Abstract
Microscale genetic differentiation of sessile organisms can arise from restricted dispersal of sexual propagules, leading to isolation by distance, or from localised cloning. Cyclostome bryozoans offer a possible combination of both: the localised transfer of spermatozoa between mates with limited dispersal of the resulting larvae, in association with the splitting of each sexually produced embryo into many clonal copies (polyembryony). We spatially sampled 157 colonies of Crisia denticulata from subtidal rock overhangs from one shore in Devon, England at a geographic scale of ca. 0.05 to 130m plus a further 21 colonies from Pembrokeshire, Wales as an outgroup. Analysis of molecular variance (AMOVA) revealed that the majority (67%) of genetic variation was distributed among individuals within single rock overhangs, with only 16% of variation among different overhangs within each shore and 17% of variation between the ingroup and outgroup shores. Despite local genetic variation, pairwise genetic similarity analysed by spatial autocorrelation was greatest at the smallest inter-individual distance we tested (5cm) and remained significant and positive across generally within-overhang comparisons (<4m). Spatial autocorrelation and AMOVA analyses both indicated that patches of C. denticulata located on different rock overhangs tended to be genetically distinct, with the switch from positive to negative autocorrelation, which is often considered to be the distance within which individuals reproduce with their close relatives or the radius of a patch, occurring at the 4-8m distance class. Rerunning analyses with twenty data sets that only included one individual of each multilocus genotype (n=97) or the single data set that contained just the unique genotypes (n=67) revealed that the presence of repeat genotypes had an impact on genetic structuring (PhiPT values were reduced when shared genotypes were removed from the dataset) but that it was not great and only statistically evident at distances between individuals of 1-2m. Comparisons to a further 20 randomisations of the data set that were performed irrespective of genotype (n=97) suggested that this conclusion is not an artefact of reduced sample size. A resampling procedure using kinship coefficients, implemented by the software package GENCLONE gave broadly similar results but the greater statistical power allowed small but significant impacts of repeat genotypes on genetic structure to be also detected at 0.125-0.5 and 4-16m. Although we predict that a proportion of the repeat multilocus genotypes are shared by chance, such generally within-overhang distances may represent a common distance of cloned larval dispersal. These results suggests that closely situated potential mates include a significant proportion of the available genetic diversity within a population, making it unlikely that, as previously hypothesised, the potential disadvantage of producing clonal broods through polyembryony is offset by genetic uniformity within the mating neighbourhood. We also report an error in the published primer note of Craig et al. (Mol Ecol Notes 1:281-282, 2001): loci Cd5 and Cd6 appear to be the same microsatellite
Pemberton, Andrew J