Parthenogenesis in Psocids (Insecta: Psocoptera)

SYNOPSIS. Obligate thelytokous parthenogenesis and, more rarely,facultative thelytoky are known to occur in psocids. The formerphenomenon is known in 28 species. The latter is known onlyin two species, both of which have obligatorily thelytokousforms as well as bisexual ones. Attempts to force thelytokousfemales to mate have been carried out for four nominal species.All have failed. Differences in color or color pattern betweenthe bisexual and thelytokous forms of a nominal species havebeen observed in two instances. In Echmepteryx hageni this seemsto involve divergence where the two forms occur sympatricallybut on different habitats. Investigations of geographic distributionof bisexual and thelytokous forms of a nominal species showin several cases that the thelytokous forms are much more widespreadthan the bisexual ones. In eastern United States, bisexual formstend to occur peripherally and in ecologically unusual areas.The areas of occurrence of bisexual populations may be relictual,and some of them may have been Pleistocene refugia. Most ofthese areas correspond to areas where coniferous trees now forman important part of the forest.     

Multiple origins of stone loach,Barbatula barbatula (Teleostei: Nemacheilidae), in Sweden based on mitochondrial DNA

Stone loach, Barbatula barbatula, occurs in scattered localities in Sweden. Some of the populations have usually been considered as feral descendants of escaped 18th Century pond stock, but historical documentation is inconclusive. Using the mitochondrial COI gene as a marker, we analyzed specimens from seven Swedish localities. One of the middle Swedish localities, in Stockholm, belongs to a haplotype found also in Poland and Lithuania. Two other samples, from near Nyköping and Lake Hjälmaren, belong to a haplotype found in northeastern Europe (Finland, Estonia, Latvia, Volga basin in Russia). Those two Swedish populations are probably descendants from at least two introductions, probably for pond rearing for human consumption. Samples from Skåne and Halland in southern Sweden belong to the haplotype found in Denmark, northern Germany and Poland; and whereas it remains possible that they also represent feral populations, they may be naturally occurring, having reached Sweden during the Ancylus period, about 8,000–10,000 years ago. A recently discovered population from the central South Swedish Highlands belongs to a mainly southeastern European haplotype. It probably represents a release of imported aquarium specimens or live bait carried by sport fishing tourists.     

Population genomics of the raccoon dog (<Emphasis Type="Italic">Nyctereutes procyonoides</Emphasis>) in Denmark: insights into invasion history and population development

The raccoon dog (Nyctereutes procyonoides) has a wide distribution in Europe and is a prominent example of a highly adaptable alien species. It has been recorded sporadically in Denmark since 1980 but observations since 2008 suggested that the species had established a free-ranging, self-sustaining population. To elucidate the origin and genetic patterns of Danish raccoon dogs, we studied the population genomics of 190 individuals collected in Denmark (n = 141) together with reference captive individuals from Poland (n = 21) and feral individuals from different European localities (Germany, Poland, Estonia and Finland, n = 28). We used a novel genotyping-by-sequencing approach simultaneously identifying and genotyping a large panel of single nucleotide polymorphisms (n = 4526). Overall, there was significant indication for contemporary genetic structuring of the analysed raccoon dog populations, into at least four different clusters, in spite of the existence of long distance gene flow and secondary admixture from different population sources. The Danish population was characterized by a high level of genetic admixture with neighbouring feral European ancestries and the presence of private clusters, non-retrieved in any other feral or captive populations sampled. These results suggested that the raccoon dog population in Denmark was founded by escapees from genetically unidentified Danish captive stocks, followed by a recent admixture with individuals migrating from neighbouring Germany.     

Picea abies- and Fraxinus excelsior-dominated wetland forest communities in Latvia

This paper reports on the syntaxonomical structure, the ecological, physiognomical and phytogeographical differentiation of wetland forests in Latvia dominated by Picea abies and Fraxinus excelsior. Data were collected following the Braun-Blanquet approach and treated by the GROUPAGE, TWINSPAN and CANOCO programs. Spruce forests (Sphagno girgensohnii-Piceetum var. Crepis paludosa, S.g.-P. myrtilletosum) form a well-defined community mostly in shallow and medium deep (0.2–1.3 m) peatland. Assemblages of spruce wetlands in NEE Europe point to amazing similarity and little variation from NE Poland and Fennoscandia over the Baltic States to the central parts of Russia. Ash-dominated wetlands (Carici elongatae-Alnetum, Carici remotae-Fraxinetum) refer only to particular localities of the country and lie on wet mineral, gleyey soils, usually along brooks and on undulating ground rich in subsoil streams, mostly in sites without an evident accumulation of peat. In Latvia, these reach the northernmost limit of distribution and are poorer in species compared with the ash wetlands of Central Europe.     

Holocene distribution and extinction of the moose (Alces alces, Cervidae) in Central Europe

Moose (Alces alces L.) were among the first large mammals to recolonize Central Europe after the last glaciation. Already during the Allerød they established themselves in most parts of the area. In the early Holocene their distribution range extended from the Pyrenees to Denmark and from Austria to Great Britain and also covered eastern Central Europe where they still occur today. In the Preboreal, the moose slowly vanished from the southwestern parts of its distribution range, leading to its extinction in France and, later, in England. During the Atlantic period, the moose died out in large parts of Denmark and population densities apparently decreased in the rest of Central Europe as well. Around the birth of Christ only relict populations were left in western Central Europe, which finally became extinct in early medieval times. In Thuringia and in the region northeast of the river Elbe as well as in central Poland, some stocks persisted until the high and late Middle Ages. The causes of the gradual extinction in Central Europe during the Holocene are complex. Changes in vegetation, climate and sea-level, the increasing fragmentation of habitat through human activities and hunting were, at different times, important factors. In the recent past, however, moose have repeatedly migrated from the east towards the west. The development of its distribution range since the end of the Second World War as well as experiences with Scandinavian populations show that moose are able to thrive in close proximity to humans and that a future expansion of its distribution range towards the west seems possible.     

Phylogeography of the common vole (Microtus arvalis) with particular emphasis on the colonization of the Orkney archipelago

To investigate the human introduction of the common vole Microtus arvalis onto the Orkney islands, the complete cytochrome b gene was sequenced in 41 specimens from both Orkney (four localities) and elsewhere in their range (26 localities). Orkney voles belonged to the same phylogenetic lineage, 'Western', as individuals from France and Spain indicating southwestern Europe as the most likely source area for the islands. This result is of interest with respect to the movement and trading links of the Neolithic people who likely transported the voles. As well as the Western lineage, our phylogenetic trees revealed three other purely European lineages: the 'Italian' (single specimen from N. Italy), the 'Central' (Germany, Netherlands, Denmark) and the 'Eastern' (Hungary, Slovakia, Poland, Ukraine, Finland, European Russia). Individuals from European Russia, W. Siberia, Georgia, Ukraine and Armenia formed a fifth distinct lineage coinciding with the distribution of the 'obscurus' chromosomal form of M. arvalis. These phylogeographical data suggest that M. arvalis occupied multiple refugia during the last glaciation.     

The biosystematic relations between a thelytokous and an arrhenotokous form ofAphytis mytilaspidis (Le Baron) [Hymenoptera: Aphelinidae]

Biological and morphological characters of the thelytokous and arrhenotokous forms ofA. mytilaspidis and their hybrids were investigated. Some biological differences between the thelytokous and arrhenotokous forms were found. The former is adapted to warmer condition and will parasitizeHemiberlesia lataniae as well asDiaspis echinocacti, while the latter will not parasitizeH. lataniae. The overall fecundity of the thelytokous form is lower than that of the arrhenotokous form, whereas the “effective fecundity”, female progeny per female parent, is higher in the thelytokous form. Apart from the no. of setae in the wings delta, there are no distinguishing morphological characters between the arrhenotokous and thelytokous forms.     

Genetic variability of central–western European pine marten (Martes martes) populations

Recent studies highlighted the potential role of cryptic glacial refugia for temperate taxa in Europe beyond the Mediterranean peninsulas. To further investigate phylogeographic features of the European pine marten (Martes martes) in previously identified cryptic refugia located in central–western Europe, we analysed the hyper-variable diagnostic fragment of the mitochondrial control region in a total of 134 specimens, allowing for reliable comparisons with previous genetic studies of the species. We included samples from eight different European countries in central–western Europe (Belgium, France, Luxembourg and the Netherlands), in south–western Europe (Spain), in north–central Europe (Denmark) and in central Europe (Germany and Poland). The sequences collapsed in 17 haplotypes, which allowed us to determine the genetic composition of the pine marten populations throughout central–western Europe. Overall, our results showed that the population genetic variation, estimated by the standardised haplotype diversity, was high (0.400?≤?Hs?≤?0.762), and it was considerably higher in Germany (0.762) and the Netherlands (0.722) compared to the other countries. The nucleotide diversity was relatively low (0.002?≤?π?≤?0.016) even in Germany and the Netherlands (0.016 and 0.014, respectively), suggesting relatively small, long-term effective population sizes or severe bottlenecks. Out of the 17 haplotypes found in our study area, 13 were unique and limited to a single country: one in Denmark, one in Spain, four in Poland and seven in the Netherlands. The pairwise genetic distance ranged from 0.001 to 0.032 and did not show any evident correlation with the geographic distances between the populations. A genealogical relationship network was constructed, which provided evidence for a recent origin of many of the unique haplotypes. Approximately 82 % of the samples analysed in this study belonged to haplotypes grouped into a previously identified central–northern European phylogroup of the species. Our results support previous findings, indicating low contribution of southern refugial populations to the postglacial recolonization of central–western Europe and a predominant contribution of pine marten populations that survived the Last Glacial Maxima in cryptic northern refugia.     

Genetic structure, habitat fragmentation and bottlenecks in Danish bank voles (Clethrionomys glareolus)

Tissue-samples from 161 bank voles (Clethrionomys glareolus) collected in three forests (five sampling localities) situated in eastern Jutland (Denmark) were analysed by nine microsatellite loci. The genetic diversity found within the populations was high (H_e=0.753–0.806).Bank voles have specific habitat requirements favouring woodlots, hedgerows and deciduous forests as their prime living area. Hence, a natural or human-induced fragmentation of the forest may cause a sub-structuring of the populations and thereby a restriction of dispersal among populations. The sub-structuring indicated by the observed significant genetic differentiation among the five geographically distinct localities (F_st =0.033, P<0.05) could either result from habitat fragmentation or a combination of home range behaviour and different tree composition in the forests. A road situated between two adjacent forests was not found to exert any barrier effect to the gene flow of bank voles. In one out of five localities investigated, genetic evidence for a recent bottleneck-like situation was found. Bank voles are known to exhibit sometimes huge density fluctuations not only from year to year but also from season to season. The bottleneck-like situation found could therefore be due to the low number of individuals during the low-density phase.     

Apochromic populations of Dactylorhiza incarnata s.l. (Orchidaceae): diversity and systematic significance as revealed by allozyme markers and morphology

Apochromic forms of the Eurasian Dactylorhiza incarnata s.l. were studied in northern Europe to reveal their genetic (allozyme) and morphological diversity and to assess their systematic significance. The study included eight localities with sympatric populations of plants with anthocyanin‐pigmented and apochromic flowers. Parallel samples of the two morphs were taken from each locality. Genetic variation was only found at the allozyme loci pgd, pgi and ugpp. Statistically significant differences in allele frequencies between the two colour morphs were found in two localities and demonstrate that the occurrence of apochromic individuals in D. incarnata s.l. is not always because of spontaneous mutation. At least in some localities the apochromic plants form distinct breeding groups (but local populations of different colour morphs may also be composed of several more or less distinct breeding groups). Based on molecular and morphometric data, it is proposed that the apochromic study populations from calcareous fens should be referred to D. incarnata var. ochroleuca, whereas the apochromic study populations from non‐calcareous fens are better treated as aberrant local populations of var. incarnata s.l. Possible evolutionary patterns and processes are discussed and guidelines for identification of var. ochroleuca from morphological features are given. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 396–407.     

Population genetics of Wolbachia‐infected,parthenogenetic and uninfected,sexual populations of Tetrastichus coeruleus (Hymenoptera: Eulophidae)

Wolbachia are endosymbiotic bacteria known to manipulate the reproduction of their hosts. These manipulations are expected to have consequences on the population genetics of the host, such as heterozygosity levels, genetic diversity and gene flow. The parasitoid wasp Tetrastichus coeruleus has populations that are infected with parthenogenesis‐inducing Wolbachia and populations that are not infected. We studied the population genetics of T. coeruleus between and within Wolbachia‐infected and uninfected populations, using nuclear microsatellites and mitochondrial DNA. We expected reduced genetic diversity in both DNA types in infected populations. However, migration and gene flow could introduce new DNA variants into populations. We therefore paid special attention to individuals with unexpected (genetic) characteristics. Based on nuclear and mitochondrial DNA, two genetic clusters were evident: a thelytokous cluster containing all Wolbachia‐infected, parthenogenetic populations and an arrhenotokous cluster containing all uninfected, sexual populations. Nuclear and mitochondrial DNA did not exhibit concordant patterns of variation, although there was reduced genetic diversity in infected populations for both DNA types. Within the thelytokous cluster, there was nuclear DNA variation, but no mitochondrial DNA variation. This nuclear DNA variation may be explained by occasional sex between infected females and males, by horizontal transmission of Wolbachia, and/or by novel mutations. Several females from thelytokous populations were uninfected and/or heterozygous for microsatellite loci. These unexpected characteristics may be explained by migration, by inefficient transmission of Wolbachia, by horizontal transmission of Wolbachia, and/or by novel mutations. However, migration has not prevented the build‐up of considerable genetic differentiation between thelytokous and arrhenotokous populations.     

TELOSCHISTES FLAVICANSIN GREAT BRITAIN: DISTRIBUTION AND ECOLOGY

Over the last century, the distribution ofTeloschistes flavicanshas contracted from being widespread in the southern half of England and Wales to being limited to South-West England with outlying populations in Pembrokeshire and North Wales. Twelve core sites have been identified where the species is well established: ten of these are saxicolous/terricolous habitats on windy coastal cliff tops; the other two are lines of sycamore trees near the coast. At 39 further localities, some inland, the species is in very small amounts (often on one tree) and vulnerable to extinction. It is normally a member of theParmelietum revolutaeor theRamalinetum scopularisassociations. The conservation of the species is discussed.     

Homogeneity and distinctiveness of Polish paternal lineages revealed by Y chromosome microsatellite haplotype analysis

Different regional populations from Poland were studied in order to assess the genetic heterogeneity within Poland, investigate the genetic relationships with other European populations and provide a population-specific reference database for anthropological and forensic studies. Nine Y-chromosomal microsatellites were analysed in a total of 919 unrelated males from six regions of Poland and in 1,273 male individuals from nine other European populations. AMOVA revealed that all of the molecular variation in the Polish dataset is due to variation within populations, and no variation was detected among populations of different regions of Poland. However, in the non-Polish European dataset 9.3% ( P<0.0001) of the total variation was due to differences among populations. Consequently, differences in R(ST)-values between all possible pairs of Polish populations were not statistically significant, whereas significant differences were observed in nearly all comparisons of Polish and non-Polish European populations. Phylogenetic analyses demonstrated tight clustering of Polish populations separated from non-Polish groups. Population clustering based on Y-STR haplotypes generally correlates well with the geography and history of the region. Thus, our data are consistent with the assumption of homogeneity of present-day paternal lineages within Poland and their distinctiveness from other parts of Europe, at least in respect to their Y-STR haplotypes. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00439-002-0728-0.     

Cranial variation in the European badger Meles meles (Carnivora,Mustelidae) in Scandinavia

Morphometric variation in 30 craniometric characters of 465 skulls of the European badgers (Meles meles) from across Europe was analysed. Multivariate analyses revealed that the populations from Norway, Sweden, and Finland differ from other European populations in having smaller skulls. The analyses also revealed significant differences between the ‘south‐western Norwegian’ and ‘main Fennoscandian’ forms. On average, badgers from south‐west Norway were smaller than those of the remaining Fennoscandia. Morphological differences between the ‘south‐western Norwegian’ and ‘main Fennoscandian’ populations of M. meles suggest a possible in situ semisympatric divergence since the beginning of the Holocene warming, or a complex history of two groups involving at least two colonization routes. The small‐sized Scandinavian badgers may be close to the ancestral form that used to be widespread in Denmark and throughout Europe. The animals from south‐west Norway may instead be descendants of ancestors that were the first to penetrate the southern parts of the Scandinavian Peninsula. The ‘main Fennoscandian’ badgers are likely to have been the descendants of the second wave of recolonization of Scandinavia. Specifically, they might have colonized the Scandinavian Peninsula from the east after the last glaciation.     

Relationship between Sex Distribution and Sporophyte Production in Pleurozium schreberi (Brid.) Mitt.

The distribution of perigonia, perichaetia, and sporophytesof Pleurozium schreberi has been investigated in Britain andin some other areas. Fruiting populations are widespread ina zone extending from northern Scotland across southern Scandinavia,but are rare in southern Britain, and appear to be infrequentin many other parts of western Europe. Less information is availableoutside western Europe: fruiting collections have been examinedfrom scattered localities throughout much of the species' northerncircumpolar range, but there is no evidence of areas where theyare abundant. The rarity of sporophytes is correlated with a rarity of plantsbearing male inflorescences. Barren specimens normally comprisedonly female and sterile plants, while most fruiting specimenswere bisexual but contained more female than male stems. Femaleplants are abundant throughout Great Britain, but plants withmale inflorescences appear to be widespread only in northernScotland and in East Anglia. The latter area is unusual in thatsporophyte production fails in many bisexual colonies. Sporophyte production was stimulated experimentally by transplantingmale plants from East Anglia into female populations in theWest Midlands. The male plants spread vegetatively within thefemale colonies, and perigonia developed during six successiveyears. At an East Anglian station sporophytes developed aftermaterial was transplanted between unisexual male and femalecolonies only 10 m apart.     

Morphometric differentiation of the moor frog (Rana arvalis Nilss.) in Central Europe

Morphometric variation of 11 characters was studied in 14 samples of Rana arvalis from allopatric population groups in Central Europe, representing two subspecies formerly recognised: R. a. arvalis and R. a. wolterstorffi. All samples from Poland (nine) were collected from the area, which is believed to be populated by the nominal form. In addition, the isolated population of R. arvalis from the Eastern Carpathians of Romania was classified to that form. All the Hungarian samples fall into the range of R. a. wolterstorffi. No significant differences in body size between sexes and among the populations from the three geographic regions (Poland, Hungary and Romanian Eastern Carpathians) were found. Multivariate analysis of the differences in body shape among groups showed that the populations from Hungary differed from the nominal form mainly in the relative hindlimb length. However, substantial overlap in overall shape differences resulted in a relatively low percentage of correct classifications to the respective geographic groups in the Discriminant Analysis. Large intrapopulation variation in the values of the two ratios (SVL/TL and TL/IMT) formerly used for differentiate the two European subspecies of R. arvalis makes them unreliable characters for distinguishing the two forms. The pattern of morphometric variation does not match the overall genetic divergence of the R. arvalis populations, which suggests that the body shape differences in this species result from the phenotypic plasticity correlated with local climatic factors.     

Nuclear genetic markers indicate Danish origin of the Norwegian beech (Fagus sylvatica L.) populations established in 500�C1,000?AD

The northernmost range of beech (Fagus sylvatica L.) is in southern Norway and consists of two distinct and isolated distributions, a single population at Seim in West Norway and several adjacent populations in Vestfold, East Norway. The modest beech pollen deposits beyond these main distributions suggest that the Norwegian beech distribution has never been an extension of the south Scandinavian range. We used genetic markers and historical sources to trace the ancestor populations for the beech at Seim and Vestfold, hypothesising Denmark as the most likely source. Nuclear inter-simple sequence repeat markers, amplified by polymerase chain reaction (PCR), were applied to estimate genetic distances between beech populations in Norway, England and Denmark. The variation in chloroplast DNA polymorphism was estimated using PCR-restriction fragment length polymorphism. The nuclear genetic data indicate Denmark as a source for the beech in Norway, although the data are less certain in the case of Seim than in that of Vestfold. The populations from South England were genetically different from most Scandinavian populations. The genetic variation within Norwegian populations was only slightly lower than that of the English and Danish populations, questioning birds as vectors for dispersal. Thus, the pollen data and our results are in accordance with the intentional introduction and documented human migrations across Skagerrak before and during the Viking Age.     

Scaling up from epidemiology to biogeography: local infection patterns predict geographical distribution in fish parasites

Aim We investigated how the spatial distribution of parasites, measured as either their geographical range size or their frequency of occurrence among localities, relates to either their average local abundance or the variance in their abundance among localities where they occur. Location We used data on the abundance of 46 metazoan parasite species in 66 populations of threespine sticklebacks, Gasterosteus aculeatus, from Europe and North America. Methods For each parasite species, frequency of occurrence was calculated as the proportion of stickleback populations in which it occurred, and geographical range size as the area within the smallest possible polygon delimited using the coordinates of the localities where it occurred. Generalized linear models were used to assess how these two measures of spatial distribution were influenced by several predictor variables: geographical region (North America or Europe), life cycle (simple or complex), average local abundance, the coefficient of variation in abundance across localities, and median prevalence (proportion of infected hosts within a locality). Results Our analyses uncovered four patterns. First, parasites in North America tend to have higher frequencies of occurrence among surveyed localities, but not broader geographical ranges, than those in Europe. Second, parasite species with simple life cycles have wider geographical ranges than those with complex cycles. Third, there was a positive relationship between average abundance of the different parasite species and their frequency of occurrence, but not between average abundance and geographical range size. Fourth, the coefficient of variation in abundance covaried positively with both the frequency of occurrence and geographical range size across the different parasite species. Thus, all else being equal, parasites showing greater site‐to‐site variability in abundance occur in a greater proportion of localities and over a broader geographical area than those with a more stable abundance among sites. Main conclusions Local infection patterns are linked with large‐scale distributional patterns in fish parasites, independently of host effects, such that local commonness translates into regional commonness. The mechanisms linking parasite success at both scales remain unclear, but may include those that maintain the continuum between specialist and generalist parasites. Regardless, the observed patterns have implications for the predicted changes in the geographical distributions of many parasites in response to climate change.     

Large and unexpected variation in expression and substrate specificity of allelic glycosyltransferase gens on three loci in Silene dioica

A pilot study with regard to the distribution and expression of isovitexin glycosylation genes in Silene dioica showed that in this species, like in S. pratensis, a large variation in expression and substrate specificty exists for these genes. From S. dioica populations from Finland, Poland, Czechoslovakia, W. Germany, France and England a single plant was tested for allelic variation on the isovitexin glycosylation loci g, f and gl. In the English population a new allele of the g-locus, gA, with a different substrate specificity was identified. In the plant taken from the population in Poland a new allele glR1 was identified which was hypostatic to the allele glR known from S. pratensis. In Germany, a new allele gX1 was identified which is hypostatic over gX from S. pratensis. With regard to the isovitexin-glycoside phenotype the populations from Finland, Poland, Czechoslovakia, and England were homogeneous, whereas the populations of W. Germany and France showed variation. The mechanisms leading to this variation and their significance were discussed.     

Aspects of the population genetics of Mytilus (Mytilidae; Mollusca) in the British Isles

Starch gel electrophoresis was used to study variation at 11 loci in mussels sampled mainly from British coastal sites. Two types of mussel were identified, Mytilus edulis, the common mussel and its southern relative Mytilus galloprovincialis. Several partially diagnostic loci were used to map the distribution of the two forms. Mytilus edulis was present at all sites sampled in Britain and Ireland but was at low frequency in SW England; M. galloprovincialis was detected in SW England, the south and west of Ireland. Scotland and NE England, but was absent from south Wales, the Irish sea coasts of Wales and Ireland, and SE England. Apart from the occurrence of M. galloprovincialis in NE England, this distribution conforms with the results of studies using morphological characters and parallels the distribution of many other southern species in Britain. At the microgeographical level, M. edulis was found to prefer more sheltered and estuarine conditions than M. galloprovincialis. Analysis using the best diagnostic loci showed that hybridization is occurring between M. edulis and M. galloprovincialis at all localities where they occur sympatrically but that the extent of hybridization varies considerably between localities. The distribution of localities having high proportions of hybrid individuals is best interpreted by assuming that hybrids have higher fitness than parental types at these localities. A study was made of variation within and between those localities where only M. edulis individuals were observed. Little significant geographic variation in allele frequency was detected, but significant deficits of heterozygotes compared with Hardy-Weinberg expectations were seen for most loci. Analysis suggests that the Wahlund effect is not involved and that the most likely cause of the deficit is low frequencies of null alleles. In M. edulis no differences in phenotypic variance in shell height and width were observed between samples of multiply heterozygous and multiply homozygous individuals and no genetic differences were found between juveniles and adults. Overall little evidence was found that balancing selection is responsible for maintenance of the polymorphisms studied in M. edulis. The pattern of geographic variation in gene frequencies in Mytilus in the British Isles is discussed in relation to variation in the south and north of Europe and North America. It is concluded that steep clines in gene frequencies in M. edulis observed by other workers in the Baltic and in Long Island Sound cannot be attributed to the presence of M. galloprovincialis.