversity. Therefore, we recommend maintaining favourable habitat conditions and connectivity of rivers or lakes, and strengthening study of life histories with host-test experiments to identify potential host fish species to strengthen the knowledge base underpinning freshwater mussel conservation.

     

Population structure in the catfish Trichogenes longipinnis: drift offset by asymmetrical migration in a tiny geographic range

Based on population genetic theory and empirical studies of small populations, we expect that species with very small ranges (narrow endemics) will exhibit reduced genetic diversity, increasing their susceptibility to the negative effects of genetic homogeneity. Although this pattern of reduced diversity applies to most narrow endemics, conservation biologists have yet to identify a general pattern for the degree of spatial population genetic structure expected in species with very small ranges. In part, this is because the degree of population structure within narrow endemics will be highly variable depending on the equilibrium between the homogenizing effects of dispersal and the diversifying effects of drift and local selection in small populations, thus precluding general predictions about the relative importance of small range, small population sizes, and habitat patchiness for maintaining genetic diversity in narrowly-distributed species. We document a striking example of high population structure in the tiny geographic range of a stream-dwelling catfish, Trichogenes longipinnis , endemic to the Atlantic Forest of Brazil. The maintenance of this diversity results from a combination of asymmetrical and limited dispersal, and drift in small populations. Our results highlight the need to understand population structure, and not only overall genetic diversity, of narrowly-distributed species for their conservation planning.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 259–274.     

Host habitat patchiness and the distance decay of similarity among gastro-intestinal nematode communities in two species of <Emphasis Type="Italic">Mastomys</Emphasis> (southeastern Senegal)

Beta-diversity, or how species composition changes with geographical distance, has seldom been studied for different habitats. We present here quantitative estimates of the relationship between geographic distance and similarity of parasitic nematode communities in two closely related rodent host species that live in habitats with very different spatial configurations. In southeastern Senegal Mastomys natalensis lives exclusively inside human villages whereas M. erythroleucus is continuously distributed outside villages. Both host species and their gastro-intestinal nematodes were sampled on the same spatial scale. Beta-diversity was found to be higher in parasite communities of M. erythroleucus than in those of M. natalensis, and significantly related to geographic distance in this first species. Even on the local spatial scale studied, host dispersal limitation, and stochastic events, may affect species turnover in nematode communities of M. erythroleucus. In M. natalensis, no relationship was found between geographic distance and nematode community similarity, however, suggesting low host dispersal rates between habitat patches. Together with previous population genetic results, this study illustrates the need for different approaches with regard to dispersal in natural populations and its effect on biodiversity.     

Comparative population structure of three snook species (Teleostei: Centropomidae) from the eastern central Pacific

Three snook species, Centropomus viridis, Centropomus medius, and Centropomus robalito, from the eastern central Pacific, representing three of the four proposed phyletic lineages in the genus, were analyzed for genetic variability by means of allozyme and RAPD to evaluate the divergence between populations at different levels of dispersal ability and to evaluate the importance of barriers to dispersal in the population subdivision and genetic diversity. Levels of genetic diversity among species estimated by allozymes were similar and consistent with the observed levels of differentiation in marine fish species. Mean heterozygosity ranged from 0.089 for C. viridis to 0.10 for C. robalito. Genetic diversity for the snook species studied was slightly higher than the mean estimation reported in allozymes for 106 marine fish (0.055) and for anadromous fish species (0.043 to 0.057). Multilocus allele frequency homogeneity tests and population-subdivision estimates for both allozyme and RAPD markers revealed the existence of population structure in C. viridis and C. medius, in coincidence with geographic separation of samples, whereas no divergence was detected in C. robalito. This finding may be attributed to the greater population size of C. robalito, which originated by a recent population range expansion, and hence the potential for dispersal is mediated by larval drift. Fluctuations in population size and population range expansion are used to explain discrepancies between levels of genetic diversity and population structure in the studied species. Supplementary material to this paper is available in electronic format at     

Genetic drift drives evolution in the bird‐pollinated,terrestrial island endemic Grevillea georgeana (Proteaceae)

Plant species distributed across terrestrial islands can show significant genetic divergence among populations if seed and pollen dispersal are restricted. We assessed the genetic connectivity between populations of Grevillea georgeana, restricted to seven disjunct inselbergs in semi‐arid Western Australia. The phylogeographical pattern and population genetics of populations were determined using sequence data from two plastid DNA intergenic spacers and ten nuclear microsatellite loci. The plastid DNA markers indicated high genetic differentiation among the majority of populations. Haplotypes were restricted to individual inselbergs, with the exception of two that were shared among three isolated populations that formed part of an elongated greenstone belt and that may be connected via inaccessible populations of G. georgeana. There was also strong differentiation within some of the populations, suggesting long‐term isolation and persistence of G. georgeana on these terrestrial islands. Overall, the genetic patterns suggest limited seed dispersal, with differentiation in the plastid DNA genome being driven by genetic drift. In contrast, pollen movement, although generally restricted, may occur between neighbouring populations, resulting in a pattern of isolation by distance in the nuclear markers. This potential for limited or no seed dispersal, but connectivity via pollen flow, should be considered, given that many of the inselbergs are under consideration for resource development. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 155–168.