Crustaceans and fish abundances and species at and around artificially introduced tetrapod fields in the southern North Sea, 2009

The micro-scale spatial distribution patterns of a demersal fish and decapod crustacean assemblage were assessed in a hard-bottom kelp environment in the southern North Sea. Using quadrats along line transects, we assessed the in situ fish and crustacean abundance in relation to substratum types (rock, cobbles and large pebbles) and the density of algae. Six fish and four crustacean species were abundant, with Ctenolabrus rupestris clearly dominating the fish community and Galathea squamifera dominating the crustacean community. Differences in the substratum types had an even stronger effect on the micro-scale distribution than the density of the dominating algae species. Kelp had a negative effect on the fish abundances, with significantly lower average densities in kelp beds compared with adjacent open areas. Averaged over all of the substrata, the most attractive substratum for the fish was large pebbles. In contrast, crustaceans did not show a specific substratum affinity. The results clearly indicate that, similar to other complex systems, significant micro-scale species-habitat associations occur in northern hard-bottom environments. However, because of the frequently harsh environmental conditions, these habitats are mainly sampled from ships with sampling gear, and the resulting data cannot be used to resolve small-scale species-habitat associations. A detailed substratum classification and community assessment, often only possible using SCUBA diving, is therefore important to reach a better understanding of the functional relationships between species and their environment in northern temperate waters, knowledge that is very important with respect to the increasing environmental pressure caused by global climate change.

(Table 1) Biomass and total standing crop of major wet sedge tundra species at five sites at Alexandra Fiord in 1981 and 2005

The global climate is changing rapidly and Arctic regions are showing responses to recent warming. Responses of tundra ecosystems to climate change have been examined primarily through short-term experimental manipulations, with few studies of long-term ambient change. We investigated changes in above- and belowground biomass of wet sedge tundra to the warming climate of the Canadian High Arctic over the past 25 years. Aboveground standing crop was harvested from five sedge meadow sites and belowground biomass was sampled from one of the sites in the early 1980s and in 2005 using the same methods. Aboveground biomass was on average 158% greater in 2005 than in the early 1980s. The belowground biomass was also much greater in 2005: root biomass increased by 67% and rhizome biomass by 139% since the early 1980s. Dominant species from each functional group (graminoids, shrubs and forbs) showed significant increases in aboveground biomass. Responsive species included the dominant sedge species Carex aquatilis stans, C. membranacea, and Eriophorum angustifolium, as well as the dwarf shrub Salix arctica and the forb Polygonum viviparum. However, diversity measures were not different between the sample years. The greater biomass correlated strongly with increased annual and summer temperatures over the same time period, and was significantly greater than the annual variation in biomass measured in 1980-1983. Increased decomposition and mineralization rates, stimulated by warmer soils, were likely a major cause of the elevated productivity, as no differences in the mass of litter were found between sample periods. Our results are corroborated by published short-term experimental studies, conducted in other wet sedge tundra communities which link warming and fertilization with elevated decomposition, mineralization and tundra productivity. We believe that this is the first study to show responses in High Arctic wet sedge tundra to recent climate change.

Table 3: Comparison of freshwater species in Ob and Yenisei during the 1997 cruise

Phytoplankton biomass distribution (chlorophyll a, chl. a) and species composition (cell numbers) were investigated during three expeditions to the Kara Sea with "Akademik Boris Petrov" (BP) in 1997, 1999, and 2000. The distribution of biomass in the estuaries of Ob and Yenisei showed a similar range in 1997 (0.2 to 3.2 µg/l) and 2000 (0.4 to 3.5 ug/l); higher chl. a concentrations during these two years were found in Yenisei than in Ob. In 1999, phytoplankton biomass in the Ob and Ob Estuary was much higher than in 1997 and 2000, with maximum values above 10.0 ug chl. a/l. In 1999, biomass in Yenisei was lower (1.5 to ~5 ug/l) than in Ob but slightly higher than in 1997 and in 2000. During the expedition in 2000, the research area extended farther to the north, here, lowest phytoplankton biomass during all three years was found. Typical summer values for integrated chl.a biomass (surface to bottom) ranged between 6 and 20 mg m**-2. Strong differences existed in species composition in both rivers, the estuaries, and the open Kara Sea. In general, three or four different populations could be distinguished in surface waters: (1) freshwater diatoms together with bluegreen algae in both rivers, (2) centric and small pennate diatoms mainly brackish species in the estuaries, (3) north of 74°N, brackish/marine species dominated, i.e. in 1999 Thalassiosira cfpunctigera and Chaetoceros spp prevailed in the phytoplankton bloom in Ob. (4) At the northernmost, almost marine stations, a region with a more heterogeneous composition of unicellular plankton was encountered. We assume, we found different seasonal signals of phytoplankton development during 1997/2000 and 1999, respectively. However, the yearly fluctuation of freshwater runoff of both rivers seems to have the strongest influence on the timing and duration of phytoplankton blooms, species compositions and biomass standing stocks during summer.

Tab. 1: Chupa Inlet, Kandalaksha Bay of the White Sea. Population density of the dominant boreal copepod species and daily pellet carbon flux

Data on the zooplankton community structure, gut evacuation rate and carbon content of zooplankton faecal pellets were used for assessing the contribution of zooplankton to vertical carbon fluxes in the White and Kara Seas. The results revealed strong regional and seasonal variations of pellet carbon input related to differences in structure and dynamics of the zooplankton communities in the regions studied. In the deep regions of the White Sea, maximum daily pellet carbon flux from the 0-50 m layer was observed in the spring. It reached 98 mg Corg m-2 day-1 and coincided with a strong predominance of the large arctic herbivorous copepod Calanus glacialis in the surface layers. In summer and fall, it decreased by 1 to 2 orders of magnitude due to migration of this copepod to its overwintering depths. In contrast, in the shallow coastal regions, the pellet production was low in spring, gradually increased during summer and reached its maximum of 138 mg Corg m-2 day-1 by late summer to beginning of autumn. Such a seasonal pattern was in accordance with the seasonal variation of abundance of major pellet producers, the small boreal copepods Acartia bifilosa, Centropages hamatus, and Temora longicornis. In the estuarine zone of the Kara Sea, the pellet flux was mostly formed by pellets of brackish-water omnivorous copepods. It varied from 35 mg Corg m-2 day-1 in 1997 to 96 mg Corg m-2 day-1 in 1999. In the central Kara Sea with its typical marine community, the daily flux reached 125 mg Corg m-2 day-1 in summer. The results of our calculations indicate that both in the White and Kara seas zooplankton pellet carbon contributes up to 30 % to the total carbon flux during particular seasons.

Climate sensitivity across marine domains of life: limits to evolutionary adaptation shape species interactions, supplementary material

Organisms in all domains, Archaea, Bacteria, and Eukarya will respond to climate change with differential vulnerabilities resulting in shifts in species distribution, coexistence, and interactions. The identification of unifying principles of organism functioning across all domains would facilitate a cause and effect understanding of such changes and their implications for ecosystem shifts. For example, the functional specialization of all organisms in limited temperature ranges leads us to ask for unifying functional reasons. Organisms also specialize in either anoxic or various oxygen ranges, with animals and plants depending on high oxygen levels. Here, we identify thermal ranges, heat limits of growth, and critically low (hypoxic) oxygen concentrations as proxies of tolerance in a meta-analysis of data available for marine organisms, with special reference to domain-specific limits. For an explanation of the patterns and differences observed, we define and quantify a proxy for organismic complexity across species from all domains. Rising complexity causes heat (and hypoxia) tolerances to decrease from Archaea to Bacteria to uni- and then multicellular Eukarya. Within and across domains, taxon-specific tolerance limits likely reflect ultimate evolutionary limits of its species to acclimatization and adaptation. We hypothesize that rising taxon-specific complexities in structure and function constrain organisms to narrower environmental ranges. Low complexity as in Archaea and some Bacteria provide life options in extreme environments. In the warmest oceans, temperature maxima reach and will surpass the permanent limits to the existence of multicellular animals, plants and unicellular phytoplankter. Smaller, less complex unicellular Eukarya, Bacteria, and Archaea will thus benefit and predominate even more in a future, warmer, and hypoxic ocean.

(Table 1) Prey species identified from the scats of Antarctic fur seals (Arctocephalus gazella) on Bouvet Island

Intraspecific differences in the diets of many species of pinnipeds are to be expected in view of the great differences in morphology, life history and foraging behaviour between the sexes of many species. We examined the diet of the Antarctic fur seal population at Bouvetøya, Southern Ocean, to assess intersexual differences. This was made possible by the analysis of prey remains extracted from scats and regurgitations collected in areas used primarily by one or the other sex. The results indicate that both males and females feed primarily on Antarctic krill Euphausia superba with several species of fish and squid being taken, likely opportunistically given their prevalence. Significant differences were identified in the frequency of occurrence of otoliths in scats and the percentage numerical abundance of the major fish prey species in the diet. Adult males ate a smaller quantity of fish overall, but ate significantly more of the larger fish species. The greater diving capabilities of males and the fact that they are not limited in the extent of their foraging area by having to return regularly to feed dependant offspring may play a role in the differences found between the diets of males and females. Additionally, females might be more selective, favouring myctophids because they are richer in energy than krill. The absence of major differences in the diet between the sexes at this location is likely due to the high overall abundance of prey at Bouvetøya.

Species-specific consequences of ocean acidification for the calcareous tropical green algae Halimeda, 2011

Ocean acidification (OA), resulting from increasing dissolved carbon dioxide (CO2) in surface waters, is likely to affect many marine organisms, particularly those that calcify. Recent OA studies have demonstrated negative and/or differential effects of reduced pH on growth, development, calcification and physiology, but most of these have focused on taxa other than calcareous benthic macroalgae. Here we investigate the potential effects of OA on one of the most common coral reef macroalgal genera,Halimeda. Species of Halimeda produce a large proportion of the sand in the tropics and are a major contributor to framework development on reefs because of their rapid calcium carbonate production and high turnover rates. On Palmyra Atoll in the central Pacific, we conducted a manipulative bubbling experiment to investigate the potential effects of OA on growth, calcification and photophysiology of 2 species of Halimeda. Our results suggest that Halimeda is highly susceptible to reduced pH and aragonite saturation state but the magnitude of these effects is species specific. H. opuntiasuffered net dissolution and 15% reduction in photosynthetic capacity, while H. taenicola did not calcify but did not alter photophysiology in experimental treatments. The disparate responses of these species to elevated CO2 partial -pressure (pCO2) may be due to anatomical and physiological differences and could represent a shift in their relative dominance in the face of OA. The ability for a species to exert biological control over calcification and the species specific role of the carbonate skeleton may have important implications for the potential effects of OA on ecological function in the future.

Radiocarbon-age differences among coexisting planktic foraminifera shells

For slowly accumulating sediments, a major contrast exists in the radiocarbon-age differences among coexisting shells of planktic foraminifera between those experiencing little dissolution and those experiencing significant dissolution. In the former, the ages generally agree to within a couple of hundred years. In the latter, age differences as large as 1000 years are common. The most likely explanation appears to be the Barker Effect, which involves the preferential fragmentation of dissolution-prone G. sacculifer and G. ruber. The whole shells of these species picked for radiocarbon dating have shorter residence times in the bioturbation zone than those for dissolution-resistant species (including benthics). As low accumulation rate sediment cores often fail to yield reliable radiocarbon-based ocean ventilation ages, where possible, such studies should be conducted on high accumulation rate cores.

Persistent organic pollutants, d13C and d15N in zooplankton, fish and bird species in Kongsfjorden according to season

Seasonality in biomagnification of persistent organic pollutants (POPs; polychlorinated biphenyls, chlorinated pesticides, and brominated flame retardants) in Arctic marine pelagic food webs was investigated in Kongsfjorden, Svalbard, Norway. Trophic magnification factors (TMFs; average factor change in concentration between two trophic levels) were used to measure food web biomagnification in biota in May, July, and October 2007. Pelagic zooplankton (seven species), fish (five species), and seabirds (two species) were included in the study. For most POP compounds, highest TMFs were found in July and lowest were in May. Seasonally changing TMFs were a result of seasonally changing POP concentrations and the d15N-derived trophic positions of the species included in the food web. These seasonal differences in TMFs were independent of inclusion/exclusion of organisms based on physiology (i.e., warm- versus cold-blooded organisms) in the food web. The higher TMFs in July, when the food web consisted of a higher degree of boreal species, suggest that future warming of the Arctic and increased invasion by boreal species can result in increased food web magnification. Knowledge of the seasonal variation in POP biomagnification is a prerequisite for understanding changes in POP biomagnification caused by climate change.

Effect of increasing sea water pCO2 on the northern Atlantic krill species Nyctiphanes couchii

Surprisingly little is known about potential effects of ocean acidification on krill of the Northern Hemisphere as ecologically very important food web component. Sub-adult individuals of the northern Atlantic krill species Nyctiphanes couchii (caught at Austevoll near Bergen, Norway, in January 2013) were exposed in the laboratory to four different levels of pCO2 (430, 800, 1,100, and 1,700 µatm) for 5 weeks in order to assess potential changes in a set of biological response variables. Survival decreased and the frequency of moulting-related deaths increased with increasing pCO2. Survival was considerably reduced at relatively high pCO2 of 1,700 µatm and tended to be negatively affected at 1,100 µatm pCO2. However, the experimental results show no significant effects of pCO2 on inter-moult period and growth at pCO2 levels below 1,100 µatm. No differences in length measurements of the carapace and uropod were observed across pCO2 levels, indicating no effect of changing carbonate chemistry on the morphology of those calciferous parts of the exoskeleton. The results suggest that sub-adult N. couchii may not suffer dramatically from predicted near-future changes in pCO2. However, potential detrimental effects on the moulting process and associated higher mortality at 1,100 µatm pCO2 cannot be excluded. Further experiments are needed in order to investigate whether early life stages of N. couchii show a different sensitivity to elevated sea water pCO2 and whether those results are transferable to other krill species of the Northern Hemisphere.

Range charts of stratigraphically important radiolarian species in sediments of ODP Leg 165 sites

The Paleogene sequences from three sites in the Caribbean were examined for radiolarians. In general, samples are highly lithified, requiring lengthy and repetitive cleaning procedures, and the assemblages are usually fragmented and/or partially dissolved. Both abundances and preservation of the assemblages vary considerably from site to site and within a single site; even within a single sample more than one degree of preservation was observed. It was possible, however, to construct at least partial stratigraphies for each of the three sites. Because the abundance of radiolarians is high even in extremely poorly preserved assemblages, we conclude that the differences in biogenic silica preservation are the result of postdepositional processes and not productivity. In both Sites 999 and 1001, near the Paleocene/Eocene boundary (Bekoma bidartensis Zone [RP7]), there is a short interval in which the abundance and preservation state of the radiolarians improves relative to overlying and underlying assemblages. In each case the intervals corresponds to the level, identified by calcareous microfossils, as representing changes in paleoceanographic conditions associated with the late Paleocene thermal maximum.

(Table 3) Leaf area and leaf nitrogen for deciduous, evergreen, forb and graminoid species at Barrow, Svalbard and Zackenberg

Arctic vegetation is characterized by high spatial variability in plant functional type (PFT) composition and gross primary productivity (P). Despite this variability, the two main drivers of P in sub-Arctic tundra are leaf area index (LT) and total foliar nitrogen (NT). LT and NT have been shown to be tightly coupled across PFTs in sub-Arctic tundra vegetation, which simplifies up-scaling by allowing quantification of the main drivers of P from remotely sensed LT. Our objective was to test the LT-NT relationship across multiple Arctic latitudes and to assess LT as a predictor of P for the pan-Arctic. Including PFT-specific parameters in models of LT-NT coupling provided only incremental improvements in model fit, but significant improvements were gained from including site-specific parameters. The degree of curvature in the LT-NT relationship, controlled by a fitted canopy nitrogen extinction co-efficient, was negatively related to average levels of diffuse radiation at a site. This is consistent with theoretical predictions of more uniform vertical canopy N distributions under diffuse light conditions. Higher latitude sites had higher average leaf N content by mass (NM), and we show for the first time that LT-NT coupling is achieved across latitudes via canopy-scale trade-offs between NM and leaf mass per unit leaf area (LM). Site-specific parameters provided small but significant improvements in models of P based on LT and moss cover. Our results suggest that differences in LT-NT coupling between sites could be used to improve pan-Arctic models of P and we provide unique evidence that prevailing radiation conditions can significantly affect N allocation over regional scales.

Species abundance and water and sediment characteristics for three ANDEEP stations on Maud Rise and the Weddell Sea

The benthic fauna was investigated during the expedition ANT-XXIV/2 (2007/08) in relation to oceanographic features, biogeochemical properties and sediment characteristics, as well as the benthic, pelagic and air-breathing fauna. The results document that Maud Rise (MR) differs distinctly from surrounding deep-sea basins investigated during previous Southern Ocean expeditions (ANDEEP 2002, 2005). Considering all taxa, the overall similarity between MR and adjacent stations was low (~20% Bray-Curtis-Similarity), and analyses of single taxa show obvious differences in species composition, abundances and densities. The composition and diversity of bivalves of MR are characterised by extremely high abundances of three species, especially the small sized Vesicomya spp. Exceptionally high gastropod abundance at MR is due to the single species Onoba subantarctica wilkesiana, a small brooder that may prey upon abundant benthic foraminiferas. The abundance and diversity of isopods also show that one family, Haplomunnidae, occurs with a surprisingly high number of individuals at MR while this family was not found at any of the 40 bathyal and abyssal ANDEEP stations. Similarly, polychaetes, especially the tube-dwelling, suspension-feeder fraction, are represented by species not found at the comparison stations. Sponges comprise almost exclusively small specimens in relatively high numbers, especially a few species of Polymastiidae. Water-column sampling from the surface to the seafloor, including observations of top predators, indicate the existence of a prospering pelagic food web. Local concentrations of top predators and zooplankton are associated with a rich ice-edge bloom located over the northern slope of MR. There the sea ice melts, which is probably accelerated by the advection of warm water at intermediate depth. Over the southern slope, high concentrations of Antarctic krill (Euphausia superba) occur under dense sea ice and attract Antarctic Minke Whales (Balaenoptera bonaerensis) and several seabird species. These findings suggest that biological prosperity over MR is related to both oceanographic and sea-ice processes. Downward transport of the organic matter produced in the pelagic realm may be more constant than elsewhere due to low lateral drift over MR.

Appendix 7: List of species with their distribution ranges restricted to the corresponding phytogeographical region

Understanding species distribution patterns and the corresponding environmental determinants is a crucial step in the development of effective strategies for the conservation and management of plant communities and ecosystems. Therefore, a central prerequisite is the biogeographical and macroecological analysis of factors and processes that determine contemporary, potential, as well as future geographic distribution of species. This thesis has been conducted in the framework of the BIOMAPS-BIOTA project at the Nees Institute of Biodiversity of Plants, which was funded by the German Federal Ministry of Education and Research (BMBF). The study investigated patterns of plants species richness and phytogeographic regions under contemporary environmental conditions and forecasted future climate change in the area of West Africa covering five countries: Benin, Burkina Faso, Côte d'Ivoire, Ghana and Togo. Firstly, geographic patterns of vascular plant species richness have been depicted at a relatively fine spatial resolution based on the potential distribution of 3,393 species. Species richness is closely related to the steep climatic gradient existing in the region with a high concentration of species in the most humid areas in the south and decreases towards the northern drier areas. The investigation of the effectiveness of the existing network of protected areas shows an overall good coverage of species in the study area. However, the proportion of covered species is considerably lower at national extent for some countries, thus calling for more protected areas in order to cover adequately a maximum number of plants species in these countries. Secondly, based on the potential distribution range of vascular plant species, seven phytogeographic regions have been delineated that broadly reflect the vegetation zones as defined by White (1983). However notable differences to the delineation of White (1983) occur at the margins of some regions. Corresponding to a general southward shifted of all regions. And expansion of the Sahel vegetation zone is observed in the north, while the rainforest zone is decreased in the very south.This is alarming since the rainforest shelters a high number of species and a high proportion of range-restricted or endemic species, despite their relatively small extent compared to the other regions. Finally, the evaluation of the potential impact of climate change on plant species richness in the study area, results in a severe loss of future suitable habitat for up to 50% of species per grid cell, particularly in the rainforest region. Moreover, the analysis of the possible shift of phytogeographic regions shows in general a strong deterioration of the West African rainforest. In contrast the drier areas are expanding continuously, although a slight gain in species number can be observed in some particular regions. The overall lesson to retain from the results of this study is that the West African rainforest should be fixed as a high priority area for the conservation of biodiversity of plants, since it is subject to severe contemporary and projected future threats.