Deep-Sea Eukaryote Ecology of the Semi-Isolated Basins Off Japan

The Japanese archipelago is surrounded by the Pacific to the east, the Okhotsk Sea to the north, the Sea of Japan to the west and the Okinawa Trough to the south. The last three seas form semi-isolated deep basins, all with potentially tectonic origin but a different primary energy source as well as hydrographic and faunistic history. The Okhotsk Sea is connected to the Pacific through the deep straits between the Kurile Islands. As a result much of the fauna has links with that fauna found at similar depths in the Pacific. By contrast, the Sea of Japan was isolated from the main Pacific during the last ice age and became anoxic. Even today the link is only through narrow shallow straits. As a result the fauna is impoverished and is believed to be composed of cold-adapted eurybathic species rather than true deep-sea species. The deep-water fauna of both these seas derive their energy from sinking surface primary production. The Okinawa Trough has a much younger tectonic history than the Okhotsk Sea or the Sea of Japan. In the Okinawa Trough the most noticeable fauna is associated with hydrothermal activity and chemosynthesis forms the base of the food chain for the bathyal community. The variable nature of these three basins offers excellent opportunities for comparative studies of species diversity, biomass and production in relation to their ambient environment. This revised version was published online in August 2006 with corrections to the Cover Date.     

Fish parasites in the Arctic deep-sea: Poor diversity in pelagic fish species vs. heavy parasite load in a demersal fish

A total of 219 deep-sea fishes belonging to five families were examined for the parasite fauna and stomach contents. The demersal fish Macrourus berglax, bathypelagic Bathylagus euryops, and mesopelagic Argentina silus, Borostomias antarcticus, Chauliodus sloani, and Lampanyctus macdonaldi were caught at 243–708 m trawling depth in the Greenland and the Irminger Sea in 2002. A total of 21 different parasite species, six Digenea, one Monogenea, two Cestoda, seven Nematoda, one Acanthocephala, and four Crustacea, were found. The parasite diversity in the meso- and bathypelagic environment was less diverse in comparison to the benthal. Macrourus berglax had the highest diversity (20 species), usually carrying 4–10 different parasite species (mean 7.1), whereas Bathylagus euryops harbored up to three and Argentina silus, Borostomias antarcticus, Chauliodus sloani and Lampanyctus macdonaldi each up to two species. Most Digenea, Cestoda, Nematoda, Acanthocephala, and Crustacea are known from a wide host range. Several of the encountered parasites occurred at a very low prevalence (<10%), indicating that the studied deep-sea fishes are most probably not instrumental to complete the parasite life cycles in the area of investigation. It is suggested that the lack of nutrients in the meso- and bathypelagial limits the abundance of potential first intermediate hosts of nematodes and cestodes, resulting in low infestation rates even of widely distributed, non-specific species. In contrast, the higher biomass in the benthic deep-sea environment increases the availability of potential intermediate hosts, such as molluscs for the digeneans, resulting in increased parasite diversity. Because many deep-sea fish have a generalistic feeding behavior, the observed different parasite diversity reflects a different depth range of the fish and not necessarily a specific fish feeding ecology.     

Colonization of habitat islands in the deep sea: recruitment to glass sponge stalks

Biogenic structures in the deep sea often act as hard substratum 'islands’ for the attachment of encrusting fauna. At an abyssal station in the NE Pacific, stalks of hexactinellid sponges in the genus Hyalonema are habitat islands for species-rich epifaunal communities. An experimental study was conducted to (1) determine the colonization rates of artificial Hyalonema stalks, (2) compare the species composition and diversity of recruits to newly available substrata to that of the natural communities, and (3) examine the vertical distribution of recruits. Four sets of six artificial sponge stalks, constructed of Hyalonema spicules, were deployed at 4100 m depth for 3- to 5-month periods. There was no difference in net colonization or immigration rate among the four deployments. Colonization rates were similar to those reported for other deep-sea, hard substratum recruitment experiments. The taxa that recruited to the artificial stalks were a subset of the taxa found in natural communities. However, several taxa important in structuring natural communities did not recruit to the artificial stalks. The two taxa with the highest invasion rates, a calcareous foraminiferan (Cibicides lobatulus) and a serpulid polychaete (Bathyvermilia sp.), also were the two taxa with greatest relative abundance in natural communities. Vertical distributions of Cibicides and an agglutinated foraminiferan (Telammina sp.) were skewed towards the top of the artificial stalks, potentially because of active habitat selection. These results have several implications for natural Hyalonema stalk communities. Most importantly, species composition and abundance of individuals in the stalk communities appear to be maintained by frequent recruitment of a few common taxa and infrequent recruitment of many rare taxa. An argument is presented for temporal-mosaic maintenance of diversity in these deep-sea, hard substratum communities.     

Spatial patterns of biodiversity in the Black Sea: An assessment using benthic polychaetes

The current study broadens the biodiversity information available for the Black Sea and neighbouring regions and improves our knowledge about the polychaete biogeographic patterns to be discerned in them. There appears to be a well-defined zoogeocline from the Marmara Sea and Bosphorus Strait to the inner parts of the region (Azov Sea), depicted both as a multivariate pattern and in terms of species (or taxa) numbers. The emergent multivariate pattern complies, to a certain extent, with Jakubova’s (1935) views: three main sectors can be defined in the basin: (a) Prebosphoric, (b) the Black Sea and, (c) the Azov Sea, whereas the Bosphorus Strait and Marmara Sea show less faunal affinities with the afore-mentioned sectors. Patterns derived both from the cosmopolitan and Atlanto-Mediterranean species closely follow the one coming from the polychaete species and genera inventories. As a general trend, species numbers decrease along with the decrease in salinity towards the inner parts of the region. The trend is homologous to that seen in the benthic invertebrate inventories of all the major European semi-enclosed regional seas. Salinity and food availability appear to be the dominant abiotic factors correlated, though weakly, with the various patterns deriving from the taxonomic/zoogeographic categories. With the exception of the Anatolia, polychaete inventories from all sectors appear to be random samples of the total inventory of the region, in terms of taxonomic distinctness values. Therefore, these sectoral inventories can be used for future biodiversity/environmental impact assessment studies. A massive invasion of Mediterranean species after the opening of the Black Sea, in the lower Quaternary period, appears to be the likely biogeographic mechanism through which the old Sarmatic fauna was almost completely replaced by species of marine origin.     

南海深海鱼类的初步报告

南海在我国濒临的四个海中面积最大，而其大陆架宽度则较狭窄，水深在200米以上的深海水域相当广阔。我国过去关于南海深海鱼类的研究报道极少，1962年《南海鱼类志》中曾记载过少数深海鱼类，但具有确切深度记录的仅有8种。在国外文献中，Gilbert和Hubbs(1920), Schultz(1938), Abe(1974 , 1975, 1976 )等曾报道过数种。 本文材料是1959-1960年在海南岛以东海域，水深200-1100米进行底栖生物拖网时所获得的标本。鉴定结果，共71种，分隶于37科，12目，其中34种为我国新记录，7种是我国在南海首次记录。 文内凡是我国新记录的种均有形态描述，并附有外形图,有些种作了分类上的讨论；对我国有过记录的种，只记有地理分布和标本采集记录。此外，就现有资料并参考有关深海鱼类文献，对这些种类的地理分布和深度分布也作了初步分析。由于调查范围较小，拖网次数不多，本文仅为南海鱼类区系提供了一些初步资料。丰富的南海深海鱼类区系，随着我国海洋调查事业的发展，还有待进一步深入研究。     

Metazoan meiofaunal communities at cold seeps along the Norwegian margin: Influence of habitat heterogeneity and evidence for connection with shallow-water habitats

Cold-seep environments and their associated symbiont-bearing megafaunal communities create islands of primary production for macro- and meiofauna in the otherwise monotonous and nutrient-poor deep-sea environment. To examine the spatial variation and distribution patterns of metazoan meiobenthos in different seepage-related habitats, samples were collected in two regions off Norway: several pockmarks associated with the Storegga Slide including the Nyegga pockmark area (730 m; 64°N), and the active, methane-venting Håkon Mosby Mud Volcano (HMMV) west of the Barents Sea (1280 m; 72°N). Based on sediment geochemistry and associated epifauna, three different habitat types were distinguished across the two regions: (1) reduced sediment with suboxic conditions, sometimes covered by bacterial mats, (2) sediment colonised by chemosynthetic, siboglinid tubeworms, and (3) sediment outside the influence of seepage and without a large chemosynthetic fauna. Meiofaunal communities varied strongly in terms of generic diversity and dominance among the different habitat types. Control sites and Siboglinidae polychaete fields both supported high nematode genus richness similar to normal deep-sea sediments, whereas the reduced sediments yielded a genus-poor nematode community dominated by one or two successful species. Meiofaunal densities in the different habitats were negatively correlated with macrobenthic densities. An extremely dense (>11,000 ind. 10 cm^–2), mono-specific nematode population appeared to be restricted to the bacterial mats at HMMV. It consisted of a new cryptic species of the Halomonhystera disjuncta complex, which has been described from intertidal habitats in the North Sea. The reduced seep sediments at Nyegga did not yield H. disjuncta but were dominated by Terschellingia longicaudata, another cosmopolitan nematode species known to be abundant in organic-rich, oxygen-poor, shallow-water environments. These observations point to a past or recent connection between margins and shallow-water habitats.     

Distribution patterns of macrobenthic fauna communities in Deukryang Bay,one of the environment conservation areas of Korea

Macrobenthic fauna were collected seasonally at 44 sites in Deukryang Bay from February to November, 2012. The species number of macrobenthic fauna was in the range of 140 to 181, and polychaetes comprised 41.4% of them. The average density of the whole study area changed seasonally from 755 to 1,507 ind. m^?2, and the most abundant fauna group was crustaceans which accounted for 55.1% of total abundance. An amphipod species Nippopisella nagatai was the most dominant species and a decapod species Xenophthalmus pinnotheroides, an amphipod species Photis longicaudata, and a polychaete species Paralacydonia paradoxa were also dominant in all seasons. The mean seasonal values of Shannon’s diversity index (H′) were in the range of 2.2–2.4, and those values for the evenness index and richness index were 0.7–0.7 and 4.6–5.7, respectively. From the cluster analysis, Deukryang Bay could be divided into 3 or 4 station groups with its specific fauna composition. The cluster analysis and an nMDS ordination revealed that local environmental factors such as water depth were related to the spatial delineation of macrobenthic fauna communities in Deukryang Bay.     

Biodiversity of nematode assemblages from deep-sea sediments of the Atacama Slope and Trench (South Pacific Ocean)

Nematode assemblages were investigated (in terms of size spectra, sex ratio, Shannon diversity, trophic structure and diversity, rarefaction statistics, maturity index, taxonomic diversity and taxonomic distinctness) at bathyal and hadal depths (from 1050 to 7800 m) in the deepest trench of the South Pacific Ocean: the Trench of Atacama. This area, characterised by very high concentrations of nutritionally-rich organic matter also at 7800-m depth, displayed characteristics typical of eutrophic systems and revealed high nematode densities (>6000 ind. 10 cm⁻²). Nematode assemblages from the Atacama Trench displayed a different composition than at bathyal depths. At bathyal depths 95 genera and 119 species were found (Comesomatidae, Cyatholaimidae, Microlaimidae, Desmodoridae and Xyalidae being dominant), whereas in the Atacama Trench only 29 genera and 37 species were encountered (dominated by Monhysteridae, Chromadoridae, Microlaimidae, Oxystominidae and Xyalidae). The genus Monhystera (24.4%) strongly dominated at hadal depths and Neochromadora, and Trileptium were observed only in the Atacama Trench, but not at bathyal depths. A reduction of the mean nematode size (by ca. 67%) was observed between bathyal and hadal depths. Since food availability was not a limiting factor in the Atacama Trench sediments, other causes are likely to be responsible for the reduction of nematode species richness and body size. The presence of a restricted number of families and genera in the Atacama Trench might indicate that hadal sediments limited nematode colonisation. Most of the genera reaching very high densities in Trench sediments (e.g., Monhystera) are opportunistic and were responsible for the significant decrease of the maturity index. The dominance of opportunists, which are known to be characterised by small sizes, might have contributed to the reduced nematode size at hadal depths. Shannon diversity and species richness decreased in hadal water depth and this pattern was more evident at genus than at species level. Epistrate feeders dominated and increased their relevance, determining a reduction of the index of trophic diversity at hadal depths. According to trophic diversity, taxonomic diversity and distinctness also decreased with depth. All diversity indices from the Atacama Slope and Trench were lower than in other equally deep areas world wide (e.g. Puerto Rico Trench). We suggest that such reduction was related to the high nutrient loading observed in this system (up to two orders of magnitude higher than in typical oligotrophic deep-sea sediments).     

Two new species of Exogone and Sphaerosyllis (Polychaeta, Syllidae)from the Huanghai Sea

-In a study of meiofauna polychaete from the Huanghai Sea, two new species belonging to Exogane and Sphaerosyllis were found and described in this paper. The two new species differ from the other species in the two genera mainly by the dorsal cirri, chaetation and distribution of papillae.     

Ichthyofauna of the Eastern Coast Bays of the Novaya Zemlya Archipelago

Based on 2014–2016 studies and published data on the ichthyofauna composition near the eastern coast of Novaya Zemlya, a revised list of fish fauna has been compiled. It includes 30 species from 23 genera and 13 families. The taxonomic diversity of ichthyofauna, its characteristics in respect to geographic range and biotopic groups of fish are considered. In general, ichthyological communities in bays off the eastern coast are similar in the species ratio to the communities in the open Kara Sea, and do not demonstrate any essential features of isolation.     

Macrofaunal succession in sediments around kelp and wood falls in the deep NE Pacific and community overlap with other reducing habitats

Sunken parcels of macroalgae and wood provide important oases of organic enrichment at the deep-sea floor, yet sediment community structure and succession around these habitat islands are poorly evaluated. We experimentally implanted 100-kg kelp falls and 200 kg wood falls at 1670 m depth in the Santa Cruz Basin to investigate (1) macrofaunal succession and (2) species overlap with nearby whale-fall and cold-seep communities over time scales of 0.25–5.5 yr. The abundance of infaunal macrobenthos was highly elevated after 0.25 and 0.5 yr near kelp parcels with decreased macrofaunal diversity and evenness within 0.5 m of the falls. Apparently opportunistic species (e.g., two new species of cumaceans) and sulfide tolerant microbial grazers (dorvilleid polychaetes) abounded after 0.25–0.5 yr. At wood falls, opportunistic cumaceans become abundant after 0.5 yr, but sulfide tolerant species only became abundant after 1.8–5.5 yr, in accordance with the much slower buildup of porewater sulfides at wood parcels compared with kelp falls. Species diversity decreased significantly over time in sediments adjacent to the wood parcels, most likely due to stress resulting from intense organic loading of nearby sediments (up to 20–30% organic carbon). Dorvilleid and ampharetid polychaetes were among the top-ranked fauna at wood parcels after 3.0–5.5 yr. Sediments around kelp and wood parcels provided low-intensity reducing conditions that sustain a limited chemoautrotrophically-based fauna. As a result, macrobenthic species overlap among kelp, wood, and other chemosynthetic habitats in the deep NE Pacific are primarily restricted to apparently sulfide tolerant species such as dorvilleid polychaetes, opportunistic cumaceans, and juvenile stages of chemosymbiont containing vesicomyid bivalves. We conclude that organically enriched sediments around wood falls may provide important habitat islands for the persistence and evolution of species dependent on organic- and sulfide-rich conditions at the deep-sea floor and contribute to β and γ diversity in deep-sea ecosystems.     

In situ video observations of benthic megafauna and fishes from the deep eastern Mediterranean Sea off Egypt

Remotely operated vehicle (ROV) video observations were used to document benthic fauna at a hydrocarbon drilling location, at 2 720 m depth, in the poorly studied deep water off northern Egypt. The decapod Chaceon mediterraneus was the most common organism at the site and the only benthic megafaunal invertebrate observed. Three species of fish, Coryphaenoides mediterraneus, Cataetyx laticeps and Bathypterois mediterraneus, were also encountered. These findings confirmed these three species as the deepest-distributed benthic fish in the eastern Mediterranean, and confirmed that the deep Mediterranean, in particular the eastern basin, is one of the world's poorest deep-sea ecosystems in terms of diversity. The behaviour exhibited by the species observed was consistent with their natatory capacity, deduced from their feeding intensity (gut fullness) and diet (mainly suprabenthic prey were consumed), and conservative life strategies adapted to an extremely low energy environment.     

First in situ observations of the deep-sea carnivorous ascidian Dicopia antirrhinum Monniot C., 1972 in the Western Mediterranean Sea

Dicopia antirrhinum C. Monniot, 1972 is a rare species of deep-sea ascidian belonging to the Family Octacnemidae, reported at depths of 1000–2500 m in European Atlantic waters. Adult individuals have never been reported before in the Mediterranean Sea, where only seven juvenile specimens were found in 1975 at 500 m water depth in the Central basin (Malta). The affinities of these specimens with D. antirrhinum were noted, but lack of some typical characters of the species in juveniles prevented a definite taxonomical identification. No other member of the Octacnemidae has ever been found in the Mediterranean. In this study we describe the sampling of an adult specimen of D. antirrhinum at around 1100 m water depth on the flank of the La Fonera (Palamós) canyon, Northwestern Mediterranean, confirming their presence in the Mediterranean Sea. We also observed 5 individuals of this species on their natural habitat with a Remotely Operated Vehicle (ROV). Our results highlight the potential occurrence of Octacnemidae, the presence of which has been largely overlooked, in several deep-sea canyon areas within the Western Mediterranean basin. These observations are important because they indicate the need for increased sampling effort with new technologies, such as ROVs, in ecologically relevant habitats such as canyons, in order to obtain a more accurate picture of deep-sea biodiversity in the Mediterranean Sea.     

Effect of the East Siberian barrier on the echinoderm dispersal in the Arctic Ocean

The distributional patterns were analyzed for 43 species and 33 genera of echinoderms in the Laptev and East Siberian seas and for 59 species and 35 genera of the asteroid species in the Arctic Ocean. The probable colonization route through the Arctic was suggested for each species based on (1) the distributional patterns of the Arctic species, (2) the distributional patterns of the closely related species, and (3) the location of the center of the diversity of the species belonging to a certain genus. The species of the Pacific origin prevailed in the asteroid fauna of the Arctic seas. The asteroid species diversity and the ratio of the species of Pacific origin decreased from the Barents towards the Laptev Sea and increased, respectively, in the East Siberian and the Chukchee seas. The species range limits were found for 19 species in the East Siberian Sea compared to only 3 species in the Laptev Sea. The East Siberian Sea was a limiting area for the dispersal of four species groups: (1) invaders from the North Pacific dispersing along the Asian coast of the Arctic (shallow-water stenobathic species), (2) invaders from the North Pacific dispersing along the American coast of the Arctic and further on back into the Arctic along the Eurasian coast (secondarily Atlantic species); (3) originally invaders from the Northern Atlantic; (4) representatives of the Arctic autochthonous fauna. A great width of the biotic boundaries (i.e., the zones of the species range boundaries crowding) was typical for the Arctic Basin, which was a sign of their young geological age.     

Patterns of bathymetric zonation of bivalves in the Porcupine Seabight and adjacent Abyssal plain,NE Atlantic

Although the organization patterns of fauna in the deep sea have been broadly documented, most studies have focused on the megafauna. Bivalves represent about 10% of the deep-sea macrobenthic fauna, being the third taxon in abundance after polychaetes and peracarid crustaceans. This study, based on a large data set, examined the bathymetric distribution, patterns of zonation and diversity–depth trends of bivalves from the Porcupine Seabight and adjacent Abyssal Plain (NE Atlantic). A total of 131,334 individuals belonging to 76 species were collected between 500 and 4866 m. Most of the species showed broad depth ranges with some ranges extending over more than 3000 m. Furthermore, many species overlapped in their depth distributions. Patterns of zonation were not very strong and faunal change was gradual. Nevertheless, four bathymetric discontinuities, more or less clearly delimited, occurred at about 750, 1900, 2900 and 4100 m. These boundaries indicated five faunistic zones: (1) a zone above ∼750 m marking the change from shelf species to bathyal species; (2) a zone from ∼750 to 1900 m that corresponds to the upper and mid-bathyal zones taken together; (3) a lower bathyal zone from ∼1900 to 2900 m; (4) a transition zone from ∼2900 to 4100 m where the bathyal fauna meets and overlaps with the abyssal fauna and (5) a truly abyssal zone from approximately 4100–4900 m (the lower depth limit of this study), characterized by the presence of abyssal species with restricted depth ranges and a few specimens of some bathyal species with very broad distributions. The ∼4100 m boundary marked the lower limit of distribution of many bathyal species. There was a pattern of increasing diversity downslope from ∼500 to 1600 m, followed by a decrease to minimum values at about 2700 m. This drop in diversity was followed by an increase up to maximum values at ∼4100 m and then again, a fall to ∼4900 m (the lower depth limit in this study).     

Local and regional species diversity of benthic Isopoda (Crustacea) in the deep Gulf of Mexico

Recent studies of deep-sea faunas considered the influence of mid-domain models in the distribution of species diversity and richness with depth. In this paper, I show that separating local diversity from regional species richness in benthic isopods clarifies mid-domain effects in the distribution of isopods in the Gulf of Mexico. Deviations from the randomised implied species ranges can be informative to understanding general patterns within the Gulf of Mexico. The isopods from the GoMB study contained 135 species, with a total of 156 species including those from an earlier study. More than 60 species may be new to science. Most families of deep-sea isopods (suborder Asellota) were present, although some were extremely rare. The isopod family Desmosomatidae dominated the samples, and one species of Macrostylis (Macrostylidae) was found in many samples. Species richness for samples pooled within sites ranged from 1 to 52 species. Because species in pooled samples were highly correlated with individuals, species diversity was compared across sites using the expected species estimator (n=15 individuals, ES₁₅). Six depth transects had idiosyncratic patterns of ES₁₅, and transects with the greatest short-range variation in topography, such as basins and canyons, had the greatest short-range disparity. Basins on the deep slope did not have a consistent influence (i.e., relatively higher or lower than surrounding areas) on the comparative species diversity. ES₁₅ of all transects together showed a weak mid-domain effect, peaking around 1200–1500 m, with low values at the shallowest and deepest samples (Sigsbee Abyssal Plain); no longitudinal (east–west) pattern was found. The regional species pool was analyzed by summing the implied ranges of all species. The species ranges in aggregate did not have significant patterns across longitudes, and many species had broad depth ranges, suggesting that the isopod fauna of the Gulf of Mexico is well dispersed. The summed ranges, as expected, had strong mid-domain patterns, contrasting with the local species richness estimates. The longitudinal ranges closely matched a randomized pattern (species ranges placed randomly, 1000 iterations), with significant deviations in the east attributable to lower sampling effort. The depth pattern, however, deviated from the mid-domain model, with a bimodal peak displaced nearly 500 m shallower than the mode of the randomized distribution. The deviations from random expectation were significantly positive above 1600 m and negative below 2000 m, with the result that regional species richness peaked between 800 and 1200 m, and decreased rapidly at deeper depths. The highest species richness intervals corresponded to the number of individuals collected. Residuals from a regression of the deviations on individual numbers, however, still deviated from the randomized pattern. In this declining depth-diversity pattern, the Gulf of Mexico resembles other partially enclosed basins, such as the Norwegian Sea, known to have suffered geologically recent extinction events. This displaced diversity pattern and broad depth ranges implicate ongoing re-colonization of the deeper parts of the Gulf of Mexico. The Sigsbee Abyssal Plain sites could be depauperate for historical reasons (e.g., one or more extinction events) rather than ongoing ecological reasons (e.g., low food supply).     

白令海和楚科奇海表层沉积中的有孔虫及其沉积环境

通过对白令海和北冰洋楚科奇海39个表层沉积样品中有孔虫的定量分析,发现表层沉积中浮游有孔虫稀少可能与该区表层生产力低、碳酸盐溶解作用较强有关,而底栖有孔虫的分布则主要受表层初级生产力以及与水深相关的碳酸盐溶解作用和水团性质所控制,其中北冰洋楚科奇海陆架区有孔虫以Elphidium spp.组合和Nonionella robusta组合为主,丰度和分异度低,受北冰洋沿岸水团控制;白令海陆坡区有孔虫以Uvigerina peregrina-Globobulimina affinis组合为主,含N.robusta较多,丰度和分异度相对高,受太平洋中层和深层水团控制,但该区碳酸盐溶跃层和补偿深度(CCD)相对浅,约分别位于2000和3800m处.此外,白令海陆坡上部表层沉积中含有北冰洋陆坡区典型深水底栖有孔虫种Stetsonia arctica,说明白令海峡两侧的海区曾有深部水交流.     

Eiders Somateria mollissima scavenging behind a lugworm boat

The eider is one of the most important molluscivorous birds in the Wadden Sea, where it feeds mainly on blue mussels Mytilus edulis and edible cockles Cerastoderma edule. These prey species are within reach of the birds at all times. Other potential prey of suitable size that are abundantly present, such as several polychaete worms, or the clam Mya arenaria, are taken to a much lesser extent, possibly because they live buried in the sediment and digging them out would take too much effort. Mya may pose another problem because they grow to sizes that prevent eiders from swallowing them. Large Mya also live too deep down in the sediment, but young (small) specimens should be available to eiders. Yet, even these have only rarely been found as prey in eiders in the Wadden Sea. However, diet studies in relation to food abundance have been few, and may have missed prey that do not leave large shell fragments (i.e. in faeces studies). This paper describes observations on eiders taking both Mya and polychaete worms. The eiders fed on these prey in a fashion reminiscent of gulls that scavenge behind fishing vessels: some eiders have learnt to follow professional worm-digging boats that supply a bycatch of molluscs (mainly Mya arenaria) and polychaete worms (mainly Arenicola marina and Nephtys hombergii). Mya and worms were also the main targets of the eiders that fed in a dense flock close to the boat's stern. Faeces found on the flats at low tide comprised mainly cockle shell fragments, a prey rarely taken by the eiders behind the boat. Faeces studies may thus give a highly biased impression of local eider diet.     

Deep-sea benthic diversity linked to seasonality of pelagic productivity

Latitudinal gradients in biodiversity are found in both terrestrial and marine environments, but little agreement exists on the mechanisms or ecological causes creating these patterns. Marine biodiversity patterns have been particularly challenging to document, because of the lack of appropriate data sets from ocean basins. We document latitudinal patterns of North Atlantic deep-sea benthic foraminifera and show that seasonality of primary productivity, as estimated from SeaWiFS satellite imagery, has a significant effect on diversity indices, with generally lower values of H(S), species ?, and species equitability found with high seasonality between 40 and 60°N. High foraminiferal diversity is not found in areas with phytodetritus deposition in the North Atlantic basin, which indicates that patch dynamics, biological disturbance, and sediment heterogeneity resulting from phytodetritus deposits do not create high deep-sea foraminiferal diversity. Annual resource stability, reflecting the timing of organic carbon flux and the mode of sedimentation, accounts for the benthic foraminiferal patterns found in this study and is an important variable structuring the deep-sea benthic foraminiferal community.