Ecological regime shift drives declining growth rates of sea turtles throughout the West Atlantic

Somatic growth is an integrated, individual‐based response to environmental conditions, especially in ectotherms. Growth dynamics of large, mobile animals are particularly useful as bio‐indicators of environmental change at regional scales. We assembled growth rate data from throughout the West Atlantic for green turtles, Chelonia mydas, which are long‐lived, highly migratory, primarily herbivorous mega‐consumers that may migrate over hundreds to thousands of kilometers. Our dataset, the largest ever compiled for sea turtles, has 9690 growth increments from 30 sites from Bermuda to Uruguay from 1973 to 2015. Using generalized additive mixed models, we evaluated covariates that could affect growth rates; body size, diet, and year have significant effects on growth. Growth increases in early years until 1999, then declines by 26% to 2015. The temporal (year) effect is of particular interest because two carnivorous species of sea turtles—hawksbills, Eretmochelys imbricata, and loggerheads, Caretta caretta—exhibited similar significant declines in growth rates starting in 1997 in the West Atlantic, based on previous studies. These synchronous declines in productivity among three sea turtle species across a trophic spectrum provide strong evidence that an ecological regime shift (ERS) in the Atlantic is driving growth dynamics. The ERS resulted from a synergy of the 1997/1998 El Niño Southern Oscillation (ENSO)—the strongest on record—combined with an unprecedented warming rate over the last two to three decades. Further support is provided by the strong correlations between annualized mean growth rates of green turtles and both sea surface temperatures (SST) in the West Atlantic for years of declining growth rates (r = −.94) and the Multivariate ENSO Index (MEI) for all years (r = .74). Granger‐causality analysis also supports the latter finding. We discuss multiple stressors that could reinforce and prolong the effect of the ERS. This study demonstrates the importance of region‐wide collaborations.     

Uncommon southwest swells trigger sea urchin disease outbreaks in Eastern Atlantic archipelagos

Recurrent sea urchin mass mortality has recently affected eastern Atlantic populations of the barren‐forming sea urchin Diadema africanum. This new episode of die‐off affords the opportunity to determine common meteorological and oceanographic conditions that may promote disease outbreaks. The population dynamics of this sea urchin species are well known—urchin barrens have persisted for many decades along most of the coastlines off the archipelagos of Madeira, Selvages, and the Canary Islands, where they limit macroalgae biomass growth. However, this new and explosive mortality event decimated the sea urchin population by 93% on Tenerife and La Palma Islands. Two severe episodes of southwestern rough sea that led to winter storms, in February 2010 (Xynthia) and February 2018 (Emma), preceded both mass mortality events. The autumn and winter months of those years were anomalous and characterized by swells with an average wave height above 2 m that hit the south and southwest sides of the islands. The amoeba Paramoeba brachiphila was the only pathogen isolated this time from the moribund and dead sea urchins, suggesting that the amoeba was the primary cause of the mortality. This new sea urchin die‐off event supports the “killer‐storm” hypothesis that has been already described for western Atlantic coasts. These anomalous southwest storms during winters generate pronounced underwater sediment movement and large‐scale vertical mixing, detected in local tide gauge, which may promote paramoebiasis. This study presents valuable insights about climate‐mediated changes in disease frequency and its impacts on the future of coastal marine ecosystems in the Atlantic.     

Invasion or invisibility: using genetic and distributional data to investigate the alien or indigenous status of the Atlantic populations of the peracarid isopod,Stenosoma nadejda (Rezig 1989)

The peracarid isopod, Stenosoma nadejda (Rezig 1989), until recently considered to be endemic of the Mediterranean region, was first reported in the Atlantic coast of southern Spain in 2001, and in 2006 abundant populations were discovered throughout the southwestern Portuguese coast. This fast expansion was intriguing because, as a direct brooder, this species has limited mechanisms for dispersal, such as rafting on seaweeds. Did S. nadejda recently extend its range into the Atlantic or was it overlooked in the past? We examined the patterns of genetic diversity and population differentiation accordingly by sequencing the cytochrome c oxidase subunit I mitochondrial gene from 75 individuals collected in five locations in Atlantic Iberia and one in the Mediterranean. Our results indicate that the newly discovered Atlantic populations of S. nadejda appear to be old and have long persisted on Atlantic shores rather than being a recent introduction. High levels of genetic diversity and geographic structure were uncovered in what was initially suspected to be an ‘invasive’ species. Recent changes in population dynamics may have made S. nadejda more conspicuous in the Atlantic shores, or a more comprehensive survey led to the recognition of this species where it was not expected.     

Phylogeography and population structure of European sea bass in the north‐east Atlantic

The European sea bass Dicentrarchus labrax represents a historically and commercially valuable species in the north‐east Atlantic, although the demographic history and the patterns of geographical structure of the species in the north‐east Atlantic remain poorly understood. The present study investigates the population genetic structure of sea bass in north‐western European waters, employing different genetic markers [a portion of the mitochondrial (mt)DNA control region and 13 nuclear microsatellites] aiming to unravel demographic history and population connectivity. The results obtained show a previously unrecognized pattern of population divergence at mtDNA, with three strikingly different lineages identified. Extant sea bass populations, including the Mediterranean lineage, derive from an Atlantic ancestor. A much increased number of nuclear microsatellite loci (comparatively to previous studies) still fail to detect biologically meaningful patterns of spatial genetic structuring in the North Atlantic. Past Pleistocene glacial and interglacial events and some degree of female philopatry might be at the basis of the current geographical separation of the Atlantic lineages that has been identified. Signatures of sudden demographic expansions are more evident in the most recent mitochondrial lineages, and their slight, yet significant, geographical segregation leads to the hypothesis that present‐day spawning grounds for European sea bass may still to some extent be linked to their most recent glacial refugia. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 364–377.     

Phylogeography and cryptic introduction of the ragworm Hediste diversicolor (Annelida,Nereididae) in the Northwest Atlantic

Many benthic marine invertebrates show striking range disjunctions across broad spatial scales. Without direct evidence for endemism or introduction, these species remain cryptogenic. The common ragworm Hediste diversicolor plays a pivotal role in sedimentary littoral ecosystems of the North Atlantic as an abundant prey item and ecosystem engineer, but exhibits a restricted dispersal capacity that may limit connectivity at both evolutionary and ecological time scales. In Europe, H. diversicolor is subdivided into cryptic taxa and genetic lineages whose distributions have been modified by recent invasions. Its origin in the northwest Atlantic has not been adequately addressed. To trace the age and origin of North American ragworm populations, we analyzed mtDNA sequence data (COI) from the Gulf of Maine and Bay of Fundy (n=73 individuals) and compared our findings with published data from the northeast Atlantic. Our results together with previous data indicate that two species of the H. diversicolor complex have independently colonized the northwest Atlantic at least three different times, resulting in two distinct conspecific assemblages in the Bay of Fundy and Gulf of Maine (respectively) that are different from the species found in the Gulf of St. Lawrence. North American populations had significantly lower genetic diversity compared with populations in the northeast Atlantic, and based on patterns of shared identity, populations in the Bay of Fundy originated from the Baltic Sea and North Sea. Populations from the Gulf of Maine were phylogenetically distinct and most likely originated from unsampled European populations. Analyses of the North American populations revealed patterns of post‐colonization gene flow among populations within the Gulf of Maine and Bay of Fundy. However, we failed to detect shared haplotypes between the two regions, and this pattern of complete isolation corroborates a strong phylogeographic break observed in other species.     

Ten years after: krill as indicator of changes in the macro-zooplankton communities of two Arctic fjords

A macro-zooplankton study from 1996 was repeated in 2006 and focused on euphausiid species as indicators of advection and warming effects in Kongsfjorden, West Spitsbergen, Svalbard. The influence of warmer Atlantic water in Kongsfjorden was indicated by the findings of three additional euphausiid species of typically Atlantic origin, relative to the previous study 10 years ago. The predominant presence of Thysanoessa inermis in Hornsund suggested persisting cold conditions in this more southerly, but more Arctic influenced fjord. In this species, moult stage analysis showed that trophic effects can override temperature forcing. Histology and lipid analysis suggest that reproductive activity should be monitored as an indication of warming and possibly a shift in food web composition.     

Predation by the carabid beetles Pterostichus madidus and Nebria brevicollis is affected by size and condition of the prey slug Deroceras reticulatum

1 Slugs are important pests in many agricultural crops but molluscicides commonly used to control slugs affect non‐target organisms. Encouraging biological control may help to reduce molluscicide use, but the efficiency of potential natural enemies needs to be investigated. 2 Serological tests have shown that certain carabid species consume slugs. These techniques, however, do not distinguish between scavenging and true predation, nor do they provide information on the size or other characteristics of the prey consumed. The study reported here was undertaken to establish whether scavenging of dead slugs might be an important factor contributing to positive serological test results. 3 Both Pterostichus madidus (Fabricius) and Nebria brevicollis (Fabricius) consumed Deroceras reticulatum (Müller) under laboratory conditions. Dead slugs were scavenged in preference to injured or healthy slugs. 4 Only small, live slugs (< 0.11 g) were killed by both beetle species, which may, therefore, be incapable of killing larger slugs. 5 These generalist beetle species appeared unable to overcome the defence mucus produced by slugs. The data suggest that positive serological results from field collected beetles may reflect scavenging rather than predation on live or injured slugs.     

Morphological and molecular clarification of the enigmatic Caulerpa floridana W.R. Taylor (Chlorophyta,Bryopsidales) from the Dry Tortugas,Florida

Morphological and molecular evidence is provided to further document the status of the enigmatic taxon known as Caulerpa floridana W.R. Taylor from White Shoal, Dry Tortugas, Florida. DNA sequencing of three historical herbarium specimens (WRT329, WRT345 and WRT349) housed at the University of Michigan Herbarium (MICH) demonstrated the molecular separation of this species based on the reconstruction of 931 nucleotides of the chloroplast gene tufA. Caulerpa floridana is sister to the western Atlantic endemic C. ashmeadii Harvey and an unknown Caulerpa taxon from the Florida Middle Grounds. Caulerpa floridana most reliably differs from C. ashmeadii by the presence of a sharp, unequivocal apiculus at the tip of each pinnule. A morphological review of southwestern Atlantic records of C. floridana from Brazil excludes these reports as representative of the species.     

Taxonomy of the southwestern Atlantic endemic kelp: Laminaria abyssalis and Laminaria brasiliensis (Phaeophyceae,Laminariales) are not different species

Two endemic species of Laminaria, Laminaria abyssalis Joly & Oliveira Filho and L. brasiliensis Joly & Oliveira Filho, from the tropical southwestern Atlantic coast have been described. The aim of this work was to determine the conspecificity of these species based on morphological and molecular analyses (ribulose 1,5‐bisphosphate carboxylase/oxgenase, large subunit (rbcL), internal transcribed spacer (ITS) and cytochrome c oxidase subunit I (coxI)). We found an overlap between the morphological characters that are considered taxonomically important for distinguishing these two species; these characters included a differing pattern of blade splitting. In the three molecular analyses, the Brazilian Laminaria specimens were grouped into one clade with maximum support. These data support the hypothesis that the individuals analyzed represent only one species, L. abyssalis. The molecular analysis also showed L. abyssalis to be sister group to L. digitata.     

An effective method for trapping scavenging seabirds at sea

ABSTRACT Most studies of seabirds that involve trapping and marking birds are carried out at breeding colonies. This bias toward the breeding period and colony‐based research is partially caused by difficulties in capturing birds at sea. From 2005 to 2007, we used a cast net thrown by hand from a fishing boat to capture albatrosses and petrels at sea in the southwestern Atlantic Ocean. About 500 birds of 13 species were captured, ranging in size from the 30‐g Wilson's Storm‐petrel (Oceanites oceanicus) to the 10‐kg Wandering Albatross (Diomedea exulans). Cast nets are potentially useful for capturing any seabird that can be attracted close to fishing vessels by bait, such as sardines, squid, offal, or shark liver, thrown into the water. Our method was most effective for capturing bold species, such as Cape (Daption capense), Spectacled (Procellaria conspicillata), and White‐chinned (Procellaria aequinoctialis) petrels and Great Shearwaters (Puffinus gravis), but was not effective for capturing shy species, such as Cory's (Calonectris diomedea), Cape Verde (Calonectris edwardsii) and Manx (Puffinus puffinus) shearwaters, species that rarely sit on the water, such as Wilson's Storm‐petrels, Thin‐billed (Pachyptila belcheri) and Antarctic (Pachyptila desolata) prions and Atlantic Petrels (Pterodroma incerta), and species with excellent diving abilities, such as Sooty Shearwaters (Puffinus griseus). For many species of seabirds, cast nets would likely be more efficient for capturing large numbers of birds than other methods, such as hoop nets.     

Polyphyly across oceans: a molecular phylogeny of the Chromodorididae (Mollusca, Nudibranchia)

The Chromodorididae is a large and colourful family of nudibranch sea slugs distributed across the world's oceans. Most diversity is centred in the Indo-Pacific, but several genera are present in multiple ocean basins, or across regions separated by biogeographical barriers. The monophyly of these widespread genera had not been tested previously. We used 18S rDNA, 16S rDNA and COI sequence data to generate a molecular phylogeny for this group. We recovered evidence of paraphyly or polyphyly in all of the widespread genera examined ( Hypselodoris , Mexichromis , Chromodoris and Glossodoris ). East Atlantic Hypselodoris and west Atlantic + east Pacific Mexichromis species were more closely related to each other than they were to their Indo-Pacific congeners. The addition of Southern Ocean species of Digidentis demonstrated an interesting alternative to this relationship, becoming the sister group for the east Atlantic Hypselodoris on the basis of 16S and 18S data, but not COI data. Sister group relationships were recovered for most monotypic or enigmatic genera. Ardeadoris is linked to Glossodoris , as is Diversidoris ; Pectenodoris is sister to the Indo-Pacific Mexichromis clade, and Verconia is the sister to Noumea haliclona . Controversy surrounding the placement of the three most basal genera was only partially resolved. Using Actinocyclus to root the mitochondrial trees, Cadlinella was the unsupported sister to the Chromodorididae (excluding Cadlina ), and Tyrinna occupied a relatively basal position, although this also did not receive significant statistical support. Adding nuclear 18S data gave support for Cadlina as the sister group to the rest of the Chromodorididae s.s. Otherwise, like previous molecular studies, mitochondrial genes supported an alternative position for Cadlina (with other dorid genera).     

Reproductive cycle of the commercially harvested sea urchin (Paracentrotus lividus) along the western coast of Portugal

Harvested populations of the sea urchin (Paracentrotus lividus) from the northwestern (Carreço) and southwestern (Aljezur) coasts of Portugal were surveyed to describe the species reproductive cycle and assess possible relationships with geographical location and seawater temperature. Individuals were sampled monthly to analyze gonad histology, mean gonadal index (GI), and gonadosomatic index (GSI) during 2 consecutive years (November 2010–November 2012). Both populations presented an annual reproductive cycle, with synchronous gonad maturation and gamete release between sexes. Gonad maturation occurred throughout autumn–winter, followed by a single but prolonged spawning season during spring–summer. The duration of the spawning season displayed a latitudinal gradient likely related to the north–south increasing trend in seawater temperature, with the northwestern population (Carreço) exhibiting a shorter spawning period compared to the southwestern population (Aljezur). The timing and duration of the spawning season was compared with several populations throughout the distributional range of P. lividus in the Atlantic Ocean and Mediterranean Sea. In the population from Carreço, the size at first sexual maturity (test diameter = 35.9 mm) was considerably smaller than the minimum conservation reference size (MCRS) of 50 mm test diameter legally established for P. lividus. This study confirms that sustainable exploitation depends on harvesters’ awareness of and compliance with the MCRS and provides useful information for the eventual establishment of a closed season in the harvesting of P. lividus.     

Slugs' last meals: molecular identification of sequestered chloroplasts from different algal origins in Sacoglossa (Opisthobranchia, Gastropoda)

Some sacoglossan sea slugs have become famous for their unique capability to extract and incorporate functional chloroplasts from algal food organisms (mainly Ulvophyceae) into their gut cells. The functional incorporation of the so-called kleptoplasts allows the slugs to rely on photosynthetic products for weeks to months, enabling them to survive long periods of food shortage over most of their life-span. The algal food spectrum providing kleptoplasts as temporary, non-inherited endosymbionts appears to vary among sacoglossan slugs, but detailed knowledge is sketchy or unavailable. Accurate identification of algal donor species, which provide the chloroplasts for long-term retention is of primary importance to elucidate the biochemical mechanisms allowing long-term functionality of the captured chloroplast in the foreign animal cell environment. Whereas some sacoglossans forage on a variety of algal species, (e.g. Elysia crispata and E. viridis) others are more selective. Hence, characterizing the range of functional sacoglossan-chloroplast associations in nature is a prerequisite to understand the basis of this enigmatic endosymbiosis. Here, we present a suitable chloroplast gene (tufA) as a marker, which allows identification of the respective algal kleptoplast donor taxa by analysing DNA from whole animals. This novel approach allows identification of donor algae on genus or even species level, thus providing evidence for the taxonomic range of food organisms. We report molecular evidence that chloroplasts from different algal sources are simultaneously incorporated in some species of Elysia. NeigborNet analyses for species assignments are preferred over tree reconstruction methods because the former allow more reliable statements on species identification via barcoding, or rather visualize alternative allocations not to be seen in the latter.     

Sea‐slug invasion of the land

Invasion of the land has been a considerable challenge for aquatic organismal lineages, primarily because of desiccation and lack of physical support in air. Gastropod molluscs are among the most successful animals in this habitat shift because of their hard exoskeleton, whereas land slugs, regarded as descendants of land snails, gradually lost their shells in sheltered, moist environments. We present solid evidence of terrestrial invasion by a lineage of sea slugs; this invasion is represented by a newly discovered species, Aiteng marefugitus (Panpulmonata: Acochlidia), from a humid tropical rainforest on a small island in the Republic of Palau, western Pacific. The slug lineage seems to have invaded the forest habitat directly from the upper littoral zone in the Cenozoic.     

Assessing the geographic scale of genetic population management with microsatellites and introns in the clam Ruditapes decussatus

The clam Ruditapes decussatus is commercially important in southwestern Europe, suffering from population decline and hybridization with exotic Manila clam (R. philippinarum). Previous studies with intronic markers showed a genetic subdivision of the species in three races (Atlantic, West Mediterranean, and Adriatic‐Aegean). However, detailed population genetic studies to help management of the main production areas in the southwest of Europe are missing. We have analyzed eight Atlantic and two Mediterranean populations from the Spanish coasts using 14 microsatellites and six intronic markers. Microsatellites confirmed the Atlantic and West Mediterranean races detected with introns and showed that genetic variability was higher in Mediterranean than in Atlantic populations. Both marker types showed that genetic differentiation of Atlantic populations was low and indicated that populations could be managed at the regional level in the case of Cantabrian and Gulf of Cadiz areas, but not in the case of Rias Baixas and the Mediterranean. This study shows the interest of including different types of markers in studies of genetic population structure of marine organisms.     

Evidence for an asymmetrical size exchange of loggerhead sea turtles between the Mediterranean and the Atlantic through the Straits of Gibraltar

The permanent eastward current at the Straits of Gibraltar may trap small Atlantic loggerhead sea turtles (Caretta caretta) inside the western Mediterranean until their swimming and diving skills improve enough to allow them counter-current swimming abilities through the current. A captivity experiment with twelve loggerhead sea turtles (straight carapace length or SCL range: 25.3-48.0 cm) revealed that the average critical velocity of this species within the considered length range was 0.38 ± 0.16 m s^− 1 or 1.01 ± 0.24 bl s^− 1. As a consequence, loggerhead sea turtles are predicted to require a minimum SCL of 36.0 cm to swim counter-current through the Straits of Gibraltar, where the water velocity ranges 0.31-0.37 m s^− 1. Genetic analysis of 105 specimens using one mitochondrial marker and seven microsatellites, as well as the recapture of three tagged individuals, support this conclusion; all Mediterranean individuals found in the Atlantic side of the Straits were not smaller than 36.0 cm SCL and the average length (47.3 cm SCL) was significantly higher than that of the Mediterranean turtles in the Mediterranean side of the Straits (31.6 cm SCL). Furthermore, the average length of the turtles of any origin moving from the Mediterranean to the Atlantic was much larger than 36.0 cm (SCL: 54.5 cm SCL), which may indicate the intervention of a different, yet unidentified mechanism restricting east-westward movement. The Algerian current, running along northern Africa, may at least partially explain the delayed departure of loggerhead sea turtles from the Mediterranean Sea to the Atlantic Ocean, as it would force the eastward drift of loggerheads occupying the southwestern Mediterranean. Exchange through the Straits is asymmetrical, and Atlantic turtles are found to enter the Mediterranean at a length of about 20.5 cm. However, once in the Mediterranean they would be retained there for up to 7.9 years, due to the surface circulation pattern. This increases the time span at which turtles are exposed to a high mortality rate, caused by fishing.     

Comparing upriver spawning migration of Atlantic salmon Salmo salar and sea trout Salmo trutta

Radio tagged wild Atlantic salmon Salmo salar(n = 30) and sea trout Salmo trutta(n = 19) were simultaneously released from a sea pen outside the mouth of the River Lærdalselva and their migration to spawning areas was recorded. The distance from the river mouth to a position held at spawning ranged from 2 to 24 km and did not differ between the species (mean ± s .d . 15·9 ± 4·3 and 14·9 ± 5·2 km for Atlantic salmon and sea trout, respectively). The duration of the migration phase, however, was significantly shorter for Atlantic salmon than for sea trout (8–12 days, respectively). All Atlantic salmon migrated straight to an area near the spawning ground, whereas 50% of the sea trout had a stepwise progression with one or more periods with erratic movements before reaching the spawning area. After the migration phase, a distinct search phase with repeated movements up‐ and downstream at or close to the position held at spawning was identified for the majority of the fishes (75%, both species). This search phase was significantly shorter for Atlantic salmon than for sea trout (mean 13–31 days, respectively). Mean ± s .d . length of the river stretch used during the search phase was larger for sea trout (3·3 ± 2·5 km) than for Atlantic salmon (1·2 ± 0·9 km). A distinct holding phase, with no movements until spawning, was also observed in the majority of the Atlantic salmon (80%, mean duration 22 days) and sea trout (65%, mean duration 12 days). For both species, a weak, non‐significant trend was observed in the relationship between time spent on the migration phase, and time spent on the search (r² = 0·43) and holding phase (r² = 0·24). There was a highly significant decrease, however, in the duration of the holding phase with an increase in the time spent on the search phase (r² = 0·67).     

Molecular phylogeny, biogeography and insights into the origin of parthenogenesis in the Neotropical genus Leposoma (Squamata: Gymnophthalmidae): Ancient links between the Atlantic Forest and Amazonia

Leposoma is a conspicuous component of leaf litter herpetofauna of South and Central American rainforests. The 15 bisexual and one parthenogenetic species are allocated to the parietale and scincoides groups based on morphology. Phylogenetic analyses of 1830 bp (mtDNA + nuclear) were performed on 63 specimens of four species from Amazonian and Panamanian rainforests, and six species and one undescribed form from the Atlantic Forest. Different methods of tree reconstruction were explored, with Anotosaura vanzolinia and Colobosauroides cearensis as outgroups. The monophyly of the parietale and scincoides groups is strongly supported. Contrary to previous hypotheses suggesting a recent contact between Atlantic and Amazon forests, our estimates point to an initial split in Miocene. The position of Leposoma baturitensis, endemic to relictual forests in the semiarid Caatingas northeastern Brazil, and its divergence from the remaining species of the Atlantic Forest, suggests an ancient isolation with no indication of a secondary contact with forests of the eastern coast. Our data do not permit unambiguous assignment of parental species of the unisexual Leposoma percarinatum or the mechanism involved in the origin of parthenogenesis, but revealed two highly divergent diploid and triploid lineages within L. percarinatum, indicating that the unisexuals represent a species complex.     

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.