Characterization of food web structure of the upper continental slope of the Celtic Sea highlighting the trophic ecology of five deep‐sea fishes

In marine ecosystems, the study of trophic relationships has extensively benefited from the development of stable isotope analyses (SIA) as dietary tracers. SIA are particularly useful in elucidating the structure of deep sea food webs given the constraints involved in obtaining gut‐content data from deep trawling. We used carbon and nitrogen stable isotope analyses and Stable Isotope Bayesian Ellipses in R (SIBER) and Stable Isotope Analysis in R (SIAR) routines, to determine the trophic ecology of five deep‐sea fishes from the upper continental slope of the Celtic Sea. SIA made it possible to deduce some general tendencies in food‐web structure and species trophic interactions and confirmed diet determined by gut‐content analysis for the same species, in other ecoregions. More specifically, mixing models revealed that the deep sea species considered are omnivorous and are able to feed on all the sampled taxa. Based on isotopic ratio, no clear differences in fish diet could be detected from one species to another except for rabbit fish, which has benthic affinities. Three species, blackbelly rosefish, greater forkbeard and softhead grenadier showed overlapping isotopic niches. This study is the first attempt to describe the trophic ecology of deep sea species on the Celtic Sea upper continental slope. In the context of the development of ecosystem integrated modeling approaches for managing fisheries in the Celtic sea, and considering the vulnerability of deep‐water species, improving the knowledge on the trophic ecology of these local species is of importance in order to allow their sustainable exploitation.     

Do some deep‐sea,sediment‐dwelling species of harpacticoid copepods have 1000‐km‐scale range sizes?

The range sizes of sediment‐dwelling deep‐sea species are largely unknown. Such knowledge is important because a deep sea composed in large part of species with 100‐km‐scale ranges would be very different from one composed predominantly of species with 1000‐km‐scale ranges. For example, the total species richness would be much greater in the first case than in the second. As a step towards the determination of the distribution of species’ range sizes in the deep sea, we asked whether harpacticoid copepods (Crustacea) on the continental rise in the northeastern Pacific had 1000‐km‐scale range sizes. We chose harpacticoids because they occur widely in deep‐sea sediments and thus are a typical deep‐sea taxon. In addition, they have no pelagic stage in their life history, so they allow a conservative test of hypotheses about species’ range sizes. We used morphology and gene‐sequence data to assign individuals to species. At least 13.3% of the species we studied had 1000‐km‐scale ranges, raising the question of how these species maintain genetic continuity.     

Biodiversity response to climate change in a warm deep sea

Climate changes are expected to induce significant modifications in biodiversity on the global scale, but little is known as to how biodiversity has been affected by recent changes in the deep sea. We have used nematodes to investigate the response of deep‐sea biodiversity to an extensive climate anomaly that modified the physico‐chemical characteristics of the deep waters of the Eastern Mediterranean. Using a decadal data set (from 1989 to 1998), we provide evidence that deep‐sea nematode diversity can be strongly and rapidly affected by temperature shifts. The abrupt decrease in temperature (of about 0.4 °C) and modified physico‐chemical conditions that occurred between 1992 and 1994 caused a significant decrease in nematode abundance and a significant increase in diversity. This temperature decrease also resulted in decreased functional diversity and species evenness, and in an increase in the similarity to colder deep‐Atlantic fauna. When the temperature recovered (after 1994–1995), the biodiversity only partially returned to previous values. We conclude that deep‐sea fauna is highly vulnerable to environmental alteration, and that deep‐sea biodiversity is also significantly affected by very small temperature changes. The results presented here provide new elements towards a better understanding of the potential large‐scale consequences of climate change.     

Dorsoventral axis inversion: Enteropneust anatomy links invertebrates to chordates turned upside down

The relationships between chordates with their dorsal nerve cord and other animal groups remain unclear. The hemichordata, specifically the enteropneusta (acorn worms), have been considered a sister group to the chordata. Enteropneusts combine various chordate features (e.g. lateral gill openings, dorsal nerve cord) with features that are usually associated with gastroneuralian invertebrates (e.g. dorsal heart, circumenteric nerve ring, ventral nerve cord). Here we analyse various morphological and functional characteristics that enteropneusts share with either invertebrates or chordates in the light of our recent proposal that the chordata may derive – by bodily dorsoventral inversion – from a gastroneuralian ancestor. We show that many seemingly non-chordate features of enteropneusts will align with similar features in the chordates – provided that we compare the ventral side of an enteropneust to the dorsal side of a chordate. This inversion proposes several interesting and new putative homologies between enteropneusts and acranian chordates, such as between their epibranchial ridge/endostyle (later thyroid gland), their postanal tails, atrial walls, and also between the chordates' dorsal notochord and the enteropneusts' posteroventral pygochord. Significantly, positional homology between notochord and pygochord is also supported by the expression domains of Brachyury orthologs in vertebrates and invertebrates: a Brachyury ortholog is active in the postero ventral mesoderm in Drosophila and in the dorsal mesoderm in chordates. In conclusion, we propose that the anatomy of enteropneusts may serve as a conceptual 'missing link' between gastroneuralian invertebrates and notoneuralian chordates. We discuss whether the enteropneust's dorsoanterior nervous centre plus their ventral trunk cord then corresponds to brain and dorsal nerve cord in the chordata.     

Adaptive Evolution of Deep-Sea Amphipods from the Superfamily Lysiassanoidea in the North Atlantic

In this study we reconstruct phylogenies for deep sea amphipods from the North Atlantic in order to test hypotheses about the evolutionary mechanisms driving speciation in the deep sea. We sequenced five genes for specimens representing 21 families. Phylogenetic analyses showed incongruence between the molecular data and morphological taxonomy, with some morphologically distinct taxa showing close molecular similarity. Approximate dating of nodes based on available calibration suggested adaptation to the deep sea around the Cretaceous-Palaeogene boundary, with three identified lineages within the deep-sea radiation dating to the Eocene–Oligocene transition. Two of those lineages contained species currently classified in multiple families. We reconstructed ancestral nodes based on the mouthpart characters that define trophic guilds (also used to establish the current taxonomy), and show a consistent transition at the earliest node defining the deep-sea lineage, together with increasing diversification at more recent nodes within the deep-sea lineage. The data suggest that the divergence of species was adaptive, with successive diversification from a non-scavenging ancestor to ‘opportunistic’, ‘obligate’ and ‘specialised’ scavengers. We propose that the North Atlantic species studied provide a strong case for adaptive evolution promoted by ecological opportunity in the deep sea.     

The island rule and the evolution of body size in the deep sea

Aim  Our goal is to test the generality of the island rule – a graded trend from gigantism in small-bodied species to dwarfism in large-bodied species – in the deep sea, a non-insular but potentially analogous system.
Location  Shallow-water and deep-sea benthic habitats in the western Atlantic Ocean from the North to South Poles.
Methods  We conducted regression analyses of body size of deep-sea gastropods species relative to their shallow-water congeners using measurements from the Malacolog ver. 3.3.3 database.
Results  Our results indicate that, consistent with the island rule, gastropod genera with small-bodied, shallow-water species have significantly larger deep-sea representatives, while the opposite is true for genera that are large-bodied in shallow water. Bathymetric body size clines within the deep sea are also consistent with predictions based on the island rule.
Main conclusions  Like islands, the deep sea is characterized by low absolute food availability, leading us to hypothesize that the island rule is a result of selection on body size in a resource-constrained environment. The body size of deep-sea species tends to converge on an optimal size for their particular ecological strategy and habitat.     

A synthesis of genetic connectivity in deep‐sea fauna and implications for marine reserve design

With anthropogenic impacts rapidly advancing into deeper waters, there is growing interest in establishing deep‐sea marine protected areas (MPAs) or reserves. Reserve design depends on estimates of connectivity and scales of dispersal for the taxa of interest. Deep‐sea taxa are hypothesized to disperse greater distances than shallow‐water taxa, which implies that reserves would need to be larger in size and networks could be more widely spaced; however, this paradigm has not been tested. We compiled population genetic studies of deep‐sea fauna and estimated dispersal distances for 51 studies using a method based on isolation‐by‐distance slopes. Estimates of dispersal distance ranged from 0.24 km to 2028 km with a geometric mean of 33.2 km and differed in relation to taxonomic and life‐history factors as well as several study parameters. Dispersal distances were generally greater for fishes than invertebrates with the Mollusca being the least dispersive sampled phylum. Species that are pelagic as adults were more dispersive than those with sessile or sedentary lifestyles. Benthic species from soft‐substrate habitats were generally less dispersive than species from hard substrate, demersal or pelagic habitats. As expected, species with pelagic and/or feeding (planktotrophic) larvae were more dispersive than other larval types. Many of these comparisons were confounded by taxonomic or other life‐history differences (e.g. fishes being more dispersive than invertebrates) making any simple interpretation difficult. Our results provide the first rough estimate of the range of dispersal distances in the deep sea and allow comparisons to shallow‐water assemblages. Overall, dispersal distances were greater for deeper taxa, although the differences were not large (0.3–0.6 orders of magnitude between means), and imbalanced sampling of shallow and deep taxa complicates any simple interpretation. Our analyses suggest the scales of dispersal and connectivity for reserve design in the deep sea might be comparable to or slightly larger than those in shallow water. Deep‐sea reserve design will need to consider the enormous variety of taxa, life histories, hydrodynamics, spatial configuration of habitats and patterns of species distributions. The many caveats of our analyses provide a strong impetus for substantial future efforts to assess connectivity of deep‐sea species from a variety of habitats, taxonomic groups and depth zones.     

Pockmarks enhance deep‐sea benthic biodiversity: a case study in the western Mediterranean Sea

Aim Pockmarks are craters on the sea floor formed by sub‐sea‐floor fluid expulsions, which occur world‐wide at all ocean depths. These habitats potentially host a highly specialized fauna that can exploit the hydrocarbons released. Pockmarks at relatively shallow depths can be easily destroyed by human activities, such as bottom trawling. In the present study, we investigated the combined effects of sea‐floor heterogeneity, rate of fluid emission and trophic conditions of different pockmarks on the biodiversity of the deep‐sea assemblages. Location Continental slope of the Gulf of Lions, western Mediterranean Sea, at water depths from 265 to 434 m. Methods We investigated the biodiversity associated with sea‐floor pockmarks that are both inactive and that have active gas emissions. Control sites were selected on the sea floor outside the influence of the gas seepage, both within and outside the pockmark fields. We examined the combined effects of: (i) sea‐floor heterogeneity; (ii) variable levels of fluid (gas) emissions; and (iii) trophic characteristics of the meiofaunal assemblage structure and nematode diversity. Results Sediments within the pockmark fields had lower meiofaunal abundance and biomass when compared with the surrounding sediments that were not influenced by the gas seepage. Although several higher taxa were absent in the pockmarks (e.g. Turbellaria, Tardigrada, Cumacea, Isopoda, Tanaidacea, Nemertina and Priapulida, which were present in the control areas), the richness of the nematode species within all of these pockmarks was very high. About 25% of the total species encountered in the deep‐sea sediments of the investigated areas was exclusively associated with these pockmarks. Main conclusions We conclude that both active and inactive pockmarks provide significant contributions to the regional (gamma) diversity of the continental slope in the western Mediterranean Sea, and thus the protection of these special and fragile habitats is highly relevant to the conservation of deep‐sea biodiversity.     

Filter feeding,deviations from bilateral symmetry,developmental noise,and heterochrony of hemichordate and cephalochordate gills

We measured gill slit fluctuating asymmetry (FA), a measure of developmental noise, in adults of three invertebrate deuterostomes with different feeding modes: the cephalochordate Branchiostoma floridae (an obligate filter feeder), the enteropneusts Protoglossus graveolens (a facultative filter feeder/deposit feeder) and Saccoglossus bromophenolosus (a deposit feeder). FA was substantially and significantly low in B. floridae and P. graveolens and high in S. bromophenolosus. Our results suggest that the gills of species that have experienced a relaxation of the filter feeding trait exhibit elevated FA. We found that the timing of development of the secondary collagenous gill bars, compared to the primary gill bars, was highly variable in P. graveolens but not the other two species, demonstrating an independence of gill FA from gill bar heterochrony. We also discovered the occasional ectopic expression of a second set of paired gills posterior to the first set of gills in the enteropneusts and that these were more common in S. bromophenolosus. Moreover, our finding that gill slits in enteropneusts exhibit bilateral symmetry suggests that the left‐sidedness of larval cephalochordate gills, and the directional asymmetry of Cambrian stylophoran echinoderm fossil gills, evolved independently from a bilaterally symmetrical ancestor.     

Evolutionary and biogeographical patterns of barnacles from deep‐sea hydrothermal vents

The characterization of evolutionary and biogeographical patterns is of fundamental importance to identify factors driving biodiversity. Due to their widespread but discontinuous distribution, deep‐sea hydrothermal vent barnacles represent an excellent model for testing biogeographical hypotheses regarding the origin, dispersal and diversity of modern vent fauna. Here, we characterize the global genetic diversity of vent barnacles to infer their time of radiation, place of origin, mode of dispersal and diversification. Our approach was to target a suite of multiple loci in samples representing seven of the eight described genera. We also performed restriction‐site associated DNA sequencing on individuals from each species. Phylogenetic inferences and topology hypothesis tests indicate that vent barnacles have colonized deep‐sea hydrothermal vents at least twice in history. Consistent with preliminary estimates, we find a likely radiation of barnacles in vent ecosystems during the Cenozoic. Our analyses suggest that the western Pacific was the place of origin of the major vent barnacle lineage, followed by circumglobal colonization eastwards through the Southern Hemisphere during the Neogene. The inferred time of radiation rejects the classic hypotheses of antiquity of vent taxa. The timing and the mode of origin, radiation and dispersal are consistent with recent inferences made for other deep‐sea taxa, including nonvent species, and are correlated with the occurrence of major geological events and mass extinctions. Thus, we suggest that the geological processes and dispersal mechanisms discussed here can explain the current distribution patterns of many other marine taxa and have played an important role shaping deep‐sea faunal diversity. These results also constitute the critical baseline data with which to assess potential effects of anthropogenic disturbances on deep‐sea ecosystems.     

Invertebrate population genetics across Earth's largest habitat: The deep‐sea floor

Despite the deep sea being the largest habitat on Earth, there are just 77 population genetic studies of invertebrates (115 species) inhabiting non‐chemosynthetic ecosystems on the deep‐sea floor (below 200 m depth). We review and synthesize the results of these papers. Studies reveal levels of genetic diversity comparable to shallow‐water species. Generally, populations at similar depths were well connected over 100s–1,000s km, but studies that sampled across depth ranges reveal population structure at much smaller scales (100s–1,000s m) consistent with isolation by adaptation across environmental gradients, or the existence of physical barriers to connectivity with depth. Few studies were ocean‐wide (under 4%), and 48% were Atlantic‐focused. There is strong emphasis on megafauna and commercial species with research into meiofauna, “ecosystem engineers” and other ecologically important species lacking. Only nine papers account for ~50% of the planet's surface (depths below 3,500 m). Just two species were studied below 5,000 m, a quarter of Earth's seafloor. Most studies used single‐locus mitochondrial genes revealing a common pattern of non‐neutrality, consistent with demographic instability or selective sweeps; similar to deep‐sea hydrothermal vent fauna. The absence of a clear difference between vent and non‐vent could signify that demographic instability is common in the deep sea, or that selective sweeps render single‐locus mitochondrial studies demographically uninformative. The number of population genetics studies to date is miniscule in relation to the size of the deep sea. The paucity of studies constrains meta‐analyses where broad inferences about deep‐sea ecology could be made.     

Abundance–occupancy relationships in deep sea wood fall communities

The generally positive relationship between the number of sites a species occupies and its average abundance within those sites provides an important link between population processes occurring at different spatial scales. Although such abundance–occupancy relationships (AORs) have been documented across a very wide range of taxa and in many different environments, little is known of such patterns in Earth's largest ecosystem, the deep sea. Wood falls – derived from natural or anthropogenic inputs of wood into the oceans – constitute an important deep‐sea habitat, habouring their own unique communities ultimately entirely dependent on the wood for chemical energy. In this study we take advantage of the unique features of an experimental wood fall deployment to examine AORs for the first time in deep‐sea invertebrates. The study design combines advantages of both experimental (tractability, control of key environmental parameters) and observational (natural colonisation by taxonomically diverse communities) studies. We show that the interspecific AOR is strongly positive across the 48 species occurring over 32 wood fall communities. The precise form of the AOR is mediated by both species‐level life history (body size) and by the colonisation stage at which communities were harvested, but not by environmental energy (wood fall size). Temporal dynamics within species are also generally consistent with positive intraspecific AORs. This support for positive AORs in the deep sea is an important extension of a macroecological generality into a new environment offering considerable potential for further testing and developing mechanistic macroecological theories.     

Taxonomic composition and zoogeographical aspects of deep sea fishes (90-540m) from the Gulf of California, Mexico

The Gulf of California has a high variety of ecosystems that allow different services and the fishery resources play a prominent role in its ecology, evolution and economics. Fish coastal species have been previously reported for most coastal areas, especially those species that are subject to fishing, however, little is known on the species from deep sea zones, due to sampling difficulties. We studied the deep sea fishes collected with trawl nets during three research surveys in the Gulf of California, Mexico in 2004-2005. We provide a systematic checklist and some notes on biogeographical aspects. For this, 74 fishing hauls were done, and a total of 9 898 fishes were captured, belonging to two classes, 15 orders, 35 families, 53 genera and 70 species. The best represented families in number of species were: Paralichthyidae (eight), Serranidae (six), and Scorpaenidae and Triglidae with five species each one. The typical families from deep waters were: Ophidiidae, Moridae, Lophiidae, Scorpaenidae, Triglidae, Paralichthydae, Pleuronectidae and Cynoglossidae. Size range varied from 13cm for the Splinose searobin (Bellator xenisma) to 234cm in the Pacific Cutlassfish (Trichiurus nitens). The biogeographical affinity showed that species with affinity to the East Tropical Pacific (ETP) dominated, followed by species from San Diego-Panamic, San Diego-Panamic-Peruvian-Chilean and Oregonian-Cortes provinces, respectively. A biogeographic overlap was found in the fauna, which reflects the Gulf of California's geographical position, with distribution limits of species from temperate, tropical and warm-temperature transition affinities, divisions that characterize the Gulf of California. Taxonomic status of fish with a focus on composition, location, characterization and zoogeography are fundamental to any subject of biodiversity and fisheries management actions.     

The Halomonhystera disjuncta population is homogeneous across the Håkon Mosby mud volcano (Barents Sea) but is genetically differentiated from its shallow‐water relatives

The deep sea has a high biodiversity and a characteristic bathyal fauna. Earlier evidence suggested that at least some shallow‐water species invaded the ecosystem followed by radiation leading to endemic deep‐sea lineages with a genetic and/or morphological similarity to their shallow‐water counterparts. The nematode Halomonhystera disjuncta has been reported from shallow‐water habitats and the deep sea [Håkon Mosby mud volcano (HMMV)], but the morphological features and the phylogenetic relationships between deep‐sea and shallow‐water representatives remain largely unknown. Furthermore, nothing is known about the genetic structure of the H. disjuncta population within the HMMV. This study is the first integrative approach in which the morphological and phylogenetic relationships between a deep‐sea and shallow‐water free‐living nematode species are investigated. To elucidate the phylogenetic relationships, we analysed the mitochondrial gene Cytochrome oxidase c subunit I (COI) and three nuclear ribosomal genes (Internal Transcribed Spacer region, 18S and the D2D3 region of 28S). Our results show that deep‐sea nematodes comprise an endemic lineage compared to the shallow‐water representatives with different morphometric features. COI genetic divergence between the deep‐sea and shallow‐water specimens ranges between 19.1% and 25.2%. Taking these findings into account, we conclude that the deep‐sea form is a new species. amova revealed no genetic structure across the HMMV, suggesting that nematodes are able to disperse efficiently in the mud volcano.     

Interannual Recruitment Dynamics for Resident and Transient Marsh Species: Evidence for a Lack of Impact by the Macondo Oil Spill

The emulsification of oil at the Deepwater Horizon (DWH) well head relegated a large proportion of resultant hydrocarbon plumes to the deep sea, facilitated the incorporation of oil droplets into microbial and planktonic food web, and limited the severity of direct, wetland oiling to coastal Louisiana. Nevertheless, many transient fish and invertebrate species rely on offshore surface waters for egg and larval transport before settling in coastal habitats, thereby potentially impacting the recruitment of transient species to coastal nursery habitats quite distant from the well site. We compared the utilization of salt-marsh habitats by transient and resident nekton before and after the DWH accident using data obtained from an oyster reef restoration project in coastal Alabama. Our sampling activities began in the summer preceding the DWH spill and continued almost two years following the accident. Overall, we did not find significant differences in the recruitment of marsh-associated resident and transient nekton in coastal Alabama following the DWH accident. Our results, therefore, provide little evidence for severe acute or persistent oil-induced impacts on organisms that complete their life cycle within the estuary and those that spent portions of their life history in potentially contaminated offshore surface waters prior to their recruitment to nearshore habitats. Our negative findings are consistent with other assessments of nekton in coastal vegetated habitats and bolster the notion that, despite the presence of localized hydrocarbon enrichments in coastal habitats outside of Louisiana the most severe oil impacts were relegated to coastal Louisiana and the deep sea. Analyzing all the information learned from this accident will undoubtedly provide a synthesis of what has or has not been affected in the Northern Gulf of Mexico, which when put in context with oil spill studies elsewhere should improve our ability to avert and manage the negative consequences of such accidents.     

Investigating the environmental drivers of deep‐seafloor biodiversity: A case study of peracarid crustacean assemblages in the Northwest Atlantic Ocean

The deep‐sea benthos covers over 90% of seafloor area and hosts a great diversity of species which contribute toward essential ecosystem services. Evidence suggests that deep‐seafloor assemblages are structured predominantly by their physical environment, yet knowledge of assemblage/environment relationships is limited. Here, we utilized a very large dataset of Northwest Atlantic Ocean continental slope peracarid crustacean assemblages as a case study to investigate the environmental drivers of deep‐seafloor macrofaunal biodiversity. We investigated biodiversity from a phylogenetic, functional, and taxonomic perspective, and found that a wide variety of environmental drivers, including food availability, physical disturbance (bottom trawling), current speed, sediment characteristics, topographic heterogeneity, and temperature (in order of relative importance), significantly influenced peracarid biodiversity. We also found deep‐water peracarid assemblages to vary seasonally and interannually. Contrary to prevailing theory on the drivers of deep‐seafloor diversity, we found high topographic heterogeneity (at the hundreds to thousands of meter scale) to negatively influence assemblage diversity, while broadscale sediment characteristics (i.e., percent sand content) were found to influence assemblages more than sediment particle‐size diversity. However, our results support other paradigms of deep‐seafloor biodiversity, including that assemblages may vary inter‐ and intra‐annually, and how assemblages respond to changes in current speed. We found that bottom trawling negatively affects the evenness and diversity of deep‐sea soft‐sediment peracarid assemblages, but that predicted changes in ocean temperature as a result of climate change may not strongly influence continental slope biodiversity over human timescales, although it may alter deep‐sea community biomass. Finally, we emphasize the value of analyzing multiple metrics of biodiversity and call for researchers to consider an expanded definition of biodiversity in future investigations of deep‐ocean life.     

一份南海深海沉积物样品中可培养细菌的多样性

海洋沉积环境蕴含丰富的微生物资源。对深海难培养微生物的分离培养,不仅有利于深海微生物资源的挖掘与利用,也有利于对深海微生物学的研究。本研究采用多种培养基分离获得细菌菌株纯培养,并通过16S r RNA基因序列鉴定,对我国南海海域1个4 000 m水深的深海表层沉积物样品的可培养细菌多样性进行初探。共设计23种分离培养基,经过选择性分离培养最终获得612株细菌菌株,分别隶属于厚壁菌门(Firmicutes)、放线菌门(Actinobacteria)和拟杆菌门(Bacteroidetes)的9目10科27个属级类群,可培养优势类群为厚壁菌门,占所有分离物种数量的85.8%,包含13个16S rRNA基因序列相似性低于98%的潜在新物种。海洋琼脂类培养基适合培养不同种类的海洋细菌类群,放线菌选择性分离类合成培养基对放线菌类群的分离效果较好。最终获得一些与具有产抗生素、细胞毒素、高效酶活、耐受不良环境、降解污染物等特殊功能微生物相近的菌株。研究结果表明,该深海沉积物样品的可培养微生物资源、潜在新物种和微生物生理特性丰富多样,研究深海环境难培养微生物的分离策略及其微生物适应生理特性对研究极端环境微生物打下了基础。     

Southern Ocean diversity: new paradigms from molecular ecology

Southern Ocean biodiversity reflects past climate, oceanographic, and tectonic changes. Molecular data from contemporary populations carry signatures of these processes. Here, we review new molecular studies on Southern Ocean benthic fauna. Many of these studies focus on species with extensive geographic or bathymetric distributions, and resolve taxonomic questions. Reviewing all available data, we show that, in addition to reflecting life-history characteristics, the molecular signals found in these studies provide an insight into how species survived the last glacial maximum (LGM). We identify molecular signatures that are characteristic of surviving glacial cycles in small refugia on the continental shelf and distinguish them from molecular signatures that are indicative of surviving glacial cycles in the deep sea.     

Spatial and temporal patterns of genetic variation in the widespread antitropical deep‐sea coral Paragorgia arborea

Numerous deep‐sea species have apparent widespread and discontinuous distributions. Many of these are important foundation species, structuring hard‐bottom benthic ecosystems. Theoretically, differences in the genetic composition of their populations vary geographically and with depth. Previous studies have examined the genetic diversity of some of these taxa in a regional context, suggesting that genetic differentiation does not occur at scales of discrete features such as seamounts or canyons, but at larger scales (e.g. ocean basins). However, to date, few studies have evaluated such diversity throughout the known distribution of a putative deep‐sea species. We utilized sequences from seven mitochondrial gene regions and nuclear genetic variants of the deep‐sea coral Paragorgia arborea in a phylogeographic context to examine the global patterns of genetic variation and their possible correlation with the spatial variables of geographic position and depth. We also examined the compatibility of this morphospecies with the genealogical‐phylospecies concept by examining specimens collected worldwide. We show that the morphospecies P. arborea can be defined as a genealogical‐phylospecies, in contrast to the hypothesis that P. arborea represents a cryptic species complex. Genetic variation is correlated with geographic location at the basin‐scale level, but not with depth. Additionally, we present a phylogeographic hypothesis in which P. arborea originates from the North Pacific, followed by colonization of the Southern Hemisphere prior to migration to the North Atlantic. This hypothesis is consistent with the latest ocean circulation model for the Miocene.