Macroinvertebrate abundance and proportion of trait categories in river biotopes across England and Wales

There is a long tradition of river monitoring using macroinvertebrate communities to assess environmental quality in Europe. A promising alternative is the use of species life-history traits. Both methods, however, have relied on the time-consuming identification of taxa. River biotopes, 1-100 m**2 'habitats' with associated species assemblages, have long been seen as a useful and meaningful way of linking the ecology of macroinvertebrates and river hydro-morphology and can be used to assess hydro-morphological degradation in rivers. Taxonomic differences, however, between different rivers had prevented a general test of this concept until now. The species trait approach may overcome this obstacle across broad geographical areas, using biotopes as the hydro-morphological units which have characteristic species trait assemblages. We collected macroinvertebrate data from 512 discrete patches, comprising 13 river biotopes, from seven rivers in England and Wales. The aim was to test whether river biotopes were better predictors of macroinvertebrate trait profiles than taxonomic composition (genera, families, orders) in rivers, independently of the phylogenetic effects and catchment scale characteristics (i.e. hydrology, geography and land cover). We also tested whether species richness and diversity were better related to biotopes than to rivers. River biotopes explained 40% of the variance in macroinvertebrate trait profiles across the rivers, largely independently of catchment characteristics. There was a strong phylogenetic signature, however. River biotopes were about 50% better at predicting macroinvertebrate trait profiles than taxonomic composition across rivers, no matter which taxonomic resolution was used. River biotopes were better than river identity at explaining the variability in taxonomic richness and diversity (40% and <=10%, respectively). Detailed trait-biotope associations agreed with independent a priori predictions relating trait categories to near river bed flows. Hence, species traits provided a much needed mechanistic understanding and predictive ability across a broad geographical area. We show that integration of the multiple biological trait approach with river biotopes at the interface between ecology and hydro-morphology provides a wealth of new information and potential applications for river science and management.

Ecological investigations in the wadden area off Borkum, East Frisian Islands, Germany

Die Bodentiergemeinschaft des Wattenmeeres ist von Frühjahr bis Herbst eines jeden Jahres durch extrem hohe Dichten von Jungtieren charakterisiert. Die Kenntnisse über die Ansiedlung von fplanktischen Larven im Wattenmeer, sowie die Dynamik postlarvaler Stadien sind aufgrund der üblicherweise verwendeten, großen Siebmaschenweiten gering. Gerade aber diesen Altersstadien kommt möglicherweise eine besondere Stellung im Energiefluß des Wattenmeeres zu. An 5 Stationen (von NWL bis HWL, B1-B5) im Rückseitenwatt der ostfriesischen Insel Borkum wurden 1986 Ansiedlung, räumliche Verteilung, Wachstum, Mortalität und Produktion der Altersklasse 0 von Macoma balthica, Mya arenaria und Cerastoderma edule untersucht. Um die Ansiedlung der planktotrophen Larven dieser Arten zu beschreiben, wurden ihre Dichten in Plankton und Bodenproben miteinander verglichen. Die Untersuchungen zur Dynamik der benthischen Stadien wurden mit zwei in der Probenfläche und der Siebmaschenweite unterschiedlichen Probenserien durchgeführt. Die Drift postlarvaler Stadien wurde durch bodennahe Planktonfänge innerhalb des Eulitorals nachgewiesen. Parallel zu den Untersuchungen an der Endofauna wurden das Vorkommen und die Größe epibenthischer Räuber im Untersuchungsgebiet erfaßt. Die Hauptansiedlung von M. balthica- und M. arenaria-Larven erfolgte nahezu gleichzeitig Ende Mai/Anfang Juni. Die meisten Larven beider Arten gingen an der prielnächsten (tiefsten) Station (B1) zum Bodenleben über, gefolgt von der nächst höher gelegenen Station B2. Während frühe Bodenstadien von M. arenaria nicht im oberen Bereich des Watts (B3,B4) gefunden wurden, ist eine geringfügige Erstansiedlung von M. balthica in diesem Gebiet nicht auszuschließen. Ein die Ansiedlung limitierender Einfluß der relativ dichten Mya arenaria-Siedlung an den Stationen B1 und B2 sowie der Alttiere von M. balthica konnte nicht festgestellt werden. Die Ähnlichkeit des Ansiedlungsprozesses bei beiden Arten, die sich im Zahlenverhältnis Larvenangebot zu Anzahl der ersten Bodenstadien widerspiegelt, kann ein Hinweis auf eine überwiegend passive Ansiedlung der Larven am Boden sein. Der Ort der Hauptansiedlung von C. edule wurde durch den Transekt nicht erfaßt. Die Station B2 war zwar durch ein Herzmuschelfeld charakterisiert, dieses war aber nach zwei Eiswintern nahezu vollständig eliminiert. Der Abundanz der planktischen Larven zufolge war der Hauptansiedlungszeitraum ebenfalls Ende Mai/Anfang Juni. Zu dieser Zeit wurden nur vereinzelt frühe Bodenstadien an den Stationen B1 und B2 gefunden, keine an den Stationen B3 und B4. Während die frühen postlarvalen Stadien von M. arenaria überwiegend am Ort der Ansiedlung blieben, verbreiteten sich die von M. balthica bis in den oberen Bereich des Untersuchungsgebietes (B3-B5). Analog zu der Besiedlung dieser Gebiete durch postlarvale M. balthica wurde die im Verlauf des Untersuchungsjahres stattfindende Kolonisierung der Station B1 durch C. edule ebenfalls postlarvalem Transport zugeschrieben. Demzufolge spielt bei beiden Muschelarten postlarvaler Transport eine wichtige Rolle bei der Besiedlung von Habitaten. Planktonfänge innerhalb der bodennahen Wasserschicht bestätigten, daß im Untersuchungsgebiet M. balthica die am stärksten verdriftende Muschelart war, gefolgt von C. edule. Mortalität, Wachstum, mittlere Biomasse, Produktion und P/B-Verhältnis wurden für M. balthica an den Stationen B1, B3 und B4 sowie für M. arenaria an der Station B1 bestimmt. Wachstum und damit auch Produktion beider Arten erwiesen sich hier - wie an den höher gelegen Stationen (nur M. balthica) - als durch größenselektiven Feinddruck beeinflußt. Der Effekt postlarvalen Transports auf Wachstum wird diskutiert. Übergreifend über die auf Artebene diskutierten Ergebnisse wird die Bedeutung der Dispersion postlarvaler Stadien und die Wirkung epibenthischen Feinddrucks im Wattenmeer erörtert. Der Vergleich postlarvalen Transportes mit der Dispersion planktischer Larvenstadien, der Dispersion von Meiofauna und der Mobilität adulter Stadien der Makrofauna verdeutlicht, daß es sich hierbei um eine Strategie handeln kann, innerhalb eines unvorhersagbaren Biotops freiwerdende Ressourcen zu nutzen und dadurch Konkurrenz zu vermeiden. Es wird die Hypothese aufgestellt, daß Initialansiedlung und Immigration einerseits sowie Feinddruck und Emigration andererseits einen Regelkreis darstellen, der in verschiedenen Teilbereichen des Watts mit unterschiedlicher Geschwindigkeit abläuft.

Benthos investigations in the Lena Delta

According to monitoring data gained between 1982-1992, macrobenthos in the Tiksi Bay is characterized by low indices of the total abundance, biomass and taxonomic diversity. 30 macrobenthic species have been recorded in the Tiksi Bay. The bottom biocenoses within the estuarine-arctic water mass consist of widespread eurybiontic boreal-arctic and brackish-water species. The maximal number of species was observed at a depth of 8.5 m. The maximum biomass was recorded on muddy grounds. The studied bottom fauna is characterized by a high population density (from 1160-600 ind/m**2) and low biomass of 15.5-22.4 g/m**2. The predominant benthic animals of the main Lena River channel 4.7 km upstream Stolb Island are Chironomidae, Plecoptera and Oligochaeta. In total, 48 species of macrobenthos were registered here. In spring the average density of macrozoobenthos in the channel is 680, in summer 770, in autumn 720 and in winter 380 ind/m**2, with the average biomass varying between 2.9 g/m**2 in spring, 7.06 in summer, 4.4 in autumn, and 2.6 in winter.

Meiofauna densities, nematode biomass, nematode genus diversity, sediment grain size, organic matter, and pigments for the surface layers of two sites in summer (November 2009) and winter (august 2010)

The meiobenthic community of Potter Cove (King George Island, west Antarctic Peninsula) was investigated, focusing on responses to summer/winter conditions in two study sites contrasting in terms of organic matter inputs. Meiofaunal densities were found to be higher in summer and lower in winter, although this result was not significantly related to the in situ availability of organic matter in each season. The combination of food quality and competition for food amongst higher trophic levels may have played a role in determining the standing stocks at the two sites. Meiobenthic winter abundances were sufficiently high to infer that energy sources were not limiting during winter, supporting observations from other studies for both shallow water and continental shelf Antarctic ecosystems. Recruitment within meiofaunal communities was coupled to the seasonal input of fresh detritus for harpacticoid copepods but not for nematodes, suggesting that species-specific life history or trophic features form an important element of the responses observed.

Nematodes and meiofauna in sediments obtained from RRS Charles Darwin cruise CD179, Portuguese continental margin

Samples collected at two different depths (ca. 3200 and ca. 4200 m) in the Setúbal and Cascais canyons off the Portuguese coast, during the HERMES RRS Charles Darwin cruise CD179, were analysed for (1) sediment biogeochemistry (TOC, TN) and (2) composition, and structural and trophic diversity of nematode communities. Multivariate PERMANOVA analysis on the nematode community data revealed differences between sediment layers that were greater than differences between canyons, water depths, and stations. This suggests that biogeochemical gradients along the vertical sediment profile are crucial in determining nematode community structure. The interaction between canyon conditions and the nematode community is illustrated by biogeochemical patterns in the sediment and the prevalence of nematode genera that are able to persist in disturbed sediments. Trophic analysis of the nematode community indicated that non-selective deposit feeders are dominant, presumably because of their non-selective feeding behaviour compared to other feeding types, which gives them a competitive advantage in exploiting lower-quality food resources. This study presents a preliminary conceptual scheme for interactions between canyon conditions and the resident fauna.

Nematodes and meiofauna in sediments obtained from RRS Discovery cruise D297, Portuguese continental margin

Samples collected in the deep Nazaré Canyon and at the adjacent slope, during the HERMES RRS Discovery D297 cruise (2005), were analysed for metazoan meiofauna, nematode structure and diversity and its relation to quality and quantity of sedimentary organic material. The amount and quality of organic matter available for direct consumption was much higher in the canyon compared to the slope and positively correlated with high nematode abundances (795-1171 ind. 10 cm**-2) and biomass (93.2-343.5 µg dry weight 10 cm**-2), thus leading to higher standing stocks. Canyon nematode assemblages also showed particular adaptations (e.g. higher trophic complexity, variability of nematode morphology, and presence of opportunistic genera) to canyon conditions, particularly in the deeper sediment layers. The Nazaré Canyon's nematode diversity was slightly lower than that of the adjacent slope and its assemblages were characterised by a higher dominance of certain genera. Still, the canyon contributes considerably to total Western Iberian Margin diversity due to different assemblages present compared to the slope. Furthermore, the harsh conditions in terms of hydrodynamic disturbance and the high organic matter flux are likely to have a negative impact on the establishment of species rich meiobenthic communities, especially in the canyon axis.

Nematodes and meiofauna in sediments obtained during Belgica cruise BG06/13, north-east Atlantic

The Gollum Channels and Whittard Canyon (NE Atlantic) are two areas that receive high input of organic matter and phytodetritus from euphotic layers, but they are typified by different trophic and hydrodynamic conditions. Sediment biogeochemistry was analysed in conjunction with structure and diversity of the nematode community and differences were tested between study areas, water depths (700 m vs 1000 m), stations, and sediment layers. The Gollum Channels and Whittard Canyon harboured high meiofauna abundances (1054-1426 ind. 10 cm**-2) and high nematode diversity (total of 181 genera). Next to enhanced meiofauna abundance and nematode biomass, there were signs of high levels of organic matter deposition leading to reduced sedimentary conditions, which in turn structured the nematode community. Striking in this respect was the presence of large numbers of 'chemosynthetic' Astomonema nematodes (Astomonema southwardorum, Order Monhysterida, Family Siphonolaimidae). This genus lacks a mouth, buccal cavity and pharynx and possesses a rudimentary gut containing internal, symbiotic prokaryotes which have been recognised as sulphur-oxidising bacteria. Dominance of Astomonema may indicate the presence of reduced environments in the study areas, which is partially confirmed by the local biogeochemical environment. The nematode communities were mostly affected by sediment layer differences and concomitant trophic conditions rather than other spatial gradients related to study area, water depth or station differences, pointing to small-scale heterogeneity as the main source of variation in nematode structure and function. Furthermore, the positive relation between nematode standing stocks, and quantity and quality of the organic matter was stronger when hydrodynamic disturbance was greater. Analogically, this study also suggests that structural diversity can be positively correlated with trophic conditions and that this relation is tighter when hydrodynamic disturbance is greater.

Results from field observations and in situ 13C feeding experiments in the Nazaré Canyon conducted during RRS James Cook 10 cruise Leg 2

Submarine canyon systems provide a heterogeneous habitat for deep-sea benthos in terms of topography, hydrography, and the quality and quantity of organic matter present. Enhanced meiofauna densities as found in organically enriched canyon sediments suggest that nematodes, as the dominant metazoan meiobenthic taxon, may play an important role in the benthic food web of these sediments. Very little is known about the natural diets and trophic biology of deep-sea nematodes, but enrichment experiments can shed light on nematode feeding selectivity and trophic position. An in-situ pulse-chase experiment (Feedex) was performed in the Nazaré Canyon on the Portuguese margin in summer 2007 to study nematode feeding behaviour. 13C-labelled diatoms and bacteria were added to sediment cores which were then sampled over a 14-day period. There was differential uptake by the nematode community of the food sources provided, indicating selective feeding processes. 13C isotope results revealed that selective feeding was less pronounced at the surface, compared to the sediment subsurface. This was supported by a higher trophic diversity in surface sediments compared to the subsurface, implying that more food items may be used by the nematode community at the sediment surface. Predatory and scavenging nematodes contributed relatively more to biomass than other feeding types and can be seen as key contributors to the nematode food web at the canyon site. Non-selective deposit feeding nematodes were the dominant trophic group in terms of abundance and contributed substantially to total nematode biomass. The high levels of 'fresh' (bioavailable) organic matter input and moderate hydrodynamic disturbance of the canyon environment lead to a more complex trophic structure in canyon nematode communities than that found on the open continental slope, and favours predator/scavengers and non-selective deposit feeders.

Antarctic shallow water benthos from three stations at Potter Cove, King George Island, West Antarctic Peninsula

The West Antarctic Peninsula is one of the fastest warming regions on the planet. Faster glacier retreat and related calving events lead to more frequent iceberg scouring, fresh water input and higher sediment loads which may affect benthic marine communities. On the other hand, the appearance of newly formed ice-free areas provides new substrates for colonization. Here we investigated the effect of these conditions on four benthic size classes (microbenthos, meiofauna and macrofauna) using Potter Cove (King George Island, West Antarctic Peninsula) as a case study. We identified three sites within the cove experiencing different levels of glacier retreat-related disturbance. Our results showed the existence of different communities at the same depth over a relatively small distance (about 1 km**2). This suggests glacial activity structures biotic communities over a relatively small spatial scale. In areas with frequent ice scouring and higher sediment accumulation rates, a patchy community, mainly dominated by macrobenthic scavengers (such as Barrukia cristata), vagile organisms, and younger individuals of sessile species (such as Yoldia eigthsi) was found. Meiofauna organisms such as cumaceans are found to be resistant to re-suspension and high sedimentation loads. The nematode genus Microlaimus was found to be successful in the newly exposed ice-free site, confirming its ability as a pioneering colonizer. In general, the different biological size classes appear to respond in different ways to the ongoing disturbances, suggesting that adaptation processes may be size related. Our results suggest that with continued deglaciation, more diverse but less patchy macrobenthic assemblages can become established due to less frequent ice scouring events.

Sublittoral and bathyal Harpacticoida (Crustacea: Copepoda) of the Magellan region

Two expeditions, undertaken in 1994 and 1996, provided quantitatively sampled material of sublittoral and bathyal meiobenthos from the Paso Ancho of the Straits of Magellan, the Beagle Channel, and the Patagonian continental slope (Chile). To investigate whether these distinct geographic areas might also be characterised by different harpacticoid assemblages, qualitative and quantitative analyses of Copepoda Harpacticoida were carried out. At supraspecific level 25 harpacticoid families were found, as well as several species that could not yet be assigned to any major harpacticoid taxon. Due to the high amount of collected Harpacticoida, detailed investigations at species level had to be restricted to six taxa, namely the Ancorabolidae, Argestidae, Cletodidae, Diosaccinae, Paramesochridae, and Paranannopinae. The corresponding specimens were assigned to 122 species in 52 genera. More than 80% of them are new to science. Qualitative comparisons of both species composition and species distribution allow the three areas to be distinguished in terms of species richness. However, statistical analyses confirm these results only partly. Similarity analyses applying non-metrical multidimensional scaling, as well as diversity analyses using the rarefaction method, suggest that the observed differences in distribution and diversity patterns are due to small-scale, local conditions, which may overlay possible large-scale ones.

Abundance and biomass of in- and epibenthic invertebrate macrofauna in the German Bight from 1969 to 2000

In order to examine the long-term development of offshore macrozoobenthic soft-bottom communities of the German Bight, four representative permanent stations (MZB-SSd, -FSd, -Slt, -WB) have been sampled continuously since 1969. Inter-annual variability and possible long-term trends were analysed based on spring-time samples from 1969 until 2000. This is part of the ecological long-term series of the AWI and is supplemented by periodic large-scale mapping of the benthos. The main factors influencing the development of the benthic communities are biological interactions, climate, food supply (eutrophication) and the disturbance regime. The most frequent disturbances are sediment relocations during strong storms or by bottom trawling, while occasional oxygen deficiencies and extremely cold winters are important disturbance events working on a much larger scale. Benthic communities at the sampling stations show a large inter-annual variability combined with a variation on a roughly decadal scale. In accordance with large-scale system shifts reported for the North Sea, benthic community transitions occurred between roughly the 1970ies, 80ies and 90ies. The transitions between periods are not distinctly marked by strong changes but rather reflected in gradual changes of the species composition and dominance structure.