The encrusting spongeHalisarca laxus: population genetics and association with the ascidianPyura spinifera

The encrusting spongeHalisarca laxus forms a seemingly obligate association with the stalked solitary ascidianPyura spinifera. In 1991 we examined spatial variation and short-term temporal variation in this association at three neighbouring sites in southeastern Australia. This sponge dominated the surface of almost all the 500 individual ascidians examined, with mean cover usually exceeding 90%. This pattern was consistent among sites and throughout the year of the study. The domination of a small isolated patch of habitable substratum by a sponge is most unusual, given that they are regarded as relatively poor recruiters. To understand how this association might be maintained, we determined the underlying genotypic diversity of the sponge population using starch-gel electrophoresis.P. spinifera is a clump-forming ascidian and usually occurs in clumps of up to 22 individuals. Electrophoretic surveys, based on six variable allozyme loci, revealed that at a total of five plots within three neighbouring New South Wales populations, single sponge genotypes may cover entire ascidian clumps; although a clump sometimes played host to more than one sponge clone. Allele frequencies (averaged across four loci that appear to conform to Mendelian inheritance) showed little variation among populations (standardised genetic variance,F _ST=0.013). Nevertheless, sponge populations were genotypically diverse, with samples from 63 of 172 individual clumps displaying unique clonal genotypes. Moreover, multi-locus genotypic diversity within all sites approached the level expected for sexual reproduction with random mating. Taken together, these data imply thatH. laxus produces sexually-derived larvae that are at least moderately widelly dispersed. Given the relatively small size of the patches that this sponge inhabits, we also conclude that these larvae are good colonists and good spatial competitors on their ascidian hosts.     

High levels of gene flow in the surf bivalve Donax deltoides (Bivalvia: Donacidae) on the east coast of Australia

We surveyed patterns of allelic variation within twelve samples of the pipi Donax deltoides Lamarck from beaches separated by up to 1200 km but connected to varying degrees by the East Australian Current. We used these data to test the prediction that the irregular patterns of water movement would cause genetic differentiation in pipis, so that there would be more genetic variation within and among the more southern regions than the northern regions. We found that six loci were at least moderately variable within all samples, and there were no clear geographic patterns in allelic frequencies. In general, genotype frequencies within samples were consistent with predictions for an outcrossed, sexually reproducing species, and we detected no evidence of population subdivision. Within samples, with the exception of the peptidase loci, single-locus genotype frequencies were in close agreement with expectations for Hardy–Weinberg equilibrium. We observed no significant linkage disequilibrium for any pairwise comparison of loci in any sample. Our hierarchical analysis of genetic variation revealed little variation among all samples (F st = 0.009). Loci showed consistently low levels of subdivision (F st from 0.003 to 0.018). We found almost no variation among the four geographic regions sampled (F rt = 0.001). All variation was therefore attributable to variation among samples within regions (F sr = 0.010). These data imply that larvae are moving between regions and that levels of present or recent gene flow are high, and support the conclusions of other studies which have inferred widespread gene flow for animals dispersing via planktonic, outcrossed larvae in parts of this region. This implies that the East Australian Current is sufficient to produce strong larval connections despite its intermittent nature. If existing levels of population subdivision reflect current levels of gene flow, then these data imply that D. deltoides represents a single fishery on the east coast of Australia. Received: 16 September 1996 / Accepted: 25 September 1996     

Mode of reproduction,recruitment, and genetic subdivision in the brooding sponge<Emphasis Type="Italic"> Haliclona</Emphasis> sp.

Sponges display a variety of reproductive strategies that have the potential to influence population genetic structure. Histological examination of ten reproductive individuals of the Western Australian sponge Haliclona sp. showed that this species broods embryonic larvae that are potentially limited in dispersal capabilities. Because sponges have the potential to propagate in a number of modes, allozyme electrophoresis was used to assess the relative importance of asexual and sexual reproduction to recruitment, and to quantify genetic subdivision over different spatial scales. Tissue samples from 227 sponges were collected from reefs within two areas 400 km apart: Hamelin Bay and Rottnest Island. Contrary to expectations for highly clonal populations, genotypic diversity within sites was high, no linkage disequilibrium was found, and there was no evidence of genotypic clustering within reefs. There was no genetic evidence that asexual reproduction is important for the maintenance of populations. Genetic comparisons were consistent with mixing of sexually produced recruits within reefs, on a scale up to a few hundred metres, but significant genetic subdivision between reefs (F_ST=0.069 at Hamelin Bay, 0.130 at Rottnest Island) indicated that water gaps of several hundred metres are effective at preventing dispersal. Subdivision between the two areas, separated by 400 km, was moderately greater (F_ST=0.142) than within, but the same alleles were predominant in the two areas. These genetic patterns are consistent with limited dispersal capabilities of brooded larvae.Communicated by G.F. Humphrey, Sydney     

DNA Fingerprinting Reveals a Lack of Genetic Variation in Northern Populations of the Western Pond Turtle (Clemmys marmorata)

I used DNA fingerprinting to provide the first analysis of the genetic composition of western pond turtle ( Clemmys marmorata ) populations in Washington, Oregon, and California. Populations of the western pond turtle in Washington and northern Oregon are rapidly approaching extinction. Genetic similarity within the largest northern populations, which are located inland, is high. An analysis of population substructure (F_st) revealed significant genetic divergence between inland populations, indicating a lack of dispersal and gene flow between sites. In contrast, northern coastal sites are not genetically distinct, but there are few if any viable populations remaining in this region. Genetic variability within southern California populations is a great deal higher than in northern inland sites. Similarly, a low F_st value indicated a lack of genetic differentiation between southern sites. An inter-regional analysis of population substructure (F_st = 0.24) revealed a significant degree of genetic divergence between geographical regions throughout the range. In addition, an estimate of western pond turtle phylogeny showed a genetic break in the species between northern and southern populations. Both population subdivision and phylogenetic analyses suggest a lack of appreciable gene flow between geographical regions for a considerable period of time. Genetic analyses support traditional subdivision based solely on the morphological variation of Clemmys marmorata into two subspecies: northern Clemmys marmorata marmorata and southern Clemmys marmorata pallida . Recovery of dwindling northern populations must combine demographic and genetic considerations. A first step should be to preserve local gene pools while augmenting population numbers, with the goal of preventing the extinction of this genetically and morphologically distinct subspecies.     

Genetic structure of a scleractinian coral, Mycedium elephantotus, in Taiwan

The genetic structure of Mycedium elephantotus (Pallas, 1766) populations from three regions around Taiwan was examined using allozyme electrophoresis. Eight loci were scored from seven enzyme systems. Seven loci were polymorphic under the 95% criterion. The high ratio of observed to expected genotypic diversities at the collection sites (G _O:G _E=0.8 to 1.0) indicate that M. elephantotus propagates predominantly by sexual reproduction. Allele frequencies of M. elephantotus differed significantly among regions (D=0.024 to 0.256, F _ST=0.032 to 0.218, p < 0.001), while populations among collection sites within each region were homogeneous (D=0.000 to 0.015, F _ST=0.010 to 0.022, p > 0.05). Genetic differentiation between populations from southern Taiwan and the Penghu Islands is greater than that between southern and northern Taiwan populations, although the former two regions are much closer geographically. The UPGMA dendrogram based on Nei's unbiased genetic distance showed a clear subdivision of populations into two groupings, northern Taiwan/Penghu Islands and southern Taiwan. A higher level of gene flow was found between M. elephantotus populations in northern Taiwan and the Penghu Islands (N _m=7.56) than that between populations in southern Taiwan and other regions (N _m=0.90 to 1.72). The pattern of genetic subdivision among regions is consistent with the pattern of ocean currents, indicating that genetic differentiation is likely driven by surface circulation vicariance. Received: 2 December 1997 / Accepted: 15 September 1998     

Interspecific variation in thermal denaturation of proteins in the congeneric musselsMytilus trossulus andM. galloprovincialis: evidence from the heat-shock response and protein ubiquitination

The genetic population structure of the precominant zooplankter, the copepod Calanus finmarchicus (Gunnerus), was examined to determine whether genetically distinct populations exist in the Gulf of Maine. C. finmarchicus was sampled in three regions of the Gulf of Maine (Great South Channel, spring 1989; northern Gulf of Maine, winter 1990; Great South Channel and Georges Bank, spring 1990). Copepods from seven locations in the Great South Channel, five in the northern Gulf of Maine and four on or near Georges Bank were assayed for allozyme variation and mitochondrial DNA variation of amplified 16S rRNA and cytochrome b genes. Restriction fragment length polymorphism (RFLP) analyses of both mitochondrial DNA genes revealed no variation among any of the individuals assayed. Analysis of five polymorphic allozyme loci revealed that genetic variation among the three geographic regions was low, and genetic identities were high between all locations (I>0.97). Most of the genetic variation was among locations regardless of region. Chi-square tests were used to examine genetic similarity between specific pairs of locations within and between regions. In the northern Gulf of Maine, genetic homogeneity occurred over larger spatial scales (hundreds of km) than in either the Great South Channel or Georges Bank (tens of km). Only copepods from the Bay of Fundy and Nova Scotian Shelf locations were genetically distinct from Wilkinson Basin copepods at two loci. Copepod populations from the northern locations may have been partially isolated or they may represent immigrant populations (e.g., from the Gulf of St. Lawrence). Several pairs of locations were genetically distinct at one or more loci in the two southern regions. Differences between locations in these regions may represent distinct populations advected into the areas at different times or from different sources (e.g., genetic variation may represent a mixture of genetically distinct northern and southern copepod populations). These results suggest extensive gene flow among populations of C. finmarchicus in the Gulf of Maine with some evidence of genetic population subdivision near the Gulf's northeastern and southern boundaries.     

Allozyme and mitochondrial DNA variation in orange roughy,Hoplostethus atlanticus (Teleostei: Trachichthyidae): little differentiation between Australian and North Atlantic populations

Allozyme and mitochondrial DNA (mtDNA) genetic variation was compared in orange roughy (Hoplostethus atlanticus Collett) collected from waters off southern Australia and from waters about 22 000 km away in the North Atlantic west of Scotland. Samples were screened for 11 polymorphic allozyme loci and with 9 restriction enzymes. Significant heterogeneity between the two areas was detected for three allozyme loci (ADA ^*, CK ^* and GPI-1 ^*), and the overall G _ST (gene-diversity statistic) value of 1% was small but significant. Significant mtDNA haplotype heterogeneity was observed after ²- of haplotype frequencies but not after a G _ST analysis. Nucleotide sequence-diversity analysis showed very low net divergence (0.0023%) between the two samples. The Australian orange roughy had a lower allozyme heterozygosity and a lower mitochondrial DNA nucleon diversity than the North Atlantic sample. The very limited, although significant, allozyme and mitochondrial DNA heterogeneity between these areas suggests that there is some gene flow between these two populations. The species appears to be widespread, with its presence reported from the southern Pacific, southern Indian, and northern and southern Atlantic Oceans, and it is likely that gene flow between the antipodes is mediated by stepping-stone exchange between adjacent populations rather than by direct migration.     

High levels of genetic subdivision in peripherally isolated populations of the atherinid fish Craterocephalus capreoli in the Houtman Abrolhos Islands,Western Australia

The atherinid fish Craterocephalus capreoli Rendahl is abundant in the Houtman Abrolhos Islands, 70 km off the Western Australia coast and 250 km south of the southern limit of the range of the species along the mainland. Electrophoretic examination of 7 allozyme loci at 17 sites in the Houtman Abrolhos revealed a substantially lower level of polymorphism than found in an earlier study of the species in its mainland distribution, with many of the uncommon alleles and some common ones missing. There is a very high degree of genetic subdivision among the populations in the Houtman Abrolhos, measured by a mean F _ST of 0.437 over a distance of 35 km. This F _ST (standardized variance in allelic frequencies) is six times that found previously among populations along the mainland coast over distances up to 850 km. The subdivision of populations in the Houtman Abrolhos is similar within one island group on a scale up to 12 km, and between two groups that are separated by 15 km of deep water. Significant differences in allelic frequencies were found between populations from the open shore and enclosed lagoons less than 800 m apart, but the overall levels of subdivision were similar for the two types of environment. Previous work had shown high levels of genetic subdivision in the Houtman Abrolhos for a gastropod with direct development. The results for C. capreoli demonstrate that the archipelago favours subdivision even for a species with potentially much greater mobility and different life history.     

Population genetics of the northern Pacific seastar Asterias amurensis (Echinodermata: Asteriidae): allozyme differentiation among Japanese,Russian, and recently introduced Tasmanian populations

The northern Pacific seastar Asterias amurensis Lütken was recently introduced to Tasmanian waters, possibly through ballast water discharged from ocean-going vessels. Three Tasmanian populations and seven native populations from Japan and eastern Russia were examined in 1994 for variation at 22 allozyme loci. There was significant spatial heterogeneity in gene frequencies among the native populations, especially for the ocus APK ^*, where G _ST was a very high 0.47. The Tasmanian populations could not be separated genetically from one another, suggesting they have a common origin from a single introduction. The average heterozygosity per locus of the Tasmanian populations (0.116 to 0.127, mean 0.123) was 30 to 40% less than that of the native populations (0.177 to 0.216, mean 0.192), suggesting that colonisation of Tasmanian waters was accompanied by a population-size bottle-neck. The Tasmanian seastars were genetically more closely related to the two populations from central Japan (Suruga and Tokyo Bays) than to populations from Vladivostok, northern Japan (Yoichi, Nemuro and Mutsu Bays) or southern Japan (Ariake Sea). However, there were significant differences between the populations from Tasmania and central Japan, which may be attributable to founder effects or may mean that the true source of the Tasmanian introduction has yet to be identified.     

Acanthochromis polyacanthus,a fish lacking larval dispersal,has genetically differentiated populations at local and regional scales on the Great Barrier Reef

Acanthochromis Gill is a monotypic genus within the damselfish family Pomacentridae, erected for an unusual species [A. polyacanthus (Bleeker)] that uniquely lacks larval dispersal. Instead, offspring are reared in the parental territory, in the manner of cichlids, and fledged into the surrounding habitat. Phenotypic and genotypic variation was surveyed on the basis of body colouration and 7 polymorphic loci in 19 populations from 5 regions of the central and southern Great Barrier Reef (GBR). Variation in both characters was found at regional and local scales. Two colour morphs were recognised: a bicoloured morph from the three northern regions and a uniform dark morph from the two southern regions. Isozyme analysis showed a similar pattern with greatest variation between the different morphs, but also with significant variation at both regional and local scales within morphotypes. Heterozygosity was maximal in the central populations, which, together with other measures of variability, suggests a mixing of separate gene pools in this region and denies species status to the two morphotypes despite numerous fixed differences in allele frequencies between the most distant populations. The presence of fixed differences in multiple alleles between populations separated by 1000 km indicates negligible gene flow over such distances and long isolation of these gene pools. These patterns may reflect recolonisation of the GBR after the last sea-level rise by fish from two stocks. Founder effects and random drift in small populations after colonisation are probably the major sources of the local and regional variations observed at smaller spatial scales. This diversity has been maintained among populations at all scales by the very low levels of gene flow possible without an effective strategy for larval dispersal between coral reefs.     

Biochemical genetic variation and evidence of speciation in Chthamalus barnacles of the tropical eastern Pacific Ocean

Gene-enzyme variation was studied electrophoretically within and between barnacle populations of the genus Chthamalus from 8 intertidal stations from central California to the Pacific coast of Panamá. Horizontal starch-gel electrophoresis separated and resolved 18 enzymes from 512 individual barnacles. A maximum of 25 gene-enzyme systems was interpretable from the resulting zymograms. Electrophoretic phenotypes and patterns of phenotypic variation generally conformed with those observed in other organisms. The amount of genetic variation within barnacle populations varied; average heterozygosity, for example, ranged from a low of 3.4% in a Mexican population of the C. fissus group to a high of 10.4% in C. anisopoma from the Gulf of California. Observed and expected average heterozygosities agreed in all population samples, indicating that these species are outbreeding. In contrast to the prediction stemming from a hypothesis that trophic stability regulates the amount of genetic variation in marine species, average heterozygosity tends to be positively correlated with latitude. Data from these and other barnacle species may support a hybrid environmental heterogeneity-trophic diversity model recently proposed to explain genetic variation in decapod crustaceans. Juxtaposition of individuals from different localities revealed numerous genetic differences among populations of the C. fissus group. At least three partially sympatric sibling species are separated by genetic distances as large as those observed between the C. fissus group and the distinct species C. dalli and C. anisopoma. A cladistic analysis places C. anisopoma close to the Mexican and Panamanian sibling species, with C. fissus from San Diego and C. dalli successively farther removed.     

No genetic differentiation of giant clam (Tridacna gigas) populations in the Great Barrier Reef,Australia

Six Tridacna gigas populations were sampled in 1990 from locations throughout the central and northern Great Barrier Reef (GBR). Despite separations in excess of 1000 km, mean Nei's unbiased genetic distances among the populations was 0.0007. The complete lack of spatial variation observed among populations did not results from lack of genetic variability. Genetic variation within populations was high, with mean heterozygosities from 0.221 to 0.250. Gene frequencies were consistent with expectations under conditions of Hardy-Weinberg equilibrium. These data suggest panmixis, or random mating, throughout the highly connected reef system of the central and northern GBR. The large gene exchange among the giant clam populations has important implications for conservation management of one of the few large populations of T. gigas in the world. Small local effects are likely to be overcome in time by inputs from other sources. However, large genetic perturbations, particularly from up-current sources, may spread rapidly through the population.Contribution No. 561 from the Australian Institute of Marine Science     

High levels of genetic subdivision of marine and estuarine populations of the estuarine catfishCnidoglanis macrocephalus (Plotosidae) in southwestern Australia

The cobblerCnidoglanis macrocephalus (Valenciennes) is an endemic marine and estuarine catfish from southern Australia. Conflicting views on the degree of isolation of the estuarine populations underscore general questions about genetic divergence in coastal species. Although estuaries are widely recognized as ecologically important, little work has been done on their role in favouring genetic divergence. In order to estimate the extent of genetic subdivision among nearshore marine and estuarine populations, electrophoretic variation of enzymes was examined in seven marine and six estuarine populations of cobbler from sites spanning 1500 km along the southwest Australian coastline. Among all populations, the mean standardized variance in allelic frequencies (F _ST) for six polymorphic loci was 0.277, a high value comparable to those of other shallow-water teleosts whose life-history characteristics and habitat preferences restrict their dispersal capability. The pattern of genetic identities between populations showed divergence between west and south coast sites. Within these regional groups, however, there was substantial heterogeneity, much of which was associated with estuaries. Among all six estuarine sites, the averageF _ST was 0.333, 40% higher than the value of 0.237 for the marine sites. Low estimates of the genetically effective number of migrants suggest population subdivision between marine and estuarine environments and between similar habitat types. This study indicates the importance of habitat in affecting the connectedness of populations, even in apparently open marine systems.     

Molecular and morphological analyses of the cuttlefish<Emphasis Type="Italic"> Sepia apama</Emphasis> indicate a complex population structure

The giant Australian cuttlefish Sepia apama Gray, 1849 annually forms a massive and unique spawning aggregation in northern Spencer Gulf, South Australia, which has attracted commercial fishing interests in recent years. However, many basic life-history characteristics of S. apama are unknown, and anecdotal evidence suggests that there is more than one species. The present study assessed the population structure and species status of S. apama using data from allozyme electrophoresis, microsatellite loci, nucleotide sequences of the mitochondrial COXIII gene, multivariate morphometrics and colour patterns. Analyses of allozyme and microsatellite allele frequencies revealed two very divergent but geographically separated populations consisting of specimens from the east coast and southern Australia. However, the presence of a heterozygote in a putative contact zone between the east coast and southern Australia suggested that these populations were not reproductively isolated. Mitochondrial haplotypes seem to have introgressed further north into the contact zone than have nuclear alleles. Differences in colour patterns that previously had been attributed anecdotally to different geographic populations were, in fact, correlated with sexual dimorphism. These data are most consistent with S. apama being one species the populations of which were geographically isolated in the past (historical vicariance) and have come into secondary contact. Comparison of microsatellite allele frequencies among four South Australian samples indicated significant deviations from panmixia. South Australian samples were also reliably diagnosed by means of multivariate morphometrics. Significant differences in mantle length were observed among populations.Communicated by M.S. Johnson, Crawley     

Characterization of disjunct populations of Zostera marina (eelgrass) from California: genetic differences resolved by restriction-fragment length polymorphisms

Comparative restriction-fragment analysis was used to analyze the nuclear ribosomal DNA, and alcohol dehydrogenase-1 loci of Zostera marina L., for variation within and among populations. Eelgrass is a perennial marine flowering plant that is widespread and ecologically significant throughout the temperate northern hemisphere. A chemical method was developed to obtain restriction-quality DNA without CsCl fractionation from experimentally relevant quantities of seagrass tissues (0.5 to 1.0 g). The yield was 25 g g^-1 fresh weight. The three morphologically distinct forms of Z. marina from disjunct populations examined in this study were found to be genetically distinct; morphologically similar populations were indistinguishable genetically. Genetic distinction also correlated with habitat depth, as subtidal and intertidal populations were clearly divergent. Homologous probes for the 17S and 28S ribosomal DNA genes were used to map 24 restriction sites on the rDNA repeat of Z. marina, which was determined to be about 14 kb in length. At least 1 length mutation and 5 restriction-site changes were identified that distinguished Z. marina populations from San Diego and Monterey Bay (Del Monte Beach) from Z. marina populations from Elkhorn Slough and Tomales Bay. Estimated sequence variation (100×p) between eelgrass populations ranged from 0.00 to 0.69. Individual plants were observed to contain as many as four different rDNA-repeat length variants. The mean number of rDNA-repeat length variants per individual in Z. marina was about two. Intrapopulation variation in rDNA-repeat type was observed in only one individual from the Tomales Bay population.     

Historical demography and contemporary spatial genetic structure of an estuarine crab in the northeast Pacific (<Emphasis Type="Italic">Hemigrapsus oregonensis</Emphasis>)

Discrete estuary subpopulations of the mud crab Hemigrapsus oregonensis (Dana, 1851) are connected via larval dispersal. Sequence variation at the mtDNA COI locus was examined in eight populations sampled in 2001–2002 from central California through northern Oregon in the northeast Pacific (36.6–45.8°N) to infer patterns of dispersal and historical connectivity in the region. Strong evidence for persistence since the mid-Pleistocene, with no range truncation resulting from southward shifting temperature isoclines, was provided by a phylogeographic pattern of haplotypes of an older clade distributed throughout the sampled range. A recently derived clade became widespread only north of Cape Blanco after the last glacial maximum. Its clear pattern of restriction to the northern area, in the absence of similarly restricted southern clades, suggests that contemporary dispersal around Cape Blanco is rare (population F _ST = 0.192). Low pairwise differentiation within Oregon and within central California, as well as contrasts between northern and southern groups in the shape of the pairwise mismatch distribution, nucleotide diversity, and Tajima’s D suggest that these regions reflect different demographic histories. Potential mechanisms explaining this latitudinal break include contemporary coastal circulation patterns, selection, and ancient patterns of larval dispersal in the California Current.     

Genetic subdivision in the subtidal,clonal sea anemone Anthothoe albocincta

Anthothoe albocincta, a common subtidal anemone along south-eastern Australia, reproduces both sexually through broadcast spawning and clonally through fission. Clones may be distinguished both by their electrophoretic genotypes and the colour of their tentacles and oral discs. Local populations typically consist of many, dense clonal aggregations. However, some clones appear to have locally extensive distributions, forming a series of separate aggregations. The capacity of clones to disperse among local populations is unknown. In this study we used an electrophoretic survey at six allozyme loci to quantify levels of variation among samples from each of 13 local populations and four geographic regions sampled between April 1992 and June 1993. These data revealed that populations of A. albocincta, separated by up to 930 km, were at least moderately subdivided. Levels of geographic variation were high and the average F _ST value (standardised genetic variation) was 0.27 based on genotypes of all individual polyps. This value reflects substantial variation both within (F _PR =0.13) and among (F _RT =0.16) regions. Estimates of gene flow among both neighbouring populations and regions are therefore low (N _e m=1.7 and 1.3, respectively). UPGMA (unweighted pair-group method using arithmetic averages) dendrograms suggest that a genetic discontinuity occurs at the very south-east corner of Australia, paralleling reports for two other south-eastern Australian marine invertebrates. In addition, our analyses and theoretical predictions imply that localised proliferation of clonal genotypes may have caused us to underestimate the potential importance of gene flow via larval dispersal. Moreover, the abundance and vast geographic range of this species suggests that widespread dispersal does occur. Collections from three populations covering the peroid December 1992 to June 1993 were examined by crude dissection along with histological sectioning, and showed A. albocincta to be dioecious, with unisexual clones. Eggs within the ovaries of six females sampled over a 3 mo period were small (96±4 m) and similar to those of related species that produce planktotrophic larvae. In contrast, we found no evidence that clones were dispersed (shared) among neighbouring local populations. An average of only 6% of six-locus genotypes were common to pairs of local populations separated by up to 125 km, this being equal to the percentage expected through sexual reproduction alone. In addition, the percentage of shared genotypes did not decline with increasing geographic separation. These data imply that although asexual reproduction may be used to maintain local populations, the sexual production of genotypically diverse larvae is the primary source of widely dispersed colonists and hence of new clones.     

Population structure in the sexually reproducing sea anemone Oulactis muscosa

The intertidal sea anemone Oulactis muscosa (Drayton) is dioecious and most individuals are sexually mature throughout the year. Biochemical genetic evidence was used to determine the genetic structure of populations and to infer the relative contributions of sexual and asexual reproduction to recruitment. Data were collected for six enzyme-encoding loci from local populations spread along 735 km of the south east coast of Australia. The genetic structure of each of the nine local populations studied was consistent with recruitment by sexually produced individuals. In almost all cases, the observed single-locus genotypic frequencies closely matched those expected for hardy-Weinberg equilibria, however, consistent deficits of heterozygotes were detected for all loci. No apparent subdivision of the population was detected within the sampling area. Low levels of genetic differentiation were found between local populations and standardised variance (F _ST ) values were similar to those for other species with widespread planktonic dispersal of larvae.Contribution No. 60 from the Ecology and Genetics Group of the University of Wollongong     

Genetic evidence for contrasting patterns of dispersal in solitary and colonial ascidians

In situ and in vitro observations indicate that brooding colonial ascidians commonly display limited larval dispersal, whilst the larvae of most solitary species are assumed to be widely dispersed. We used allozyme data to determine the population genetic consequences of reproduction and dispersal in a broadcast-spawning solitary ascidian and two brooding colonial species along the central and southern coast of New South Wales, Australia. We surveyed genetic variation at 2 to 9 variable loci for samples collected from 6 to 8 local populations of each of the stalked solitary species Pyura gibbosa gibbosa Heller, 1878; the social Stolonica australis Michaelsen, 1927 and the compound Botrylloides magnicoecum Hartmeyer, 1912. Samples from each local population displayed levels and patterns of genotypic diversity that were consistent with expectations for sexually-derived recruitment of both solitary zooids and separate colonies. However, we found clear differences in the structure of the populations of solitary and colonial species. Genotype frequencies within all nine samples of P. gibbosa gibbosa conformed to expectations for random mating (i.e. Hardy–Weinberg equilibria). Moreover, allele frequencies showed little variation among samples [mean standardised genetic variance (F _{S T} ) =0.002], which implies that local populations are strongly connected by larval dispersal. We estimate (via Wright's “island model”) that gene flow (N _e m) within this set of local populations is 125 effective migrants per generation, which is very similar to estimates obtained for other broadcast-spawning taxa in this region. In contrast, genotype frequencies within samples of both colonial species were characterised by large and statistically significant deficits of heterozygotes, consistent with expectations for highly limited dispersal of larvae or sperm. Moreover, local populations were highly differentiated (F _{S T} =0.201 and 0.202 for S. australis and B. magnicoecum, respectively) and N _e m was estimated to be ∼1.0 in each case. These values of F _{S T} and subsequent estimates of N _e m lie within the range of values reported for other New South Wales taxa with direct larval development, and imply that local populations are effectively closed to immigration. Received: 13 February 1997 / Accepted 18 July 1997     

Effects of isolation by distance and geographical discontinuity on genetic subdivision of Littoraria cingulata

Littoraria cingulata (Philippi, 1846) is a Western Australian, mangrove littorine snail, represented by two morphologically distinct subspecies, whose distributions are separated by >300 km. The southern subspecies, L. cingulata pristissini, is distinguished from the northern subspecies, L. cingulata cingulata, by having a thinner, keelless shell with more primary grooves, and lower and much more numerous ribs. In contrast with these striking differences, L. cingulata cingulata is morphologically very similar to another species, L. sulculosa, with which it also shares a nearly coincident geographic range. Allozyme comparisons at 22 presumptive loci confirmed a large genetic distance between L. cingulata and L. sulculosa, and the apparent conspecificity of the morphologically divergent subspecies of L. cingulata. Based on geological evidence, the geographical separation of the morphologically divergent forms of L. cingulata has developed within the past 5000 to 10 000 yr. The extensive continuous distribution of the northern subspecies, L. cingulatacingulata, and the large geographic disjunction between the northern and Shark Bay subspecies, L. cingulata pristissini, allowed a test of the genetic importance of this relatively recent disjunction. Within the continuous distribution of the two subspecies, a pattern of isolation by distance was visible up to distances of 300 km. Beyond 300 km, genetic subdivision, measured by pairwise G _ST (the proportion of genetic diversity due to differences between populations), averaged 0.028, whereas subdivision between Shark Bay and northern populations averaged 0.055 over the same range of distances. Although the relative paucity of barriers to gene flow tends to limit genetic subdivision in marine species with planktotrophic larvae, the results for L. cingulata suggest that subdivision can occur within a continuous distribution, but that special events leading to major disjunctions can substantially increase divergence, even over a relatively short period of time. Received: 16 February 1998 / Accepted: 23 April 1998