Habitat heterogeneity influences cold‐seep macrofaunal communities within and among seeps along the Norwegian margin – Part 2: contribution of chemosynthesis and nutritional patterns

The relative contribution of chemosynthesis in heterotrophic fauna at seeps is known to be influenced by depth and by habitat. Using stable isotopes of carbon and nitrogen, we investigated macro‐ and megafaunal nutritional patterns in Norwegian margin cold seeps by comparing food webs both among habitats within a seep site and between different sites. The very active Håkon Mosby mud volcano (HMMV) is characterized by geochemical gradients, microbial activity and faunal zonation from the centre to the periphery. The Storegga Slide (600–900 m depth) has pockmarks with patchy less active seeps, and also shows concentric zonation of habitats but at much smaller spatial scale. The dominant carbon source for macrofaunal nutrition in both areas was chemosynthetically fixed and the bulk of organic carbon was derived from sulphur‐oxidizing bacteria. In HMMV, food chains were clearly separated according to habitats, with significantly lighter δ¹³C signatures on microbial mats and adjacent sediment (?33.06 to ?50.62‰) than in siboglinid fields (?19.83 to ?35.03‰). Mixing model outputs revealed that the contribution of methane‐derived carbon was small in siboglinid fields (0–17%) but significant (39–61%) in the microbial mats. Moreover, the variability of macrofauna signatures within this later habitat suggests the co‐occurrence of two food chains, one based on primary production via methanotrophy and the other via sulphide oxidation. The length of the food chains also varied among habitats, with at least one more trophic level in the siboglinid fields located at the periphery of the volcano. Conversely, in Storrega pockmarks, faunal δ¹³C signatures did not vary among habitats but among species, although separate food chains seem to co‐occur. The small size of the seepage areas and their lower fluxes compared to HMMV allow more background species to penetrate the seep area, increasing the range of δ¹⁵N and the trophic level number. Probably due to the higher flux of photosynthetic particulate organic carbon, the overall chemosynthesis‐based carbon contribution in invertebrate nutrition was lower than that in HMMV.     

Demographic Characteristics of Lytechinus variegatus (Echinoidea: Echinodermata) from Three Habitats in a North Florida Bay, Gulf of Mexico

Abstract. The population densities, spatial distributions, size frequencies, growth rates, longevity and reproductive activities of sub‐populations of the sea urchin Lytechinus variegatus were investigated over a two‐year period. Sea urchins were examined in three habitats in Saint Joseph Bay, Florida, which is within the northern limits of their distribution. Densities of sea urchins, which ranged as high as 35 individuals ·^?2, fluctuated seasonally at all sites and were higher in seagrass beds comprised of Thalassia testudinum than Syringodium filiforme or on a sand flat. A cold front caused large‐scale, catastrophic mortality among adult, and especially juvenile, sea urchins in nearshore habitats of the Bay in the spring of 1993, leading to a dramatic decline in sea urchin densities at the Thalassia seagrass site. The population recovered over 6 months at this site and was attributable to immigration of new adults. Juvenile recruitment displayed both interannual and site‐specific variability, with recruitment being highest in seagrass habitats in fall and spring. The most pronounced recruitment event occurred in fall 1993 at the Thalassia site. Spatial distributions of adult individuals ascertained monthly never varied from random in the seagrass beds (T. testudinum and S. filiforme) or during spring, summer or fall months on the sand flat. Nonetheless, aggregations of adult sea urchins were observed on the sand flat in the winter months and were associated with patchy distributions of plant food resources. Juvenile sea urchins (< 25 mm test diameter) exhibited aggregations at all sites and 67 % of all juveniles under 10 mm test diameter (91 of 165 individuals observed) were found under the spine canopies of adults. Measurements of the inducibility of spawning indicated peak gametic maturity in all three sub‐populations in spring and summer. Gonad indices varied between habitats and years, but distinct maxima were detected, particularly in spring 1993 and late summer 1994. The mean gonad index of individuals at the Syringodium seagrass site was 2‐ to 4‐fold higher than the other sites in spring 1993 and gonad indices were much higher at all sites in spring of 1993 than 1994. Estimates of growth based on changes in size frequency cohorts coupled with measurements of growth bands on lantern demipyramids indicated that L. variegatus in three habitats of Saint Joseph Bay have similar growth rates and attain a mean test diameter of approximately 35 mm in one year. In contrast to populations within the central biogeographical range of the species, which may attain test diameters up to 90 mm, the largest individuals recorded in Saint Joseph Bay were 60 mm in test diameter, and almost all individuals were no more than 45 mm in test diameter or two years of age. The demographics of L. variegatus in the northern limits of their distribution appear to be strongly influenced by latitudinally driven, low‐temperature events and secondarily by local abiotic factors, especially springtime low salinities, which may negatively impact larval development and recruitment.     

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).     

Petrographic and geochemical characterization of seep carbonate from Bush Hill (GC 185) gas vent and hydrate site of the Gulf of Mexico

Authigenic carbonates are common at cold seep sites as a result of microbial oxidation of hydrocarbons. Seep carbonate samples were collected from the surface of the Bush Hill (Green Canyon Block 185, Gulf of Mexico), a mound containing gas hydrate. The carbonates consisted of oily, porous limestone slabs and blocks containing bioclasts and matrix. Analysis by X-ray diffraction shows that aragonite is the dominant mineral (89–99 wt% with an average of 94 wt%) in the matrix of seep carbonate. This cement occurs in microcrystalline, microspar, and sparite forms. The moderate ¹³C depletion of the seep carbonate (the most depleted one has δ¹³C value of −29.4‰, and 26 of 38 subsamples have δ¹³C values >−20.0‰) indicates that the non-methane hydrocarbons was incorporated during seep carbonate precipitation. Relative enrichment of ¹⁸O may be related to localized destabilization of gas hydrate or derived from ¹⁸O-enriched pore water originated from smectite–illite transition in the deep sediments. The total content of rare earth elements (REE) of the 5% HNO₃-treated solution of the carbonates is from 0.40 ppm to 30.9 ppm. The shale-normalized REE patterns show varied Ce anomalies from significantly negative, slightly negative, and no to positive Ce anomalies. Variable content of trace elements, total REE, and Ce anomalies in different samples and even in the different carbonate mineral forms (microcrystalline, microspar and sparite) of the same sample suggest that the formation condition of the Bush Hill seep carbonate is variable and complex, which is possibly controlled by the rate of fluid flux.     

Microhabitat preferences and depth distribution of combtooth blennies (Blenniidae) in the Gulf of Trieste (North Adriatic Sea)

The microhabitat preferences and depth distribution of blenniid species (Blenniidae) in the Gulf of Trieste (North Adriatic Sea) were surveyed using the all‐occurrence sampling method, a non‐destructive visual census method, aided by SCUBA diving. Fourteen species were identified during the entire survey. Four species showed to be indiscriminate in their microhabitat choice while 10 species were classified as infrequent in the surveyed coastal area. A statistically significant correlation was found between the blenniid assemblage and nine microhabitat variables. The blenniid assemblage was divided in two main groups, by the use of canonical correspondence analysis, electivity index, and the depth distribution analysis. The first group comprises species that dwell in surface waters and show a high positive correlation with boulders, the presence of Mytilus galloprovincialis, cirripeds and empty holes bored by Lithophaga lithophaga. The second group includes species that mostly inhabit deeper waters and show a high positive correlation with rocks covered by precoralligenous bioformations.     

Distribution and Diversity of Ibla cumingi Darwin [Crustacea, Cirripedia) from the Gulf of Elat (Red Sea)

Abstract. The distribution of Ibla cumingi DARWIN on different types of hard substrata along 30 km of coast in the Gulf of Elat was studied. I. cumingi was found among and underneath Tetraclita squamosa rufotincta , in oyster beds, underneath slabs of beachrock, and in fissures in igneous rocks. In the Tetraclita belt, I. cumingi is most frequent in empty shells of dead Tetraclita while in the oyster bed it is common inside the meshwork made by the oysters. The distribution is regarded as clustered distribution. The density and size of the clusters of I. cumingi is determined by the available sheltered space within the midlittoral zone. The form of the habitat determines the shape of the animals. Those found in the oyster bed and fissures are usually longer than those found in the Tetraclita belt due to the depth of the fissures or to the bigger spaces within the oyster bed. I. cumingi is orientated so that the capitulum with the cirri is projecting towards the open water, and the water current. It is concluded that I. cumingi can withstand the physical stress encountered in a tropical intertidal zone, but that the distribution pattern is caused by predation.     

北太平洋中西部剑鱼(Xiphias gladius)管理策略评价

剑鱼（Xiphias gladius）属于大洋性中上层高度洄游性鱼种,是远洋渔业重要的目标渔获物和兼捕种类,具有较高经济价值。本文在北太平洋金枪鱼与类金枪鱼国际科学委员会（ISC）对剑鱼资源评估结果基础上,使用1951年到2018年渔获量数据,通过先验参数设置构造操作模型模拟北太平洋中西部剑鱼的种群动态和渔业动态,通过不同管理策略捕捞规则计算对数据和参数要求筛选出9种备选管理策略（DD,DD4010,CC1,SBT1,GB_slope,ICI,ICI2,SPmod,MCD）并对其管理效果进行量化分析,并对短期和长期产量进行预测。通过对各先验参数时间序列分布及各管理策略的权衡,使用Kobe图表达实施管理策略前后剑鱼资源状况的变化,最终确定使用根据时间序列平均值和标准误差指数来调整渔获量的管理策略（ICI）为最佳的管理策略。对9种管理策略其中4种输出型管理策略（MCD,ICI,ICI2,SPmod）进行总可捕量（TAC）计算,分别拟合中国台湾CPUE序列和日本CPUE序列,最终得到ICI管理策略对未来50年TAC控制量在10 404.48 t,50%置信区间在6 678.51~18 743.22 t。对最终所选择的ICI管理策略进行敏感性测试,结果表明其对渔获量和丰度指数较为敏感。     

Fauna of whale falls: systematics and ecology of a new polychaete (Annelida: Chrysopetalidae) from the deep Pacific Ocean

Vigtorniella flokati, a new species of polychaete worm associated with decaying whale remains, is described. Three separate V. flokati populations were sampled using submersibles: two associated with implanted gray whale carcasses in the San Diego Trough (1240 m depth) and the Santa Cruz Basin (1670 m) off California, and one from sperm whale and balaeanopterid bones implanted on the slope of Oahu, Hawaii at 1000 m. Extraordinarily large numbers of live specimens were observed and videotaped in situ using submersibles in the San Diego Trough and the Santa Cruz Basin. The populations on the carcass implanted for 4 months in San Diego Trough, and on the bones implanted for 2 years off Oahu, were composed only of small sized individuals (including juveniles). V. flokati most closely resembles the poorly known V. zaikai Kiseleva, 1992, described from the Black Sea. Forty-six morphological characters were used in a phylogenetic analysis of selected nereidiform polychaetes. The resulting most-parsimonious trees indicate sister taxon status between V. flokati and V. zaikai, and that this clade is sister to the putatively ancient polychaete clade Chrysopetalidae. Whale falls, which are intense point sources of organic enrichment at the deep-sea floor, pass through three successional stages. V. flokati appears to colonize the middle, “enrichment opportunist” stage, inhabiting organic-rich bones and sediments 4–24 months after carcass arrival. V. flokati exhibits remarkable behavior, clinging posteriorly to whale bones or nearby sediments to form a writhing carpet at densities exceeding 8000 m⁻². Its extraordinary abundance on whale falls, and apparent absence from other habitats, suggests it to be a whale-fall specialist. The precise feeding mechanism of the worm remains unknown, but we hypothesize that it may utilize dissolved organic carbon derived from the organic-rich setting of whale falls. The widespread occurrence of V. flokati on ephemeral, food-rich habitat islands in the Pacific suggests life-history strategies analogous to those for hydrothermal-vent and cold-seep species.     

Surficial Hydrocarbon Seep Infauna from the Blake Ridge (Atlantic Ocean, 2150 m) and the Gulf of Mexico (690–2240 m)

Abstract.  Infauna, including foraminifera and metazoans, were enumerated and identified from five types of seep habitats and two adjacent non-seep habitats. Collections were made with the deep submergence research vessel 'Alvin' from three areas of active seepage in the Gulf of Mexico (Alaminos Canyon [2220 m], Atwater Canyon [1930 m], and Green Canyon lease block 272 [700 m]) and on the Blake Ridge Diapir [2250 m], which is located off the southeastern coast of the United States. The seep habitats sampled included four types of microbial mats ( Beggiatoa , Thioploca , thin and thick Arcobacter ) and the periphery of a large mussel bed. Sediments under large rhizopod protists, xenophyophores, were sampled adjacent to the mussel bed periphery. A non-seep site, which was >1 km away from active seeps, was also sampled for comparison. Densities of most taxa were higher in the Gulf of Mexico seeps than in Blake Ridge samples, largely because densities in the thick microbial mats of Blake Ridge were significantly lower. Diversity was higher in the Thioploca mats compared to other microbial-mat types. Within an ocean basin ( i.e. , Atlantic, Gulf of Mexico) we did not observe significant differences in meiofaunal or macrofaunal composition in Beggiatoa versus Thioploca mats or thin versus thick Arcobacter mats. Foraminifera represented up to 16% of the seep community, a proportion that is comparable to their contribution at adjacent non-seep communities. In general, the observed densities and taxonomic composition of seep sites at the genus level was consistent with previous observations from seeps ( e.g. , the foraminifers Bolivina and Fursenkoina , the dorvilleid polychaete Ophryotrocha ).     

Connectivity and stock composition of loggerhead turtles foraging on the North African continental shelf (Central Mediterranean): implications for conservation and management

The loggerhead turtle, Caretta caretta, is a highly migratory species with a complex life cycle that involves a series of ontogenetic habitat shifts and migrations. Understanding the links amongst nesting populations and foraging habitats is essential for the effective management of the species. Here we used mixed stock analysis to examine the natal origin of loggerhead turtles foraging on the North African continental shelf off Tunisia, one of the most important Mediterranean neritic habitats. An 815‐bp fragment of the mitochondrial DNA control region was sequenced from 107 individuals sampled from 2007 to 2009. No temporal variation in haplotype frequencies was detected. Juveniles (n = 87) and adults (n = 23) exhibited weak but significant genetic differentiation that resulted in different stock compositions. Libya was the main source population but the proportion of turtles from this rookery was higher in adults (median = 80%) than in juveniles (median = 35%). Western Greece was the second most important contributing population. Juvenile stock composition derived from mixed stock analysis and the estimates produced by numerical simulation of hatchling dispersion in the Mediterranean Sea were significantly correlated, supporting the recent theory that loggerheads imprint on possible future neritic habitats during the initial phase of their life. This association was not significant for adults, suggesting that other factors contribute to shaping their distribution. Overall, our results show that human activities on the South Tunisian continental shelf pose an immediate threat to the survival of the Libyan rookery.     

Molecular weight and chemical reactivity of dissolved trace metals (Cd,Cu, Ni) in surface waters from the Mississippi River to Gulf of Mexico

It is generally assumed that estuarine mixing is continuous for metals from terrestrial sources, gradually decreasing towards the open ocean endmember. Here we show that, chemical reactivity, determined by ion exchange method, and molecular weight distributions, obtained using cross-flow ultrafiltration, of dissolved Cd, Cu, and Ni in the surface waters of the Gulf of Mexico varied systematically across the estuarine mixing zone of the Mississippi River. Most size or chemical affinity fractions of dissolved metals (<0.4 μm) were linearly related to salinity (10.8–36.6), suggesting that the distribution of these elements was mainly controlled by continuous mixing processes. Dissolved concentrations across the salinity gradient ranged for Cd: 87–187 pM; Cu: 1.4–18.3 nM; and Ni: 2.6–18.8 nM, with highest values near the Mississippi river mouth, and lowest concentrations in a warm core ring in the Gulf of Mexico. Dissolved Cd was mostly present as a truly dissolved (<10 kDa, 97 ± 1%) and cationic fraction (Chelex-100 extractable, 94 ± 4%). A novel observation across the estuarine mixing zone was that colloidal metal concentrations were identical to either inert (for Cu, Ni) or AMPG-labile anionic (Cu, Cd) fractions. The difference in behavior between Cu and the other two metals might indicate differences in the biopolymeric nature of the metal–organic chelates. In particular, the anionic-organic Cd fractions accounted for just 3 ± 1%, on average. However, for Cu, it was 24 ± 4%, and for Ni, it was 9 ± 6%. The fractions of the total dissolved metal fractions that were “inert” averaged 31 ± 10% for Cu and 29 ± 12% for Ni. Small but noticeable amounts (6 ± 3%) of dissolved inert Cd fractions were also present. Apparent non-local transport processes, likely associated with cross-shelf sediment resuspension processes, could have been responsible for the relatively high concentrations of ‘inert’ and ‘anionic’ metal fractions in high salinity coastal waters, and accounting for the persistence of metals bound to humic substances in the Gulf of Mexico.     

Distribution of Trace Bioelements in the Subarctic North Pacific Ocean and the Bering Sea (the R/V Hakuho Maru Cruise KH-97-2)

A column concentration-high resolution inductively coupled plasma mass spectrometry (ICP-MS) determination was applied to measure the total dissolved concentrations of Fe, Co, Ni, Cu and Zn in seawater collected from the subarctic North Pacific (~45°N) and the Bering Sea in July–September 1997. Total adsorbable Mn was determined on board by column electrolysis preconcentration and chemiluminescence detection. The vertical profiles for Fe, Ni and Zn were nutrient-like. The deep water concentration of Fe was ~0.5 nM in the northeast Pacific (18°-140°W) and increased to ~1 nM in the northwest Pacific (161°E) and ~2 nM in the Bering Sea (57°N, 180°E). The deep water concentrations for Ni and Zn in the Bering Sea were also 1.3–2 times higher than in the North Pacific. The profiles for Co and Cu were examined in the subarctic North Pacific, and results obtained were consistent with previous reports. There was a significant correlation between the concentrations of Co and Mn except for surface mixed layer. The profiles for total adsorbable Mn were similar to the reported profiles for total dissolvable Mn. The deep water concentration of Mn in the Bering Sea was also 4 times higher than in the North Pacific. Iron and zinc were depleted in surface water of the subarctic North Pacific. The relationship between these trace elements and nutrients suggests that these elements could be a limiting factor of phytoplankton productivity. In the Bering Sea, surface water contained ~0.3 nM of Fe. The Zn concentration, which was less than the detection limit in surface water, increased at shallower depths (~30 m) compared with the subarctic North Pacific. These results imply a higher flux of Fe and Zn to surface water in the Bering Sea. This in turn may cause the ecosystem in the Bering Sea characterized by a dominance of diatoms and high regenerated production.     

Genetic diversity and population structure of three dominant myctophid fishes (<Emphasis Type="Italic">Diaphus theta,Stenobrachius leucopsarus</Emphasis>, and <Emphasis Type="Italic">S. nannochir</Emphasis>) in the North Pacific Ocean

We compared the genetic diversity of three dominant myctophid fishes in the North Pacific Ocean that have different diel vertical migration patterns on the basis of the nucleotide sequences of the mitochondrial gene for cytochrome b. No genetic structure was detected for each of these three species. The genetic diversity progressively increased for Diaphus theta, a diel migrant species showing clear diel vertical migration; Stenobrachius leucopsarus, a semi-diel migrant, in which only part of the population migrates vertically; S. nannochir, a non-diel migrant. All three species were suggested to have experienced a recent, sudden population expansion. Interspecific differences in genetic diversity might be attributable to differences in the degree of population size reduction during the glacial periods; this degree in turn corresponds to the energy demand of the fishes.