The genus Grimpoteuthis (Octopoda: Grimpoteuthidae) in the north-east Atlantic, with descriptions of three new species

The systematics and distribution of the cirrate octopod genus Grimpoteuthis in the north-east Atlantic are reviewed. Three new species are described and Grimpoteuthis wuelkeri (Grimpe, 1920) is redescribed. A new generic diagnosis is proposed. Five species of Grimpoteuthis are recognized in the north-east Atlantic. The type species, G. umbellata (Fisher, 1883) is known only from the type specimen, which is in such poor condition that comparison with recently captured material was not possible. G. wuelkeri is a large, slope species, caught between 1600 m and 2200 m in the north-east and north-west Atlantic. Of the three new species, both G. boylei and G. challengeri are large abyssal species. G. boylei is found in the north-east Atlantic at the Porcupine Abyssal Plain (PAP) and the Madeira Abyssal Plain and may be found at abyssal depths throughout the north-east Atlantic. G. challengeri is known from the PAP, with a single specimen from the north-west Atlantic. G. discoveryi is a small, lower slope and abyssal species found in the north-east Atlantic. The Grimpoteuthis species can be separated based on shell form, presence of a radula and posterior salivary glands, arrangement of suckers and cirri and gill morphology. Two species, G. megaptera Verrill and G. plena Verrill, have been described from the north-west Atlantic, but the types are either lost ( G. megaptera ) or in poor condition ( G. plena ), hindering comparisons. Material examined from the north-west Atlantic included G. wuelkeri , G. challengeri and at least two other species.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 139 , 93−127.     

A new deep-sea pennatulacean (Anthozoa: Octocorallia: Chunellidae) from the Porcupine Abyssal Plain (NE Atlantic)

During the BENGAL cruises, an important collection of deep-sea benthic organisms was sampled. Among the pennatulacean colonies, a previously undescribed species of chunellid was collected. That material is here described as the type species of a new genus, Porcupinella gen. nov. The new genus and species are described based on material collected in the Porcupine Abyssal Plain (NE Atlantic), 4,839–4,847 m in depth. This is the first time that a chunellid is reported from the Atlantic Ocean. The new genus is compared with the other genera in the family, and some phylogenetic remarks about the families Chunellidae and Umbellulidae are also provided.     

The Lepocreadiidae (Digenea) of fishes from the north-east Atlantic: Profundivermis intercalarius n. g., n. sp. from the marine fish Coryphaenoides (Nematonurus) armatus (Hector) (Macrouridae) from the Porcupine Abyssal Plain

A new genus and species, Profundivermis intercalarius, is described from the deep-sea fish Coryphaenoides (Nematonurus) armatus from 4,850 m deep on the Porcupine Abyssal Plain, NE Atlantic. The genus is characterized by an inter-testicular ovary and is the only member of the subfamily Lepidapedinae to show this feature. The relationships of the members of the subfamily are analysed phylogenetically. Members of the subfamily are often found in deep-sea gadiforms.     

Deep-sea scavenging demersal fish fauna of the Nazaré Canyon system, Iberian coast, north-east Atlantic Ocean

A baited imaging lander was deployed six times in the Nazaré Canyon at depths from 909 to 4361 m during August 2005 to investigate the demersal scavenging fishes. Species observed and lander-derived abundance estimates were similar to previous data from the Porcupine Seabight and abyssal plain, north-east Atlantic Ocean.     

Ecology and Gametogenic Biology of the Genus Umbellula (Pennatulacea) in the North Atlantic Ocean

Four putative species of the seapen genus Umbellula, U. lindahli, U. thomsonii, U. durissima and U. monocephalus, are found at bathyal and abyssal depths in the Porcupine Seabight and Abyssal Plain in the NE Atlantic Ocean and at bathyal depths off the Bahamas. The most common species, U. lindahli, has a cosmopolitan distribution in the world ocean. In situ observations, from bathyal depths in the Bahamas, of the posture of U. lindahli show that the polyps are spread out to maximize particle capture from the water column. In addition, the individuals of U. lindahli from the Bahamas suggest that the tentacles are cropped by a predator, but there is no evidence of predation in the NE Atlantic specimens. Reproduction in U. lindahli is typical of that previously observed in pennatulids. Fecundity is high, although most of the oocytes present are previtellogenic with only a few developing through to maximum size at any one time. Some specimens of U. lindahli appear to be infested with a small flatworm.     

Micro- and Nanoforaminifera from Abyssal Northeast Atlantic Sediments: A Preliminary Report

Rose Bengal stained benthic foraminifera which pass through a 63 μm mesh (microforaminifera and nanoforaminifera) have been extracted by handsorting the fine sieve residues (> 45 μm, 31 μm, 28 μm, 20 μm, 15 μm) of abyssal sediment samples. The samples were collected using a multiple corer in four areas of the northeast Atlantic between 31° N and 59° N. The abundance of these minute foraminifera varied from 2 specimens per 1 cm² (Madeira Abyssal Plain) to > 110 per 1 cm² (BIOTRANS area). They include a variety of taxa, the most common being certain rotaliid species, hormosinaceans and other multilocular agglutinated forms, the unilocular agglutinated genus Lagenammina, soft-bodied agglutinated sphaeres and flasks (saccamminids and psammosphaerids) and allogromiids. Some specimens are < 63 μm in maximum dimension but others belonging to elongate taxa are longer. Two samples taken 10 cm apart on the Porcupine Abyssal Plain suggest that minute foraminifera may be patchily distributed on a small scale. One sample, which was overlain by substantial amounts of phytodetritus, contained > 100 stained specimens (> 30 per 1 cm²) while the other, in which much less phytodetritus was present, yielded only 10 specimens (2.9 per 1 cm²). This observation suggests that some micro- and nanoforaminifera may flourish in the presence of decaying organic matter, perhaps consuming the associated bacteria. The presence of phytodetritus may also explain why two of our samples from the Madeira Abyssal Plain (MAP) contained an order of magnitude more stained tiny foraminifera than two other MAP samples in which phytodetritus was absent.     

Deep-Sea Macrobenthic Communities at Contrasting Sites off Portugal,Preliminary Results: I Introduction and Diversity Comparisons

Species diversity, faunal composition and abundance in the deep-sea macrobenthos are investigated at two, adjoining deep-sea areas off Portugal experiencing contrasting hydrodynamic conditions in order to test for the importance of disturbance in structuring the community. The first was located at 5,035 m depth on a presumably low current-energy site on the Tagus Abyssal Plain (TAP) and the second was centered in the lower end of the Setubal Canyon (SC) at 3,400 m depth where high levels of hydrodynamic activity are inferred from ripple bedforms visible in sea bed photographs. Closely grouped, transponder-mapped, “vegematic” box core samples were taken at each site. Agglutinated foraminifers numerically dominate the macrofauna (retained by 300 μm sieve) at both sites. Seagrass content in the SC cores indicates downslope detrital input from shallow water which is absent in the TAP samples. This must contribute, possibly with high levels of resuspended particles in near-bed flow, to the almost tenfold difference in abundance of metazoan macrofauna at the two sites. Expected species diversity measured from rarefaction is highest at TAP, both for pooled data and when foraminifers. tanaids and bivalve are considered separately. Evenness (equitability) in species' abundances is also higher at TAP than at SC. These differences are compared with benthic community data from the hydrodynamically disturbed High Energy Benthic Boundary Layer (HEBBLE) site on the Nova Scotia continental rise, and from the Rockall Trough (N.E. Atlantic). In common with these sites, strong bed flow at SC is thought to result in relatively high abundances, lower species richness and greater unevenness by acting to periodically disturb and reduce populations. Differences in the degree to which different faunal groups seem to be affected might be related to differences in lifestyle, particularly burrowing depth.     

Comparison of the Vertical Distribution of Nematodes from Two Contrasting Abyssal Sites in the Northeast Atlantic Subject to Different Seasonal Inputs of Phytodetritus

The vertical distribution of nematodes in the sediments of three cores from the phytodetritus-influenced Porcupine Abyssal Plain station is compared with three cores from the Madeira Abyssal Plain station in the DEEPSEAS programme. Nematode vertical distributions are compared with sediment organic chemistry data sampled at the same time from the same sites. The results support the two hypotheses erected by Thiel (1983) that the penetration of meiofauna into deep-sea sediments might be correlated with the input of organic matter and that a greater biological activity of larger organisms would increase the penetration of nematodes into the sediment through increase downward transport of food material.     

Walking with worms: coral‐associated epifaunal nematodes

Aims To study the community structure and habitat preferences of the Epsilonematidae and Draconematidae in coral degradation zones. To assess the contribution of different localities and microhabitats to meiobenthic diversity in such ecosystems. To discuss dispersive capacities and the occurrence of cryptic species in meiobenthic organisms. Location Porcupine Seabight (north‐east Atlantic Ocean; continental slope) and a transect along the Kenyan coast (Indian Ocean; shallow lagoon). Methods In the north‐east Atlantic, dead coral fragments, sponge skeletons and sediment were collected with a boxcorer. Along the Kenyan coast, dead coral fragments and coral gravel were collected during snorkelling and skin diving. Only nematodes belonging to the families Epsilonematidae and Draconematidae were considered. Community structure was analysed using multivariate techniques. Biodiversity was represented via rarefaction curves. Additive partitioning of species diversity was conducted. Turnover between microhabitats within locations and between locations within microhabitats were compared in a ternary plot. Results Twelve epsilonematid and five draconematid species were found in the Porcupine Seabight. In Kenya, 39 epsilonematid and 20 draconematid species were distinguished. Three species were found at both sampling locations. A table with the known distribution of all currently described species encountered in our study area is provided. At both sampling locations, the communities on coral fragments were significantly different from those in the other microhabitats, and were most diverse. In Kenya, species richness was mainly determined by local diversity and by turnover between localities. The contribution of β‐diversity decreased when abundance data were analysed. Turnover between microhabitats and between coral samples from different localities was higher than turnover between locations for gravel samples. Main conclusions Coral fragments were recognized as favourable substrata for typically epifaunal nematodes. Species‐specific habitat preferences were explained by finely tuned morphological adaptations. Our results suggest that cosmopolitan species could well be cryptic species, and this explanation for the existence of morphologically identical nematodes in geographically distant areas is weighed up against other plausible explanations. Coral degradation zones are an important source for new species of Epsilonematidae and Draconematidae. The addition of sampling locations contributed to the total number of species, although the added species were generally rare.     

The reproductive biology of two deep-water,reef-building scleractinians from the NE Atlantic Ocean

The reproductive ecology of colonies of Lophelia pertusa (Linné 1758) and Madrepora oculata Linné (1758) from the Porcupine Seabight (Thérèse Mound and South Porcupine Seabight site) and the Darwin Mounds (NE Rockall Trough—L. pertusa only) was investigated using histological techniques. Samples of L. pertusa exhibited seasonal reproduction, whereas the evidence for M. oculata is equivocal but suggests multiple cohorts of gamete production. L. pertusa produces a single cohort of around 3,000 oocytes, whereas M. oculata produces two cohorts, with a total fecundity of around 60 oocytes. The maximum observed oocyte size in L. pertusa was 140 μm and in M. oculata was 405 μm. From these oocyte sizes and the timing of reproduction, a lecithotrophic larva is expected, though not observed. This seasonality of reproduction fits with the phytodetrital food fall occurring around July in the Seabight area. L. pertusa was found to be non-reproductive at the Darwin Mound site. Though unable to be specifically tested, this may suggest that the increased trawling activity in this area might be keeping colonies below sexually viable sizes, as seen in numerous shallow water situations. All areas in the NE Atlantic are coming under threat from increased fishing and commercial exploration practices. This study shows that these highly seasonal reproducers could be sensitive to these fishing operations and care must be taken so as not to repeat the destruction that has occurred on shallower reefs.     

In situ comparison of activity in two deep-sea scavenging fishes occupying different depth zones

The activity of two scavenging deep-sea fishes occupying the same niche in overlapping depth zones were compared by in situ measurements of swimming speeds, tail-beat frequencies and by arrival time at baits. At 4800 m on the Porcupine Abyssal Plain, the grenadier Coryphaenoides (Nematonurus) armatus was the dominant scavenger, arriving at baits after 30 min, and swimming at relatively slow speeds of 0.17 body lengths (BL) sec^-1. At 2500 m in the relatively food rich Porcupine Seabight both C. (N.) armatus and the blue-hake, Antimora rostrata, were attracted to bait, but A. rostrata was always the first to arrive and most of the bait was consumed before the C. (N.) armatus arrived. A. rostrata swam at mean speeds of 0.39 BL sec^-1, similar to related shallow water species at equivalent temperatures. Observations on tail-beat frequency from video sequences confirmed the greater activity of A. rostrata. The data indicate that, given sufficient food supply, high pressure and low temperature do not limit activity levels of demersal deep-sea fishes. Low activity of C. (N.) armatus is an adaptation to poor food supply in the abyss, where these fishes dominate, but prevents it competing with the more active A. rostrata in shallower depths.     

The family Myriotrochidae Théel (Echinodermata: Holothurioidea) in the deep northeast Atlantic Ocean

Recent sampling in the Rockall Trough, Porcupine Seabight and Porcupine Abyssal Plain, in the NE Atlantic, has yielded 200 specimens of apodous holothurians belonging to seven species of the family Myriotrochidae Théel from depths between 1000 and 4310 m. These include the type species of a new genus and two new species of existing genera. Parvotrochus belyaevi gen. et sp. nov . is described from some minute specimens from the Rockall Trough that possess both exceptionally small wheel-like deposits and numerous, large curved rods in the body wall and tentacles. Myriotrochus clarki sp. Nov . has affinities to M. vitreus (M. Sars) but shows differences in the shape of the plates of the calcareous ring and in having larger, frequently abnormally formed, wheels. Siniotrochus myriodontus sp. Nov . is similar to the type species of this genus, S. phoxus, Pawson, but differs in the arrangement of the teeth on the wheels. A single small specimen of Prototrochus Belyaev et Mironov from the Whittard Canyon, northern Bay of Biscay, is similar to P. minutus (Östergren), a species known only from the Sea of Japan, but differs in some characteristics of the wheels and tentacles. A new subspecies, P. zenkevitchi rockallensis subsp. nov. , is proposed for a form of P. zenkevitchi (Belyaev) that was the most common myriotrochid encountered. P. zenkevitchi was known previously only from a few localities in the Pacific and S Atlantic deep-sea trenches. Similarly, two species previously known only from the NE Pacific, Myriotrochus bathybius H. L. Clark and M. giganteus H. L. Clark, are also recorded from the N Atlantic for the first time. The synonymy of M. giganteus and M. sp. ex gr. macquariensis-giganteus Belyaev et Mironov is proposed. The greatest number both of specimens and species came from the areas most intensively sampled. The present records show that several species have world-wide distributions. The wide geographic separation of many records is the result of both poor sampling effort in the deep sea, particularly with fine-meshed gear, and the difficulties in sampling infaunal animals in the deep sea. Box core samples in the Rockall Trough suggest that the myriotrochids are more common in this area than would be supposed from epibenthic sledge data. Hence, myriotrochids may be more prevalent in the deep sea than previously thought.     

Abyssal Megafaunal Necrophages: Latitudinal Differences in the Eastern North Atlantic Ocean

A pop-up time-lapse camera system (BATHYSNACK) was used to study megafaunal organisms attracted to bait at three abyssal localities in the eastern North Atlantic Ocean, a northern site on the Porcupine Abyssal Plain and two sub-tropical sites on the Madeira Abyssal Plain. Fish and decapod crustaceans appeared on 47% of photographs taken during deployments of up to seven days duration. The fishes Coryphaenoides (Nematonurus) armatus and Pachycara bulbiceps and the decapod Munidopsis spp. dominated appearances at the northern locality whereas over 95% of records at the two subtropical localities were of the decapod Plesiopenaeus armatus. Cyclical variations in necrophage abundance did not correlate with tidal cycles of near-bottom currents, although high P. armatus numbers were inversely related to current speed. Most of the frequently recorded entities showed contagious intraspecific distributions. In contrast, many of the interspecies relationships were non-random, with co-occurrences lower than expected, probably as a result of physical disturbance or potential predation. Population densities based on times of first arrival were calculated. Assemblages dominated by fish at higher latitudes and crustaceans at lower latitudes are discussed in terms of organic carbon supply to the deep sea and pelagic community structures.     

Postembryonic Morphology in Epsilonematidae, with a Discussion on the Variability of Caudal Gland Outlets

A new species of Akanthepsilonema and the first-stage juvenile of Glochinema trispinatum are described. Furthermore, additional morphological information is provided for Triepsilonema tripapillata. Animals originate from a cold-water coral degradation zone in the Porcupine Seabight area (North-East Atlantic Ocean). Akanthepsilonema sinecornibus sp. n. differs from A. heUeouetae in number of body annules, sexual dimorphism in amphid size, absence of copulatory thorns in males, absence of large spines and horns, shape of the copulatory apparatus, and position of ambulatory setae relative to vulva in females. The genus diagnosis for Akanthepsilonema is adjusted to incorporate the new species. Akanthepsilonema mainly differs from every other genus in the family by the combination of six rows of ambulatory setae situated around the vulva in females and eight subcephalic setae not displaced toward the anterior part of the head capsule. Small differences between the Papua New Guinea and the Porcupine Seabight populations of T. tripapillata indicate minimal intraspecific variability. Second-stage juveniles from Papua New Guinea have two rows of three ambulatory setae, whereas Porcupine Seabight specimens have two rows of four ambulatory setae. First- and fourth-stage juveniles of T. tripapillata are described for the first time. Literature data and personal observations showed that the molting of first-stage juveniles into second-stage juveniles and of third-stage juveniles into fourth-stage juveniles involves a decrease in the number of body rings, resulting in a loss of flexibility which is possibly compensated for by the development (I-II) or the doubling of the number of rows (III-IV) of ambulatory setae. This decrease is also linked with the formation of the head capsule and the smooth tail tip, although intergeneric variability is evident. The molting of second-stage juveniles into third-stage juveniles and of fourth-stage juveniles into adults is also subject to intergeneric variability. The variability in the number and orientation of caudal gland outlets among different nematode taxa is discussed. The presence of separate outlets for the caudal glands seems to be widespread within the family Epsilonematidae and has also been observed in various other, unrelated taxa of free-living aquatic nematodes, although their arrangement in Epsilonematidae is opposite. This aberrant arrangement is probably related to the aberrant locomotory pattern in this family.     

Subsurface microbiology and biogeochemistry of a deep, cold-water carbonate mound from the Porcupine Seabight (IODP Expedition 307)

The Porcupine Seabight Challenger Mound is the first carbonate mound to be drilled (∼270 m) and analyzed in detail microbiologically and biogeochemically. Two mound sites and a non-mound Reference site were analyzed with a range of molecular techniques [catalyzed reporter deposition-fluorescence in situ hybridization (CARD-FISH), quantitative PCR (16S rRNA and functional genes, dsrA and mcrA ), and 16S rRNA gene PCR-DGGE] to assess prokaryotic diversity, and this was compared with the distribution of total and culturable cell counts, radiotracer activity measurements and geochemistry. There was a significant and active prokaryotic community both within and beneath the carbonate mound. Although total cell numbers at certain depths were lower than the global average for other subseafloor sediments and prokaryotic activities were relatively low (iron and sulfate reduction, acetate oxidation, methanogenesis) they were significantly enhanced compared with the Reference site. In addition, there was some stimulation of prokaryotic activity in the deepest sediments (Miocene, > 10 Ma) including potential for anaerobic oxidation of methane activity below the mound base. Both Bacteria and Archaea were present, with neither dominant, and these were related to sequences commonly found in other subseafloor sediments. With an estimate of some 1600 mounds in the Porcupine Basin alone, carbonate mounds may represent a significant prokaryotic subseafloor habitat.     

New Deep-sea Leptognathiid Tanaids (Crustacea, Tanaidacea) from the North-east Atlantic

Three new genera ( Araphura, Leptognathioides and Stenotanais ) and six new species of tanaid crustaceans from below 1000 m depth in the north-east Atlantic are described. Two of the new species belong to the genus Leptognathiella Hansen, 1913, which is redescribed and removed from synonymy with Leptognathia Sars, 1882 ( sensu Lang 1968). These species are relatively common in the tanaid material examined and have a widespread distribution in the area under study, i.e. the Rockall Trough, Porcupine Seabight and Bay of Biscay. All six species are found at depths between 2070 and 2892 m. A further nine species previously belonging to the genus Leptognathia are removed to the new genera or to Leptognathiella. New records of Leptognathioides polita (Hansen, 1913) comb.n. and Leptognathiella abyssi (Hansen, 1913) have been obtained.     

Draconematidae (Nematoda) from cold-water corals in the Porcupine Seabight: The genus Cygnonema Allen & Noffsinger, 1978

Two new and closely related species of the genus Cygnonema Allen & Noffsinger, 1978 are described from a cold-water coral degradation zone in the Porcupine Seabight (NE Atlantic). Both species differ from C. steineri Allen & Noffsinger, 1978 by more pronounced pharyngeal and posterior swellings, a smaller body, a shorter pharynx in relation to body length, a higher number of CAT, and by the absence of a dorsal tooth. Cygnonema verum sp. n. differs from C. belgicae sp. n. by its greater body length, the relatively larger head capsule, a higher number of CAT, a more anteriorly positioned anteriormost laterodorsal CAT, a higher number of PAT, by the external labial sensilla being setiform, a higher number of subcephalic setae, and by a more anterior position of the amphidial fovea on the head capsule. Males of C. verum sp. n. are easily recognised by the presence of two large subventral, precloacal corniform setae. They also differ from males of C. belgicae sp. n. in the smaller amount of cytoplasm in the sperm cells, a knob-like capitulum, and a relatively shorter tail tip. The diagnosis of Cygnonema is emended, a dichotomic identification key to the three species is provided, and their biogeography is discussed.     

Hydrogenase Activity in Deeply Buried Sediments of the Arctic and North Atlantic Oceans

The goal of this study was to measure hydrogenase activity in oceanic sediments of the Lomonosov Ridge (Arctic Ocean) and of the Porcupine Seabight (North Atlantic), and to understand how its distribution varied depending on geochemical and lithological characteristics of the sediment. Hydrogenase activity was found at all sites, and absolute values and downhole distributions varied widely within and between sites. At the Lomonosov Ridge, hydrogenase was below detection in the top 190 meters of sediment, but high levels were measured in the organic carbon-rich layers below this depth. Activity at Challenger Mound, Porcupine Seabight (Site 1317) was one to two orders of magnitude higher than the upslope or downslope sites (1318 and 1316, respectively), and was higher in the mound than below the mound base. Consistent with the interpretation of hydrogenase as an indicator of microbial activity, cells were present in all hydrogenase-positive samples or in nearby horizons. Cell-specific activity values were as much as 1000 fold lower than those of cultured Clostridium pasteurianum, with the values from the highest activity sediments approaching those of C. pasteurianum more closely. This suggests that little adaptation in hydrogenase activity might be necessary to support H₂ metabolism in deeply buried sediments that contain relatively high levels of substrates.