Food of Iago omanensis, a deep water shark from the northern Red Sea

Stomach contents of 252 lago omanensis were examined. Sharks were collected in deep water between 150 and 450 m depth off the Marine Laboratory of Eilat. This species feeds on benthopelagic fishes, deep water cephalopods, crustaceans, benthic molluscs and polychaetes. Cephalopods were the main food of I. omanensis , except during the summer when fishes (mainly myctophids) were ingested in greater quantity. Stomachs also contained terrigenous plant matter, animal remains and large quantities of mud. Prey taxa included two new cephalopod records for the Red Sea.     

The complex effects of the invasive polychaetes Marenzelleria spp. on benthic nutrient dynamics

The effects of the polychaetes Marenzelleria sp. (Polychaeta, Spionidae), nonindigenous, rapidly increasing species in the Baltic Sea, on benthic nutrient fluxes, denitrification and sediment pore water nutrient concentration were studied in laboratory experiments using a flow-through setup with muddy sediment from coastal regions of the Gulf of Finland. In addition, different forms of sediment phosphorus (P), separated by chemical fractionation, were studied in three sediment layers. At a population density corresponding to about half the highest measured in the northern Baltic Sea, Marenzelleria sp. increased the fluxes of P and ammonium to the water column. No effect could be recorded for denitrification. Since the previously dominant species of the area, Monoporeia affinis, can enhance denitrification and lower the amount of dissolved P in the pore water, the replacement of M. affinis with Marenzelleria spp. may lead to increased P flux to the water column and decreased denitrification, further increasing the ammonium flux to the water column. However, sediment reworking by Marenzelleria spp. also oxidizes the surface sediment in the long run, improving its ability to retain P and support nitrification. Therefore, the impact of Marenzelleria spp. on sediment nutrient release may not be as drastic as the initial reactions seen in our experiments suggest.     

Oxygen minimum expansion in the Sulu Sea,western equatorial Pacific,during the last glacial low stand of sea level

The Sulu Sea in the western equatorial Pacific is presently a shallowly-silled, dysaerobic, deep-marine basin. Deep waters in the Sulu Sea are ventilated through a single sill at 420 m depth which connects it to the China Sea. Benthic and planktonic foraminiferal oxygen and carbon isotope records, benthic and planktonic foraminiferal census data and total organic carbon measurements have been used to evaluate changes in water mass conditions in the Sulu Sea between the last glacial maximum (18,000 yrs. B.P.) and the present day.An increase in the abundance of the planktonic foraminiferaNeogloboquadrina dutertrei and relatively light planktonic foraminiferal δ¹⁸O values suggest that during the last glacial maximum surface water salinities were reduced in the Sulu Sea. Enhanced isolation of the basin due to glacio-eustatic lowering of sea level and reduced surface salinities resulted in stagnation of deep water and an expansion of the mid-water oxygen minimum layer. Increased organic carbon preservation at mid-water depths occurs at this time. Benthic carbon isotope data and an increase in the abundance of benthic foraminiferal species considered to prefer low oxygen environments support the conclusion of an oxygen-minimum expansion at mid-water depths during the last glacial maximum. At water depths greater than 4000 m, bottom waters appear to have maintained some degree of oxygenation during the last glacial maximum. Stronger Pacific Ocean trade winds at this time may have caused the influx of denser Celebes Sea surface water into the southern part of the Sulu Sea. The slow sinking of this water would have then ventilated bottom waters in this part of the basin.At the transition from glacial to interglacial conditions, rising sea level caused denser water to flow over the deepest sill into the Sulu Sea. Vertical circulation increased, resulting in a greater downward flux of oxygen and a dissipation of the oxygen minimum. Continued post-glacial sea level rise caused periodic ventilation of deep water until the present dysaerobic conditions were established.     

High-resolution SAR11 ecotype dynamics at the Bermuda Atlantic Time-series Study site by phylogenetic placement of pyrosequences

Advances in next-generation sequencing technologies are providing longer nucleotide sequence reads that contain more information about phylogenetic relationships. We sought to use this information to understand the evolution and ecology of bacterioplankton at our long-term study site in the Western Sargasso Sea. A bioinformatics pipeline called PhyloAssigner was developed to align pyrosequencing reads to a reference multiple sequence alignment of 16S ribosomal RNA (rRNA) genes and assign them phylogenetic positions in a reference tree using a maximum likelihood algorithm. Here, we used this pipeline to investigate the ecologically important SAR11 clade of Alphaproteobacteria. A combined set of 2.7 million pyrosequencing reads from the 16S rRNA V1–V2 regions, representing 9 years at the Bermuda Atlantic Time-series Study (BATS) site, was quality checked and parsed into a comprehensive bacterial tree, yielding 929 036 Alphaproteobacteria reads. Phylogenetic structure within the SAR11 clade was linked to seasonally recurring spatiotemporal patterns. This analysis resolved four new SAR11 ecotypes in addition to five others that had been described previously at BATS. The data support a conclusion reached previously that the SAR11 clade diversified by subdivision of niche space in the ocean water column, but the new data reveal a more complex pattern in which deep branches of the clade diversified repeatedly across depth strata and seasonal regimes. The new data also revealed the presence of an unrecognized clade of Alphaproteobacteria, here named SMA-1 (Sargasso Mesopelagic Alphaproteobacteria, group 1), in the upper mesopelagic zone. The high-resolution phylogenetic analyses performed herein highlight significant, previously unknown, patterns of evolutionary diversification, within perhaps the most widely distributed heterotrophic marine bacterial clade, and strongly links to ecosystem regimes.     

Novel use of burrow casting as a research tool in deep-sea ecology

Although the deep sea is the largest ecosystem on Earth, its infaunal ecology remains poorly understood because of the logistical challenges. Here we report the morphology of relatively large burrows obtained by in situ burrow casting at a hydrocarbon-seep site and a non-seep site at water depths of 1173 and 1455 m, respectively. Deep and complex burrows are abundant at both sites, indicating that the burrows introduce oxygen-rich sea water into the deep reducing substrate, thereby influencing benthic metabolism and nutrient fluxes, and providing an oxic microhabitat for small organisms. Burrow castings reveal that the solemyid bivalve Acharax johnsoni mines sulphide from the sediment, as documented for related shallow-water species. To our knowledge, this is the first study to examine in situ burrow morphology in the deep sea by means of burrow casting, providing detailed information on burrow structure which will aid the interpretation of seabed processes in the deep sea.     

Novel lineages of Prochlorococcus and Synechococcus in the global oceans

Picocyanobacteria represented by Prochlorococcus and Synechococcus have an important role in oceanic carbon fixation and nutrient cycling. In this study, we compared the community composition of picocyanobacteria from diverse marine ecosystems ranging from estuary to open oceans, tropical to polar oceans and surface to deep water, based on the sequences of 16S-23S rRNA internal transcribed spacer (ITS). A total of 1339 ITS sequences recovered from 20 samples unveiled diverse and several previously unknown clades of Prochlorococcus and Synechococcus. Six high-light (HL)-adapted Prochlorococcus clades were identified, among which clade HLVI had not been described previously. Prochlorococcus clades HLIII, HLIV and HLV, detected in the Equatorial Pacific samples, could be related to the HNLC clades recently found in the high-nutrient, low-chlorophyll (HNLC), iron-depleted tropical oceans. At least four novel Synechococcus clades (out of six clades in total) in subcluster 5.3 were found in subtropical open oceans and the South China Sea. A niche partitioning with depth was observed in the Synechococcus subcluster 5.3. Members of Synechococcus subcluster 5.2 were dominant in the high-latitude waters (northern Bering Sea and Chukchi Sea), suggesting a possible cold-adaptation of some marine Synechococcus in this subcluster. A distinct shift of the picocyanobacterial community was observed from the Bering Sea to the Chukchi Sea, which reflected the change of water temperature. Our study demonstrates that oceanic systems contain a large pool of diverse picocyanobacteria, and further suggest that new genotypes or ecotypes of picocyanobacteria will continue to emerge, as microbial consortia are explored with advanced sequencing technology.     

Inshore–offshore comparison of parasite infestation in Mytilus edulis: implications for open ocean aquaculture

Offshore production offers a new perspective for mussel aquaculture in the German Bight (North Sea) as no expansion is possible in the intertidal and subtidal zone of the Wadden Sea because of restrictions on the number of licenses. The development of offshore wind farms offers a unique opportunity because of the associated infrastructure. Service platforms, as well as the pylons themselves, offer perfect structures for mooring longlines and other culture units. One of the advantages of offshore culture may be a lower parasite load in offshore mussels compared with mussels produced under traditional inshore bottom culture. By sampling mussel spat from offshore suspended buoys or collectors, we simulated an offshore culture situation and compared parasite infestation rates with those in mussel spat obtained from suspended inshore buoys or collectors, in mussels from inshore benthic subtidal beds and from inshore benthic intertidal mussel beds. Mussels from offshore sites were free of trematodes and shell‐boring polychaetes. Parasitic copepods only occurred at a single offshore site, on a 20‐year‐old research platform, but not on buoys or collectors exposed for shorter time periods. All three monitored parasite taxa were present at all other sites. The highest prevalence was found for trematodes in inshore benthic intertidal mussels (78.7 ± 6.4%) and locally reached 100%. Through a variety of detrimental effects, trematodes, parasitic copepods and shell‐boring polychaetes are known to affect growth performance and product quality. We therefore propose that offshore mussel production could be a promising culture procedure because it seems to result in lower parasite burden than at traditional culture sites. Whether offshore production also results in better survival and growth, compared with inshore mussel culture on a commercial scale, needs to be investigated further.     

Lineage divergence of a freshwater snail clade associated with post-Tethys marine basin development

The complex evolutionary history of the Eurasian gastropod lineage Theodoxus reflects the evolution of marine basins following the breakup of the Tethys Sea. Today, this clade inhabits the lakes, rivers, streams, and estuaries of Europe, southwestern Asia, and North Africa. Here we present the first phylogenetic hypothesis for this clade. Based upon extensive geographic and taxonomic sampling, portions of the mitochondrial genes for cytochrome c oxidase subunit I and 16S rRNA were sequenced and analysed using maximum likelihood and maximum parsimony methods. Results from bootstrap analyses, Bayesian analysis, and sensitivity analyses lend support to six deep phylogenetic subdivisions within Theodoxus. These major clades are geographically associated with the major post-Tethyan marine basins. Estimates of divergence times using a penalized likelihood approach indicate that divergence of these major lineages occurred during the Miocene, simultaneous with the breakup of the Mediterranean and Paratethys Seas. The resulting major subclades later diversified during the Pliocene, primarily within geographic regions associated with the eastern and western Mediterranean Sea, the Pannonian Basin, and the Black Sea, thus producing the extant species assemblages. Finally, these phylogenetic results imply that much of the current taxonomy is flawed, therefore we offer recommendations for revising the classification of Theodoxus species based on phylogenetic systematics.     

Polychaete diversity in the Scotia Arc benthic realm: Are polychaetes tracers for faunal exchange?

The Scotia Arc is the only shallow-water and island bridge linking nowadays Patagonia and the Antarctic. The Antarctic Circumpolar Current as an oceanographic peculiarity makes this region an interesting biogeographic transition zone, because this frontal system traditionally is said to isolate the Antarctic fauna from that of the adjacent northern ecosystems. Based on benthos samples from three expeditions onboard R/V Polarstern, we studied distribution patterns of 200 polychaete species and 34 major benthic taxa in order to evaluate the role of polychaetes in the benthic realm of this part of the Southern Ocean. ANOSIM test distinguished three station groups: the central eastern Scotia Sea, the continental shelf off South America and stations at the tip of the Antarctic Peninsula. These station groups differed in organism densities and diversities with stations at the tip of the Antarctic Peninsula hosting the most diverse and dense community. The polychaete diversity patterns in the three assemblages evidenced closer connectivity between the tip of the Antarctic Peninsula and the central eastern Scotia Sea than between the continental shelf off South America with either the stations off the tip of the Peninsula or the central eastern Scotia Sea. This is probably supported by the Polar Front, which divides the island chain into two branches. Species distribution and community patterns of polychaetes appear to be associated with oceanographic and sediment conditions in this region. Most of the shared species showed the capability to tolerate differences in hydrostatic pressure. We suggest that the islands of the Scotia Sea may constitute a bridge for exchange of benthic species, particularly for polychaetes with eurybathic distribution and high dispersal capabilities.     

Periodic and episodic benthos recruitment in southwest Mecklenburg Bay (western Baltic Sea)

Temporal changes in the zoobenthos of southwest Mecklenburg Bay were studied using routine survey data collected in 1980–94. Earlier studies suggested that long-term oxygen deficiency on the scale of several months affects the zones below 20 m depth. Episodic benthic recolonization occurred during periods with improved oxygen conditions, as a rule by opportunistic polychaetes followed by some actively immigrating adult species. However, these observations were restricted to central and southern Lübeck Bay. Our recent data indicate that benthic recolonization exhibits greater regularity along the northern shore of Lübeck Bay. Special wind conditions and water exchange processes in the western Baltic Sea are likely to displace the deep water in northern Lübeck Bay more rapidly than in the southern zones. By means of multidimensional scaling, two types of episodic recruitment are identified in the benthos that are probably related to enhanced nitrogen discharge through river runoff in the winter and exceptionally weak water stratification in the summer.     

Evolution of seahorses' upright posture was linked to Oligocene expansion of seagrass habitats

Seahorses (Syngnathidae: Hippocampus) are iconic marine teleosts that are readily identifiable by their upright posture. The fossil record is inadequate to shed light on the evolution of this trait because it lacks transitional forms. There are, however, extant syngnathid species (the pygmy pipehorses) that look like horizontally swimming seahorses and that might represent a surviving evolutionary link between the benthic seahorses and other, free-swimming members of the family Syngnathidae. Using sequence data from five nuclear loci, we confirm the sister taxon relationship between seahorses and pygmy pipehorses. Molecular dating indicates that the two taxa diverged during the Late Oligocene. During this time, tectonic events in the Indo-West Pacific resulted in the formation of vast amounts of new shallow-water areas and associated expansion of seagrass habitats that would have favoured the seahorses’ upright posture by improving their camouflage while not affecting their manoeuvrability negatively. The molecular techniques employed here provide new insights into the evolution of a taxon whose fossil record is incomplete, but whose evolutionary history is so recent that the major stages of morphological evolution are still represented in extant species.     

Historical changes in the benthos of the Wadden Sea around the island of Sylt in the North Sea

The situation regarding the distribution and abundance of seagrass, macroalgae and benthic fauna near the island of Sylt in the south-eastern North Sea during the period 1923 to 1940 is compared with that of the 1980s. Evidence of organic enrichment in recent times is provided by (1) massive growth of green algal mats on sheltered tidal flats, (2) a decline of red algae in the subtidal zone, (3) an expansion of mussel beds along low water line and down to 20 m depth, (4) increased abundance of polychaetes inhabiting intertidal and subtidal sandy bottoms. Seagrass beds have undergone complex changes which remain unexplained. Intensified erosion has contributed to the loss of habitats in the intertidal zone, and probably affected sessile epifauna in the deep channels. Here, direct removal and disturbance by the bottom-trawling fishery may also have contributed to the observed species impoverishment. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Warm-water decapods and the trophic amplification of climate in the North Sea

A long-term time series of plankton and benthic records in the North Sea indicates an increase in decapods and a decline in their prey species that include bivalves and flatfish recruits. Here, we show that in the southern North Sea the proportion of decapods to bivalves doubled following a temperature-driven, abrupt ecosystem shift during the 1980s. Analysis of decapod larvae in the plankton reveals a greater presence and spatial extent of warm-water species where the increase in decapods is greatest. These changes paralleled the arrival of new species such as the warm-water swimming crab Polybius henslowii now found in the southern North Sea. We suggest that climate-induced changes among North Sea decapods have played an important role in the trophic amplification of a climate signal and in the development of the new North Sea dynamic regime.     

Gambierdiscus balechii sp. nov (Dinophyceae), a new benthic toxic dinoflagellate from the Celebes Sea (SW Pacific Ocean)

A new benthic toxic dinoflagellate is described from the Celebes Sea. Gambierdiscus balechii sp. nov. was isolated from seaweeds growing in tidal ponds. Its morphology was studied by means of LM and SEM; G. balechii has a very ornamented theca, a hatchet shaped second apical plate, a narrow second antapical plate and an asymmetrical third precigular plate, a unique combination of characters among Gambierdiscus species. It has a very wide size range with widths from 36 to 88 μm. Phylogenetic analyses of two G. balechii strains, based on LSU rRNA (D8–D10) and partial SSUrRNA sequences confirmed that these clustererd in its’ own group, separated from the rest of Gambierdiscus species and with G. pacificus, G. belizeanus and G. scabrosus as its closest relatives. Thecate cysts are described from culture as non motile vegetative-like cells which germinated after being isolated and transferred to fresh medium. Mouse tests showed that this species is toxic and hence it is a potential cause of ciguatera in the Celebes Sea.     

Genome scan of the mitten crab Eriocheir sensu stricto in East Asia: population differentiation, hybridization and adaptive speciation

We examine the genetic structure and evolutionary history of the mitten crab Eriocheir sensu stricto in East Asia by employing a genome scan - amplified fragment length polymorphism. Population analysis reveals three divergent clades in Eriocheir s. s., which dominate the East China Sea-Yellow Sea, the Sea of Japan (plus Okinawa) and the South China Sea, respectively, mostly in agreement with our previous mtDNA analysis. With the tropical South China Sea inferred as the origin, the East China Sea-Yellow Sea and the Sea of Japan clades in the north diverged successively from the ancestral clade during the mid-Pleistocene. The divergence of the three clades likely resulted from isolation of the three marginal seas caused by sea level change in the Pleistocene. Two sympatric zones, one of the East China Sea-Yellow Sea and the South China Sea clades in southeast China and the other of the East China Sea-Yellow Sea and the Sea of Japan clades in Vladivostok, are demonstrated to be hybrid zones, with hybridization occurring currently in the former but historically in the latter. Adaptive speciation is observed in the divergence process of the three clades, possibly because of selection from accumulated temperature. Our study indicates that the genetic structure and evolutionary history of Eriocheir s. s. have been primarily affected by Pleistocene glacial cycles, secondarily by divergent selection and drainage isolation, but only minimally by human activities.     

Phylogeography of the mottled spinefoot Siganus fuscescens: Pleistocene divergence and limited genetic connectivity across the Philippine archipelago

Historical isolation during Pleistocene low sea level periods is thought to have contributed to divergence among marine basin populations across the Coral Triangle. In the Philippine archipelago, populations in the South China Sea, Sulu Sea–inland seas, and Philippine Sea‐Celebes Sea basins might have been partially isolated. Meanwhile, present‐day broadscale oceanographic circulation patterns suggest connectivity between these basins. To evaluate hypotheses regarding the influence of historical and contemporary factors on genetic structure, phylogeographic patterns based on mitochondrial control region sequences for a reef‐associated fish, Siganus fuscescens, were analysed. Three distinct lineages were recovered. One lineage was identified as the morphologically similar species Siganus canaliculatus, while two lineages are monophyletic with S. fuscescens. Clade divergence and demographic expansion in S. fuscescens occurred during the Pleistocene. A strong signal of latitudinal structure was detected (Φ_CT = 0.188), driven by marked differences in clade distribution: one clade is widely distributed (clade A), while a second clade (clade B) has a restricted northern distribution. Regional structure of clade A is consistent with the basin isolation hypothesis (Φ_CT = 0.040) and suggests isolation of the South China Sea (Φ_CT = 0.091). Fine‐scale structure was observed in the South China Sea and south Philippine Sea, while Sulu Sea and inland seas were unstructured. Genetic structure across multiple spatial scales (archipelagic, regional, and fine‐scale within basins) suggests the influence of vicariant barriers and contemporary limits to gene flow in S. fuscescens that may be influenced by oceanographic circulation, geographical distance between available habitats, and latitudinal temperature differences.     

Experimental study on the influence of bioturbation performed by Nephtys caeca (Fabricius) and Nereis virens (Sars) annelidae on the distribution of dinoflagellate cysts in the sediment

Dinoflagellates include noxious microalgae responsible for the formation of toxic red tides and the poisoning of molluscs and crustaceans, resulting in important economic losses. As a consequence, the life cycle of these algae has been extensively studied, but the dormancy phase (cyst) in the sediment record is little known. In the intertidal zone, bioturbation, an important biological process resulting from the activities of benthic fauna, significantly influences the movement of particles in the sediments. Laboratory experiments have allowed comparing and quantifying the movements of fluorescent microspheres resulting from the activity of two polychaetes annelidae, Nereis virens and Nephtys caeca. The particles, which simulate 45 µm diameter dinoflagellate cysts, are deposited in flat aquaria at the surface or deep in the sediment. Photographs of the aquaria were taken at regular intervals, to observe, in a non-destructive manner, the movement of the particles and to calculate, using adapted software, the optical reworking coefficient (ORC) over time. A difference appears between the movements of the particles generated by both species of polychaetes. Nereis virens create “permanent” galleries that carry the microspheres deeply in the sediment during the digging, bioirrigation and feeding, and Nephtys caeca homogenize the particles in the first centimetres of sediment during its erratic movements. The study shows that the bioturbation generated by these organisms can modify the distribution of the 45 µm diameter dinoflagellate cysts in the sedimentary column, burying them or raising them back to the water-sediment interface.     

Environmental Drivers of Benthic Flux Variation and Ecosystem Functioning in Salish Sea and Northeast Pacific Sediments

The upwelling of deep waters from the oxygen minimum zone in the Northeast Pacific from the continental slope to the shelf and into the Salish Sea during spring and summer offers a unique opportunity to study ecosystem functioning in the form of benthic fluxes along natural gradients. Using the ROV ROPOS we collected sediment cores from 10 sites in May and July 2011, and September 2013 to perform shipboard incubations and flux measurements. Specifically, we measured benthic fluxes of oxygen and nutrients to evaluate potential environmental drivers of benthic flux variation and ecosystem functioning along natural gradients of temperature and bottom water dissolved oxygen concentrations. The range of temperature and dissolved oxygen encountered across our study sites allowed us to apply a suite of multivariate analyses rarely used in flux studies to identify bottom water temperature as the primary environmental driver of benthic flux variation and organic matter remineralization. Redundancy analysis revealed that bottom water characteristics (temperature and dissolved oxygen), quality of organic matter (chl a:phaeo and C:N ratios) and sediment characteristics (mean grain size and porosity) explained 51.5% of benthic flux variation. Multivariate analyses identified significant spatial and temporal variation in benthic fluxes, demonstrating key differences between the Northeast Pacific and Salish Sea. Moreover, Northeast Pacific slope fluxes were generally lower than shelf fluxes. Spatial and temporal variation in benthic fluxes in the Salish Sea were driven primarily by differences in temperature and quality of organic matter on the seafloor following phytoplankton blooms. These results demonstrate the utility of multivariate approaches in differentiating among potential drivers of seafloor ecosystem functioning, and indicate that current and future predictive models of organic matter remineralization and ecosystem functioning of soft-muddy shelf and slope seafloor habitats should consider bottom water temperature variation. Bottom temperature has important implications for estimates of seasonal and spatial benthic flux variation, benthic–pelagic coupling, and impacts of predicted ocean warming at high latitudes.     

Disjunct distribution of highly diverged mitochondrial lineage clade and population subdivision in a marine bivalve with pelagic larval dispersal

Mitochondrial DNA sequence data for 295 individuals of the marine bivalve Macoma balthica (L.) were collected from 10 sites across the European distribution, and from Alaska. The data were used to infer population subdivision history and estimate current levels of gene flow. Inferred historical biogeography was expected to be congruent with colonization of the Atlantic Ocean from the Pacific Ocean after the opening of the Bering Strait 3.5 Ma. In addition, the last glacial maximum, about 18000 years ago, was expected to have been responsible for most of the present-day distribution of molecular variation within Europe, because the area must have been recolonized after confinement to France and the south of the British Isles during the last glacial maximum. Current gene flow was hypothesized to be high, because the larvae of M. balthica spend 2-5 weeks drifting in the water column. The geographical distribution of one highly diverged haplotype clade was found to be disjunct and was encountered exclusively in samples from the Baltic Sea and Alaska. A molecular clock calibration for marine bivalve cytochrome-c-oxidase I dates this clade as having split off from the other haplotypes 9.8-39 Ma. Multiple colonizations of the Atlantic Ocean from the Pacific by M. balthica may explain the strong differences found between Baltic Sea and other European populations of this species. The sympatric occurrence of the highly diverged mitochondrial lineages in western parts of the Baltic Sea points to secondary admixture. With the use of coalescent analysis, population divergence times for French vs. other non-Baltic European populations ('Atlantic population assemblage') were estimated at a minimum of about 110000 years ago, well before the last glacial maximum 18000 years ago. Signatures of population divergence of M. balthica that appear to have originated during the Pleistocene have thus survived the last glacial maximum. Some of the populations within the Atlantic assemblage are currently isolated, while others appear to be connected by gene flow. Apparently, populations of this species can remain highly subdivided in spite of the potential for high gene flow, implying that their population and evolutionary dynamics can be independent.