Henning Reiss

A bioturbation classification of European marine infaunal invertebrates

Bioturbation, the biogenic modification of sediments through particle reworking and burrow ventilation, is a key mediator of many important geochemical processes in marine systems. In situ quantification of bioturbation can be achieved in a myriad of ways, requiring expert knowledge, technology, and resources not always available, and not feasible in some settings. Where dedicated research programmes do not exist, a practical alternative is the adoption of a trait-based approach to estimate community bioturbation potential (BPc). This index can be calculated from inventories of species, abundance and biomass data (routinely available for many systems), and a functional classification of organism traits associated with sediment mixing (less available). Presently, however, there is no agreed standard categorization for the reworking mode and mobility of benthic species. Based on information from the literature and expert opinion, we provide a functional classification for 1033 benthic invertebrate species from the northwest European continental shelf, as a tool to enable the standardized calculation of BPc in the region. Future uses of this classification table will increase the comparability and utility of large-scale assessments of ecosystem processes and functioning influenced by bioturbation (e.g., to support legislation). The key strengths, assumptions, and limitations of BPc as a metric are critically reviewed, offering guidelines for its calculation and application.

Influence of macrofauna long-term natural variability on benthic indices used in ecological quality assessment

An essential prerequisite for the assessment of the ecological quality of marine ecosystems is the understanding of the natural variability and its effect on the performance of quality indices. This study is focused on the long-term natural variability of diversity, biotic and multimetric indices by using long-term macrofauna data of a coastal area in the southern North Sea (1978-2005). The univariate and most biotic and multimetric indices respond significantly on specific natural disturbance events such as cold winters, but the strength of response varied between indices as well as between events. As a result, the ecological quality status can decrease over a range of 3 (out of 5) classification units. The overall ecological quality was good to high, but an increase of indices occurred from the mid 1980s onwards due to changes in the climate regime. This long-term variability has to be considered within ecological quality assessment schemes.

20 years of the German Small-Scale Bottom Trawl Survey (GSBTS): A review

AbstractThe German Small-scale Bottom Trawl Survey (GSBTS) was initiated in 1987 in order to provide complementary investigations to the International Bottom Trawl Survey (IBTS) in the North Sea, using the same methodology but focussing high-intensity sampling on selected survey areas. Over the last 20 years, the initial number of 4 survey areas (10 × 10 nautical miles; “Boxes”) has been increased to 12, which are distributed over the entire North Sea. This paper describes the survey methods of the GSBTS, summarizes the scientific outcome of the first 20 years, and suggests that international fisheries research institutions would join the GSBTS.The major outcomes of the survey include to date:— Documentation changes in the distribution of fish species and in species assemblages (e.g. changes in species richness, shifts in the southern species component).— Geostatistical evaluation of GSBTS data.— Analysis of spatial scale effects: the relevance of GSBTS survey results for interpreting large-scaled abundance and distribution data from the IBTS.— Description of benthic habitats, composition of invertebrate fauna and its variability.— Process studies, especially investigation of predator-prey interactions between fish through analyses of stomach contents.— Characterization of the typical hydrographic conditions in the survey areas and their variability, and description of the nutrient supply.— Observations of seabirds and their feeding habits.— Analysis of the effects of different parameters on catch rates for bottom fish and on the estimates of abundance indices (e.g. vessel and gear effects, towing time, hydrographic conditions, time of day, number of hauls per area). In continuing this interdisciplinary survey with simultaneous sampling of all faunal and environmental compartments and especially in making it an international effort, we see the possibility of contributing data for the implementation of the ecosystems approach to fisheries management. Particularly, the following aspects can be addressed and would further increase the scientific value of the GSBTS:— Combining the survey data with highly resolved data from the commercial fishery to separate the effects of fishing from natural variability.— Further interdisciplinary analyses of the entire data set. Main aspects include benthos-fish-bird-community changes over time and their relation to historic fisheries impacts, and the coupling of biological and physical habitat characterisation.— Collection of accompanying data (phyto-, zoo- and ichthyoplankton data) in order to make the GSBTS a true ecosystem survey in detecting temporal changes in nearly all major levels of the food web.

Invertebrate associations with gastropod shells inhabited by Pagurus­bernhardus (Paguridae) – secondary hard substrate increasing biodiversity­in North Sea soft‐bottom communities

The crustacean Pagurus bernhardus is an abundant hermit crab species in the soft‐bottom benthic communities of the North Sea. The gastropod shells inhabited by the hermit crab were found to provide substrate for a diverse epizoic fauna. In order to describe the epizoan community as well as spatial differences in epizoan communities of the North Sea, hermit crabs were sampled during summer 2001 at five stations in the North Sea. In total, 51 epizoic species with up to 15 species and 427 individuals per crab were found. The most abundant epizoans were obligate associated species such as the polychaete Circeis amoricani paguri, the cnidaria Hydractinia echinata and the crustacean Trypetesa lampas, as well as sessile epizoans also found in hard‐bottom habitats such as balanids Balanus crenatus and Verruca stroemia or free‐living species such as the amphipod Gammaropsis nitida. The epizoic community structure on shells of the stations north of the 50 m contour differs from the southern communities on the Dogger Bank and in the German Bight. The northern stations were dominated by sessile species, whereas a higher proportion of free‐living epizoans occurred at the southern stations. The results are discussed concerning the ecological importance of shells inhabited by hermit crabs as a hard substrate in soft‐bottom habitats.

Biodiversity Trends along the Western European Margin

The seas along the western European margin encompass a vast geographical area comprising numerous different habitats, and are home to more than 10,000 metazoan species. Although research in this extensive region has been undertaken since the early 1800s, many new species are being described and distributional patterns identified. Recent studies incorporating the most extensive data series ever used in such European studies have failed to find any relationship between latitude and infaunal shelf biodiversity. Along the European shelf, species richness generally increases to a depth of 200 m and then decreases from 300–500 m. In the deep Northeast Atlantic, a unimodal curve illustrates how macrofaunal species diversity changes with depth whilst the megafauna appear to have a bimodal distribution. Regional studies are equivocal in that poleward increases in species diversity have been observed in some studies or taxa, but not in others. In the North Sea, arguably the best studied system in European waters, there appears to be a distinct increase in diversity with increasing latitude. Since this trend is confounded by similar latitudinal gradients in depth and trawling intensity, there is no clear explanation for the biodiversity pattern. Climatic shifts in diversity patterns and species ranges have recently been observed. Here we report previously unpublished data on changes in species richness that have been observed along the Norwegian coast over the past two decades, with the most northerly region seeing more than a 15% increase in the number of species being discovered there. This review synthesizes published and new biodiversity data across multiple spatial and temporal scales, and from the coast to the deep-sea, to provide an overview of what is known along the western European margin. Threats to the biodiversity of the region are highlighted, as well as identifying where there are still gaps in our knowledge.

On the Myths of Indicator Species : Issues and Further Consideration in the Use of Static Concepts for Ecological Applications

The use of static indicator species, in which species are expected to have a similar sensitivity or tolerance to either natural or human-induced stressors, does not account for possible shifts in tolerance along natural environmental gradients and between biogeographic regions. Their indicative value may therefore be considered at least questionable. In this paper we demonstrate how species responses (i.e. abundance) to changes in sediment grain size and organic matter (OM) alter along a salinity gradient and conclude with a plea for prudency when interpreting static indicator-based quality indices. Six model species (three polychaetes, one amphipod and two bivalves) from the North Sea, Baltic Sea and the Mediterranean Sea region were selected. Our study demonstrated that there were no generic relationships between environment and biota and half of the studied species showed different responses in different seas. Consequently, the following points have to be carefully considered when applying static indicator-based quality indices: (1) species tolerances and preferences may change along environmental gradients and between different biogeographic regions, (2) as environment modifies species autecology, there is a need to adjust indicator species lists along major environmental gradients and (3) there is a risk of including sibling or cryptic species in calculating the index value of a species.

Diet overlap among flatfish species in the southern North Sea.

The diets of two non-commercial flatfish species (solenette Buglossidium luteum and scaldfish Arnoglossus laterna) and two commercial flatfish species (dab Limanda limanda and plaice Pleuronectes platessa) were compared in a study area in the German Bight (southern North Sea) to investigate prey-resource partitioning between these species. The diets of A. laterna and B. luteum mainly comprised crustaceans (harpacticoids, amphipods, cumaceans and decapods), whereas the diet of L. limanda and P. platessa consisted mainly of polychaetes. The Schoener index, calculated for different fish size classes between these flatfish species, showed a biologically significant diet overlap between small-sized L. limanda and P. platessa and B. luteum and A. laterna, using similar prey resources of smaller prey (e.g. amphipods, harpacticoids and juvenile bivalves). In contrast, with increasing body size, a change in the diet of L. limanda and P. platessa towards larger prey occurred (e.g. polychaetes and decapods), resulting in low diet overlap values with B. luteum and A. laterna. Due to these size-related differences in resource use, it is assumed that there is reduced interspecific competition for prey between larger L. limanda and P. platessa and both non-commercial flatfishes, probably facilitating resource partitioning within the same area. In contrast, smaller L. limanda and P. platessa may compete directly for the same prey resources with B. luteum and A. laterna. Furthermore, prey availability of most important prey items of the studied flatfishes was relatively low in the study area. Therefore, increasing abundances of B. luteum and A. laterna in the southern North Sea since the late 1980s, owing to fishing effects and climate change, might affect the population dynamics of L. limanda and P. platessa.

Linking prey composition of haddock Melanogrammus aeglefinus to benthic prey availability in three different areas of the northern North Sea

The feeding strategy and prey selection of haddock Melanogrammus aeglefinus related to the benthic infauna in the field were investigated in three different study areas (boxes) in the northern North Sea in 2007. The stomach contents of M. aeglefinus were dominated by invertebrates in all three boxes, mainly echinoderms and polychaetes, similar to the benthic communities in the field. Prey densities in the field significantly determined prey selection and, thus the feeding strategy of M. aeglefinus appeared highly opportunistic. Other prey characteristics, such as the position in the sediment or its palatability, had no significant effect on the selection process although trends were apparent that tubicolous polychaetes of the family spionids, especially Spiophanes kröyeri and Spiophanes bombyx as well as the tentaculate Phoronis spp., were not consumed and potentially avoided due to their ability to withdraw below the feeding depths or due to chemical deterrents. High abundances of echinoderm species with hard calcareous shells in the M. aeglefinus stomachs, such as the ophiurid Ophiocten affinis and the echinoid Echinocyamus pusillus, indicated their use as grinding elements. A gradual shift from benthos to fish feeding with increasing M. aeglefinus size was not found.

Benthos and demersal fish habitats in the German Exclusive Economic Zone (EEZ) of the North Sea

We compiled data from different monitoring surveys to analyse and compare community and diversity patterns of fish, epi- and infauna in the German Exclusive Economic Zone (EEZ) of the North Sea in order to identify benthic habitats common to all faunal components. We found congruent community patterns of fish, epi- and infauna for the coastal waters, the Oysterground and the area called “Duck’s Bill”, which coincided with specific abiotic characteristics of these regions. The three regions were defined as special habitats for fish, epi- and infauna species in the German EEZ. The differences in the seasonal variability of abiotic factors seem to be the most important discriminating abiotic characteristic for the three habitats. The spatial distribution of fish, epifauna and infauna communities remained stable over time although habitat characteristics such as sea surface temperature increased due to climate change. However, it is expected that the coastal habitat will be more sensitive to future climate change effects in contrast to the Oysterground and Duck’s Bill habitat.

Spatial and temporal distribution of macrofauna in the Otzumer Balje (East Frisian Wadden Sea, Germany)

The spatial and temporal distribution of the macrofauna in a subtidal area of the East Frisian Wadden Sea (Otzumer Balje) was investigated. Grab samples were taken at 30 stations covering the whole tidal inlet area. Focusing on the seasonal variability sampling was carried out in March and September 1998 using a 0.1 m2 van Veen grab.During the survey, a total of 108 macrobenthic species was recorded, dominated by polychaetes and crustaceans contributing 47 and 36 species, respectively. Highest mean species numbers and abundances were found in the deeper central inshore parts of the channel, whereas the lowest were found in the highly dynamic inlet region. With a mean of 3.2 g AFDW/m2 the biomass of the subtidal macrofauna was relatively low compared to the intertidal fauna.Multivariate data analysis (MDS and clusteranalysis) showed three distinct faunal assemblages: (1) a community in the inlet region on sandy sediments characterized bySpio martinensis, Scoloplos armiger, Bathyporeia pelagica and, additionally, in September by the polychaetesMagelona johnstoni andScolelepis squamata; (2) a community in the shallow and deep inshore channels on muddy sediments dominated byTubificoides benedeni, Heteromastus filiformis andTharyx killariensis; and (3) an ‘intermediate community’ in shallower regions of the tidal channel and smaller gullies on muddy sands characterized bySpio martinensis, Tubificoides benedeni, Scoloplos armiger andTharyx killariensis.The seasonal spatial community structure was more or less stable, whereas the seasonal variability in the species composition within the communities was high, due to recruitment and import processes during the summer.The community structure is discussed in terms of ecosystem properties such as sediment composition and hydrography.