Cryptic diversity of Melanochlamys sea slugs (Gastropoda,Aglajidae) in the North Pacific

North Pacific specimens of Melanochlamys sea slugs were examined morphologically (including the male reproductive morphology, shell and external coloration) and were sequenced for three genes (mitochondrial COI and 16S and nuclear H3). Phylogenetic and species delimitation analyses were used to determine the evolutionary relationships and species diversity among the specimens examined. Both molecular and morphological data revealed an unexpected high level of cryptic diversity. At least four distinct species occur on the Northwestern Pacific. Melanochlamys ezoensis occurs in Russia and temperate and cold areas in Japan. Three additional undescribed species occur in Japan and/or South Korea. One of the undescribed species occurs both in South Korea and in Japan, but only in Tokyo Bay, suggesting that it could be non‐native in Japan. Two distinct species occur on the Northeastern Pacific coast; Melanochlamys diomedea is widespread from Southern California to Alaska, whereas M. ezoensis was found only in San Francisco Bay, suggesting a human‐mediated introduction. This is further supported by the absence of records of M. ezoensis in San Francisco prior to 2001. The species diversity of Melanochlamys in the Northwestern Pacific is much greater than in the Northeastern Pacific; it is hypothesized that differences in geographic and ocean current system complexity might account for different responses to glacial extinction and postglacial expansion.     

Geographic range and structure of cryptic genetic diversity among Pacific North American populations of the non-native amphipod Grandidierella japonica

Reconstructing the invasion history of aquatic invasive species can enhance understanding of invasion risks by recognizing areas most susceptible to invasion and forecasting future spread based on past patterns of population expansion. Here we reconstruct the invasion history of the Japanese amphipod Grandidierella japonica Stephensen 1938 combining information from historical collection data with molecular genetic data to better understand post-invasion range expansion and anthropogenic connectivity across the Pacific coast of North America. Compilation of collection data from bays and estuaries of the Pacific North American coast show many new localities have been colonized in the last two decades, moving outward from harbors and bays with high commercial traffic into smaller coastal locations dominated by local recreational traffic. DNA barcode sequence data for G. japonica reveals two distinct clades: one found in San Francisco Bay and sites to the north, and one also found in San Francisco Bay and sites to the south. The two clades differ by an average 7.28 % genetic distance, large enough to consider these invasive amphipods two separate species. Both northern and southern clades exhibit low levels of genetic diversity, suggesting a single introduction event for each. The presence of cryptic diversity within this invasive amphipod highlights the need for more extensive study of the invasive and native populations of aquatic invasive invertebrates to address questions of taxonomy, diversity, and invasion history.     

Global distribution of cryptic native,introduced and hybrid lineages in the widespread estuarine amphipod Ampithoe valida

Biological invasions can pose a severe threat to coastal ecosystems, but are difficult to track due to inaccurate species identifications and cryptic diversity. Here, we clarified the cryptic diversity and introduction history of the marine amphipod Ampithoe valida by sequencing a mtDNA locus from 683 individuals and genotyping 10,295 single-nucleotide polymorphisms (SNPs) for 349 individuals from Japan, North America and Argentina. The species complex consists of three cryptic lineages: two native Pacific and one native Atlantic mitochondrial lineage. It is likely that the complex originated in the North Pacific and dispersed to the north Atlantic via a trans-arctic exchange approximately 3 MYA. Non-native A. valida in Argentina have both Atlantic mitochondrial and nuclear genotypes, strongly indicating an introduction from eastern North America. In two eastern Pacific estuaries, San Francisco Bay and Humboldt Bay, California, genetic data indicate human-mediated hybridization of Atlantic and Pacific sources, and possible adaptive introgression of mitochondrial loci, nuclear loci, or both. The San Francisco Bay hybrid population periodically undergoes population outbreaks and profoundly damages eelgrass Zostera marina thalli via direct consumption, and these ecological impacts have not been documented elsewhere. We speculate that novel combinations of Atlantic and Pacific lineages could play a role in A. valida’s unique ecology in San Francisco Bay. Our results reinforce the notion that we can over-estimate the number of non-native invasions when there is cryptic native structure. Moreover, inference of demographic and evolutionary history from mitochondrial loci may be misleading without simultaneous survey of the nuclear genome.

     

Introduced or glacial relict? Phylogeography of the cryptogenic tunicate Molgula manhattensis (Ascidiacea,Pleurogona)

Aim The tunicate Molgula manhattensis has a disjunct amphi‐Atlantic distribution and a recent history of world‐wide introductions. Its distribution could be the result of regional extinctions followed by post‐glacial recolonization, or anthropogenic dispersal. To determine whether the North Atlantic distribution of M. manhattensis is natural or human‐mediated, we analysed mtDNA cytochrome c oxidase subunit I (COI) sequence variation in individuals from cryptogenic and introduced ranges. Location North Atlantic Europe and America; Black Sea; San Francisco Bay; Osaka Bay. Methods Nuclear 18S rDNA sequences were used to resolve phylogenetic relationships and mtDNA COI sequences for phylogeographic analyses. Results Phylogenetic analyses confirmed that M. manhattensis and M. socialis, which are frequently confused, are distinct species. MtDNA haplotype diversity was nearly three times higher with deeper relationships among haplotypes on the North‐east American coast than in Europe. Diversity declined from south to north in America but not in Europe. In areas of known introductions (Black Sea, Japan, San Francisco Bay), M. manhattensis showed variable levels of haplotype diversity. Medium‐to‐high‐frequency haplotypes originating from the North‐west Atlantic were present in two locations of known introductions, but not in Europe. Private haplotypes were found on both sides of the Atlantic and in introduced populations. The mismatch distribution for the North‐east Atlantic coast indicates a recent expansion. Main conclusions Molgula manhattensis is native in North‐east America. However, whether it was introduced or is native to Europe remains equivocal. Additional sampling might or might not reveal the presence of putative private European haplotypes in America. The low European diversity may be explained by low effective population size and a recent expansion, or by low propagule pressure of anthropogenic introduction. Absence of medium‐to‐high‐frequency American haplotypes in Europe may be the result of exclusive transport from southern ports, or long‐term residence. These arguments are ambiguous, and M. manhattensis remains cryptogenic in Europe.     

Global population genetic structure of the starlet anemone <Emphasis Type="Italic">Nematostella vectensis</Emphasis>: multiple introductions and implications for conservation policy

Distinguishing natural versus anthropogenic dispersal of organisms is essential for determining the native range of a species and implementing an effective conservation strategy. For cryptogenic species with limited historical records, molecular data can help to identify introductions. Nematostella vectensis is a small, burrowing estuarine sea anemone found in tidally restricted salt marsh pools. This species’ current distribution extends over three coast lines: (i) the Atlantic coast of North America from Nova Scotia to Georgia, (ii) the Pacific coast of North America from Washington to central California, and (iii) the southeast coast of England. The 1996 IUCN Red List designates N. vectensis as “vulnerable” in England. Amplified fragment length polymorphism (AFLP) fingerprinting of 516 individuals from 24 N. vectensis populations throughout its range and mtDNA sequencing of a subsample of these individuals strongly suggest that anthropogenic dispersal has played a significant role in its current distribution. Certain western Atlantic populations of N. vectensis exhibit greater genetic similarity to Pacific populations or English populations than to other western Atlantic populations. At the same time, F-statistics showing high degrees of genetic differentiation between geographically proximate populations support a low likelihood for natural dispersal between salt marshes. Furthermore, the western Atlantic harbors greater genetic diversity than either England or the eastern Pacific. Collectively, these data clearly imply that N. vectensis is native to the Atlantic coast of North America and that populations along the Pacific coast and in England are cases of successful introduction.     

Evidence for multiple introductions of Centaurea stoebe micranthos (spotted knapweed,Asteraceae) to North America

Invasive species’ success may depend strongly on the genetic resources they maintain through the invasion process. We ask how many introductions have occurred in the North American weed Centaurea stoebe micranthos (Asteraceae), and explore whether genetic diversity and population structure have changed as a result of introduction. We surveyed individuals from 15 European native range sites and 11 North American introduced range sites at six polymorphic microsatellite loci. No significant difference existed in the total number of alleles or in the number of private alleles found in each range. Shannon–Weaver diversity of phenotype frequencies was also not significantly different between the ranges, while expected heterozygosity was significantly higher in the invasive range. Population structure was similar between the native range and the invasive range, and isolation by distance was not significant in either range. Traditional assignment methods did not allocate any North American individuals to the sampled European populations, while Bayesian assignment methods grouped individuals into nine genetic clusters, with three of them shared between North America and Europe. Invasive individuals tended to have genetically admixed profiles, while natives tended to assign more strongly to a single cluster. Many North American individuals share assignment with Romania and Bulgaria, suggesting two separate invasions that have undergone gene flow in North America. Samples from three other invasive range sites were genetically distinct, possibly representing three other unique introductions. Multiple introductions and the maintenance of high genetic diversity through the introduction process may be partially responsible for the invasive success of C. stoebe micranthos.     

Genetic diversity of the black mangrove (Avicennia germinans L.) in Colombia

This study analyzed the genetic diversity and patterns of genetic structure in Colombian populations of Avicennia germinans L. using microsatellite loci. A lower genetic diversity was found on both the Caribbean (Ho = 0.439) and the Pacific coasts (Ho = 0.277) than reported for the same species in other locations of Central American Pacific, suggesting the deterioration of genetic diversity. All the populations showed high inbreeding coefficients (0.131–0.462) indicating heterozygotes deficience. The genetic structure between the Colombian coasts separated by Central American Isthmus was high (F_RT = 0.39) and the analyses of the genetic patterns of A. germinans revealed a clear differentiation of populations and no-recent gene flow evidence between coasts. Genetic structure was found within each coast (F_ST = 0.10 for the Caribbean coast and F_ST = 0.22 for the Pacific coast). The genetic patterns along the two coasts appear to reflect a forcing by local geomorphology and marine currents. Both coasts constitute a different Evolutionary Significant Unit, so we suggest for future transplantations plans that propagules or saplings of the populations of the Caribbean coast should not be mixed with those of the Pacific Colombian coast. Besides, we suggest that reforestation efforts should carefully distinguish propagules sources within each coast.     

Tracing the introduction history of the invasive swimming crab <Emphasis Type="Italic">Charybdis hellerii</Emphasis> (A. Milne-Edwards, 1867) in the Western Atlantic: evidences of high genetic diversity and multiple introductions

The swimming crab Charybdis hellerii is an invader with global distribution in warm waters. Native to the Indo-Pacific, this species invaded the eastern Mediterranean Sea after the Suez Canal opening. In 1987, it was first reported in the Western Atlantic, probably transported via ballast water of ships. Since then, it has been registered from many localities along the American coast from the USA to southern Brazil where it has rapidly established reproducing populations. Our main aim was to investigate the introduction history of this species along the American coast, using a phylogeographic approach. Additionally, we attempted to clarify the identity of this invasive species by molecular analyses and morphological assessment in order to provide a basis for our main investigation. C. hellerii was confirmed as a single species, but both cytochrome c oxidase subunit I and 16S rDNA revealed a genetic structure, splitting the potential source populations of American introductions into two groups: “western Indian Ocean and Mediterranean Sea” (WIO + MS) versus “eastern Indian + western Pacific oceans” (EIO + WPO). Most specimens from America clustered with the former group, supporting the hypothesis that the Mediterranean Sea represented the main source of Western Atlantic populations. However, the clustering of animals from southern Brazil with the latter group indicates that introductions from the eastern Indian or Pacific oceans must also have occurred. The existence of a third group, found exclusively within the American range and genetically related to EIO + WPO, also indicates an independent introduction from an unsampled locality from the native range. The haplotype and nucleotide diversities of American localities were comparable to those of source populations, contradicting a founder effect prediction. This finding might be related to the high propagule pressure associated with introductions via ballast water and the occurrence of multiple introductions from genetic distinct sources. The direct comparison of the haplotype numbers suggested no genetic bottleneck during introduction from the Mediterranean Sea, but a bottleneck might have occurred during introductions from the eastern Indian or Pacific oceans.     

Understanding invasion history: genetic structure and diversity of two globally invasive plants and implications for their management

Aim Resolving the origin of invasive plant species is important for understanding the introduction histories of successful invaders and aiding strategies aimed at their management. This study aimed to infer the number and origin(s) of introduction for the globally invasive species, Macfadyena unguis‐cati and Jatropha gossypiifolia using molecular data. Location Native range: Neotropics; Invaded range: North America, Africa, Europe, Asia, Pacific Islands and Australia. Methods We used chloroplast microsatellites (cpSSRs) to elucidate the origin(s) of introduced populations and calculated the genetic diversity in native and introduced regions. Results Strong genetic structure was found within the native range of M. unguis‐cati, but no genetic structuring was evident in the native range of J. gossypiifolia. Overall, 27 haplotypes were found in the native range of M. unguis‐cati. Only four haplotypes were found in the introduced range, with more than 96% of introduced specimens matching a haplotype from Paraguay. In contrast, 15 haplotypes were found in the introduced range of J. gossypiifolia, with all invasive populations, except New Caledonia, comprising multiple haplotypes. Main conclusions These data show that two invasive plant species from the same native range have had vastly different introduction histories in their non‐native ranges. Invasive populations of M. unguis‐cati probably came from a single or few independent introductions, whereas most invasive J. gossypiifolia populations arose from multiple introductions or alternatively from a representative sample of genetic diversity from a panmictic native range. As introduced M. unguis‐cati populations are dominated by a single haplotype, locally adapted natural enemies should make the best control agents. However, invasive populations of J. gossypiifolia are genetically diverse and the selection of bio‐control agents will be considerably more complex.     

High genetic diversity is not essential for successful introduction

Some introduced populations thrive and evolve despite the presumed loss of diversity at introduction. We aimed to quantify the amount of genetic diversity retained at introduction in species that have shown evidence of adaptation to their introduced environments. Samples were taken from native and introduced ranges of Arctotheca populifolia and Petrorhagia nanteuilii. Using microsatellite data, we identified the source for each introduction, estimated genetic diversity in native and introduced populations, and calculated the amount of diversity retained in introduced populations. These values were compared to those from a literature review of diversity in native, confamilial populations and to estimates of genetic diversity retained at introduction. Gene diversity in the native range of both species was significantly lower than for confamilials. We found that, on average, introduced populations showing evidence of adaptation to their new environments retained 81% of the genetic diversity from the native range. Introduced populations of P. nanteuilii had higher genetic diversity than found in the native source populations, whereas introduced populations of A. populifolia retained only 14% of its native diversity in one introduction and 1% in another. Our literature review has shown that most introductions demonstrating adaptive ability have lost diversity upon introduction. The two species studied here had exceptionally low native range genetic diversity. Further, the two introductions of A. populifolia represent the largest percentage loss of genetic diversity in a species showing evidence of substantial morphological change in the introduced range. While high genetic diversity may increase the likelihood of invasion success, the species examined here adapted to their new environments with very little neutral genetic diversity. This finding suggests that even introductions founded by small numbers of individuals have the potential to become invasive.     

Low genetic diversity contrasts with high phenotypic variability in heptaploid Spartina densiflora populations invading the Pacific coast of North America

Species can respond to environmental pressures through genetic and epigenetic changes and through phenotypic plasticity, but few studies have evaluated the relationships between genetic differentiation and phenotypic plasticity of plant species along changing environmental conditions throughout wide latitudinal ranges. We studied inter‐ and intrapopulation genetic diversity (using simple sequence repeats and chloroplast DNA sequencing) and inter‐ and intrapopulation phenotypic variability of 33 plant traits (using field and common‐garden measurements) for five populations of the invasive cordgrass Spartina densiflora Brongn. along the Pacific coast of North America from San Francisco Bay to Vancouver Island. Studied populations showed very low genetic diversity, high levels of phenotypic variability when growing in contrasted environments and high intrapopulation phenotypic variability for many plant traits. This intrapopulation phenotypic variability was especially high, irrespective of environmental conditions, for those traits showing also high phenotypic plasticity. Within‐population variation represented 84% of the total genetic variation coinciding with certain individual plants keeping consistent responses for three plant traits (chlorophyll b and carotenoid contents, and dead shoot biomass) in the field and in common‐garden conditions. These populations have most likely undergone genetic bottleneck since their introduction from South America; multiple introductions are unknown but possible as the population from Vancouver Island was the most recent and one of the most genetically diverse. S. densiflora appears as a species that would not be very affected itself by climate change and sea‐level rise as it can disperse, establish, and acclimate to contrasted environments along wide latitudinal ranges.     

Molecular diversity of species of the Elymus trachycaulus species complex and their relationships to non-North American taxa

Species of the E. trachycaulus complex species are known for their morphological variability, but little is known about their genetic basis. The delimitation of taxa within the complex has been controversial and difficult. E. trachycaulus is predominantly self-pollinating, and lacks clear morphological boundaries between it and E. alaskanus. Another controversial taxonomic issue of E. trachycaulus is the relationships of this complex species to non-North American E. caninus. The objectives of this study were to examine genetic diversity and the systematic relationships among the species of the E. trachycaulus complex and their relationships with E. caninus, E. alaskanus and E. mutabilis. Random amplified polymorphic DNA method was used to study 35 accessions of E. trachycaulus complex and other Elymus species. Higher genetic variation was detected within species of E. trachycaulus complex. Eurasian accessions are as variable as the North American ones. Both UPGMA and NJ analyses did not show clearly separation among species of the E. trachycaulus complex. No clear association between geographic origin and genetic grouping among these species was found. Eurasian E. trachycaulus probably originated from multiple North American populations.     

Genetic variation in photosynthetic characteristics among invasive and native populations of reed canarygrass (Phalaris arundinacea)

With the extensive spread of invasive species throughout North America and Europe there is an urgent need to better understand the morphological and physiological characteristics of successful invasive plants and the evolutionary mechanisms that allow introduced species to become invasive. Most ecological studies have focused on morphological differences and changes in community dynamics, and physiological studies have typically explored the differences between native and invasive species. In this study, 15 different genotypes of Phalaris arundinacea from both its native (European) and invasive (North American) range were grown in a common garden experiment to monitor the physiological differences between native and invasive genotypes. Here we present data that suggests high variability exists in the physiological traits among genotypes of P. arundinacea, yet genotypes from the native range are not necessarily physiologically inferior to the hybridized invasive genotypes. Previous work has shown that multiple introductions of P. arundinacea from various European locations to the United States resulted in numerous hybridization events, yielding more genetic variability and phenotypic plasticity in the invasive range. Of the genotypes studied, both morphological and physiological traits of genotypes with French origin were significantly different from the plants from the Czech Republic, North Carolina, and Vermont. The lack of clear differences between native and invasive genotypes indicates that physiological traits may be highly conserved in P. arundinacea and enhanced photosynthetic rates are not indicative of successful invasive genotypes. Instead, morphological traits and defensive secondary compound metabolism may play a more important role in the success of P. arundinacea within its invasive range, and patterns of genetic variation in physiological traits between invasive and native range may be more important than the mean traits of each region when explaining reed canarygrass’ invasive potential in North America.     

Tri-oceanic connectivity of the supposedly cosmopolitan polychaete,Harmothoe imbricata (Annelida: Polynoidae): insights from the COI marker

ABSTRACT

In this study, we investigated the global population genetic structure of the polychaete, Harmothoe imbricata, to determine connectivity patterns within the species. We sequenced the mtDNA marker, cytochrome c oxidase I (COI) from 29 specimens sampled across three sites from the New England coast of the United States. These were supplemented with 145 archived sequences from GenBank and the Barcode of Life Database, representing 16 global populations, which encompassed three broadly defined marine biogeographic regions: the Atlantic, Pacific and Arctic. The resulting haplotype network and pairwise AMOVA results showed marked structure across all the major biogeographic regions and also provides evidence for cryptic diversity in the species. Haplotypes from Arctic populations were more closely related to each other than those from the northwestern and northeastern Atlantic. Two evolutionary divergent lineages were recovered from Los Angeles, California and Manitoba, Canada. The highest genetic diversity was observed in the Arctic populations, providing evidence for an Arctic origin for H. imbricata. While human-mediated introductions may have likely contributed to some of the genetic patterns observed in this study, future work should incorporate a nuclear DNA component which could shed more light on contemporary movement of this species across large spatial scales.     

High genetic diversity in French invasive populations of common ragweed, Ambrosia artemisiifolia, as a result of multiple sources of introduction

Ambrosia artemisiifolia is an aggressive North American annual weed, found particularly in sunflower and corn fields. Besides its economic impact on crop yield, it represents a major health problem because of its strongly allergenic pollen. Ragweed was imported inadvertently to Europe in the 18th century and has become invasive in several countries, notably in the Rhône Valley of France. It has recently expanded in both the Provence-Alpes-Côte-d’Azur and Bourgogne regions. As first steps towards understanding the causes and mechanisms of ragweed invasion, genetic variability of French and North American populations was analysed using microsatellites. Overall genetic variability was similar in North America and in the Rhône-Alpes region, but within-population levels of genetic variability were surprisingly lower in native than in invasive French populations. French populations also exhibited lower among-population differentiation. A significant pattern of isolation by distance was detected among North American populations but not among French populations. Assignment tests and distribution of rare alleles did not point to a single origin for all French populations, nor for all individuals within populations and private alleles from different North American populations were found in the same French populations. Indeed, within all French populations, individual plants were roughly equally assigned to the different North American populations. Altogether, these results suggest that the French invasive populations include plants from a mixture of sources. Reduced diversity in populations distant from the original area of introduction indicated that ragweed range expansion probably occurred through sequential bottlenecks from the original populations, and not from subsequent new introductions.     

European green crabs (Carcinus maenas) in the northeastern Pacific: genetic evidence for high population connectivity and current-mediated expansion from a single introduced source population

Aim The European green crab (Carcinus maenas) expanded dramatically after its introduction to the west coast of North America, spreading over 1000 km in < 10 years. We use samples of Carcinus maenas collected over time and space to investigate the genetic patterns underlying the species’ initial establishment and spread, and discuss our findings in the context of the species’ life history characteristics and demography. Location The central west coast of North America, encompassing California, Oregon, and Washington (USA) and British Columbia (Canada). Methods We collected 1040 total samples from 21 sites representing the major episodes of population establishment and expansion along the west coast of North America. Microsatellite markers were used to assess genetic diversity and structure at different time points in the species’ spread, to investigate connectivity between embayments and to estimate both short‐term effective population sizes and the number of original founders. Assignment testing was performed to determine the likely source of the introduction. Results Carcinus maenas in western North America likely derived from a single introduction of a small number of founders to San Francisco Bay, CA from the east coast of North America. Throughout its western North American range, the species experiences periodic migration between embayments, resulting in a minor loss of genetic diversity in more recently established populations versus the populations in the area of initial establishment. Main conclusions Low genetic diversity has not precluded the ability of C. maenas to successfully establish and spread on the west coast of North America. An efficient oceanographic transport mechanism combined with highly conducive life history traits are likely the major drivers of C. maenas spread. Evidence for a single introduction underscores the potential utility of early detection and eradication of high‐risk invasive species.     

Patterns of genetic diversity reveal multiple introductions and recurrent founder effects during range expansion in invasive populations of Geranium carolinianum (Geraniaceae)

Genetic diversity, and thus the adaptive potential of invasive populations, is largely based on three factors: patterns of genetic diversity in the species'' native range, the number and location of introductions and the number of founding individuals per introduction. Specifically, reductions in genetic diversity (‘founder effects'') should be stronger for species with low within-population diversity in their native range and few introductions of few individuals to the invasive range. We test these predictions with Geranium carolinianum, a winter annual herb native to North America and invasive in China. We measure the extent of founder effects using allozymes and microsatellites, and ask whether this is consistent with its colonization history and patterns of diversity in the native range. In the native range, genetic diversity is higher and structure is lower than expected based on life history traits. In China, our results provide evidence for multiple introductions near Nanjing, Jiangsu province, with subsequent range expansion to the west and south. Patterns of genetic diversity across China reveal weak founder effects that are driven largely by low-diversity populations at the expansion front, away from the introduction location. This suggests that reduced diversity in China has resulted from successive founder events during range expansion, and that the loss of genetic diversity in the Nanjing area was mitigated by multiple introductions from diverse source populations. This has implications for the future of G. carolinianum in China, as continued gene flow among populations should eventually increase genetic diversity within the more recently founded populations.     

Cryptic species in Pacific sipunculans (Sipuncula: Phascolosomatidae): east‐west divergence between non‐sister taxa

Cryptic diversity, i.e. diversity observable in genetic but not in morphological traits, is prevalent in marine invertebrates and presents one of the greatest obstacles to obtaining accurate estimates of species richness. Sipunculans, commonly called peanut worms, are marine annelids in which high levels of cryptic diversity have previously been documented. In this study, we use genetic identification techniques to examine divergence of two lineages of Pacific sipunculans, both known under the name Phascolosoma agassizii. One lineage is isolated to the eastern Pacific coast while the other one inhabits the western Pacific coast. These clades are reciprocally monophyletic and are not recovered as sister taxa, suggesting relatively early divergence within Phascolosoma. Furthermore, we did not find support for a genetic distinction between the western Pacific Phascolosoma agassizii agassizii and Phascolosoma agassizii kurilense, a subspecies reported from the Kuril Islands. Considering that the type locality for P. agassizii is in the Eastern Pacific, we suggest that the western clade, including the samples from the Kuril islands, represent a new, undescribed species.