Allozyme variability in brown trout Salmo trutta in Chile

1. Brown trout ova were imported during the last century from different locations in Europe to establish populations in Chilean rivers (South America). The rivers are currently occupied by naturalized populations that have adapted to very different environmental conditions, such as areas of semi-desert in the north, or rainy and cold areas in the south.
2. In this first study in this geographical area, electrophoretic variability of proteins encoded by twenty-five loci was screened in seven populations from northern to southern Chile.
3. The results show significant heterogeneity of allelic frequencies between populations in seven of eleven polymorphic loci detected. The estimated value of genetic diversity 0.1274 ( H _T) is higher than that observed in populations from areas of natural distribution of this species. However, only 12.64% of this genetic diversity was found between samples ( G _ST), indicating a low genetic divergence among Chilean populations. The observed associations among the Chilean and 'modern' group of European populations suggests the probable origin of the new populations.     

The voyage of an invasive species across continents: genetic diversity of North American and European Colorado potato beetle populations

The paradox of successful invading species is that they are likely to be genetically depauperate compared to their source population. This study on Colorado potato beetles is one of the few studies of the genetic consequences of continent-scale invasion in an insect pest. Understanding gene flow, population structure and the potential for rapid evolution in native and invasive populations offers insights both into the dynamics of small populations that become successful invaders and for their management as pests. We used this approach to investigate the invasion of the Colorado potato beetle (Leptinotarsa decemlineata) from North America to Europe. The beetles invaded Europe at the beginning of the 20th century and expanded almost throughout the continent in about 30 years. From the analysis of mitochondrial DNA (mtDNA) and amplified fragment length polymorphism (AFLP) markers, we found the highest genetic diversity in beetle populations from the central United States. The European populations clearly contained only a fraction of the genetic variability observed in North American populations. European populations show a significant reduction at nuclear markers (AFLPs) and are fixed for one mitochondrial haplotype, suggesting a single successful founder event. Despite the high vagility of the species and the reduction of genetic diversity in Europe, we found a similar, high level of population structure and low gene flow among populations on both continents. Founder events during range expansion, agricultural management with crop rotation, and selection due to insecticide applications are most likely the causes partitioning genetic diversity in this species.     

危害茄科蔬菜的外来入侵害虫——马铃薯甲虫

马铃薯甲虫是世界著名的入侵害虫,是我国的重要检疫性有害生物,主要危害马铃薯、茄子和西红柿等茄科蔬菜。该甲虫目前分布于美洲、欧洲和亚洲等30多个国家和地区,于1993年入侵我国新疆,到2010年扩散到36个县市,最前沿为木垒县。该文介绍了马铃薯甲虫的扩散历史、危害情况、发生规律以及防治措施,指出我国在西北方向遭受新疆马铃薯甲虫入侵内地威胁的同时,也在东北方向遭受俄罗斯滨海边区马铃薯甲虫对黑龙江及东北地区入侵的严重威胁,应加强防范。     

A few north Appalachian populations are the source of European black locust

The role of evolution in biological invasion studies is often overlooked. In order to evaluate the evolutionary mechanisms behind invasiveness, it is crucial to identify the source populations of the introduction. Studies in population genetics were carried out on Robinia pseudoacacia L., a North American tree which is now one of the worst invasive tree species in Europe. We realized large‐scale sampling in both the invasive and native ranges: 63 populations were sampled and 818 individuals were genotyped using 113 SNPs. We identified clonal genotypes in each population and analyzed between and within range population structure, and then, we compared genetic diversity between ranges, enlarging the number of SNPs to mitigate the ascertainment bias. First, we demonstrated that European black locust was introduced from just a limited number of populations located in the Appalachian Mountains, which is in agreement with the historical documents briefly reviewed in this study. Within America, population structure reflected the effects of long‐term processes, whereas in Europe it was largely impacted by human activities. Second, we showed that there is a genetic bottleneck between the ranges with a decrease in allelic richness and total number of alleles in Europe. Lastly, we found more clonality within European populations. Black locust became invasive in Europe despite being introduced from a reduced part of its native distribution. Our results suggest that human activity, such as breeding programs in Europe and the seed trade throughout the introduced range, had a major role in promoting invasion; therefore, the introduction of the missing American genetic cluster to Europe should be avoided.     

Microsatellite DNA analysis of northern pike (Esox lucius L.) populations: insights into the genetic structure and demographic history of a genetically depauperate species

The northern pike Esox lucius L. is a freshwater fish exhibiting pronounced population subdivision and low genetic variability. However, there is limited knowledge on phylogeographical patterns within the species, and it is not known whether the low genetic variability reflects primarily current low effective population sizes or historical bottlenecks. We analysed six microsatellite loci in ten populations from Europe and North America. Genetic variation was low, with the average number of alleles within populations ranging from 2.3 to 4.0 per locus. Genetic differentiation among populations was high (overall θ_ST = 0.51; overall ρ_ST = 0.50). Multidimensional scaling analysis of genetic distances between populations and spatial analysis of molecular variance suggested a single phylogeographical race within the sampled populations from northern Europe, whereas North American and southern European populations were highly distinct. A population from Ireland was monomorphic at all loci, presumably reflecting founder events associated with introduction of the species to the island in the sixteenth century. Bayesian analysis of demographic parameters showed differences in θ (a product of effective population size and mutation rate) among populations from large and small water bodies, but the relative differences in θ were smaller than expected, which could reflect population subdivision within the larger water bodies. Finally, the analyses showed drastic population declines on a time scale of several thousand years within European populations, which we ascribe to either glacial bottlenecks or postglacial founder events.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 91–101.     

Integrating stable isotopes,parasite, and ring‐reencounter data to quantify migratory connectivity—A case study with Barn Swallows breeding in Switzerland,Germany, Sweden,and Finland

Ecosystems around the world are connected by seasonal migration. The migrant animals themselves are influenced by migratory connectivity through effects on the individual and the population level. Measuring migratory connectivity is notoriously difficult due to the simple requirement of data conveying information about the nonbreeding distribution of many individuals from several breeding populations. Explicit integration of data derived from different methods increases the precision and the reliability of parameter estimates. We combine ring‐reencounter, stable isotope, and blood parasite data of Barn Swallows Hirundo rustica in a single integrated model to estimate migratory connectivity for three large scale breeding populations across a latitudinal gradient from Central Europe to Scandinavia. To this end, we integrated a non‐Markovian multistate mark‐recovery model for the ring‐reencounter data with normal and binomial mixture models for the stable isotope and parasite data. The integration of different data sources within a mark‐recapture modeling framework enables the most precise quantification of migratory connectivity on the given broad spatial scale. The results show that northern‐breeding populations and Southern Africa as well as southern‐breeding populations and Western–Central Africa are more strongly connected through Barn Swallow migration than central European breeding populations with any of the African wintering areas. The nonbreeding distribution of Barn Swallows from central European breeding populations seems to be a mixture of those populations breeding further north and south, indicating a migratory divide.     

Douglas‐fir plantations in Europe: a retrospective test of assisted migration to address climate change

We evaluate genetic test plantations of North American Douglas‐fir provenances in Europe to quantify how tree populations respond when subjected to climate regime shifts, and we examined whether bioclimate envelope models developed for North America to guide assisted migration under climate change can retrospectively predict the success of these provenance transfers to Europe. The meta‐analysis is based on long‐term growth data of 2800 provenances transferred to 120 European test sites. The model was generally well suited to predict the best performing provenances along north–south gradients in Western Europe, but failed to predict superior performance of coastal North American populations under continental climate conditions in Eastern Europe. However, model projections appear appropriate when considering additional information regarding adaptation of Douglas‐fir provenances to withstand frost and drought, even though the model partially fails in a validation against growth traits alone. We conclude by applying the partially validated model to climate change scenarios for Europe, demonstrating that climate trends observed over the last three decades warrant changes to current use of Douglas‐fir provenances in plantation forestry throughout Western and Central Europe.     

Invasion history of North American Canada thistle,Cirsium arvense

Aim Canada thistle (Cirsium arvense– Cardueae, Asteraceae) is one of the worst invasive plants world‐wide. Native to Eurasia, its unintentional introduction into North America now threatens the native flora and is responsible for enormous agricultural losses. The goals of this study are to: (1) reconstruct the evolutionary history of C. arvense and estimate how often it may have colonized North America, (2) compare the genetic diversity between European and North American populations to detect signs of demographic bottlenecks and/or patterns of population admixture, and (3) conduct bioclimatic comparisons to infer eventual niche shifts following this species’ introduction into North America. Location Europe and North America. Methods A total of 1522 individuals from 58 populations were investigated with six microsatellite markers. Estimates of heterozygosity (H_E) and allelic richness (R_S) were quantified for each population, and population structure was inferred via analyses of molecular variance (AMOVAs), principal components analyses (PCAs), Mantel tests and Bayesian clustering analyses. Climatic niche spaces were based on 19 bioclimatic variables extracted from approximately 32,000 locations covering the entire range, and compared using PCA and hierarchical cluster analysis. Results Although there is evidence of multiple introductions from divergent European lineages, North American populations of C. arvense exhibited significantly lower levels of genetic diversity than their putative ancestors. Bioclimatic comparisons pointed to a high degree of niche conservatism during invasion, but indicated that genotypes from the former USSR and Central European mountain chains were probably best adapted to invade North America upon entry into the continent. Main conclusions Genetic and historical data suggest that C. arvense first entered North America from Western Europe with the first European settlers, and was later introduced from Eastern Europe into the prairie states during the agricultural boom. The species went through a significant bottleneck following its introduction into the New World, but the level of genetic diversity remained high owing to admixture between genetically differentiated lineages and to a highly efficient outcrossing breeding system.     

Genetic variability in European black grouse (<Emphasis Type="Italic">Tetrao tetrix</Emphasis>)

We studied microsatellite genetic variation in 14 different geographic populations of black grouse (Tetrao tetrix) across the European range. Populations were grouped in three different fragmentation categories: isolated, contiguous and continuous, respectively. Genetic diversity, measured as observed heterozygosity (H _O), expected heterozygosity (H _E) and allelic richness, were lower in isolated populations as compared to the other two categories that did not differ amongst one another. These results imply that lowered genetic variability in black grouse populations is negatively affected by population isolation. Our results suggest that the connectivity of small and isolated populations in Western Europe should be improved or else these face an increased risk of extinction due to genetic and demographic stochasticity.     

Molecular and pathobiological characterization of 61 Potato mop‐top virus full‐length cDNAs reveals great variability of the virus in the centre of potato domestication,novel genotypes and evidence for recombination

The evolutionary divergence of Potato mop‐top virus (PMTV), a tri‐partite, single‐stranded RNA virus, is exceptionally low, based on the analysis of sequences obtained from isolates from Europe, Asia and North America. In general, RNA viruses exist as dynamic populations of closely related and recombinant genomes that are subjected to continuous genetic variation. The reason behind the low genetic variation of PMTV remains unclear. The question remains as to whether the low variability is a shared property of all PMTV isolates or is a result of the limited number of isolates characterized so far. We hypothesized that higher divergence of the virus might exist in the Andean regions of South America, the centre of potato domestication. Here, we report high variability of PMTV isolates collected from 12 fields in three locations in the Andean region of Peru. To evaluate PMTV genetic variation in Peru, we generated full‐length cDNA clones, which allowed reliable comparative molecular and pathobiological characterization of individual isolates. We found significant divergence of the CP‐RT and 8K sequences. The 8K cistron, which encodes a viral suppressor of RNA silencing, was found to be under diversifying selection. Phylogenetic analysis determined that, based on the CP‐RT sequence, all PMTV isolates could be categorized into three separate lineages (clades). Moreover, we found evidence for recombination between two clades. Using infectious cDNA clones of the representatives of these two clades, as well as reassortants for the RNA‐CP genomic component, we determined the pathobiological differences between the lineages, which we coined as S (for severe) and M (for mild) types. Interestingly, all isolates characterized previously (from Europe, Asia and North America) fall into the S‐type clade, whereas most of the Peruvian isolates belong to the M‐type. Taken together, our results support the notion of the single introduction of PMTV from the centre of potato origin to Europe, and subsequent spread of the S‐type into Asia and USA. This is also supported by the suggested novel classification of isolates based on genetic constellations.     

Population Structure and the Colonization Route of One of the Oldest North American Invasive Insects: Stories from the Worn Road of the Hessian Fly,Mayetiola destructor (Say)

An integral part to understanding the biology of an invasive species is determining its origin, particularly in pest species. As one of the oldest known invasive species, the goals of this study were to evaluate the evidence of a westward expansion of Hessian fly into North America, from a potential singular introduction event, and the population genetic structure of current populations. Levels of genetic diversity and population structure in the Hessian fly were compared across North America, Europe, North Africa, Western Asia, and New Zealand. Furthermore, Old World populations were evaluated as possible sources of introduction. We tested diversity and population structure by examining 18 microsatellite loci with coverage across all four Hessian fly chromosomes. Neither genetic diversity nor population genetic structure provided evidence of a westward movement from a single introduction in North America. Introduced populations in North America did not show identity or assignment to any Old World population, likely indicating a multiple introduction scenario with subsequent gene flow between populations. Diversity and selection were assessed on a chromosomal level, with no differences in diversity or selection between chromosomes or between native and introduced populations.     

Past and current gene flow in the selfing, wind-dispersed species Mycelis muralis in western Europe

The distribution of genetic diversity in Mycelis muralis, or wall lettuce, was investigated at a European scale using 12 microsatellite markers to infer historical and contemporary forces from genetic patterns. Mycelis muralis has the potential for long-distance seed dispersal by wind, is mainly self-pollinated, and has patchily distributed populations, some of which may show metapopulation dynamics. A total of 359 individuals were sampled from 17 populations located in three regions, designated southern Europe (Spain and France), the Netherlands, and Sweden. At this within-region scale, contemporary evolutionary forces (selfing and metapopulation dynamics) are responsible for high differentiation between populations (0.34 < F(ST) < 0.60) but, contrary to expectation, levels of within-population diversity, estimated by Nei's unbiased expected heterozygosity (H(E)) (0.24 < H(E) < 0.68) or analyses of molecular variance (50% of the variation found within-populations), were not low. We suggest that the latter results, which are unusual in selfing species, arise from efficient seed dispersal that counteracts population turnover and thus maintains genetic diversity within populations. At the European scale, northern regions showed lower allelic richness (A = 2.38) than populations from southern Europe (A = 3.34). In light of postglacial colonization hypotheses, these results suggest that rare alleles may have been lost during recolonization northwards. Our results further suggest that mutation has contributed to genetic differentiation between southern and northern Europe, and that Sweden may have been colonized by dispersers originating from at least two different refugia.     

Geographical origin of an introduced pest species, Delia radicum (Diptera: anthomyiidae), determined by RAPD analysis and egg micromorphology

The origin of introduction of the cabbage root fly, Delia radicum Linnaeus to the north-eastern coast of North America in the 19th century has been assumed to be from Europe. From that point of introduction, D. radicum gradually spread westward to occupy available ecological niches. DNA fingerprinting and egg micromorphology were used to determine the most likely geographical origin of the North American populations of this species. Forty-five informative RAPD loci obtained from ten primers and three criteria for egg micromorphology were studied. These characters indicated a common origin for the North American populations and a high similarity between populations from North America and north-western Europe. The results suggest a single entrance point of D. radicum into North America, probably via the north-eastern coast (New York area) from north-western Europe. The implications of this study in assisting selection of natural enemies of this important agricultural pest are discussed.     

RFLP diversity and relationships among traditional European maize populations

Given the large extent of hybrid cultivation, the importance of conserving the diversity of crop genetic resources has given birth to numerous collections of old races. In the present paper, we conduct a molecular characterisation of a large collection of 488 European maize populations using the bulk RFLP analysis. The analysis of 23 RFLP loci showed a high allelic richness of 11.5 alleles per locus. Populations from eastern Europe (Poland, Austria, Germany, etc.) showed the lowest genetic diversity, a lower number of unique alleles and a higher percentage of fixed loci than populations from southern Europe. In fact, genetic diversity appeared higher in Southern regions where the first maize populations are thought to have been introduced. Molecular classification based on Rogers' distance (i.e. alleles frequencies) allowed us to distinguish three main clusters which were highly consistent with geographic origins. A Northeastern cluster grouped together early or intermediate populations from Northeastern countries and the Balkans, a southeastern cluster joined late and partially dent populations from Greece and Italy, and, a southwestern cluster was made up of early flint populations from northern Spain, Portugal and the Pyrenees. A correlation between allelic frequencies at some loci and latitude and/or longitude was observed. Such tendencies may reflect the direction of gene flow between different races of maize: for instance, North American (Northern flint) and Caribbean populations were introduced, respectively, to northern and southern Europe, in the past.     

Warming increases plant biomass and reduces diversity across continents,latitudes, and species migration scenarios in experimental wetland communities

Atmospheric warming may influence plant productivity and diversity and induce poleward migration of species, altering communities across latitudes. Complicating the picture is that communities from different continents deviate in evolutionary histories, which may modify responses to warming and migration. We used experimental wetland plant communities grown from seed banks as model systems to determine whether effects of warming on biomass production and species richness are consistent across continents, latitudes, and migration scenarios. We collected soil samples from each of three tidal freshwater marshes in estuaries at three latitudes (north, middle, south) on the Atlantic coasts of Europe and North America. In one experiment, we exposed soil seed bank communities from each latitude and continent to ambient and elevated (+2.8 °C) temperatures in the greenhouse. In a second experiment, soil samples were mixed either within each estuary (limited migration) or among estuaries from different latitudes in each continent (complete migration). Seed bank communities of these migration scenarios were also exposed to ambient and elevated temperatures and contrasted with a no‐migration treatment. In the first experiment, warming overall increased biomass (+16%) and decreased species richness (?14%) across latitudes in Europe and North America. Species richness and evenness of south‐latitude communities were less affected by warming than those of middle and north latitudes. In the second experiment, warming also stimulated biomass and lowered species richness. In addition, complete migration led to increased species richness (+60% in North America, + 100% in Europe), but this higher diversity did not translate into increased biomass. Species responded idiosyncratically to warming, but Lythrum salicaria and Bidens sp. increased significantly in response to warming in both continents. These results reveal for the first time consistent impacts of warming on biomass and species richness for temperate wetland plant communities across continents, latitudes, and migration scenarios.     

Late glacial history of the cold-adapted freshwater fish Cottus gobio,revealed by microsatellites

The distribution of genetic diversity at 10 highly polymorphic microsatellite loci within the European freshwater fish, Cottus gobio, L. was examined. The sampling range comprised a large geographical scale including lineages known to be highly divergent at both mitochondrial DNA (mtDNA) and allozymes. An analysis of genetic variability within populations showed that expected heterozygosity and allelic richness could be explained largely by current effective population sizes. Evidence was found, however, that historical processes predating the last major glaciation affected allelic richness. In addition to confirming the large-scale patterns from earlier studies, the microsatellite data revealed new insights into recent processes by analysing genetic structure within ancient lineages defined by mtDNA data. Stepwise mutation model (SMM) and nonSMM-based methods demonstrated a clear genetic structuring within the Northwestern European lineage comprising populations from Britain and Belgium, and within the Central European lineage populations from the rivers Danube, Elbe and Main. Supported by an analysis of genetic variability within populations these results showed that the bullhead populations most probably persisted throughout the last major glaciation within the British Isles and within the drainages of the rivers Elbe and Main. Such observations provide the first genetic evidence for a glacial refugium in such close proximity to the European glacial margins.     

Genetic diversity and population structure identify the potential source of the invasive red clover casebearer moth, <Emphasis Type="Italic">Coleophora deauratella</Emphasis>, in North America

The red clover casebearer, Coleophora deauratella, is an invasive pest of red clover grown for seed in North America. In 2006, an outbreak in Alberta, Canada was discovered that resulted in significant seed losses, while further invasion threatens the world’s largest red clover forage seed production region in Oregon, USA. Prior to the recent outbreak, C. deauratella was thought to be restricted to eastern North America in its invasive range. We sequenced a 615-bp fragment of the mitochondrial cytochrome c oxidase subunit 1 gene, and developed three microsatellite markers to assess the genetic diversity and population structure of C. deauratella in North America and its native range in Europe. We observed signatures of a founder effect in North American populations and a further loss of genetic diversity within Alberta populations. Most genetic differentiation was found between continents, with no evidence of isolation-by-distance within each continent. From the limited number of European populations sampled, a single introduction from Switzerland is the most probable source of North American populations based on similar mitochondrial diversity and lack of population differentiation. Within North America, based on increased genetic diversity compared to the rest of the continent, the first North American record from Ithaca, NY, and the first documented outbreak in southern Ontario in 1989, the initial C. deauratella invasion most likely occurred in southern Ontario, Canada or adjacent states in the USA, followed by transport throughout the continent. This study provides insight into the phylogeographic history of C. deauratella in North America and Europe and may help to identify a regional source of future classical biological control agents.     

Historical and present distribution of coyote (Canis latrans) in Mexico and Central America

Aim Coyote (Canis latrans) distribution in Mexico and Central America has expanded recently reaching the Yucatan peninsula, Belize and Panama, probably promoted by deforestation of tropical areas. Historically, the southern distribution of coyotes prior to European settlement in America was described as reaching only as far south as central Mexico and that introduction of livestock favoured migration of coyotes to southern Mexico and Central America. However, coyote fossil records in Central America and Yucatan, as well as observational records of travellers during the sixteenth century suggest that the coyote's arrival to the region was earlier. Because of the uncertainty of past coyote distribution and the possible economic and ecological impacts due to recent range expansion, the objectives of this study were to confirm if paleontological and historical evidence support the hypothesis that the southernmost limit of coyote distribution before the arrival of European settlers was the centre of Mexico, to discuss the possible factors that have influenced historical shifts in coyote distribution, and to model the present distribution of the coyote in Mexico and Central America, determining the areas where they could invade in the near future. Location The research area comprises continental Mexico and the Central American Isthmus countries: Guatemala, Belize, El Salvador, Honduras, Nicaragua, Costa Rica and Panama. Methods The historical distribution (Pleistocene–Early Holocene, Pre‐Columbian, sixteenth to nineteenth centuries and twentieth century) was established from coyote records obtained from museum collections and specialized literature. Present coyote distribution for Mexico and Central America was modelled using the Genetic Algorithms for Rule‐set Prediction (GARP). Results Historical coyote records show that this species was distributed in southern Mexico and Central America during the Pleistocene–Early Holocene, the Pre‐Columbian period, and during the arrival of Europeans in the sixteenth century. Coyote records indicate a continuous range expansion during the twentieth century. Historical advance and regression of tropical forests in southern Mexico and Central America produced by natural and human events such as climatic changes and variation in human densities could help us understand the historical coyote distribution. The modelled present‐day coyote distribution included the north of Belize, the north of Panama, the north of the Yucatan Peninsula and a corridor on the Gulf costal plain of Campeche in Mexico. Also, the model predicted a region north of the Darien in southern Panama as appropriate for the presence of coyotes, although they have not been detected there so far. Main conclusion Coyote records in southern Mexico and Central America during the Pleistocene–Early Holocene, the Pre‐Columbian period, and early arrival of European settlers to the area indicated that coyotes were probably already present there and did not recently disperse from the north of Mexico to the south due to livestock introduction.     

Significance of caribou (Rangifer tarandus) ecotypes from a molecular genetics viewpoint

Three caribou ecotypes are present in easternNorth America: the mountain caribou which isfound south of the St. Lawrence River, thebarren-ground caribou which calves in thetundra, and in between, the forest-dwellingecotype which lives all year long in the borealforest. Blood and muscle samples were collectedfrom seven populations and characterized ateight microsatellite loci to test thehypotheses that forest-dwelling andbarren-ground ecotypes constitute a singlemetapopulation and that geographical isolationresults in reduced genetic diversity. The meannumber of alleles per locus, allelic richness,and observed and expected heterozygositydeclined from north to south and were thesmallest in isolated forest-dwellingpopulations. Correspondence analysis showedthree groups of samples corresponding to thethree ecotypes. Gene flow estimates weremoderate or high among all forest-dwellingpopulations and particularly between those <200 km apart. Our results suggest that thethree caribou ecotypes represent three distinctgenetic entities and that the forest-dwellingpopulations in the continuous range form ametapopulation. Genetic diversity was lower inisolated populations but does not seem to be ofimmediate concern for conservation. We proposethat management strategies should favorincrease in caribou numbers in order to avoidextinction due to stochastic events and tomaintain local biodiversity. In the continuousrange, conservation strategies of cariboupopulations must be planned on a large scale tomaintain occasional exchanges betweenpopulations, thus preserving genetic diversity.     

Some observations on shell-shape variation in North American populations of Nucella lapillus (L.)

This paper considers shell-shape variation in samples of the dog-whelk/dog-winkle, ucella lapillus , from the southern half of its North American range, between 41 and 46N, and compares it with the pattern seen in Europe. At the extreme southern limit of its American range, to the south of Cape Cod, the species exhibits features to be expected in an animal close to an environmental limit. It is generally rare and has a patchy distribution. There is little variation in the shell: almost all adult individuals have rather large, thick, white, elongated shells. This contrasts sharply with the situation in Portugal, at the southern limit in Europe, where the animals have small thin, coloured shells of intermediate shape.
Away from the marginal situation, north of Nahant (42^CN) in America, the species shows much the same range of variability in shell size, shape and colour on both sides of the Atlantic. Most populations appear to show the same pattern of shape variation in asociation with the exposure of their habitat to wave action. Animals from exposed sites tend to have shorter, squatter shells than their compatriots in shelter. However, in America as in some parts of Europe, there are occasional enclaves which do not fit the usual pattern. It is interesting to note that the form normally associated with extremely exposed shores from Brittany to Faroe in Europe is found in Atlantic Canada but is apparently absent from southern populations in both the U.S.A and the Iberian Peninsula.