Microgeographic Scale Research Articles

Larval vertical migration and hierarchical selectivity of settlement in a brooding marine sponge

Knowledge of larval behaviours of sessile marine invertebrates from release to recruitment and of the role these behaviours play in determining adult distributions is limited. In manipulative experiments using larvae from the Great Barrier Reef sponge Rhopaloeides odorabile, we quantified larval behaviours associated with vertical migration, phototaxis and swimming ability. We also measured settlement responses to cues associated with light, settlement surface micro-topography, coral rubble and biofilms. Following an afternoon release, the majority of larvae (72%) migrated vertically to the surface (light) for 6 to 18 h. After 24 h, 55% of active larvae had moved from the surface to the bottom and maintained this position for up to 54 h before settling. Larvae did not display gregarious settlement patterns, or a preference for settlement surface topographies, but did preferentially settle to light-exposed surfaces. Initial settlement to biofilms or coral rubble was higher than in controls with no cue. However, the transition from initial settlement and attachment to metamorphosis was much higher when treatments comprised a combination of biofilm and coral rubble compared to biofilm-only treatments (49 vs. 9%). Overall, this demonstrates that hierarchical cues contribute to selective settlement. Vertical migration to surface waters facilitates passive dispersal via wind-driven surface currents and contributes to wide-scale dispersal, while a subsequent demersal phase, where larvae actively explore the benthos for settlement sites, enables dispersal over fine, micro-geographic spatial scales.

Scale-independent criteria and scale-dependent agents determining the structure of a ground cricket mosaic hybrid zone (Allonemobius socius - Allonemobius fasciatus)

The structure of the hybrid zone between the ground crickets, Allonemobius socius and Allonemobius fasciatus, approximates an environmental mosaic based on temperature/moisture at regional spatial scales. In the present study, we show that the micro-geographic spatial structure (i.e. single fields) of this hybrid zone is governed by the same criteria. Thus, the criteria that structure this hybrid zone are scale independent, even though the agents that implement these criteria may differ at various scales (climate, latitude, and elevation at regional scales; grass height, slope aspect, and field use at micro-geographic scales). Additionally, the study demonstrates a previously unknown barrier to genetic exchange in this system that acts before conspecific sperm precedence. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94, 777–796.

Song dialects do not restrict gene flow in an urban population of the orange-tufted sunbird, Nectarinia osea

Geographic variation in vocalizations is widespread in passerine birds, but its origins and maintenance remain unclear. In this study, we test the hypothesis that song dialect, a culturally transmitted trait, is related to the population genetic structure of the orange-tufted sunbird, Nectarinia osea. To address this, we compared mitochondrial DNA (mtDNA) sequence variation together with allele frequencies at five microsatellite loci from an urban population of sunbirds exhibiting two distinct song dialects on a microgeographic scale. Our findings reveal no association between dialect membership and genetic composition. All genetic measures, from both mitochondrial and nuclear DNA, indicate high levels of gene flow between both dialect populations. The low F ST values obtained from mtDNA and microsatellite analysis imply that the variation among dialects does not account for more than 2%, at best, of the overall genetic variation found in the entire population. These measures fall well within the range of similar measures obtained in other studies of species exhibiting vocal dialects, most of which fail to detect any dialect-based genetic differentiation. The persistence of dialects in the orange-tufted sunbird may thus best be explained by dispersal of individuals across dialect boundaries and possibly from surrounding areas, followed by postdispersal vocal matching. Because genetic structuring appears weaker than cultural structure in this species, we discuss the behavioral mechanisms underlying dialect maintenance in the presence of apparent gene flow.

Small-scale spatial genetic structure in an ant species with sex-biased dispersal

In a population of the monogynous, polyandrous ant Cataglyphis cursor, we analysed the spatial genetic structure of queens, colony fathers and workers at a microgeographical scale to infer the extent of sex-biased dispersal and to assess the impact of limited dispersal on the patterns of relatedness within the colony. To this end, four microsatellite markers were scored for the queen and an average of 26 workers from each of 35 mapped colonies. We used pair-wise kinship coefficients between all pairs of genotypes, including the reconstructed colony father genotypes (1) to test and quantify isolation by distance patterns within each sex or caste through the analysis of kinship–distance curves, and (2) to compute the average relatedness between categories of colony members. The kinship–distance curve was much steeper for colony queens than colony fathers, indicating male-biased dispersal. However, colony fathers also displayed a non-random spatial genetic structure, so that even males show some dispersal limitation at the scale of the population, which extends over less than 250 m. The degree of relatedness between the different sexes and castes of colonies was well predicted from the number of mates per queen and the inbreeding of queens, and the impact of limited dispersal was very weak at this scale of observation. We discuss the interest of kinship–distance curves to assess sex-biased dispersal on a local scale and we compare our results with large-scale analyses of genetic structure in Cataglyphis cursor and other monogynous ant species.

Population structure of <i>Magnaporthe oryzae</i> isolates from green foxtail in Japan examined by DNA fingerprint analysis

The population structure of Magnaporthe oryzae from green foxtail (Setaria viridis) in Japan was examined by DNA fingerprint analyses using the transposable elements MGR586 and MAGGY as probes. Fifteen M. oryzae isolates from green foxtail were collected from 11 Japanese prefectures so that a macrogeographic population of this pathogen is represented. All the 15 isolates were sorted into distinct haplotypes by DNA fingerprint analyses with both probes. Furthermore, similarities between the DNA fingerprint profiles of the 15 isolates were exclusively low; i.e., if lineages are arbitrarily established based on greater than 70% similarities in isolates, the 15 isolates could be categorized into 13 distinct lineages by DNA fingerprinting with both probes. We also examined the MGR586 DNA fingerprint variations of this pathogen in 9 microgeographic populations each of which contained 20 to 24 isolates collected from a 1 m2 or 50 m2 area. In all the 9 populations, more than 2 haplotypes, which shared less than 70% similarities, were identified in the DNA fingerprint profiles. These results suggested that M. oryzae isolates from the green foxtail in Japan possessed a complex lineage structure, even at the microgeographic scale.

Spatially and temporally fluctuating selection at non-MHC immune genes: evidence from TAP polymorphism in populations of brown trout (Salmo trutta, L.)

Temporal samples of Danish brown trout (Salmo trutta) from populations representing varying geographical scales were analysed using eight putatively neutral microsatellite loci and two microsatellite loci embedded in TAP genes (Transporter associated with Antigen Processing). These genes encode molecules that are central to the major histocompatibility complex (MHC) class I restricted antigen presentation and thus integral components in the adaptive immune system. As such, they could be influenced by selection, driven by pathogens and parasites in a manner similar to MHC genes. Analysis of allele frequencies at presumably neutral microsatellite loci revealed a temporally unstable population structure within regions, while the population structure was stable over time among regions. Analyses of the two TAP markers indicated an effect of selection at both a regional and micro-geographical spatial scale. Moreover, signals of divergent selection among temporal samples within localities suggest that selection also might fluctuate at a temporal scale. These results suggest that immune genes other than the classical MHC class I and II might be subject to selection and warrant further studies of functional polymorphism of such genes in natural populations.

Risky prey behavior evolves in risky habitats

Longstanding theory in behavioral ecology predicts that prey should evolve decreased foraging rates under high predation threat. However, an alternative perspective suggests that growth into a size refuge from gape-limited predation and the future benefits of large size can outweigh the initial survival costs of intense foraging. Here, I evaluate the relative contributions of selection from a gape-limited predator (Ambystoma opacum) and spatial location to explanations of variation in foraging, growth, and survival in 10 populations of salamander larvae (Ambystoma maculatum). Salamander larvae from populations naturally exposed to intense A. opacum predation risk foraged more actively under common garden conditions. Higher foraging rates were associated with low survival in populations exposed to free-ranging A. opacum larvae. Results demonstrate that risky foraging activity can evolve in high predation-risk habitats when the dominant predators are gape-limited. This finding invites the further exploration of diverse patterns of prey foraging behavior that depends on natural variation in predator size-selectivity. In particular, prey should adopt riskier behaviors under predation threat than expected under existing risk allocation models if foraging effort directly reduces the duration of risk by growth into a size refuge. Moreover, evidence from this study suggests that foraging has evolved over microgeographic scales despite substantial modification by regional gene flow. This interaction between local selection and spatial location suggests a joint role for adaptation and maladaptation in shaping species interactions across natural landscapes, which is a finding with implications for dynamics at the population, community, and metacommunity levels.

Age-adjusted Plasmodium falciparum antibody levels in school-aged children are a stable marker of microgeographical variations in exposure to Plasmodium infection.

BackgroundAmongst school-aged children living in malaria endemic areas, chronic morbidity and exacerbation of morbidity associated with other infections are often not coincident with the presence or levels of Plasmodium parasitaemia, but may result from long-term exposure to the parasite. Studies of hepatosplenomegaly associated with Schistosoma mansoni infection and exposure to Plasmodium infection indicate that differences that occur over 1–2 km in levels of Plasmodium transmission are related to the degree of exacerbation of hepatosplenomegaly and that Plasmodium falciparum schizont antigen (Pfs)-IgG3 levels may be a marker for the differing levels of exposure.MethodsTo investigate the validity of Pfs-IgG3 measurements as a tool to assess these comparative exposure levels on a microgeographical scale, cross-sectional community surveys were conducted over a 10 × 6 km study site in Makueni District, Kenya, during low and high malaria transmission seasons. During both high and low malaria transmission seasons, thick blood smears were examined microscopically and circulating Pfs-IgG3 levels measured from dried blood spot elute. GIS techniques were used to map prevalence of parasitaemia and Pfs-IgG3 levels.ResultsMicrogeographical variations in prevalence of parasitaemia were observed during the high but not the low transmission season. Pfs-IgG3 levels were stable between high and low transmission seasons, but increased with age throughout childhood before reaching a plateau in adults. Adjusting Pfs-IgG3 levels of school-aged children for age prior to mapping resulted in spatial patterns that reflected the microgeographical variations observed for high season prevalence of parasitaemia, however, Pfs-IgG3 levels of adults did not. The distances over which age-adjusted Pfs-IgG3 of school-aged children fluctuated were comparable with those distances over which chronic morbidity has previous been shown to vary.ConclusionAge-adjusted Pfs-IgG3 levels of school-aged children are stable and when mapped can provide a tool sensitive enough to detect microgeographical variations in malaria exposure, that would be useful for studying the aetiology of morbidities associated with long-term exposure and co-infections.

Active dispersal by wild Triatoma infestans in the Bolivian Andes

Triatoma infestans is the main vector of Chagas disease and target of control programmes in the Southern Cone countries. So far Bolivia is the only country where true T. infestans wild foci are documented. The dispersal ability for wild T. infestans was studied at microgeographical scale in Bolivian Andes, to assess the possibility for wild populations to actively recolonize insecticide-treated villages. Nine microsatellite loci were used to detect the extent of gene flow between neighbouring collecting sites. The detection of restricted gene flow between close but distinct sylvatic sites supports the hypothesis that wild T. infestans does not disperse by flying at high altitude (2,750 m asl). It gradually disperses over small distances by walking within a 'patch' of continuous land cover. The genetic differentiation detected between sylvatic and domestic populations suggests a limited short-term role of wild insects in the process of recolonization of insecticide-treated houses in the Andes.

Fine-scale genetic structure overrides macro-scale structure in a marine snail: nonrandom recruitment, demographic events or selection?

The planktotrophic littorinid species Littoraria flava occupies a continuous habitat on rocky shores close to brackish and freshwater sources. Previous studies of this species have shown a moderate genetic structure over a broad geographical scale, with high deviations from Hardy–Weinberg expectations in many allozymic loci. Local-scale subdivision in marine species with a long dispersal phase is unexpected, but occasionally found. Using a horizontal transect at three locations, we examined whether microscale and short-term subdivision also occurred in L. flava populations and, if so, whether this could explain the Hardy–Weinberg deviations. Littoraria flava showed even more structuring on a microgeographical scale (4–300 m) than on a large-scale (> 200 km). The Ewens–Watterson neutrality test showed that 18% of the tests deviated significantly from the neutrality model. A homogeneity test for each locus across samples within transects showed homogeneous and high FIS values in many loci. These results and the apparent genetic patchiness within transects suggest that asynchronous spawning associated with recurrent colonizations in L. flava can explain the local differentiation without a recognizable pattern. In addition, there could be a balance between these factors and diversifying selection acting on different loci at different times and localities.

REPRODUCTIVE ISOLATION BETWEENPOGONOMYRMEX RUGOSUSAND TWO LINEAGES WITH GENETIC CASTE DETERMINATION

Hybrid speciation occurs when combination of two interspecific genomes results in individuals that are of high fitness but reproductively incompatible with the parental species. Although hybrid speciation is a relatively common source of new species in plants, it appears to be a much rarer occurrence in animal taxa. Here we report on reproductive isolation and range overlap between the rough harvester ant Pogonomyrmex rugosus and two lineages with hybrid genotypes (H 1 and H2). Both lineages obligately interbreed and produce genetically distinct queen and worker offspring, a phenomenon referred to as genetic caste determination (GCD). Diploid offspring produced by gametes of the same lineage develop only into queens, whereas diploid offspring derived from gametes of distinct lineages develop into workers. We investigated small-scale patterns of gene flow between the parent and the two H lineages by sampling along an 80-km transect between a pure P. rugosus population and a two-lineage population. Microsatellite and mitochondrial markers both indicated virtually no gene flow between the parent species and either lineage even at sites where parental and H-lineage colonies co-occurred. The geographic ranges of the parental species and the two-lineage population were essentially parapatric, with a surprisingly narrow band of overlap and evidence of spatial structuring even at microgeographic scales within the transition zone. This suggests that ecological competition with the parent species plays a significant role in determining the evolutionary persistence and current distribution of the hybrid lineages and the genetic caste system.

Genetic population structure across a range of geographic scales in the commercially exploited marine gastropod Buccinum undatum

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 317:157-169 (2006) - doi:10.3354/meps317157 Genetic population structure across a range of geographic scales in the commercially exploited marine gastropod Buccinum undatum D. Weetman1,*, L. Hauser1,2, M. K. Bayes3, J. R. Ellis4, P. W. Shaw3 1Department of Biological Sciences, University of Hull, Hull HU6 7RX, UK 2School of Aquatic and Fishery Sciences, University of Washington, 1122 NE Boat Street, Box 355020, Seattle, Washington 98195-5020, USA 3School of Biological Sciences, Royal Holloway, University of London, Egham, Surrey TW20 0EX, UK 4Centre for Environmental Fisheries and Aquaculture Sciences, (CEFAS) Lowestoft Laboratory, Lowestoft, Suffolk NR33 0HT, UK *Email: dweetman@hotmail.co.uk ABSTRACT: Marine invertebrates are exploited increasingly as food resources worldwide, but knowledge of genetic population structure is limited for most commercially valuable taxa. We investigated genetic structure in the widely distributed, subtidal gastropod Buccinum undatum by screening samples from 28 locations at 5 microsatellite loci. Multidimensional scaling and cluster analysis were applied to study macrogeographic structure, and revealed 4 well-supported groupings of populations: Canada, Iceland, Swedish Skaggerak, and a cluster containing most samples from the European continental shelf (hereafter ‘Shelf’). A fifth cluster comprising all 5 samples from the Solent (UK) relied on exceptional differentiation of just 2 alleles, which is consistent with significant evidence for recent bottlenecks in this area. Within the Shelf cluster most pairwise tests of differentiation were significant, although the global FST of 0.014 was very low for a direct-developing species. Our data suggest that use of highly polymorphic markers caused a relatively minor downward bias to FST, although historical connectivity of populations that are not in migration-drift equilibrium might be more important. However, significant isolation by distance among British North Sea coast samples (FST = 0.010) is consistent with approach to equilibrium and suggests recent gene flow, probably between semi-continuous populations. At a microgeographic scale, we found migration to be consistently higher from inshore to offshore within 3 separate areas; a factor that may underpin the lower diversity and greater differentiation observed for bay and inlet populations. Such populations might serve as important sources of genetic diversity, but are likely to be particularly vulnerable to exploitation. KEY WORDS: Fishery genetics · Whelk · Microsatellites · Isolation by distance · Asymmetric migration · Bottleneck Full text in pdf format PreviousNextExport citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 317. Online publication date: July 18, 2006 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2006 Inter-Research.

Running to stand still: adaptation and the response of plants to rapid climate change.

Climate is a potent selective force in natural populations, yet the importance of adaptation in the response of plant species to past climate change has been questioned. As many species are unlikely to migrate fast enough to track the rapidly changing climate of the future, adaptation must play an increasingly important role in their response. In this paper we review recent work that has documented climate-related genetic diversity within populations or on the microgeographical scale. We then describe studies that have looked at the potential evolutionary responses of plant populations to future climate change. We argue that in fragmented landscapes, rapid climate change has the potential to overwhelm the capacity for adaptation in many plant populations and dramatically alter their genetic composition. The consequences are likely to include unpredictable changes in the presence and abundance of species within communities and a reduction in their ability to resist and recover from further environmental perturbations, such as pest and disease outbreaks and extreme climatic events. Overall, a range-wide increase in extinction risk is likely to result. We call for further research into understanding the causes and consequences of the maintenance and loss of climate-related genetic diversity within populations.

The utility of the 16S gene in investigating cryptic speciation within the Brachionus plicatilis species complex

Recent reports indicate an extensive amount of molecular evolution separating cryptic taxa as well as significant population structure at a microgeographical scale. Appropriate molecular markers are particularly suitable for distinguishing cryptic biological species. In this study, we examine the phylogenetic utility of 16S rRNA in elucidating the evolutionary relationships within the recently described euryhaline Brachionus plicatilis species complex. In addition, we assess the applicability of this marker in the genetic identification and monitoring of rotifer populations. We have sequenced a 378-bp fragment of the mitochondrial 16S rRNA gene in laboratory reference strains, hatchery clones as well as collections from a wild population of the subsaline Lake Koroneia (Northern Greece). Also, restriction fragment length polymorphism (RFLP) analysis was performed with eight restriction endonucleases. Rotifer samples are distinguished into six genetically divergent lineages. Average sequence divergence between lineages is 0.1038. The evolutionary relationships and divergence time-scales revealed with the 16S sequence data are in agreement with previous analyses using different mitochondrial and nuclear markers. The 16S region appears to have several advantages over other regions of the genome regarding use of species-specific primers, ease of amplification from single specimens and undiluted informational content over both recent and more ancient separations. It has also exhibited maximum discriminatory power (100% success) between lineages during RFLP analysis. The 16S assayed region has proven especially informative and consistent in detecting, supporting and establishing the lineage status within the B. plicatilis species complex both from a phylogenetic perspective and as an identification tool.

Amplified fragment length polymorphism among Rhynchosporium secalis isolates collected from a single barley field in Syria

SummaryAFLP markers were used to measure the amount and distribution of genetic variation among Rhynchosporium secalis isolates on a microgeographical scale in Syria. Forty isolates hierarchically sampled from a single barley field were assayed for AFLP variation using primer combinations not previously tested in populations of the pathogen from Syria. In contrast to a previous study, which showed high clonality within field populations of R. secalis in Syria, the present study revealed a much higher level of genetic diversity, stressing the important roles that sampling strategies and the choice of primers/primer combinations play in the evaluation of genetic variation in R. secalis populations at a microgeographical scale. A high level of genetic variation was found to occur on a fine scale throughout the pathogen population examined, with 40 different haplotypes being identified among the 40 isolates sampled. Data were consistent with the hypothesis that the primary inoculum originated from a genetically diverse founding population, which may have consisted of ascospores of an as yet undescribed teleomorph and/or asexual spores of a highly mutable local population.

Cytotype segregation on regional and microgeographic scales in snow buttercups (Ranunculus adoneus: Ranunculaceae)

Polyploid speciation is an important source of angiosperm diversity. Insights into the origin and establishment of new polyploid species may be gained by studying the distributions of ancestral and derivative cytotypes at multiple spatial scales. Diploid (2n = 16) and tetraploid (2n = 32) snow buttercups (Ranunculus adoneus: Ranunculaceae) occur in the alpine of the central and southern Rocky Mountains. Root-tip squashes and flow cytometry were used to determine the ploidy of 1618 individuals from 35 populations. Samples from 31 of the 35 sites were entirely of one cytotype, either diploid or tetraploid. Diploid and tetraploid snow buttercups have nonoverlapping regional distributions. Where both cytotypes occur on the same site, the two are spatially segregated despite no apparent change in habitat. Triploid snow buttercups were only found at a diploid/tetraploid contact zone, while two hexaploid plants were found in tetraploid populations. Tetraploid establishment once or twice in the history of the species complex could account for the regional distribution of the two cytotypes. Habitat differentiation between cytotypes or reproductive exclusion of minority cytotypes may explain the observed segregation at both microgeographic and regional scales.

Landscape genetics of alpine-snowbed plants: comparisons along geographic and snowmelt gradients.

The genetic structure of three snowbed-herb species (Peucedanum multivittatum, Veronica stelleri, and Gentiana nipponica) was analyzed using allozymes across nine populations arranged as a matrix of three snowmelt gradients x three geographic locations within 3 km in the Taisetsu Mountains, northern Japan. Phenologically asynchronous populations are packed within a local area in alpine snowbeds, because flowering season of alpine plants depends strongly on the timing of snowmelt. Moderate genetic differentiation was detected among local populations in every species (FST=0.03-0.07). There was a significant correlation between the geographic distance and genetic distance in the P. multivittatum populations, but not in the V. stelleri and G. nipponica populations. On the other hand, a significant correlation between the phenological distance caused by snowmelt timing and genetic distance was detected in the V. stelleri and G. nipponica populations, but not in the P. multivittatum populations. The snowmelt gradient or geographic separation influenced hierarchical genetic structure of these species moderately (FRT <0.04). Restriction of gene flow due to phenological separation and possible differential selection along the snowmelt gradient may produce genetic clines at microgeographic scale in these species.

Microgeographic heterogeneity in spatial distribution and mtDNA variability of gray mouse lemurs (Microcebus murinus, Primates: Cheirogaleidae)

The objective of our study was to investigate the spatial distribution and genetic structure of a solitary primate at the microgeographical scale of adjacent local populations. We obtained spatial data and tissue samples for mtDNA analysis from 205 gray mouse lemurs (Microcebus murinus) captured along transects and within 3 grid systems within a 12.3 km2 area in Kirindy Forest, western Madagascar. Our capture data revealed that, even though the forest was continuous, gray mouse lemurs were not evenly distributed, and that daily and maximum dispersal distances were significantly greater in males. The frequency distribution of 22 mtDNA D-loop haplotypes was highly skewed. Nine haplotypes were unique to males, indicating male-mediated gene flow from surrounding areas. The geographic distribution of haplotypes revealed that males were also more dispersed than females. Females with the same haplotype showed a tendency towards spatial aggregation, and the correlation between genetic and geographic distances was higher in females. In several areas of the forest, however, spatially clustered females were not of the same haplotype, and females were not always found in clusters. Hence, in contrast to suggestions from previous studies, matrilineal clustering is not the only way females are socially organized. In addition, our study revealed heterogeneity and patterns in population structure that were not evident at smaller spatial scales, some of which may be relevant for designing conservation strategies.

MICROGEOGRAPHIC COUNTERGRADIENT VARIATION IN THE WOOD FROG, RANA SYLVATICA

Prior studies have shown that macrogeographic gradients in temperature associated with latitude and altitude can lead to countergradient patterns of variation in a number of taxa: individuals from colder environments are known to grow or develop faster than their conspecifics from warmer environments when placed in a common setting. In this study, I hypothesized that countergradient variation also is important at microgeographic scales. The wood frog, Rana sylvatica, breeds in open-canopied, temporary wetlands as well as those heavily shaded by vegetation. Shading leads to cooler thermal environments that are associated with embryonic development rates as much as 50% slower than those in unshaded wetlands. Wetlands with contrasting canopy environments are often found within tens or hundreds of meters of each other. In a common garden experiment, embryos from nearby natural wetlands displayed countergradient variation: individuals collected from shaded wetlands developed up to 12% faster than those collected from relatively unshaded wetlands. The results of this study suggest that the concept of countergradient variation may be extended to small scales of space. In addition, the rate and scale of vegetation dynamics (the agent of wetland shading) imply that divergence in development among residents of nearby wetlands may be relatively rapid, on the order of decades.

Mating system and gene flow in the red seaweed Gracilaria gracilis: effect of haploid-diploid life history and intertidal rocky shore landscape on fine-scale genetic structure.

The impact of haploid-diploidy and the intertidal landscape on a fine-scale genetic structure was explored in a red seaweed Gracilaria gracilis. The pattern of genetic structure was compared in haploid and diploid stages at a microgeographic scale (< 5 km): a total of 280 haploid and 296 diploid individuals located in six discrete, scattered rock pools were genotyped using seven microsatellite loci. Contrary to the theoretical expectation of predominantly endogamous mating systems in haploid-diploid organisms, G. gracilis showed a clearly allogamous mating system. Although within-population allele frequencies were similar between haploids and diploids, genetic differentiation among haploids was more than twice that of diploids, suggesting that there may be a lag between migration and (local) breeding due to the long generation times in G. gracilis. Weak, but significant, population differentiation was detected in both haploids and diploids and varied with landscape features, and not with geographic distance. Using an assignment test, we establish that effective migration rates varied according to height on the shore. In this intertidal species, biased spore dispersal may occur during the transport of spores and gametes at low tide when small streams flow from high- to lower-shore pools. The longevity of both haploid and diploid free-living stages and the long generation times typical of G. gracilis populations may promote the observed pattern of high genetic diversity within populations relative to that among populations.