Cross-taxa generalities in the relationship between population abundance and ambient temperatures.

Identifying patterns in the effects of temperature on species' population abundances could help develop a general framework for predicting the consequences of climate change across different communities and realms. We used long-term population time series data from terrestrial, freshwater, and marine species communities within central Europe to compare the effects of temperature on abundance across a broad range of taxonomic groups. We asked whether there was an average relationship between temperatures in different seasons and annual abundances of species in a community, and whether species attributes (temperature range of distribution, range size, habitat breadth, dispersal ability, body size, and lifespan) explained interspecific variation in the relationship between temperature and abundance. We found that, on average, warmer winter temperatures were associated with greater abundances in terrestrial communities (ground beetles, spiders, and birds) but not always in aquatic communities (freshwater and marine invertebrates and fish). The abundances of species with large geographical ranges, larger body sizes, and longer lifespans tended to be less related to temperature. Our results suggest that climate change may have, in general, positive effects on species' abundances within many terrestrial communities in central Europe while the effects are less predictable in aquatic communities.

Commonness and rarity in the marine biosphere.

Explaining patterns of commonness and rarity is fundamental for understanding and managing biodiversity. Consequently, a key test of biodiversity theory has been how well ecological models reproduce empirical distributions of species abundances. However, ecological models with very different assumptions can predict similar species abundance distributions, whereas models with similar assumptions may generate very different predictions. This complicates inferring processes driving community structure from model fits to data. Here, we use an approximation that captures common features of "neutral" biodiversity models--which assume ecological equivalence of species--to test whether neutrality is consistent with patterns of commonness and rarity in the marine biosphere. We do this by analyzing 1,185 species abundance distributions from 14 marine ecosystems ranging from intertidal habitats to abyssal depths, and from the tropics to polar regions. Neutrality performs substantially worse than a classical nonneutral alternative: empirical data consistently show greater heterogeneity of species abundances than expected under neutrality. Poor performance of neutral theory is driven by its consistent inability to capture the dominance of the communities' most-abundant species. Previous tests showing poor performance of a neutral model for a particular system often have been followed by controversy about whether an alternative formulation of neutral theory could explain the data after all. However, our approach focuses on common features of neutral models, revealing discrepancies with a broad range of empirical abundance distributions. These findings highlight the need for biodiversity theory in which ecological differences among species, such as niche differences and demographic trade-offs, play a central role.

Baseline Inventory of Benthic Macrofauna in German Marine Protected Areas (2020-2022) before Closure for Bottom-Contact Fishing.

The response of benthic habitats and organisms to bottom-contact fishing intensity is investigated in marine protected areas (MPAs) of the German EEZ in the North and Baltic Seas. We examined the current state of macrofauna biodiversity in 2020-2022. Comparative analysis for macrofauna (in- and epifauna) inhabiting nine Natura 2000 MPAs constitutes a baseline to assess the effects of bottom-contact fishing exclusion in the future. Aspects of spatial and temporal variability are briefly summarized and discussed. We provide a species list for each region, including 481 taxa, of which 79 were found in both regions, 183 only in the North Sea, and 219 only in the Baltic Sea. The Baltic Sea dataset surprisingly included higher numbers of taxa and revealed more Red List species. The share of major taxonomic groups (polychaetes, bivalves and amphipods) in species richness showed peculiar commonalities between the two regions. In the North Sea, multivariate analysis of community structure revealed significantly higher within-similarity and stronger separation between the considered MPAs compared to the Baltic MPAs. Salinity, temperature and sediment fractions of sand were responsible for over 60% of the variation in the North Sea macrofauna occurrence data. Salinity, mud fraction and bottom-contact fishing were the most important drivers in the Baltic Sea and, together with other considered environmental drivers, were responsible for 53% of the variation. This study identifies aspects of macrofauna occurrence that may be used to assess (causes of) future changes.

Application of equilibrium passive sampling to assess the influence of anthropogenic activities and bioturbation on the distribution of hydrophobic organic chemicals in North Sea sediment cores.

The pollution state in the German Bight was investigated by determination of pollutant concentrations of sediment samples using equilibrium passive sampling. Polychlorinated biphenyls (PCBs) and polycyclic aromatic hydrocarbons (PAH) were determined in the pore water of North Sea sediment. The freely dissolved pore water concentration (Cfree) was measured applying Solid Phase Microextraction (SPME) by using PDMS-coated glass fibers. The obtained results show that the North Sea contamination level with the investigated pollutants is relatively low. However, the stations close to the sediment-dumping site were higher contaminated. A macrofauna analysis showed that bioturbation activities were mostly present in the upper sediment layers, but a direct bioturbation influence on the sediment concentration distribution could not be shown. Overall, the contamination load was below baseline toxicity, but considering that several other priority pollutants will also make a contribution to the baseline toxicity, it can be counted as relatively high.

Abundance, biomass and species richness of macrozoobenthos along an intertidal elevation gradient.

Ecology aims to comprehend species distribution and its interaction with environmental factors, from global to local scales. While global environmental changes affect marine biodiversity, understanding the drivers at smaller scales remains crucial. Tidal flats can be found on most of the world's coastlines and are particularly vulnerable to anthropogenic disturbances. They are important transient ecosystems between terrestrial and marine ecosystems, and their biodiversity provides important ecosystem services. Owing to this unique, terrestrial-marine transition, strong environmental gradients of elevation, sediment composition and food availability prevail. Here, we investigated which regional and local environmental factors drive the spatio-temporal dynamics of macrozoobenthos communities on back-barrier tidal flats in the East Frisian Wadden Sea. On the regional level, we found that species composition changed significantly from west to east on the East Frisian islands and that total abundance and species richness decreased from west to east. On the local abiotic level, we found that macrozoobenthos biomass decreased with higher elevation towards the salt marsh and that the total abundance of organisms in the sediment significantly increased with increasing mud content, while biodiversity and biomass were not changing significantly. In contrast to expectations, increasing Chl a availability as a measure of primary productivity did not result in changes in abundance, biomass or biodiversity, but extremely high total organic carbon (TOC) content was associated with a decrease in biomass and biodiversity. In conclusion, we found regional and local relationships that are similar to those observed in previous studies on macrozoobenthos in the Wadden Sea. Macrozoobenthos biomass, abundance and biodiversity are interrelated in a complex way with the physical, abiotic and biotic processes in and above the sediment.

Benthic ecosystem functioning under climate change: modelling the bioturbation potential for benthic key species in the southern North Sea.

Climate change affects the marine environment on many levels with profound consequences for numerous biological, chemical, and physical processes. Benthic bioturbation is one of the most relevant and significant processes for benthic-pelagic coupling and biogeochemical fluxes in marine sediments, such as the uptake, transport, and remineralisation of organic carbon. However, only little is known about how climate change affects the distribution and intensity of benthic bioturbation of a shallow temperate shelf sea system such as the southern North Sea. In this study, we modelled and projected changes in bioturbation potential (BPp) under a continuous global warming scenario for seven southern North Sea key bioturbators: Abra alba, Amphiura filiformis, Callianassa subterranea, Echinocardium cordatum, Goniada maculata, Nephtys hombergii, and Nucula nitidosa. Spatial changes in species bioturbation intensity are simulated for the years 2050 and 2099 based on one species distribution model per species driven by bottom temperature and salinity changes using the IPCC SRES scenario A1B. Local mean bottom temperature was projected to increase between 0.15 and 5.4 °C, while mean bottom salinity was projected to moderately decrease by 1.7. Our results show that the considered benthic species are strongly influenced by the temperature increase. Although the total BP remained rather constant in the southern North Sea, the BPp for four out of seven species was projected to increase, mainly due to a simultaneous northward range expansion, while the BPp in the core area of the southern North Sea declined for the same species. Bioturbation of the most important species, Amphiura filiformis and Echinocardium cordatum, showed no substantial change in the spatial distribution, but over time. The BPp of E. cordatum remained almost constant until 2099, while the BPp of A. filiformis decreased by 41%. The northward expansion of some species and the decline of most species in the south led to a change of relative contribution to bioturbation in the southern North Sea. These results indicate that some of the selected key bioturbators in the southern North Sea might partly compensate the decrease in bioturbation by others. But especially in the depositional areas where bioturbation plays a specifically important role for ecosystem functioning, bioturbation potential declined until 2099, which might affect the biochemical cycling in sediments of some areas of the southern North Sea.

Climate change effects on marine protected areas: Projected decline of benthic species in the North Sea.

Climate change is a global threat for marine ecosystems, their biodiversity and consequently ecosystem services. In the marine realm, marine protected areas (MPAs) were designated to counteract regional pressures, but they might be ineffective to protect vulnerable species and habitats, if their distribution is affected by global climate change. We used six Species Distribution Models (GLM, MARS, FDA, RF, GBM, MAXENT) to project changes in the distribution of eight benthic indicator and key species under climate change in the North Sea MPAs for 2050 and 2099. The projected distribution area of most species will be stable or even increase within the MPAs between 2001 and 2050. Thereafter, the distribution area decreased, especially within MPAs in the central North Sea by 2099, and some key species even disappeared from the MPAs. Consequently, the monitoring and protection of benthic species might not be possible within static MPA borders under climate change.

The role of artificial material for benthic communities - Establishing different concrete materials as hard bottom environments.

Concrete is used in marine coastal constructions worldwide. These structures are colonized by specialized hard-bottom biota consisting of macroalgae and benthic macrofauna. As concrete manufacturers face challenges such as limited natural resources and high CO2-emissions, the need for supplementary materials increases. Still, there has been little research on the reaction of species to the differences in concrete composition and what ecological impact these reactions could have. This study addresses the questions (1) if there are differences in settlement communities, depending on differences in concrete constitutes and (2) if so, what are the consequences for the usability of alternative concretes in marine constructions. For the experiment 15 cubes (15 × 15 × 15 cm) made of five different concretes, containing different cements (Portland cement and blast furnace cements) and aggregates (sand, gravel, iron ore and metallurgical slags) were deployed in a natural hard bottom experimental field near Helgoland Island (German Bight) in April 2016. After 12 months, all cubes were examined regarding species composition and coverage, followed by statistical analysis (PERMANOVA, SIMPER, DIVERSE). Results indicate differences in settlement communities for different surface orientation (Top, Front/Back) of the cubes. Significant differences in settlement communities of the Front/Back side were present depending on the used material type. However, the found differences in settlement between the concrete types tested are not sufficiently clear to provide recommendations for their usability in coastal constructions.

Meta-analysis of multidecadal biodiversity trends in Europe.

Local biodiversity trends over time are likely to be decoupled from global trends, as local processes may compensate or counteract global change. We analyze 161 long-term biological time series (15-91 years) collected across Europe, using a comprehensive dataset comprising ~6,200 marine, freshwater and terrestrial taxa. We test whether (i) local long-term biodiversity trends are consistent among biogeoregions, realms and taxonomic groups, and (ii) changes in biodiversity correlate with regional climate and local conditions. Our results reveal that local trends of abundance, richness and diversity differ among biogeoregions, realms and taxonomic groups, demonstrating that biodiversity changes at local scale are often complex and cannot be easily generalized. However, we find increases in richness and abundance with increasing temperature and naturalness as well as a clear spatial pattern in changes in community composition (i.e. temporal taxonomic turnover) in most biogeoregions of Northern and Eastern Europe.

Shifts in trait-based and taxonomic macrofauna community structure along a 27-year time-series in the south-eastern North Sea.

Current research revealed distinct changes in ecosystem functions, and thus in ecosystem stability and resilience, caused by changes in community structure and diversity loss. Benthic species play an important role in benthic-pelagic coupling, such as through the remineralization of deposited organic material, and changes to benthic community structure and diversity have associated with changes in ecosystem functioning, ecosystem stability and resilience. However, the long-term variability of traits and functions in benthic communities is largely unknown. By using abundance and bioturbation potential of macrofauna samples, taken along a transect from the German Bight towards the Dogger Bank in May 1990 and annually from 1995 to 2017, we analysed the taxonomic and trait-based macrofauna long-term community variability and diversity. Taxonomic and trait-based diversity remained stable over time, while three different regimes were found, characterised by changes in taxonomic and trait-based community structure. Min/max autocorrelation factor analysis revealed the climatic variables sea surface temperature (SST) and North Atlantic Oscillation Index (NAOI), nitrite, and epibenthic abundance as most important environmental drivers for taxonomic and trait-based community changes.

Full-coverage spatial distribution of epibenthic communities in the south-eastern North Sea in relation to habitat characteristics and fishing effort.

Full-coverage spatial data of occurrence and a detailed description of habitat requirements of epibenthic communities are needed in many conservation and management contexts. In the North Sea the focus has so far been on small benthic infauna, whereas structure and ecosystem functions of larger epifaunal communities have been largely ignored. This study provides a comprehensive analysis of epibenthic community structure in the south-eastern North Sea, including a detailed inventory of species, diversity and spatially contiguous distribution of communities. Data from nearly 400 stations were compiled for the study, enabling us to describe epibenthic community structure at unprecedented spatial resolution. Eight distinct epibenthic communities were found in the south-eastern North Sea by using multivariate analysis. Distribution modelling with eight environmental variables (bottom temperature and salinity, temperature differences between summer and winter, mud content of sediments, maximum bottom shear stress, stratification, water depth and annual primary production) and one human pressure (fishing effort) was used to extrapolate probable spatial distributions and to identify associated habitat characteristics of the communities in the south-eastern North Sea. Three large epibenthic communities "Coast", "Oysterground" and "Tail End" reflect a gradual habitat change from the coast towards offshore regions, expressed in gradients of bottom salinity, seasonal temperature differences and stratification as the dominant environmental factors. Five smaller communities ("Amrum Bank", "Frisian Front", "Deeps", "Dogger Bank" and "Dogger Slope") outline specific habitats in the south-eastern North Sea. The "Dogger Slope" community has not been recognized before, but has a predicted spatial extent of 7118 km2. Due to the high occurrence of long-lived, sessile species such as sponges this community is very sensitive to demersal fishing.

Cross-realm assessment of climate change impacts on species' abundance trends.

Climate change, land-use change, pollution and exploitation are among the main drivers of species' population trends; however, their relative importance is much debated. We used a unique collection of over 1,000 local population time series in 22 communities across terrestrial, freshwater and marine realms within central Europe to compare the impacts of long-term temperature change and other environmental drivers from 1980 onwards. To disentangle different drivers, we related species' population trends to species- and driver-specific attributes, such as temperature and habitat preference or pollution tolerance. We found a consistent impact of temperature change on the local abundances of terrestrial species. Populations of warm-dwelling species increased more than those of cold-dwelling species. In contrast, impacts of temperature change on aquatic species' abundances were variable. Effects of temperature preference were more consistent in terrestrial communities than effects of habitat preference, suggesting that the impacts of temperature change have become widespread for recent changes in abundance within many terrestrial communities of central Europe.

Seasonal variability of benthic indices: an approach to test the applicability of different indices for ecosystem quality assessment.

In recent studies, several benthic biological indices were developed or used to assess the ecological quality status of marine environments. In the present study the seasonal variability of several univariate and multimetric indices was studied on a monthly scale (September 2000 until May 2002) in different areas of the North Sea such as the German Bight, the Oyster Ground and the Dogger Bank. The stations were chosen to reflect a gradient in the hydrographic regime, temperature and organic matter supply. The seasonal variability was highest for the univariate indices such as the Shannon-Wiener Index and the Hurlbert Index. Thus, due to sensitivity to recruitment the corresponding ecological status ranged from 'good' to 'poor' depending on the season. For the multimetric indices such as the AMBI or the BQI the seasonal variability and the corresponding ecological status were low. The results are discussed concerning possible consequences for ecological quality assessment especially related to the Water Framework Directive (WFD).

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.