Deep-sea Taxa Research Articles

Sexually dimorphic eye size in dragonfishes, a response to a bioluminescent signalling gap.

Deep-sea fishes must overcome extremely large nearest-neighbour distances and darkness to find mates. Sexual dimorphism in the size of luminescent structures in many deep-sea taxa, including dragonfishes (family Stomiidae), indicates reproductive behaviours may be mediated by visual signalling. This presents a paradox: if male photophores are larger, females may find males at shorter distances than males find females. Solutions to this gap may include females closing this gap or by males gathering more photons with a larger eye. We examine the eye size of two species of dragonfishes (Malacosteus niger and Phostomias guernei) for sexual dimorphism and employ a model of detection distance to evaluate the potential for such dimorphism to bridge the detection gap. This model incorporates the flux of sexually dimorphic postorbital photophores and eye lens size to predict detection distances. In both species, we found a significant visual detection gap in which females find males before males find females and that male lens size is larger, marking the second known case of size dimorphism in the actinopterygian visual system. Our results indicate the larger eye affords males a significant improvement in detection distance. We conclude that this dimorphic phenotype may have evolved to close the detection gap.

A new mid-Cretaceous hermit crab (Crustacea, Anomura) from Central Russia sheds new light on paguroid evolution

Paguroid anomurans constitute a large portion of modern-day decapod crustacean assemblages in all oceans and seas across the globe. However, their palaeontological record is still rather patchy, which partly explains why their phylogeny remains poorly understood. A newly collected specimen of an extinct paguroid is here shown to have a major impact on the evolutionary history of this group of anomurans. A new genus and species, Mutotylaspis tripudium, are here erected to accommodate a well-preserved, extraordinarily crab-like hermit crab from the lower Albian (mid-Cretaceous) of Vladimir Oblast (Central Russia) that appears closely related to the extant enigmatic deep-sea taxon Tylaspis anomala Henderson, 1885, as well as to other members of the extant family Probeebeidae Boone, 1926. Ongoing palaeontological research during the last two decades has provided ample evidence that paguroid evolution was much more intricate and complex than gleaned solely from extant hermit crabs. The newly collected specimen named herein is the first fossil member of the family Probeebeidae and constitutes an additional key species for unravelling paguroid evolution and extending the stratigraphical range of the family downwards into the mid-Cretaceous.

Effects of fixation and long-term preservation on finned octopods (Cephalopoda: Cirrata)

Among mollusks, the octopods stand out due to the almost entire absence of a stabilizing shell, potentially rendering these organisms susceptible to deformation caused by fixation and preservation. Such artifacts have previously been shown to occur especially in specimens of the deep-sea taxon Cirrata, the finned octopods. As an adaptation to their extreme habitat, many cirrates are composed of (semi-)gelatinous tissues, which are particularly prone to distortion, shrinkage, and deformation following fixation and preservation. Using one or more species from each of the eight currently recognized cirrate genera, the present study provides pre- and post-fixation color imagery of entire specimens as well as measurements of taxonomically relevant characters. These novel data illustrate the sometimes drastic effects that short- and long-term chemical treatment entails for this cephalopod taxon. Both structural and color changes may occur when finned octopods are fixed and preserved using routine protocols. However, shrinkage does not occur uniformly in all taxonomically relevant structures. The consequences of such a treatment for species descriptions and identification are discussed and existing anaesthetization, fixation and preservation protocols are summarized.

Adaptation to the shallow sea floor environment of a species of marine worms, Oligobrachia mashikoi, generally inhabiting deep-sea water

Beard worms from the family Siboglinidae, are peculiar animals and are known for their symbiotic relationships with sulfur bacteria. Most Siboglinids inhabit the deep-sea floor, thus making difficult to make any observations in situ. One species, Oligobrachia mashikoi, occurs in the shallow depths (24.5 m) of the Sea of Japan. Taking advantage of its shallow-water habitat, the first ecological survey of O. mashikoi was performed over a course of 7 years, which revealed that its tentacle-expanding behavior was dependent on the temperature and illuminance of the sea water. Furthermore, there were significantly more O. mashikoi with expanding tentacles during the nighttime than during the daytime, and the prevention of light eliminated these differences in the number of expending tentacles. These results confirmed that the tentacle-expanding behavior is controlled by environmental light signals. Consistent with this, we identified a gene encoding a photoreceptor molecule, neuropsin, in O. mashikoi, and the expression thereof is dependent on the time of day. We assume that the described behavioral response of O. mashikoi to light signals represent an adaptation to a shallow-water environment within the predominantly deep-sea taxon.

Gross anatomy of the Pacific hagfish, Eptatretus burgeri, with special reference to the coelomic viscera.

Hagfish (Myxinoidea) are a deep-sea taxon of cyclostomes, the extant jawless vertebrates. Many researchers have examined the anatomy and embryology of hagfish to shed light on the early evolution of vertebrates; however, the diversity within hagfish is often overlooked. Hagfish have three lineages, Myxininae, Eptatretinae, and Rubicundinae. Usually, textbook illustrations of hagfish anatomy reflect the morphology of the Myxininae lineage, especially Myxine glutinosa, with its single pair of external branchial pores. Here, we instead report the gross anatomy of an Eptatretinae, Eptatretus burgeri, which has six pairs of branchial pores, especially focusing on the coelomic organs. Dissections were performed on fixed and unfixed specimens to provide a guide for those doing organ- or tissue-specific molecular experiments. Our dissections revealed that the ventral aorta is Y-branched in E. burgeri, which differs from the unbranched morphology of Myxine. Otherwise, there were no differences in the morphology of the lingual apparatus or heart in the pharyngeal domain. The thyroid follicles were scattered around the ventral aorta, as has been reported for adult lampreys. The hepatobiliary system more closely resembled those of jawed vertebrates than those of adult lampreys, with the liver having two lobes and a bile duct connecting the gallbladder to each lobe. Overall, the visceral morphology of E. burgeri does not differ significantly from that of the known Myxine at the level of gross anatomy, although the branchial morphology is phylogenetically ancestral compared to Myxine.

Challenges and Advances in the Taxonomy of Deep-Sea Peracarida: From Traditional to Modern Methods

As one of the oldest branches of biology, taxonomy deals with the identification, classification and naming of living organisms, using a variety of tools to explore traits at the morphological and molecular level. In the deep sea, particular challenges are posed to the taxonomic differentiation of species. Relatively limited sampling effort coupled with apparent high diversity, compared to many other marine environments, means that many species sampled are undescribed, and few specimens are available for each putative species. The resulting scarce knowledge of intraspecific variation makes it difficult to recognize species boundaries and thus to assess the actual diversity and distribution of species. In this review article, we highlight some of these challenges in deep-sea taxonomy using the example of peracarid crustaceans. Specifically, we offer a detailed overview of traditional as well as modern methods that are used in the taxonomic analysis of deep-sea Peracarida. Furthermore, methods are presented that have not yet been used in peracarid taxonomy, but have potential for the analysis of internal and external structures in the future. The focus of this compilation is on morphological methods for the identification, delimitation and description of species, with references to molecular analysis included where relevant, as these methods are an indispensable part of an integrative taxonomic approach. The taxonomic impediment, i.e. the shortage of taxonomists in view of a high undescribed biodiversity, is discussed in the context of the existing large taxonomic knowledge gaps in connection with the increasing threat to deep-sea ecosystems. Whilst peracarid crustaceans are used here as an exemplary taxon, the methodology described has broad relevance to many other deep-sea taxa, and thus will support broader research into deep-sea biodiversity and ecology more widely.

Regional-scale genetic differentiation of the stony coral Desmophyllum dianthus in the southwest Pacific Ocean is consistent with regional-scale physico-chemical oceanography

Deep-sea corals are diverse and abundant in New Zealand’s EEZ and the southwest Pacific Ocean. We assessed genetic diversity and gene flow of the deep-sea scleractinian (stony cup coral) Desmophyllum dianthus in five areas (Kermadec Ridge, Louisville Seamount Chain, Chatham Rise, Campbell Plateau, Macquarie Ridge), and applied dispersal kernels estimated from drifters observations and seascape genetics to compare with observed connectivity patterns and regional physico-chemical data. We observed pronounced between-region genetic differentiation, high levels of self-recruitment, but also high connectivity between distant sites (e.g., at the Kermadec Ridge inside the New Zealand EEZ, and the Louisville Seamount Chain outside the New Zealand EEZ). Genetic diversity was marginally highest at the Kermadec Ridge, whilst populations on the Chatham Rise showed evidence of unique genetic diversity that may be driven by converging ocean currents. Overall, patterns of genetic connectivity were consistent with oceanographic predictions of dispersal routes. Seascape genetic analyses indicated that environmental variables most often related to physical seafloor habitat characteristics were important predictors of regional-scale genetic differentiation of D. dianthus. Our research shows that current spatial closures within the New Zealand EEZ and surrounding high seas areas do not encompass the extent of genetic diversity and connectivity for D. dianthus, which parallels research for other deep-sea taxa in the region. Future measures implemented to protect deep-sea coral biodiversity require consideration of complex connectivity patterns and migration routes amongst taxa, at various spatial scales. This study demonstrates that a combined gene flow, seascape genetics and dispersal kernel modelling approach can provide a robust evaluation of connectivity in deep-sea environments to inform management decision making at the scale of the southwest Pacific Ocean.

Two new xylophile cytheroid ostracods (Crustacea) from Kuril-Kamchatka Trench, with remarks on the systematics and phylogeny of the family Keysercytheridae, Limnocytheridae, and Paradoxostomatidae

Abstract Keysercythere reticulatasp. nov. and Redekea abyssalissp. nov., collected from the wood fall submerged in the Kuril-Kamchatka Trench (Northwestern Pacific), are only the second records of the naturally occurring, wood-associated ostracod fauna from a depth of over 5000 m. At the same time, K. reticulata is the second and R. abyssalis is the third representative of their respective genera. While Keysercythere Karanovic and Brandão, 2015 species are free-living, deep-sea taxa, all Redekea de Vos, 1953 live symbiotically on the body surface of wood-boring isopods, Limnoria spp. Since R. abyssalis is the only genus representative found in the deep sea, we hypothesize that its ancestor colonized this ecosystem as a result of the symbiotic relationship. Newly collected material enabled us to update molecular phylogeny of Cythreoidea based on 18S rRNA gene sequences, especially to clarify the current systematics of the families Keysercytheridae, Limnocytheridae, and Paradoxostomatidae. The resulting phylogenetic tree supports a close relationship between Keysercythere and Redekea and a distant relationship between two Limnocytheridae lineages, Timiriaseviinae and Limnocytherinae. Consequently, we propose a transfer of Redekea from Paradoxostomatidae to Keysercytheridae, and erecting of the two limnocytherid subfamilies onto the family level. The phylogenetic analysis also implies a close relationship between the nominal Limnocytherinae genus and Keysercythere+Redekea clade, albeit with a low posterior probability, requiring further studies to clarify this.

New data on crinoid assemblages from the oldest sedimentary rocks of the Polish Outer Carpathians (Jurassic-Cretaceous)

Crinoids (Crinoidea) preserved in the so-called Cieszyn Beds (Tithonian-Hauterivian; southern Poland) are relatively numerous, especially in some levels, but not highly diversified. They are absent in the Lower Cieszyn Shale (Tithonian) and the Upper Cieszyn Shale (Valanginian-Hauterivian). The Lower Cieszyn Limestone (Tithonian) yielded crinoid remains classified as Isocrinus cf. amblyscalaris (Thurmann) and Isocrinida indet. In the Upper Cieszyn Limestone (Berriasian) crinoids are much more numerous and diversified. The following taxa are recorded: Isocrinus? annulatus (Roemer), Balanocrinus subteres (Münster), B. cf. smithi Hess and Gale, and Hemicrinus sp. The presence of specimens resembling Isocrinus amblyscalaris (Thurmann) in the Tithonian strata is surprising since the last occurrence of this taxon was reported from the Kimmeridgian. The Lower Cieszyn Limestone crinoidal assemblage is dominated by isocrinids (Isocrinida). This probably reflects the much shallower sedimentary environment of these sediments compared to that of the Upper Cieszyn Limestone, where excepting isocrinids, hemicrinids (Hemicrinidae, Cyrtocrinida) typical for deep marine environments were found. It is consistent with the analytical results from the foraminiferal assemblages, among which the benthic forms known from shelfal and paralic epicontinental environments first dominated, and then were supplanted by deep-sea taxa at the turn of the Jurassic and Cretaceous.

Functional diversity and nutritional content in a deep-sea faunal assemblage through total lipid, lipid class, and fatty acid analyses.

Lipids are key compounds in marine ecosystems being involved in organism growth, reproduction, and survival. Despite their biological significance and ease of measurement, the use of lipids in deep-sea studies is limited, as is our understanding of energy and nutrient flows in the deep ocean. Here, a comprehensive analysis of total lipid content, and lipid class and fatty acid composition, was used to explore functional diversity and nutritional content within a deep-sea faunal assemblage comprising 139 species from 8 phyla, including the Chordata, Arthropoda, and Cnidaria. A wide range of total lipid content and lipid class composition suggested a diversified set of energy allocation strategies across taxa. Overall, phospholipid was the dominant lipid class. While triacylglycerol was present in most taxa as the main form of energy storage, a few crustaceans, fish, jellyfishes, and corals had higher levels of wax esters/steryl esters instead. Type and amount of energy reserves may reflect dietary sources and environmental conditions for certain deep-sea taxa. Conversely, the composition of fatty acids was less diverse than that of lipid class composition, and large proportions of unsaturated fatty acids were detected, consistent with the growing literature on cold-water species. In addition, levels of unsaturation increased with depth, likely suggesting an adaptive strategy to maintain normal membrane structure and function in species found in deeper waters. Although proportions of n-3 fatty acids were high across all phyla, representatives of the Chordata and Arthropoda were the main reservoirs of these essential nutrients, thus suggesting health benefits to their consumers.

Desk-based deep sea exploration

e-ISSN: 1948-6596 https://escholarship.org/uc/fb doi:10.21425/F59235175 Book Review Desk-based deep sea exploration Biological Sampling in the Deep Sea, by Malcolm R. Clark, Mireille Consalvey, and Ashley A. Rowden (editors), 2016 Wiley-Blackwell, 466 pp., ISBN: 9780470656747 “Well if you go one-on-one with a limpet you can learn an enormous amount. You can do that. How do you do that with a great white shark or blue whale? There’s this barrier to what I would call natural history.” Bob Paine, interviewed in 2013 1 As a macroecologist, I don't get out much. Grow- ing fat on Other People’s Data, the temptation is simply to sit and process and analyse and publish on whatever system provides the best API (Application Programming Interface). However, I have always shared Paine’s view that it is im- portant to have a feel for the natural history of your subject too. Starting out, I worked -- as was more or less obligatory at the time, for a macroe- cologist -- on birds. I'm no twitcher, but I do get birds. I enjoy watching them, and have a sense of their lives and of their interactions with the envi- ronment. There is no barrier to natural history there, especially working with a nice colour field guide at hand to remind me of the feathered real- ity of my rows of data. Shifting my focus to the macroecology of marine ecosystems, I maintained a similar rou- tine: greedily hoovering up data, keeping a stock of glossy pictures and natural history monographs close by. This was all well and good when working on the shelf seas. I had never been out for more than a few hours of boat based sampling, but the general process -- whether sampling by trawl, box core, or scuba -- was at least familiar enough that the idea of going ‘one-on-one’ with the organisms did not feel outlandish. But then I started to ven- ture off the shelf, into the world’s biggest ecosys- tems in the deep sea (e.g. Webb et al. 2010). In this alien setting, even something as apparently straightforward as casting and hauling in a net starts to seem unfeasible as you realise that the cables involved must be passing 3, 4, 5 km in length… I approached Clark et al.’s book, then, probably not as a member of its main target audi- ence. I had no intention of putting any of the de- tailed protocols for different sampling gears into practice 2 , but read it rather as someone hoping to get a feel for the labour that has gone into the data I analyse, and thus of its potential and limi- tations for macroecological research. Even viewed from this somewhat oblique angle, the book stands up very well. The first chapters pre- sent a broad overview of the ‘deep sea’, and in so doing reveal its complexities and heterogeneities. Tyler et al. (Chapter 1) provide an excellent pri- mer on the habitats of the deep sea floor, as well as a very readable history of key points in its dis- covery, from the early ‘heroic’ age through the first quantitative studies and on to the finer scale, more systematic approaches now possible using new imaging and geolocation technologies. For instance, they show how recent, high resolution exploration of the abyssal plain -- previously thought to be flat and boring -- has revealed in- teresting biogenic and abiotic structuring down to sub-metre scales. In the next chapter, Schiaparelli et al. move from habitats to organisms, with an overview of key deep sea taxa as well as of broad patterns in their distribution. This covers several topics likely to be of interest to macroecologists, including the widespread occurrence of both miniaturisation and of gigantism in the deep sea, driven largely by energetics. They also describe the feature that has really drawn me to deep sea macroecology: 1 http://www.biodiverseperspectives.com/2013/09/10/diverse-introspectives-a-conversation-with-bob-paine/ 2 Or so I thought: by a strange twist of fate I write this on board the RV Pelican in the Gulf of Mexico, a mile and more of cable stretching below us. But that’s another story. Frontiers of Biogeography 2017, 9.2, e35175 © the authors, CC-BY license

Bathymetric distributions of notothenioid fishes

There has been no comprehensive study of the bathymetry of notothenioid fishes. Therefore, I analyzed minimum and maximum depths and depth ranges for 128 of 142 species that collectively range from 0 to ≈3000 m. Means (and medians) for maximum depths are 176 m (75 m) for non-Antarctic, 511 m (360 m) for sub-Antarctic, and 963 m (899 m) for Antarctic species; medians are significantly different. Means (and medians) for depth ranges for the three groups are 140 (55 m), 470 (345 m), and 727 m (714 m), respectively, with significantly different medians. The mean maximum depths for the Cryonotothenioidea are: Nototheniidae—525 m, Harpagiferidae—88 m, Artedidraconidae—906 m, Bathydraconidae—1165 m, and Channichthyidae—910 m. If five species of Bathydraco, with a mean of 2098 m are excluded, the mean for the Bathydraconidae is 741 m. With the exceptions of the harpagiferids and Bathydraco, there is overlap of the 95% confidence intervals for the means of other families. Thirteen Antarctic and sub-Antarctic species with maximum depths and/or depth ranges ≥1500 m are especially deep-living: four nototheniids, four species of Bathydraco, three species of Pogonophryne, and two channichthyids. The most common depth range is 0–50 m for non-Antarctic, 200–400 m for sub-Antarctic and 600–800 m for Antarctic species. Three species have depth ranges >2000 m. Species diversity peaks at 50 m for non-Antarctic, 100 m for sub-Antarctic, and 500 m for Antarctic species. Notothenioids are not eurybathic compared with older deep-sea taxa but are eurybathic relative to marine percomorphs.

The pace of life in deep-dwelling squids

Coastal and epipelagic cephalopods are among the fastest growing invertebrates, with life cycles of typically 1 year or less. Evidence is accumulating that deep-sea taxa often live longer and grow slower than their shallow water relatives. We test the hypothesis that deep-sea squid show increased longevity and reduced growth rates compared to coastal and epipelagic species, by validation experiments and quantification of statolith increments of three deep-sea squids from the Monterey Submarine Canyon. The periodicity of statolith increment formation in coastal species is daily, but is unknown for deep-sea squid. Between 2010 and 2013, specimens of Chiroteuthis calyx, Galiteuthis phyllura and Octopoteuthis deletron were captured by remotely operated vehicles and trawl nets off California. ROV-captured living squid were immersed in tetracycline and kept alive in the lab for between 3 and 14 days. Correlating the number of elapsed days with the number of newly deposited statolith increments, and statolith growth after the fluorescent tetracycline mark, provided evidence of regular and daily increment deposition, in C. calyx and O. deletron. This relationship was less strong in G. phyllura and the one-increment-per-day hypothesis was not accepted for this species. Reconstructing growth rates based on statolith counts and wet weights from animals of a wide size range suggest that O. deletron is a slower growing squid (0.59% BW/day) than C. calyx (1.3% BW/day) and G. phyllura (1.2% BW/day). Octopoteuthis deletron matures at around two years, the oldest C. calyx was a mature male of 1.5 years and the eldest G. phyllura was 10 months and still immature. Maximum reported sizes for G. phyllura and C. calyx exceed those of our examined specimens, and therefore their longevity likely exceeds 2 years, in particular if the females brood their eggs. Our study supports the hypothesis that deeper living squid exhibit reduced growth rates and an increased longevity compared to shallow living species. We discuss these traits in the context of a life in the deep pelagic ocean.

The unique deep sea-land connection: interactive 3D visualization and molecular phylogeny of Bathyhedyle boucheti n. sp. (Bathyhedylidae n. fam.)-the first panpulmonate slug from bathyal zones.

The deep sea comprises vast unexplored areas and is expected to conceal significant undescribed invertebrate species diversity. Deep waters may act as a refuge for many relictual groups, including elusive and enigmatic higher taxa, but the evolutionary pathways by which colonization of the deep sea has occurred have scarcely been investigated. Sister group relationships between shallow water and deep sea taxa have been documented in several invertebrate groups, but are unknown between amphibious/terrestrial and deep-sea species. Here we describe in full and interactive 3D morphoanatomical detail the new sea slug species Bathyhedyle boucheti n. sp., dredged from the continental slope off Mozambique. Molecular and morphological analyses reveal that it represents a novel heterobranch gastropod lineage which we establish as the new family Bathyhedylidae. The family is robustly supported as sister to the recently discovered panpulmonate acochlidian family Aitengidae, which comprises amphibious species living along the sea shore as well as fully terrestrial species. This is the first marine-epibenthic representative among hedylopsacean Acochlidiida, the first record of an acochlidian from deep waters and the first documented panpulmonate deep-sea slug. Considering a marine mesopsammic ancestor, the external morphological features of Bathyhedyle n. gen. may be interpreted as independent adaptations to a benthic life style in the deep sea, including the large body size, broad foot and propodial tentacles. Alternatively, the common ancestor of Bathyhedylidae and Aitengidae may have been a macroscopic amphibious or even terrestrial species. We hypothesize that oophagy in the common ancestor of Aitengidae and Bathyhedylidae might explain the impressive ecological and evolutionary flexibility in habitat choice in the Acochlidiida.

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.

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 0.24km to 2028km with a geometric mean of 33.2km 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.

A starfish bed in the Middle Miocene Grand Bay Formation of Carriacou, The Grenadines (West Indies)

AbstractThe first starfish bed to be recognized from the Antilles is a lensoid body in the middle Miocene Grand Bay Formation of Carriacou, The Grenadines (West Indies). This unit was deposited in a turbidite basin in a region of active volcanism fed from one centre and preserves common deep-water taxa more typical of the Palaeozoic, such as crinoids and brachiopods. The starfish bed is a channel-fill deposit laid down in at least 150–200 m water depth, although the specimens may have been derived from shallower water. A goniasterid asteroid and an ophiacanthid ophiuroid have been recognized. The first articulated asteroid from the Antillean fossil record isParagonaster(?)haldixonisp. nov. In all skeletal features it appears close to the extant Atlantic speciesParagonaster grandisH. L. Clark andP. subtilis(Perrier), but differs in having a single row of rectangular abactinal ossicles extending to the arm tip; these are longer than wide. The brittlestar,Ophiocamax ventosasp. nov., is described on the basis of a fragmentary disc and arms from this deposit. The closest similarities are with the extant tropical western Atlantic speciesOphiocamax hystrixLyman andO. austeraVerrill. However, the new species has thorns covering the entire surface of dorsal arm plates, while arm spines have a multitude of small thorns, loosely arranged in numerous rows and dorsal arm plate shape differs markedly. The occurrence ofO. ventosasp. nov. suggests thatOphiocamaxhas been a deep-sea taxon at least since the Miocene.

The ink sac clouds octopod evolutionary history

Difficulties in elucidating the evolutionary history of the octopods have arisen from problems in identifying informative morphological characters. Recent classifications have divided the largest group, the incirrate octopods, into five groups. These include the pelagic superfamily Argonautoidea and three gelatinous pelagic families (Vitreledonellidae, Bolitaenidae, Amphitretidae). All benthic incirrate octopods have been accommodated in the family Octopodidae, itself divided into four subfamilies, Octopodinae, Eledoninae, Bathypolypodinae and Graneledoninae, which are defined by the presence or absence of an ink sac, and uniserial or biserial sucker arrangements on the arms. We used relaxed clock models in a Bayesian framework and maximum likelihood methods to analyse three nuclear and four mitochondrial genes of representatives from each of the previous subfamilies. Strong evidence indicates that the family Octopodidae is paraphyletic and contains the gelatinous pelagic families. The subfamilies of Octopodidae recognised in earlier works do not reflect evolutionary history. The following clades were supported in all analyses: (1) Eledone/Aphrodoctopus, (2) Callistoctopus/Grimpella/Macroctopus/Scaeurgus, (3) Abdopus/Ameloctopus/Amphioctopus/Cistopus/Hapalochlaena/Octopus, (4) Enteroctopus/Muusoctopus/Vulcanoctopus, (5) Vitreledonella/Japetella, (6) Southern Ocean endemic and deep-sea taxa with uniserial suckers. These clades form the basis for a suite of taxa assigned family taxonomic rank: Amphitretidae, Bathypolypodidae, Eledonidae, Enteroctopodidae, Megaleledonidae and Octopodidae sensu nov. They are placed within the superfamily Octopodoidea.

Molecular phylogeny of the Forcipulatacea (Asteroidea: Echinodermata): systematics and biogeography

We present a comprehensively sampled three-gene phylogeny of the monophyletic Forcipulatacea, one of three major lineages within the crown-group Asteroidea. We present substantially more Southern Hemisphere and deep-sea taxa than were sampled in previous molecular studies of this group. Morphologically distinct groups, such as the Brisingida and the Zoroasteridae, are upheld as monophyletic. Brisingida is supported as the derived sister group to the Asteriidae (restricted), rather than as a basal taxon. The Asteriidae is paraphyletic, and is broken up into the Stichasteridae and four primary asteriid clades: (1) a highly diverse boreal clade, containing members from the Arctic and sub-Arctic in the Northern Hemisphere; (2) the genus Sclerasterias; (3) and (4) two sister clades that contain asteriids from the Antarctic and pantropical regions. The Stichasteridae, which was regarded as a synonym of the Asteriidae, is resurrected by our results, and represents the most diverse Southern Hemisphere forcipulatacean clade (although two deep-sea stichasterid genera occur in the Northern Hemisphere). The Labidiasteridae is artificial, and should be synonymized into the Heliasteridae. The Pedicellasteridae is paraphyletic, with three separate clades containing pedicellasterid taxa emerging among the basal Forcipulatacea. Fossils and timing estimates from species-level phylogeographic studies are consistent with prior phylogenetic hypotheses for the Forcipulatacea, suggesting diversification of basal taxa in the early Mesozoic, with some evidence for more widely distributed ranges from Cretacous taxa. Our analysis suggests a hypothesis of an older fauna present in the Antarctic during the Eocene, which was succeeded by a modern Antarctic fauna that is represented by the recently derived Antarctic Asteriidae and other forcipulatacean lineages.

Low endemism, continued deep-shallow interchanges, and evidence for cosmopolitan distributions in free-living marine nematodes (order Enoplida)

BackgroundNematodes represent the most abundant benthic metazoa in one of the largest habitats on earth, the deep sea. Characterizing major patterns of biodiversity within this dominant group is a critical step towards understanding evolutionary patterns across this vast ecosystem. The present study has aimed to place deep-sea nematode species into a phylogenetic framework, investigate relationships between shallow water and deep-sea taxa, and elucidate phylogeographic patterns amongst the deep-sea fauna.ResultsMolecular data (18 S and 28 S rRNA) confirms a high diversity amongst deep-sea Enoplids. There is no evidence for endemic deep-sea lineages in Maximum Likelihood or Bayesian phylogenies, and Enoplids do not cluster according to depth or geographic location. Tree topologies suggest frequent interchanges between deep-sea and shallow water habitats, as well as a mixture of early radiations and more recently derived lineages amongst deep-sea taxa. This study also provides convincing evidence of cosmopolitan marine species, recovering a subset of Oncholaimid nematodes with identical gene sequences (18 S, 28 S and cox1) at trans-Atlantic sample sites.ConclusionsThe complex clade structures recovered within the Enoplida support a high global species richness for marine nematodes, with phylogeographic patterns suggesting the existence of closely related, globally distributed species complexes in the deep sea. True cosmopolitan species may additionally exist within this group, potentially driven by specific life history traits of Enoplids. Although this investigation aimed to intensively sample nematodes from the order Enoplida, specimens were only identified down to genus (at best) and our sampling regime focused on an infinitesimal small fraction of the deep-sea floor. Future nematode studies should incorporate an extended sample set covering a wide depth range (shelf, bathyal, and abyssal sites), utilize additional genetic loci (e.g. mtDNA) that are informative at the species level, and apply high-throughput sequencing methods to fully assay community diversity. Finally, further molecular studies are needed to determine whether phylogeographic patterns observed in Enoplids are common across other ubiquitous marine groups (e.g. Chromadorida, Monhysterida).