Inheritance of a nuclear DNA polymorphism assayed in single bivalve larvae

We examined the inheritance of theMytilus edulis CaM-1 Intron 3 locus, a non-coding DNA locus with two potentially neutral length-variants. The polymerase chain reaction (PCR) was used to determine theCaM-1 genotype for 799 larvae obtained from 11 laboratory crosses. Larvae were typed singly only 8 to 24 h after fertilization. We find evidence that each allele can be inherited from either sex, that there are no barriers to fertilization between gametes of different genotypes, and that most larvae have genotypes compatible with Mendelian inheritance of the locus. Deviations from expected genotype frequencies were found in some crosses; we suggest that a contributing factor is aneuploidy in early larvae, but a major cause is more likely to be selection at a locus linked toCaM-1, occurring under the artificial laboratory culture conditions. This study demonstrates the feasibility of applying molecular genetic techniques to early larval stages of marine bivalves, and presents a new non-destructive biopsy method for DNA analysis from living adult mussels.     

Patterns of genetic subdivision in populations of a clonal cnidarian,Zoanthus coppingeri,from the Great Barrier Reef

Samples of an intertidal zoanthid, Zoanthus coppingeri, Haddon and Shackelton, 1891, were collected from three localities in the Great Barrier Reef region during 1992–1993, and subjected to allozyme electrophoretic analysis at seven polymorphic loci. The reduced ratio of observed to expected genotypic diversity indicated that populations were partly clonal, but they were not dominated by a few clones as occurs in some other cnidarians. Regular disturbance by wave action is postulated to prevent the formation of large stands of particular clones by clearing space and mixing genotypes over small scales. The sexual origin of clonal genotypes was confirmed by conformance to Hardy-Weinberg predictions of genotype frequencies at all but one locus. Values of the standardised genetic variance among populations, F _ST , were highly significant between localities and between replicate sites within localities separated by only 50 m. Strong genetic structure has not previously been described in a Great Barrier Reef invertebrate species, and is considered to be the consequence of stochastic changes in gene frequencies as a result of low levels of gene flow. High clonal longevity and low recruitment rates may maintain genetic differences over long periods. Similar effects may be seen in other Great Barrier Reef invertebrate species with comparable reproductive patterns.     

Genotype-dependent fecundity and temporal variation of spawning in hybrid mussel (Mytilus) populations

Mussel samples were collected at 4 to 6 wk intervals throughout 1987 from two hybridMytilus edulis/M. galloprovincialis populations, at Croyde Bay and Whitsand Bay, in southwest England. These were analyzed at two polymorphic loci which are diagnostic for allozyme differences which typifyM. edulis andM. galloprovincialis. Dry mantle weight as a function of shell length was determined for all individuals of each sample. Size-frequency data for the two populations was obtained in September 1987 and March 1988. For all genotypes at both sites, fecundity was a function of shell length, and in both populations the frequency ofM. galloprovincialis alleles was positively correlated with shell length. At both sites, allozyme genotype explained a significant amount of variation in mantle weight either when assessed as a main effect or when assessed as an interaction with shell length or time of collection. At Croyde,M. galloprovincialis mussels had greater estimated fecundity per unit length than theM. edulis mussels. Differences in the timing of spawning activity between theM. edulis and theM. galloprovincialis mussels were inferred, and these differences might act to reduce the amount of interbreeding at Croyde. At Whitsand, a reduced level of variability in the timing of spawning activity and fecundity between the genotypes was observed and explained by a higher degree of genetic mixing. Because theM. galloprovincialis mussels had (1) a greater estimated fecundity at any length, and (2) a greater mean length than theM. edulis mussels, the mean genotypic annual fecundity perM. galloprovincialis mussel was 2.8 times greater than an individualM. edulis mussel at Croyde, and 2.2 times greater than an individualM. edulis mussel at Whitsand. This evidence thatM. galloprovincialis mussels have an advantage in fecundity, and thus perhaps in fertility, taken together with the evidence thatM. galloprovincialis also has a higher viability, indicates directional selection in favour of theM. galloprovincialis phenotype. Because of the observed temporal stability of the population it seems likely that this selection is counterbalanced by a massive imigration ofM. edulis spat from neighbouring populations.     

Comparisons of genotype-tolerance responses in populations of <Emphasis Type="Italic">Hediste diversicolor</Emphasis> (Polychaeta: Nereididae) exposed to copper stress

The analysis of genotypic shifts in field-exposed populations has been proposed as a tool for monitoring the environmental impacts of contaminants. Previous laboratory experiments showed the occurrence of genotypic shifts induced by copper exposure in samples of Hediste diversicolor from the Pialassa estuary (North Adriatic Sea, Italy). In order to test if genotype-tolerance responses observed were consistent at larger spatial scales, populations of H. diversicolor were sampled in three haphazardly chosen estuaries along the Italian coast (Comacchio, Pialassa, Serchio) and exposed to 0.48 mg l⁻¹ Cu²⁺ under laboratory conditions. Survival analysis procedures were used to test for possible relationships between time-to-death and genotypes at three allozyme loci (ALD, FH, PGI). Genotype-tolerance responses observed at locus ALD were consistent among populations up to 2500 km distant, with genotype ALD ^102/102 associated with the shortest survival times. Comparisons with previous laboratory experiments showed that responses to copper stress at locus ALD were also consistent in time, with genotypes ALD ^102/102 and ALD ^100/102 associated with higher 96 h mortality ratios. Results suggested the occurrence of stable relationships between genotypes at locus ALD and tolerance responses of H. diversicolor to copper stress. Conversely, genotype-tolerance responses at loci PGI and FH were not consistent either in space (among populations from the three estuaries) or in time (between laboratory experiments), being possibly affected by site-specific features of populations or stochastic factors. Results show that genotype-tolerance responses should be carefully verified at different spatial and temporal scales, before considering genotypic shifts among the possible tools for monitoring the effects of environmental stressors.     

Genetic differentiation of populations of the planktonic copepod Labidocera aestiva

Planktonic populations of the calanoid copepod Labidocera aestiva show significant biochemical genetic heterogeneity along the Atlantic coast of the USA. In summer, 1981, copepods were collected by surface tows at Beaufort Inlet, North Carolina; Fort Pierce Inlet, Florida; and Vineyard Sound, Massachusetts. Genetic variation within each population and genetic differentiation among the three populations were studied by micro-acrylamide gel electrophoresis of six loci encoding four enzymes. All six enzyme loci were polymorphic when all populations were considered together, but the North Carolina population was monomorphic at two loci. High genetic variability was indicated by the following: (1) the number of alleles per locus averaged over all loci was 2.57±0.26 SD; (2) the average proportion of loci for which the frequency of the most common allele was not greater than 0.95 was 0.78±0.10; (3) the frequency of heterozygous individuals was 0.25±0.07. Genetic differentiation among population samples in the three regions was demonstrated in several ways: allele frequencies at one aminopeptidase-I locus, Lap-1, differed significantly among samples of the three populations, and there were unique alleles of high frequency at this locus in two population samples. Values of the statistic of genetic distance (D) averaged 0.20±0.08 for pairwise comparisons between all samples. Compared to expected heterozygosity if individuals across the entire range sampled mated at random, there were highly significant heterozygote deficiencies at five of the six loci. Genetic differentiation of populations of L. aestiva may result from (1) differential selection on populations in the three regions, or (2) restricted gene flow between the populations. Gene flow may be limited by geographic separation or differences in life history, such as seasonal presence in the plankton and diapause egg production.Contribution No. 5810 of Woods Hole Oceanographic Institution     

Genetic subdivision within the eastern Australian population of the sea anemoneActinia tenebrosa

An electrophoretic survey of allozyme variation revealed substantial genetic differentiation within the eastern Australian population ofActinia tenebrosa. This differentiation appears to reflect the effects of both asexual reproduction and limited gene flow among local populations separated by up to 1050 km. Variation was assessed within groups of 27 to 55 adults sampled between September 1985 and December 1988 collected from small areas of shore within each of 24 local populations. All individuals were collected from stable rock platforms, with the exception of Boulder Bay, where some sea anemones were removed from small mobile boulders. High levels of variability were detected for each of seven enzyme-encoding loci. The patterns of genotypic variation detected imply that local populations are maintained by predominantly asexually generated recruitment. Levels of multi-locus genotypic diversity within samples were consistently less than 50% of the level expected for sexual reproduction with free recombination. This was reflected by the detection of relatively low numbers of multi-locus genotypes and significant departures from expectations for single-locus Hardy-Weinberg equilibria within 17 of the 24 local populations. Standardised genetic variances (F _ST ), calculated from the genotypes of all individual adults were typically much greater than those expected for marine organisms with widely dispersed larvae. The former values were reduced, but were still extremely large when clonal genotype frequencies were substituted into the calculation. These data imply that although widely dispersed larvae may be an important source of initial colonists, levels of gene flow among established local populations are low. Furthermore, cluster analysis revealed a clear subdivision of the population into northern and southern groups. However, this subdivision was largely explained by strong clinal variation at a GPI-encoding locus. For this locus, allele frequencies ranged from fixation of the A allele in samples from the 12 most northern sites to near fixation of the alternative B allele in southern samples. Subdivision of the eastern Australian population is consistent with the predicted off-shore movement of the Eastern Australian Current close to the border between Victoria and New South Wales. However, the split into northern and southern regions, as evidenced by the variation forGpi, could reflect patterns of gene flow and/or other factors such as natural selection or the recent patterns of colonisation.Contribution No. 78 from the Ecology and Genetics Group of the University of Wollongong     

Calanoid copepod,cladoceran, and rotifer eggs in sea-bottom sediments of northern Californian coastal waters: Identification,occurrence and hatching

In Ireland, mussels on exposed rocky shores constitute an interbreeding mixture of two forms of mussels,Mytilus edulis L. and the Mediterranean musselM. galloprovincialis Lmk. This paper presents an in-depth analysis, carried out between October 1984 and December 1986, of genetic variability at two partially diagnostic loci,Odh andEst-D, in two exposed-shore populations ofMytilus spp. in the west of Ireland. Significant differences at theOdh locus were observed in the genetic composition of adult mussels from different tidal levels. These differences were repeatable whether one was analysing replicate samples at a single point in time, samples collected at different points in time, i.e., in different years, or samples collected from different shores. Mussels recruiting to artificial substrates set out for a period of one month at different tidal levels at one of these sites were also observed to be genetically different; mussels higher up the shore exhibited higher frequencies of those alleles characteristically at high frequency inM. galloprovincialis for both theOdh andEst-D loci. Hence, the genetic differences observed in adult mussels are much more exaggerated in juveniles and are already apparent within the first month of benthic life. Possible reasons for the observed microgeographic differentiation are discussed. It is concluded that the observed genetic differences between mussels at different tidal levels arise either in the pelagic/attachment stage or very shortly after settlement.     

Genetic variability and differentiation of sporophytes and gametophytes in populations of Gelidium arbuscula (Gelidiaceae: Rhodophyta) determined by isozyme electrophoresis

Genetic differentiation and genetic variability of sporophytic and gametophytic populations of Gelidium arbuscula (Bory) from three localities sampled in 1989 and 1990 in the Canary Islands (Spain) were examined by isozyme electrophoresis. Twenty-three to 29 putative alleles corresponding to 22 gene loci, were compared. High deviations in Hardy-Weinberg equilibrium, and significant differences between allelic frequencies of sporophytic and gametophytic subpopulations at the same locality were found, suggesting a predominant asexual reproduction of G. arbuscula. The genetic variability (percentage of polymorphic loci, mean number of alleles per locus and average gene diversity) of haploid subpopulations was lower than that of diploid subpopulations at all three localities, being the lowest described for seaweeds. No correlation between genetic and geographical distance was found. The high genetic differentiation coefficient between all subpopulations suggests a very reduced genetic flow between subpopulations of the same and of different localities. These results suggest that the genetic structure of the populations of G. arbuscula from the Canary Islands is due to a founder-effect combined with a predominance of asexual reproduction. This is the first report comparing allelic frequencies between sporophytic and gametophytic subpopulations of seaweeds.     

Genetic structure and allozyme variation of sea bass (Dicentrarchus labrax and D. punctatus) in the Mediterranean Sea

This paper reports data on 28 allozyme loci in wild and artificially reared sea bass (Dicentrarchus labrax) samples, originating from either coastal lagoon or marine sites in the Mediterranean Sea. F _ST analysis (θ estimator) indicated strong genetic structuring among populations; around 34% of the overall genetic variation is due to interpopulation variation. Pairwise θ estimates showed that, on average, the degree of genetic structuring was much higher between marine populations than between samples from lagoons. Six polymorphic loci showed differences in allele frequencies between marine and lagoon samples. Multivariate analyses of individual allozymic profiles and of allele frequencies suggested that different arrays of genotypes prevail in lagoons compared to marine samples, particularly at those loci that, on the basis of previous acclimation experiments, had been implicated in adaptation to freshwater. On the other hand, variation at “neutral” allozyme loci reflects to a greater extent the geographic location of populations. Allozyme differentiation was also studied in a D. labrax population from the Portuguese coast. Average genetic distance between this population and the Mediterranean populations was quite high (Nei's D = 0.236) and calls into question the taxonomic status of the Portuguese population. Finally, genetic relationships between D. labrax and D. punctatus were evaluated. Average Nei's D was 0.648, revealing high genetic differentiation between the two species, even for two sympatric populations of these species in Egypt; thus gene flow was not indicated between species. Received: 24 October 1996 / Accepted: 27 November 1996     

Genetic variation of mitochondrial DNA in mussel (Mytilus edulis and M. galloprovincialis) populations from South West England and South Wales

Mitochondrial DNA (mtDNA) and allozyme variation were analysed in samples of mussels collected in 1984 and 1985 from four localities in South West England and one locality in South Wales, a region of Britain where the common mussel (Mytilus edulis) occurs sympatrically and hybridises with the Mediterranean mussel (M. galloprovincialis). Significant differences in mtDNA genotype frequencies for three restriction enzymes (BstEII, XbaI, and EcoRI) were observed between mussels from M. galloprovincialis populations (Padstow and Bude) and those from an M. edulis population (Swansea). Some mtDNA genotypes at high-frequency in M. galloprovincialis were not observed in M. edulis, although there was no indication that mtDNA variation provides greater overall diagnostic power than allozyme variation in distinguishing between the two forms of mussel. Construction of a phylogenetic tree of multiple mtDNA genotypes revealed small mutational distances between the genotypes characterising M. edulis and M. galloprovincialis. The results were consistent with predominant mtDNA flow from M. edulis to M. galloprovincialis. This can be explained by the dispersal of larvae to South West England from M. edulis regions to the north and east, but little dispersal in the opposite directions. Samples from two hybrid populations (Whitsand and Croyde) were analysed. mtDNA genotype frequencies at Croyde were in line with predictions made on the basis of two partially diagnostic allozyme loci (Est-D and Odh), mtDNA frequencies at Whitsand were not. Frequencies of some mtDNA genotypes at Whitsand were characteristic of M. edulis, others of M. galloprovincialis. Differential selective mortality or flow of different mtDNA genotypes and allozyme variation are proposed as possible causes of these results.     

Short-term stability of genetic structure in populations of the sea anemone Metridium senile

Populations of the sea anemone Metridium senile (L.) were sampled from several locations in eastern North America in two series, one collected from 1977–1979 and the other from 1981–1985. Fourteen populations were sampled twice at one- to six-year intervals. Samples were analyzed for temporal differences in genetic composition at both the single locus and multiple locus levels. Overall patterns of geographic variation in allele frequency did not change between series. Regressions describing clines did not differ significantly, and loci not showing clinal variation in the first series remained similar in the second. Analysis of populations sampled twice produced no systematic evidence of change in allele frequency with time for any of four polymorphic loci. Comparisons with computer simulations of repeated sampling of multiple locus genotypes from panmictic populations with free recombination also revealed little temporal change at this level. One population showed possible evidence of recruitment from a different gene pool. Other significant departures from expectations reflected more reproducibility of genotype distributions between samples than expected for sexual populations. This excess stability likely results jointly from clonal reproduction and little sexual recruitment. Despite these indications of genetic stability in adult populations, newly settled juveniles were genetically different from resistant adults in one population, demonstrating the potential for genetic change by immigration. Successful sexual recruitment seems to be rare, even though larvae regularly settle from the plankton. Although interpretation of these results is somewhat limited by lack of knowledge of longevities and generation times, to the extent that they reflect longer-term trends, they suggest that at least some of the observed patterns of geographic variation in allele frequency probably result from natural selection.     

Lignocellulose and lignin in the salt marsh grass Spartina alterniflora: initial concentrations and short-term,post-depositional changes in detrital matter

Individuals of Mytilus edulis of the same age (ca 2 months) were collected as spat from natural populations. Relative growth rates were determined among individuals differing in heterozygosity at five enzyme loci. Growth rate was positively correlated with individual heterozygosity and each of the five loci contributed about equally to the relatinship. More heterozygous individuals also achieved more uniform average growth rates. Although there was a deficiency of heterozygotes at each locus, relative to Hardy-Weinberg expectations, the magnitude of the deficiency, measured as F_IS, was less among faster growing mussels. Our results conform closely with those of Zouros et al. (1980) on the American oyster. We conclude that the relationship between multiple locus heterozygosity and growth rate is one that is general to a diversity of outbreeding plant and animal populations. Other studies indicate that this relationship is due to a greater average metabolic efficiency of more heterozygous individuals. This relationship does not emerge from experimental designs in which there has been limited genetic sampling of the natural genetic variation.     

Population genetics of the blue crabCallinectes sapidus: modest population structuring in a background of high gene flow

To determine the genetic population structure of blue crabs (Callinectes sapidus Rathbun), electrophoretic allozyme analysis was performed on 750 individuals collected from 16 nearshore locations ranging from New York to Texas, USA. Twenty enzymes and non-enzymatic proteins coded by 31 presumptive loci were examined. Twenty-two loci were either monomorphic or polymorphic at less than theP ₉₅ level; alleles for these polymorphic loci were geographically dispersed. Allele frequencies for three of the remaining polymorphic loci were homogeneous over all populations, as were levels of polymorphism and heterozygosity. Phenograms generated by the UPGMA (unweighted pair-group method using arithmetic averages) and distance Wagner methods exhibited no geographic pattern in the clustering of populations. Estimates ofN _em (effective number of migrants per generation between populations) indicated substantial gene flow, with aalues sufficiently high to infer panmixia between all blue crab populations from New York to Texas. However, despite this high level of gene flow, two striking patterns of geographic differentiation occurred: genetic patchiness and clinal variation. Allele frequencies atEST-2, GP-1, IDHP-2, DPEP-1, DPEP-2, andTPEP exhibited genetic patchiness on local and range-wide geographic scales, and allele frequencies atEST-2 varied temporally. Genetic patchiness in blue crabs is likely to be the result of the pre-settlement formation and subsequent settlement of genetically heterogeneous patches of larvae; allele frequencies of those larval patches may then be further modified through ontogeny by localized selection. In the Atlantic Ocean, a regional latitudinal cline ofEST-2 allele frequencies was superimposed on the range-wide genetic patchiness exhibited by that locus. This pattern against a background of high gene flow is highly likely to be maintained by selection. In estuaries along the Atlantic Ocean coast, a combination of low adult long-distance migration and a high retention rate of locally spawned larvae could serve to segregate populations and allow for the development of the geographic cline inEST-2. The Gulf of Mexico showed no apparent cline, perhaps due to long-distance migration of females in some regions of the Gulf, or to masking by genetic patchiness. These results emphasize the importance of both ecological and evolutionary time scales and structuring mechanisms in determining genetic population structure.     

Glucosephosphate isomerase genotype frequencies,homozygous excess and size relationships in the sand flounder Rhombosolea plebeia

Genetic variation was studied at two polymorphic loci for 3 yr (November 1978–February 1982) in juvenile sand flounder (Rhombosolea plebeia) in Wellington Harbour, New Zealand. The Gpi-1 locus showed a heterogeneity in genotype frequencies, but this was not clearly related to season or to fish length. At the Gpi-2 locus, genotype frequencies were dependent upon length and showed a significant excess of homozygotes in juveniles but not in adults. This latter phenomenon has been reported previously in several species of marine molluscs, and so may be common to marine organisms producing large numbers of pelagic larvae. Reasons for the excess of homozygotes are discussed, and it is suggested that assortative mating is the most likely explanation and that this is an area of marine population genetic research which has been over-looked.     

Allozyme variation in populations of the dog-whelkNucella lapillus (Prosobranchia: Muricacea) from the South West peninsula of England

Four populations of the predatory gastropodNucella lapillus were sampled at sites around the South West Peninsula of England in 1986, and analysed for allozyme variation at 18 enzyme loci. Two of these loci, Gpd-1 andHk-1, exhibited sex-specific phenotypes. An absolute locus association was observed between two other loci,Mdh-1 andEst-3. This association was only found at one site (Prawle), and it is suggested that the presence of chromosomal polymorphisms could explain this finding. As a measure of overall similarity, Nei's genetic identity statistic,I, was calculated; the mean for all populations was 0.989, with values ranging from 0.981 to 0.997. Although similar on this gross level, considerable interpopulation variation was evident. Observed mean heterozygosity (per locus) ranged from 0.043 to 0.104 (mean 0.074). Populations differed also in the loci at which significant heterozygote deficits were seen (of the seven deficits recorded only those at thePep-1 locus were consistent across sites) and in the presence of rare alleles undetected elsewhere. The variation observed showed no correlation to shell morphology or geographical distance and confirmed the conclusion that species of the genusNucella show considerable disjunct variation.     

Genetic structure of hybrid mussel populations in the west of Ireland: two hypotheses revisited

In Ireland, mussels on exposed rocky shores constitute an interbreeding mixture of two forms of mussels, the blue mussel, Mytilus edulis, and the Mediterranean mussel, M. galloprovincialis. Results from an Irish study in the 1980s, using partially diagnostic allozyme markers, indicated that mussels higher up the shore were more galloprovincialis-like than those lower down. In this study we set out to test two hypotheses: (a) recruits arriving on the shore are composed of genetically distinct cohorts that settle preferentially at different levels on the shore, and maintain genetic distinctiveness into adulthood; (b) recruits are genetically homogeneous, but once settled they diverge genetically over time, due to within-habitat site specific-selection. The diagnostic Me 15/16 DNA marker was used to analyse the genetic composition of newly-settled spat recruiting to artificial substrates, which were placed at two-week intervals from May–October 2002, on the mid- and low shore areas of two exposed sites in Galway Bay. Adult mussels were also collected on each sampling date. Results did not support the preferential settlement hypothesis, i.e., the genetic composition of primary settlers (≤ 500 μm) was similar between tidal heights and shores. Neither was there evidence of post settlement selective mortality, as adults were genetically similar to settling spat. In spat and adults the frequency of the M. galloprovincialis allele was high (0.56–0.80), due to high frequencies of M. galloprovincialis (> 37%) and hybrid (> 33%) genotypes, and correspondingly low frequencies of the M. edulis genotype (< 11%). Adult mussels from a nearby sheltered estuarine site, while significantly different to exposed shore mussels, still had low frequencies of the M. edulis genotype (< 17%), indicating no apparent advantage for the genotype in this environment. There are indications that the genetic composition of mussels may be changing on the Atlantic coasts of Ireland.     

Size-dependent, spatial and temporal genetic variation at a leucine aminopeptidase (LAP) locus among blue mussel (Mytilus galloprovincialis) populations along a salinity gradient

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus is related to salinity variation in several marine bivalve molluscs. This paper details an investigation of the Long Island Sound model of LAP selection (LAP genotype-dependent mortality occurring among newly settled Mytilus edulis mussels) carried out in 1997 among three M. galloprovincialis mussel populations along the salinity gradient of Wellington Harbour, New Zealand. Significant LAP genotypic heterogeneity was observed at the LAP locus between small (<25 mm shell length) and large (>25 mm shell length) M. galloprovincialis from Petone and Eastbourne (the two sites experiencing the greatest salinity variation), whereas genotypic heterogeneity was not significantly different between small and large mussels from Seatoun (the site experiencing the least salinity variation). The Lap ³ allele decreased in frequency and the Lap ⁴ allele increased in frequency at Petone and Eastbourne, whereas the Lap ³ and Lap ⁴ allele frequencies remained effectively constant at Seatoun. Both these findings are consistent with the Long Island Sound model of selection. At all three locations, the Lap ^3,3 genotype decreased in frequency from small to large mussels, whereas the Lap ^3,4 genotype increased in frequency from small to large mussels. All other LAP genotypes occurred at low frequencies (<0.10) at all three locations and showed no evidence of frequency change from small to large-size mussels nor evidence of clinal change among the three locations. These genotype frequency data possibly indicate that the Lap ^3,3 genotype is at a selective disadvantage compared to the Lap ^3,4 genotype at all three locations, and that this selective disadvantage is related to the extent of salinity variation which exists at each location. Further investigation is required before it can be determined if the Long Island Sound model of selection best describes the size-dependent and location-specific changes in LAP allele and genotype frequencies along this salinity gradient. Comparison of the population genetic structure at the LAP locus in 1995 and in 1997 revealed a profound change from heterozygote excesses to heterozygote deficiencies for all three M. galloprovincialis populations. The reason for the change is unknown, but the change indicates that population genetic structure at the LAP locus is highly variable in time, but consistent in space, among these M.␣galloprovincialis populations. Received: 5 February 1998 / Accepted: 27 May 1998     

Genetic divergence between colour morphs in populations of the common intertidal sea anemones Actinia equina and A. prasina (Anthozoa: Actiniaria) in the Isle of Man

A distinctive morph ofActinia equina (L.) is found at low frequency among populations on intertidal hard substrata on some shores on the Isle of Man. This morph is red with rows of green oval spots or elongate markings running longitudinally down the column. Starch gel electrophoresis of 21 allozyme loci was used to compare samples of this morph from two localities with other sympatricActinia spp. Collections were made in early summer 1986. For one sampling site the comparison was with redA. equina and greenA. prasina Gosse, whilst at the other site with redA. equina only. At both sites significant genetic differences in allele frequencies at several loci were found between the new morph and sympatricA. equina andA. prasina. The results also confirm the reproductive isolation ofA. prasina. As with much previous work on sea anemones, levels of mean heterozygosity per locus (H) were found to be high (H=0.157 to 0.342). A surprising feature of the results, although bearing out earlier unpublished data, is that high levels of genetic differentiation are found between populations of what appear to be the same morphs collected from shores only a few km apart. Among theActinia spp. studied, differentiation between populations of the same morph on different shores was of the same order as between sympatric, reproductively isolated populations.     

Genetic variance in Ophiomusium lymani (Ophiuroidea: Echinodermata) from lower bathyal depths in the Rocklall Trough (northeast Atlantic)

Specimens of the deep-sea brittle-star Ophiomusium lymani were collected from six sites in the Rockall Trough (northeast Atlantic_. Four monomorphic and four polymorphic loci were detected, with up to 75 individuals screened at any one locus. The results showed little difference in allele frequencies between sites and, consequently, estimates of genetic identity indicated no significant genetic differentiation between the populations sampled. Deficits of heterozygotes were observed at all polymorphic loci, but the deficiencies were only significant at the phosphoglucose isomerase locus from two locations. The deficit of heterozygotes was not the same across loci, suggesting that inbreeding is not the cause of the excess homozygosity. No relationship between heterozygosity and depth was observed.     

Muscle esterase variation and size variation in the sand launce Ammodytes dubius

Esterase variation detected in homogenates of muscle tissue taken from the sand launce Ammodytes dubius in April, 1970 and 1971 is described. Eight alleles were found segregating at a single locus, and the frequencies of these alleles were homogenous in population samples taken on the Emerald Bank, off Nova Scotia, Canada, in successive years. When ordered by their electrophoretic mobility, the profile of the frequencies of these alleles was unimodal and symmetric. In both of the population samples, there were significant excesses of homozygotes and deficencies of heterozygotes. There was an association between genotype and size of individuals among homozygous genotypes; individual homozygous for the most common, intermediately-migrating allele were larger than homozygotes bearing alleles with relatively fast or slow mobilities.