The economic valuation of biological resources and ecosystem services in the Sea of Okhotsk

The total monetary value of the harvestable stocks of commercial aquatic species in the Sea of Okhotsk, based on the Forecast of the Total Allowable Catch for 2014 and world prices for seafood, is estimated at $23.5 × 10⁹ USD/year. The potential value of all the surveyed biological resources in the sea (including potential ones) in the case of their more complete economic use, $58.5 × 10⁹ USD/year, is determined using the databases on macrofauna in the pelagic and benthic zones of the Far Eastern seas. Taking the mean value of ecosystem services per unit area in some parts of the World Ocean into account, the total value of ecosystem services for the Sea of Okhotsk may constitute $294.4 × 10⁹ USD/year. The ecosystem services whose market values cannot be determined have a much higher cost than the traditionally exploited biological resources. The value of ecosystem services expressed in monetary terms can be considered as a tool to enhance the conservation importance of natural complexes in the course of the implementation of various industrial projects.     

Structure and diversity of shallow soft-bottom benthic macrofauna in the Gulf of Lions (NW Mediterranean)

Samples of soft-sediment macrobenthos from 92 sites between 10 and 50 m depth were used to assess (1) the main soft-bottom macrofauna communities in the Gulf of Lions, (2) the different components of the diversity of benthic macrofauna in this area, and (3) the relevance of the use of major taxonomic groups as surrogates for the analysis of the structure and diversity of total macrofauna. Three main communities were identified by cluster analysis and associated procedures. These communities corresponded well to the assemblages recently identified on the basis of polychaete composition. The α-diversity indices were in accordance with those reported for similar communities in the Mediterranean. Conversely, the β-diversity value was higher than the few other data available in the literature for marine soft-bottom macrofauna. The total number of species in the studied area estimated by the “total species accumulation curve” (TS) method was 2,319, which was only 10% higher than the number obtained by extrapolation of the species–area curve. The similarity matrix based on polychaetes correlated best with the one based on total macrofauna. Polychaetes and crustaceans were also the best surrogates of total macrofauna when assessing α-diversity (except in the case of Δ*). Conversely, molluscs were the best surrogates of total macrofauna β-diversity. Our results show that the choice of an optimal surrogate for total benthic macrofauna depends on the characteristic of the benthic macrofauna to be studied. Moreover, this choice is also dependent on the environment to be studied.     

The meagre future of benthic fauna in a coastal sea—Benthic responses to recovery from eutrophication in a changing climate

Nutrient loading and climate change affect coastal ecosystems worldwide. Unravelling the combined effects of these pressures on benthic macrofauna is essential for understanding the future functioning of coastal ecosystems, as it is an important component linking the benthic and pelagic realms. In this study, we extended an existing model of benthic macrofauna coupled with a physical–biogeochemical model of the Baltic Sea to study the combined effects of changing nutrient loads and climate on biomass and metabolism of benthic macrofauna historically and in scenarios for the future. Based on a statistical comparison with a large validation dataset of measured biomasses, the model showed good or reasonable performance across the different basins and depth strata in the model area. In scenarios with decreasing nutrient loads according to the Baltic Sea Action Plan but also with continued recent loads (mean loads 2012–2014), overall macrofaunal biomass and carbon processing were projected to decrease significantly by the end of the century despite improved oxygen conditions at the seafloor. Climate change led to intensified pelagic recycling of primary production and reduced export of particulate organic carbon to the seafloor with negative effects on macrofaunal biomass. In the high nutrient load scenario, representing the highest recorded historical loads, climate change counteracted the effects of increased productivity leading to a hyperbolic response: biomass and carbon processing increased up to mid‐21st century but then decreased, giving almost no net change by the end of the 21st century compared to present. The study shows that benthic responses to environmental change are nonlinear and partly decoupled from pelagic responses and indicates that benthic–pelagic coupling might be weaker in a warmer and less eutrophic sea.     

Interrelationships of bacteria,meiofauna and macrofauna in a Mediterranean sedimentary beach (Maremma Park,NW Italy)

Collelungo beach (Maremma Park, NW Italy), was sampled quantitatively for macrofauna, meiofauna and bacteria in May 2003; several physicochemical variables and variables associated with food availability and sediment structure were also measured. Replicated samples were collected from three sites representing natural conditions, an erosion regime, and the influence of the Ombrone River, respectively, as well as from four stations each located in the surf and sublittoral zones. Both uni- and multivariate techniques were used to assess the benthic community structure and the associated environmental variables. Different diversity indices revealed no pattern; in contrast, multivariate techniques applied on the macrobenthic fauna and the polychaete taxocommunity distinguished between the sites located in natural and eroding conditions from the one located nearby the discharges of the Ombrone river. Τhe community patterns deriving from meio- and macrofauna are clearly divergent. The overall benthic faunal community appears to be influenced by both groups of organisms. The patterns of the meio- and macrofaunal communities seem to be affected synergistically by a number of environmental variables, in accordance with the multicausal environmental severity hypothesis. Meiofaunal patterns are more often correlated with bacteria and the protein concentration than are macrofaunal patterns, indicating a potential utilization of bacteria as a food source by the meiofaunal organisms. Total bacterial numbers are associated with the macrofaunal pattern under the erosion regime, probably as a consequence of competition for food between macrofauna and meiofauna.     

Influence of benthic macrofauna on microbial production of methylmercury in sediments on the New England continental shelf

Microbial methylation processes in sediment are an important source of toxic monomethylmercury (MMHg) to aquatic ecosystems. Although bioturbation activities (feeding, digging of galleries, excavations, bioirrigation) by benthic fauna are known to affect many biogeochemical processes, their influence on benthic MMHg production is poorly understood. We investigated the effect of benthic fauna on the microbial production of MMHg in sediments on the continental shelf of the northwest Atlantic Ocean in September 2009. Replicate cores of sieved (control) and unaltered sediment containing native macrofauna were incubated to examine the influence of benthic macrofauna on net MMHg production, potential gross rates of Hg methylation, sediment reworking, dissolved oxygen and organic carbon concentrations, and microbial metabolic activities. The presence of macrofauna stimulated aerobic microbial respiration and net MMHg production, but had no observed effect on short-term gross rates of Hg methylation. This suggests that bioturbation may promote net MMHg production by inhibiting demethylating microorganisms, although overall community metabolism was increased. Results from this work emphasize the need to enhance our knowledge and understanding of the interactions among benthic fauna, microorganisms, and geochemistry in affecting MMHg production.     

深圳福田红树林区底栖大型动物群落的空间分带及灌污的可能影响

本文自１９９１年４月至１９９３年１月对深圳福田红树林中底栖大型动物的空间分带及灌污的可能影响进行了研究。结果表明,该红树林湿地中主要出现的底栖动物为拟沼螺科,黑螺科,汇螺科,沙蟹科,方蟹科和弹涂鱼科种类。红树区内底栖动物从高潮位到低潮位可分为３个群落分布带：亮泽拟沼螺带;拟黑螺－褶痕相手蟹带;弧边招潮－印尼拟蟹守螺－刻纹拟沼螺带。群落的分带可能主要由潮位线,食物适应性及底质结构因素决定。林前泥滩底栖动物种类多样性最大,生物量最高。林内动物群落则表现出低种类多样性,高种群个体数的特点。群落总栖息密度的变化基本上由软体动物所主导。生活污水排灌对红树林中底栖大型动物的影响不明显,仅在排污口端引起少数污水动物种类的出现及群落总生物量轻微的升高。     

The comparative statuses of Far Eastern seas and the Northwestern Pacific based on the range of integral characteristics of pelagic macrofauna

Several characteristics, such as macrofauna population density, species richness and diversity, evenness of species by abundance, and mean weight of individuals were used to make a comparison among the pelagic zones of the Far Eastern seas (mainly within the Russian EEZ) and the adjacent waters of the Pacific Ocean.     

海南清澜港红树林滩涂大型底栖动物初步研究

对海南清澜港红树林滩涂大型底栖动物进行了定性及定量调查,记录到大型底栖动物４５种,隶属于４个门,２６个科。其中软体动物３１种,占记录到物种数的６８９％;甲壳动物１０种,占记录到物种数的２２２％。优势种是珠带拟蟹手螺（ＣｅｒｉｔｈｉｄｅａｃｉｎｇｕｌａｔａＧｍｅｌｉｎ）、古氏滩栖螺（ＢａｔｉｌａｒｉａｃｕｍｉｎｇｉＣｒｏｓｅ）、中国紫蛤（ＨｉａｔｕｌａｃｈｉｎｅｎｓｉｓＭｏｒｃｈ）及奥莱彩螺（ＣｌｉｔｈｏｎｏｕａｌａｎｉｅｎｓｉｓＬｅｓｏｎ）。大型底栖动物的生物量平均１９８５ｇ／ｍ２、栖息密度３５２４个／ｍ２。软体动物的生物量占到９９％,这可能与取样设在中、低潮区有关。     

Macrofauna communities of the Dogger Bank (central North Sea) in the late 1990s: spatial distribution,species composition and trophic structure

Macrofauna samples were taken in May 1996–1998 at 28 stations on the Dogger Bank (central North Sea) to describe species composition, spatial distribution and trophic structure of the macrofaunal communities. In general, five communities could be distinguished depending on the influence of different water masses, depth, type of sediment and food availability. Except for those species which feed in the sediment, macrofauna communities did not show any correlation to the measured sediment variables. In fact, macrofauna communities of the Dogger Bank were mainly structured by lateral food supply via frontal systems. This was indicated by the dominance of interface feeders foraging in the benthic boundary layer, which are able to switch from surface deposit to suspension feeding depending on flow conditions. The importance of sandlickers in the shallowest parts of the Dogger Bank hints at an important contribution of benthic primary production to the nutritional and energetic needs of the benthic community. Electronic Publication     

Non-Linear Interactions Determine the Impact of Sea-Level Rise on Estuarine Benthic Biodiversity and Ecosystem Processes

Sea-level rise induced by climate change may have significant impacts on the ecosystem functions and ecosystem services provided by intertidal sediment ecosystems. Accelerated sea-level rise is expected to lead to steeper beach slopes, coarser particle sizes and increased wave exposure, with consequent impacts on intertidal ecosystems. We examined the relationships between abundance, biomass, and community metabolism of benthic fauna with beach slope, particle size and exposure, using samples across a range of conditions from three different locations in the UK, to determine the significance of sediment particle size beach slope and wave exposure in affecting benthic fauna and ecosystem function in different ecological contexts. Our results show that abundance, biomass and oxygen consumption of intertidal macrofauna and meiofauna are affected significantly by interactions among sediment particle size, beach slope and wave exposure. For macrofauna on less sloping beaches, the effect of these physical constraints is mediated by the local context, although for meiofauna and for macrofauna on intermediate and steeper beaches, the effects of physical constraints dominate. Steeper beach slopes, coarser particle sizes and increased wave exposure generally result in decreases in abundance, biomass and oxygen consumption, but these relationships are complex and non-linear. Sea-level rise is likely to lead to changes in ecosystem structure with generally negative impacts on ecosystem functions and ecosystem services. However, the impacts of sea-level rise will also be affected by local ecological context, especially for less sloping beaches.     

海南东寨港红树林滩涂大型底栖动物多样性的初步研究

记录到海南东寨港红树林滩涂大型底栖动物68种,其中软体动物39种,占记录到物种数的57.4%;甲壳动物19种,占记录到物种数的27.9%。冬季优势种是珠带拟蟹手螺(Cerithidea cingulata )、古氏滩栖螺(Batillaria cumingi )和环肋樱蛤(Cyclotellina remies );夏季优势种是珠带拟蟹手螺、环肋樱蛤和红肉河蓝蛤(Potamocorbula rubromuscula )。大型底栖动物的生物量夏季平均为133.0 g/m²,冬季平均为63.0 g/m²; 栖息密度夏季平均为106.4 个/m²,冬季平均为103.5 个/m²。以密度为计量单位的物种多样性指数夏季为1.841,冬季为0.380,均匀度指数夏季为0.514,冬季为0.112;以生物量为计量单位的物种多样性指数夏季为1.994,冬季为1.751, 均匀度指数夏季为0.556,冬季为0.520。大型底栖动物的生物量、栖息密度、物种多样性指数和均匀度指数大都有季节变化及底质差异,基本趋势是夏季明显高于冬季,沙泥底的滩涂高于泥底质的滩涂。     

Evaluating the importance of predation on subtidal benthic assemblages in sandy habitats around rocky reefs

It was proposed that predation could explain differences in the structure of the benthic macrofaunal assemblages in subtidal sandy sediments close to compared with those far from rocky reefs. This hypothesis was tested using experimental exclusion cages and partial cages at two sites at two distances at two different rocky reefs. Undisturbed uncaged assemblages of macrofauna close to the rocky reefs were generally different from those in the partial cages and full cages. However, caging artifacts could not be detected and there were no strong correlations between the macrofauna and the proportions of different grain size and organic content. The structure of the macrofaunal assemblages close to the rocky reefs was, nevertheless, different from those far from the reefs and the sediments were finer far from than close to the rocky reefs. The results indicated that factors other than predation or grain size caused the differences in the macrofauna. For the spatial and temporal scales used in this study, it was clear that, although predation maybe intense, on its own it cannot explain the differences in the structure of the assemblages close and far from rocky reefs. The importance of adequate replication on caging experiments is discussed and it is suggested that alternative ways need to be found to test predictions about the influence of predation on soft sediment benthic assemblages.     

Influence of the mode of macrofauna-mediated bioturbation on the vertical distribution of living benthic foraminifera: First insight from axial tomodensitometry

We investigated the influence of bioturbation by macrofauna on the vertical distribution of living (stained) benthic foraminifera in marine intertidal sediments. We investigated the links between macrofaunal bioturbation and foraminiferal distribution, by sampling from stations situated on a gradient of perturbation by oyster-farming, which has a major effect on benthic faunal assemblages. Sediment cores were collected on the French Atlantic coast, from three intertidal stations: an oyster farm, an area without oysters but affected by oyster biodeposits, and a control station. Axial tomodensitometry (CT-scan) was used for three-dimensional visualization and two-dimensional analysis of the cores. Biogenic structure volumes were quantified and compared between cores. We collected the macrofauna, living foraminifera, shells and gravel from the cores after scanning, to validate image analysis. We did not investigate differences in the biogenic structure volume between cores. However, biogenic structure volume is not necessarily proportional to the extent of bioturbation in a core, given that many biodiffusive activities cannot be detected on CT-scans. Biodiffusors and larger gallery-diffusors were abundant in macrofaunal assemblage at the control station. By contrast, macrofaunal assemblages consisted principally of downward-conveyors at the two stations affected by oyster farming. At the control station, the vertical distribution of biogenic structures mainly built by the biodiffusor Scorbicularia plana and the large gallery-diffusor Hediste diversicolor was significantly correlated with the vertical profiles of living foraminifera in the sediment, whereas vertical distributions of foraminifera and downward-conveyors were not correlated at the station affected by oyster farming. This relationship was probably responsible for the collection of foraminifera in deep sediment layers (> 6 cm below the sediment surface) at the control station. As previously suggested for other species, oxygen diffusion may occur via the burrows built by S. plana and H. diversicolor, potentially increasing oxygen penetration and providing a favorable microhabitat for foraminifera in terms of oxygen levels. By contrast, the absence of living foraminifera below 6 cm at the stations affected by oyster farming was probably associated with a lack of biodiffusor and large gallery-diffusor bioturbation. Our findings suggest that the effect of macrofaunal bioturbation on the vertical distribution of foraminiferal assemblages in sediments depends on the effects of the macrofauna on bioirrigation and sediment oxidation, as deduced by Eh values, rather than on the biogenic structure volume produced by macrofauna. The loss of bioturbator functional diversity due to oyster farming may thus indirectly affect infaunal communities by suppressing favorable microhabitats produced by bioturbation.     

Species size distributions in marine benthic communities

Summary Species body size distributions from eight temperate benthic communities show a highly conservative pattern with two separate lognormal distributions, corresponding to the traditional categories of meiofauna and macrofauna. The meiofaunal mode occurs at a dry body weight of 0.64 g and the macrofaunal mode at 3.2 mg, with a trough between them at 45 g. It is suggested that there is a particular body size at which meiofaunal life-history and feeding traits can be optimised, and another for macrofaunal traits. As size departs in either direction (larger or smaller) from these optima, fewer species of the same size are able to co-exist. The split occurs at 45 g because many life history and feeding characteristics switch more or less abruptly at about this body size, compromise traits being either non-viable or disadvantageous. Meiofauna and macrofauna therefore comprise two separate evolutionary units each with an internally coherent set of biological characteristics.The expression of this conservative pattern is modified by water depth: the proportion of macrofauna species increases from intertidal situations to deeper water, and it is suggested that mechanisms of resource partitioning and diversity maintainence in the meiofauna and macrofauna are affected differentially by sediment disturbance. Salinity does not affect this proportionality, and so does not differentially affect mechanisms for maintaining species diversity in any particular size category of animals. Meiofauna species size distributions may be modified in sandy sediments because of physical impositions on interstitial or burrowing lifestyles.Brief discussion of some implications of these observations includes speculations on the larval ecology of macrofauna, on gigantism in Antarctic invertebrates, and on the benthic Sheldon spectrum.     

Benthic taxa as potential indicators of a deep-sea oil spill

The effect of the Deepwater Horizon oil spill on benthic macrofauna in the deep-sea Gulf of Mexico was measured in September–October 2010. Macrofauna community diversity and abundance were lowest closest to the wellhead and increased with distance from the wellhead up to 10 km. The macrofauna loss was primarily in surface sediments, which could be due to the deposition of oil and other toxic chemicals. Crustacean taxa appeared to be sensitive to the deep-sea blowout. Polychaete taxa varied in their sensitivity, but Dorvilleidae which is often associated with organic enrichment, was responsible for the largest amount of dissimilarity between stations close and far from the wellhead. Several other taxa were classified as sensitive or tolerant to the deep-sea blowout by comparing their distributions among impacted and non-impacted zones. The macrobenthic communities in the deep Gulf of Mexico exhibit a toxic response to the blowout on the Deepwater Horizon well, and this is correlated with barium and petroleum hydrocarbons.     

Reef fish community structure in the Tropical Eastern Pacific (Panamá): living on a relatively stable rocky reef environment

We compared the community structure of reef fish over different physical complexities in 12 study zones of Bahía Honda, Gulf of Chiriquí (BH-GCH), Tropical Eastern Pacific (TEP), Panama, aiming at an analysis of the importance of the physical structure provided by corals, rocks and benthic sessile organisms. This was the first region that emerged in the Isthmus of Panama; it exhibits the oldest benthic fauna and has constant conditions in terms of temperature and salinity. Two hundred and eighty-eight visual fish censuses were conducted on 48 benthic transects from February to July 2003. One hundred and twenty-six fish species of 44 families were found. Plankton feeding pomacentrids and labrids along with haemulids that feed on mobile invertebrates were the most abundant, particularly in shallow areas. Fourteen species showed size-segregations between zones, suggesting ontogenetic migrations (smaller fishes in shallow high-complexity zones, larger-sized fishes in deeper habitats). Highly mobile and site-attached genera were abundant in most shallow, wave-exposed zones particularly on exposed rocky substrates. Planktivores were the most abundant, followed by carnivores, feeders on mobile invertebrate and piscivores. Herbivores and feeders on sessile invertebrate were lower in abundance. Species richness exceeds that of any other studied region close to the mainland in the TEP and correlates with substrate diversity, increasing size-heterogeneity of holes and structural complexity. Species diversity increases with habitat complexity and benthic diversity. It seems that water current strength, tides and waves which select for swimming, play an important role in the community organization. The study region has been proposed as a refuge-centre in the TEP, where reef fishes that evolved on coral reefs have shifted their distribution onto rocky reef habitats.     

Zonation of intertidal macrobenthos in the estuaries of Schelde and Ems

Based on data, collected in 1980–1990, the intertidal benthic macrofauna of the Schelde and Ems estuaries was compared. The spatial occurrence of the benthic macrofauna along the salinity gradient, including the freshwater tidal area was emphasized. Both estuaries appeared to have a very similar species composition, especially at genus level. The higher number of species observed in the Schelde estuary was probably due to a greater habitat diversity. In both estuaries species diversity decreased with distance upstream. The total density did not vary along the estuarine gradient, whereas biomass is highest in the polyhaline zone.In both estuaries distinct intertidal benthic communities were observed along the salinity gradient: a marine community in the polyhaline zone, a brackish community in the mesohaline zone, and a third community in the oligohaline and freshwater tidal zones of the estuary. These three communities were very similar between both estuaries. Their main characteristics were discussed together with the occurrence and distribution of the dominant species.For the Schelde estuary and to a lesser extent also for the Ems estuary, there was evidence that anthropogenic stress had a negative effect on the intertidal macrobenthic communities of the oligohaline/freshwater tidal zone. Only Oligochaeta were dominating, whereas the very euryhaline and/or true limnetic species were missing. In the mesohaline zone, the Schelde estuary was dominated by large numbers of short-living, opportunistic species, whereas in the Ems estuary relatively more stable macrobenthic communities were observed. A comparison with some other European estuaries showed in general similar trends as those observed for the Schelde and Ems estuaries.     

Effects of Deposit-Feeding Macrofauna on Benthic Bacteria,Viruses, and Protozoa in a Silty Freshwater Sediment

In microcosm experiments, we simultaneously tested the effects of increased numbers of deposit-feeding macrofauna (chironomids, oligochaetes and cladocerans) on the standing stock, activities and interactions of heterotrophic bacteria, viruses, and bacterivorous protozoa (heterotrophic nanoflagellates and ciliates) in the aerobic layer of a silty littoral freshwater sediment. On average, bacterial secondary production was stimulated between 11 and 29% by all macrofaunal groups compared to control experiments without macrofauna addition. Bacterial standing stock increased significantly by 8 and 13% in case of chironomids and cladocerans, respectively. Oligochaetes and chironomids produced significant negative effects on viral abundance while the results with cladocerans were inconsistent. The addition of oligochaetes and chironomids resulted in a significant decrease by on average 68 and 32% of viral decay rates, respectively, used as a measure of viral production. The calculated contribution of virus-induced lysis to benthic bacterial mortality was low, with 2.8 to 11.8% of bacterial secondary production, and decreased by 39 to 81% after the addition of macrofauna compared to the control. The abundances of heterotrophic nanoflagellates were significantly reduced by 20% by all tested macrofauna groups, while ciliates showed inconsistent results. The importance of heterotrophic nanoflagellate grazing on benthic bacteria was very low (<1% of bacterial secondary production) and was further reduced by elevated numbers of macrofauna. Thus, the selected deposit feeding macrofauna groups seem to have several direct and indirect and partly antagonistic effects on the benthic bacterial compartment through the enhancement of bacterial production and the reduction of virus-induced cell lysis and protozoan grazing.     

Patterns and processes in abundance-body size relationships for marine benthic invertebrates

1. The nature of abundance-body size relationships in animal communities, and especially the drivers behind the observed patterns, have been a focus of persistent debate in animal ecology. In a recent review, Allen et al. (2006) categorized five mechanistic explanations behind the commonly observed polymodality in these relationships: energetic constraints; phylogenetic constraints; biogeographical determinants; habitat structure; and community interactions. Progress in understanding of these patterns and the processes underlying them have been hindered by the use of a range of methods that differ in their validity and robustness. 2. Here, we used data on invertebrate body sizes from a variety of sandy beaches in the UK to test the hypothesis that these communities display modality in their abundance-body size relationships. We quantified modality in the relationships using kernel density estimation and smoothed bootstrap resampling and then evaluated the competing explanations for this modality based on the patterns identified in conjunction with measurements of the physical beach environment. 3. We found bimodal distributions in the body size spectrum for benthic invertebrates at nine of 16 sites. There was a consistent trough in the spectrum at around 0·5-1 mm diameter, which reflected the traditional split between meiofauna and macrofauna. Beaches with finer particle sizes and more heterogeneous macrofauna hosted communities with more than two modes. 4. Our results suggest that modality in sandy beach benthic communities is unlikely to be explained by any single hypothesis. There will be an interplay between physical and biological factors, with different explanations accounting for modality at different scales.