Southern Ocean biogeography and taxonomic resolution: what's in the name?

We examined the taxonomic resolution of zooplankton data required to identify ocean basin scale biogeographic zonation in the Southern Ocean. A 2,154 km transect was completed south of Australia. Sea surface temperature (SST) measured at 1 min intervals showed that seven physical zones were sampled. Zooplankton were collected at a spatial resolution of similar to 9.2 km with a continuous plankton recorder, identified to the highest possible taxonomic resolution and enumerated. Zooplankton assemblage similarity between samples was calculated using the Bray-Curtis index for the taxonomic levels of species, genus, family, order and class after first log(10)(x + 1) (LA) and then presence/absence (PA) transformation of abundance data. Although within and between zone sample similarity increased with decreasing taxonomic resolution, for both data transformations, cluster analysis demonstrated that the biogeographic separation of zones remained at all taxonomic levels when using LA data. ANOSIM confirmed this, detecting significant differences in zooplankton assemblage structure between all seven a priori determined physical zones for all taxonomic levels when using the LA data. In the case of the PA data for the complete data set, and both LA and PA data for a crustacean only data set, no significant differences were detected between zooplankton assemblages in the Polar frontal zone (PFZ) and inter-PFZ at any taxonomic level. Loss of information at resolutions below the species level, particularly in the PA data, prevented the separation of some zones. However, the majority of physical zones were biogeographically distinct from species level to class using both LA and PA transformations. Significant relationships between SST and zooplankton community structure, summarised as NMDS scores, at all taxonomic levels, for both LA and PA transformations, and complete and crustacean only data sets, highlighted the biogeographic relevance of low resolution taxonomic data. The retention of biogeographic information in low taxonomic resolution data shows that data sets collected with different taxonomic resolutions may be meaningfully merged for the post hoc generation of Southern Ocean time series.

Monitoring marine plankton ecosystems: Identification of the most relevant indicators of the state of an ecosystem

The number of variables involved in the monitoring of an ecosystem can be high and often one of the first stages in the analysis is to reduce the number of variables. We describe a method developed for geological purposes, using the information theory, that enables selection of the most relevant variables. This technique also allows the examination of the asymmetrical relationships between variables. Applied to a set of physical and biological variables (plankton assemblages in four areas of the North Sea), the method shows that biological variables are more informative than physical variables although the controlling factors are mainly physical (sea surface temperature in winter and spring). Among biological variables, diversity measures and warm-water species assemblages are informative for the state of the North Sea pelagic ecosystems while among physical variables sea surface temperature in late winter and early spring are highly informative. Although often used in bioclimatology, the utilisation of the North Atlantic Oscillation (NAO) index does not seem to provide a lot of information. The method reveals that only the extreme states of this index has an influence on North Sea pelagic ecosystems. The substantial and persistent changes that were detected in the dynamic regime of the North Sea ecosystems and called regime shift are detected by the method and corresponds to the timing of other shifts described in the literature for some European Systems such as the Baltic and the Mediterranean Sea when both physical and biological variables are considered.

Marine plankton phenology and life history in a changing climate: current research and future directions

Increasing availability and extent of biological ocean time series (from both in situ and satellite data) have helped reveal significant phenological variability of marine plankton. The extent to which the range of this variability is modified as a result of climate change is of obvious importance. Here we summarize recent research results on phenology of both phytoplankton and zooplankton. We suggest directions to better quantify and monitor future plankton phenology shifts, including (i) examining the main mode of expected future changes (ecological shifts in timing and spatial distribution to accommodate fixed environmental niches vs. evolutionary adaptation of timing controls to maintain fixed biogeography and seasonality), (ii) broader understanding of phenology at the species and community level (e.g. for zooplankton beyond Calanus and for phytoplankton beyond chlorophyll), (iii) improving and diversifying statistical metrics for indexing timing and trophic synchrony and (iv) improved consideration of spatio-temporal scales and the Lagrangian nature of plankton assemblages to separate time from space changes.

Long-term variability and environmental preferences of calycophoran siphonophores in the Bay of Villefranche (North-Western Mediterranean)

Long-term variability of the main calycophoran siphonophores was investigated between 1974 and 1999 in a coastal station in the north-western Mediterranean. The data were collected at weekly frequency using a macroplankton net (680 μm mesh size) adapted to quantitatively sample delicate gelatinous plankton. A 3-year collection (1967–1969) of siphonophores from offshore waters using the same methodology showed that the patterns of variability observed inshore were representative of siphonophores’ changes at a regional scale. The aims of the study were: (i) to investigate the patterns of variability that characterised the dominant calycophoran species and assemblages; (ii) to identify the environmental optima that were associated with a significant increase in the dominant siphonophore species and (iii) to verify the influence of hydroclimatic variability on long-term changes of siphonophores. Our results showed that during nearly 3 decades the standing stock of calycophoran siphonophores did not show any significant change, with the annual maximum usually recorded in spring as a result of high densities of the dominant species Lensia subtilis, Muggiaea kochi and Muggiaea atlantica. Nevertheless, major changes in community composition occurred within the calycophoran population. Since the middle 1980s, M. kochi, once the most dominant species, started to decrease allowing other species, the congeneric M. atlantica and Chelophyes appendiculata, to increasingly dominate in spring and summer–autumn, respectively. The comparison of environmental and biotic long-term trends suggests that the decrease of M. kochi was triggered by hydrological changes that occurred in the north-western Mediterranean under the forcing of large-scale climate oscillations. Salinity, water stratification and water temperature were the main hydroclimatic factors associated with a significant increase of siphonophores, different species showing different environmental preferences.

Zooplankton associations in a Mediterranean long-term time-series

Coastal zooplankton have been investigated since 1984 at a Long Term Ecological Research station MC (LTER-MC) in the inner Gulf of Naples (Tyrrhenian Sea, Western Mediterranean). The sampling site, located between the littoral and the open sea systems, has very active hydrography that affects plankton communities. The present work was aimed at establishing whether, in such a dynamic and variable environment, species associations and homogeneous periods could be identified as characteristic and stable features of the mesozooplankton over the period 1984–2006. Hierarchical clustering was applied to assess species associations based on a matrix of similarities between species (R-mode), and homogeneous periods based on a matrix of similarities between observations (Q-mode). The Indicator Value index [IndVal, Dufrene and Legendre (1997) Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecol. Monogr., 67, 345–366] was calculated to identify species characterizing each period. Five taxonomic groups with well-defined composition and abundance were identified as robust associations that likely reflect different modes of community functioning. The temporal course of these associations was largely shaped by strong seasonal forcing comprising both physical and biological (e.g. trophic) signals. These associations persisted over the long term, thus indicating some stable characters in the Naples zooplankton time-series, providing evidence of resilience in communities in highly variable coastal conditions.

In hot water: zooplankton and climate change

An overview is provided of the observed and potential future responses of zooplankton communities to global warming. I begin by describing the importance of zooplankton in ocean ecosystems and the attributes that make them sensitive beacons of climate change. Global warming may have even greater repercussions for marine ecosystems than for terrestrial ecosystems, because temperature influences water column stability, nutrient enrichment, and the degree of new production, and thus the abundance, size composition, diversity, and trophic efficiency of zooplankton. Pertinent descriptions of physical changes in the ocean in response to climate change are given as a prelude to a detailed discussion of observed impacts of global warming on zooplankton. These manifest as changes in the distribution of individual species and assemblages, in the timing of important life-cycle events, and in abundance and community structure. The most illustrative case studies, where climate has had an obvious, tangible impact on zooplankton and substantial ecosystem consequences, are presented. Changes in the distribution and phenology of zooplankton are faster and greater than those observed for terrestrial groups. Relevant projected changes in ocean conditions are then presented, followed by an exploration of potential future changes in zooplankton communities from the perspective of different modelling approaches. Researchers have used a range of modelling approaches on individual species and functional groups forced by output from climate models under future greenhouse gas emission scenarios. I conclude by suggesting some potential future directions in climate change research for zooplankton, viz. the use of richer zooplankton functional groups in ecosystem models; greater research effort in tropical systems; investigating climate change in conjunction with other human impacts; and a global zooplankton observing system.

Nematode diversity in different microhabitats in a mangrove region

Mangroves are highly productive environments that play important ecological and socioeconomic roles; however, they have been impacted to different degrees in most countries worldwide. The knowledge of which organisms inhabit this environment and their ecological interactions is the first step towards its conservation. The natural variability of environmental factors in mangroves provides numerous niches available to different species. Meiofauna have patchy patterns of distribution that are related to the availability of resources. Hence, meiofauna are expected to present a high diversity of different taxa occupying the different microhabitats offered by mangroves. This work aims to test the hypothesis that the assemblage structure of Nematoda varies significantly among mangrove microhabitats and to contribute knowledge on the meiofauna diversity in mangrove environments. This work was carried out in a mangrove region at Pernambuco state, Northeastern Brazil. Qualitative samples were collected in nine microhabitats which show different characteristics mainly in terms of presence of vegetation or another organism and sediment grain size. Univariate and multivariate analysis were applied to Nematoda genera abundance data. Our results demonstrate the existence of significant differences among microhabitats regarding nematode assemblage structure corroborating the hypothesis. Different Nematoda assemblages are present in at least seven microhabitats. These assemblages are composed of nematode genera with different trophic and morphological features, demonstrating a strong relationship between morphological diversity and ecological plasticity. Furthermore, this study also demonstrates the importance of the conservation of this ecosystem and its attributes.

Temporal and spatial variation in the Nazaré Canyon (Western Iberian margin): Inter-annual and canyon heterogeneity effects on meiofauna biomass and diversity

The Nazaré Canyon on the Portuguese Margin (NE Atlantic) was sampled during spring-summer for three consecutive years (2005–2007), permitting the first inter-annual study of the meiofaunal communities at the Iberian Margin at two abyssal depths (~3500 m and ~4400 m). Using new and already published data, the meiofauna standing stocks (abundance and biomass) and nematode structural and functional diversity were investigated in relation to the sediment biogeochemistry (e.g. organic carbon, nitrogen, chlorophyll a, phaeopigments) and grain size. A conspicuous increase in sand content from 2005 to 2006 and decrease of phytodetritus at both sites, suggested the occurrence of one or more physical disturbance events. Nematode standing stocks and trophic diversity decreased after these events, seemingly followed by a recovery/recolonisation period in 2007, which was strongly correlated with an increase in the quantity and bioavailability of phytodetrital organic matter supplied. Changes in meiofauna assemblages, however, also differed between stations, likely because of the contrasting hydrodynamic and food supply conditions. Higher meiofauna and nematode abundances, biomass and trophic complexity were found at the shallowest canyon station, where the quantity, quality and bioavailability of food material were higher than at the deeper site. The present results suggest that even though inter-annual variations in the sedimentary environment can regulate the meiofauna in the abyssal Nazaré Canyon, heterogeneity between sampling locations in the canyon were more pronounced.

Interactions between multiple large macrofauna species and nematode communities — Mechanisms for indirect impacts of trawling disturbance

Highlights •We exposed meiofauna to 7 different large macrofauna species at high and low densities. •Macrofauna presence altered nematode community structure and reduced their abundance. •Macrofauna species had similar effects by reducing the few dominant nematode species. •Meio–macrofauna resource competition and spatial segregation are the main drivers. •Trawling effects on macrofauna affect nematode communities indirectly. Diverse assemblages of infauna in sediments provide important physical and biogeochemical services, but are under increasing pressure by anthropogenic activities, such as benthic trawling. It is known that trawling disturbance has a substantial effect on the larger benthic fauna, with reductions in density and diversity, and changes in community structure, benthic biomass, production, and bioturbation and biogeochemical processes. Largely unknown, however, are the mechanisms by which the trawling impacts on the large benthic macro- and megafauna may influence the smaller meiofauna. To investigate this, a mesocosm experiment was conducted whereby benthic nematode communities from a non-trawled area were exposed to three different densities (absent, low, normal) of 7 large (> 10 mm) naturally co-occurring, bioturbating species which are potentially vulnerable to trawling disturbance. The results showed that total abundances of nematodes were lower if these large macrofauna species were present, but no clear nematode abundance effects could be assigned to the macrofauna density differences. Nematode community structure changed in response to macrofauna presence and density, mainly as a result of the reduced abundance of a few dominant nematode species. Any detectable effects seemed similar for nearly all macrofauna species treatments, supporting the idea that there may be a general indirect, macrofauna-mediated trawling impact on nematode communities. Explanations for these results may be, firstly, competition for food resources, resulting in spatial segregation of the meio- and macrobenthic components. Secondly, different densities of large macrofauna organisms may affect the nematode community structure through different intensities of bioturbatory disturbance or resource competition. These results suggest that removal or reduced densities of larger macrofauna species as a result of trawling disturbance may lead to increased nematode abundance and hints at the validity of interference competition between large macrofauna organisms and the smaller meiofauna, and the energy equivalence hypothesis, where a trade-off is observed between groups of organisms that are dependent on a common source of energy.

The effect of pressure on leucine and thymidine incorporation by free-living bacteria and by bacteria attached to sinking oceanic particles

The effect of pressure on upper ocean free-living bacteria and bacteria attached to rapidly sinking particles was investigated through studying their ability to synthesize DNA and protein by measuring their rate of 3H-thymidine and 3H-leucine incorporation. Studies were carried out on samples from the NE Atlantic under the range of pressures (1–430 atm) encountered by sinking aggregates during their journey to the deep-sea bed. Thymidine and leucine incorporation rates per bacterium attached to sinking particles from 200 m were about six and ten times higher, respectively, than the free-living bacterial assemblage. The ratio of leucine incorporation rate per cell to thymidine incorporation rate per cell was significantly different between the larger attached (18.9:1) and smaller free-living (10.4:1) assemblages. The rates of leucine and thymidine incorporation decreased exponentially with increasing pressure for the free-living and linearly for attached bacteria, while there was no significant influence of pressure on cell numbers. At 100 atm leucine and thymidine incorporation rate per free-living bacterium was reduced to 73 and 20%, respectively, relative to that measured at 1 atm. Pressure of 100 atm reduced leucine and thymidine incorporation per attached bacterium to 94 and 70%, and at 200 atm these rates were reduced to 34 and 51%, respectively, relative to those measured at 1 atm. There was no significant uncoupling of thymidine and leucine incorporation for either the free-living or attached bacterial assemblages with increasing pressure, indicating that the processess of DNA and protein synthesis may be equally affected by increasing pressure. It is therefore unlikely that bacteria, originating from surface waters, attached to rapidly sinking particles play a role in particle remineralization below approximately 1000–2000 m. These results may help to explain the occurrence of relatively fresh aggregates on the deep-sea bed that still contain sufficient organic carbon to fuel the rapid growth of benthic micro-organisms; they also indicate that the effect of pressure on microbial processes may be important in oceanic biogeochemical cycles.

The potential of offshore windfarms to act as marine protected areas – A systematic review of current evidence

As offshore windfarm (OWF) construction in the UK is progressing rapidly, monitoring of the economic and ecological effects of these developments is urgently needed. This is to enable both spatial planning and where necessary mitigation in an increasingly crowded marine environment. One approach to mitigation is co-location of OWFs and marine protected areas (MPAs). This systematic review has the objective to inform this co-location proposal and identify areas requiring further research. A limited number of studies addressing marine renewable energy structures and related artificial structures in coastal waters were found. The results of these studies display a change in species assemblages at artificial structures in comparison to naturally occurring habitats. An increase in hard substrata associated species, especially benthic bivalves, crustaceans and reef associated fish and a decrease in algae abundance were the dominant trends. Assemblages associated with complex concrete structures revealed greater similarity to natural hard substrata compared to those around steel structures. To consider marine renewable energy sites, especially large scale OWFs as MPAs, the dissimilar nature of assemblages on the structures themselves to natural communities should be considered. However positive effects were recorded on the abundance of commercially important crustacean species. This suggests potential for incorporation of OWFs as no fishing, or restricted activity zones within a wider MPA to aid fisheries augmentation. The limited available evidence highlights a requirement for significant further research involving long term monitoring at a variety of sites to better inform management options.

Living in warmer, more acidic oceans retards physiological recovery from tidal emersion in the velvet swimming crab, Necora puber

The distribution patterns of many species in the intertidal zone are partly determined by their ability to survive and recover from tidal emersion. During emersion, most crustaceans experience gill collapse, impairing gas exchange. Such collapse generates a state of hypoxemia and a hypercapnia-induced respiratory acidosis, leading to hyperlactaemia and metabolic acidosis. However, how such physiological responses to emersion are modified by prior exposure to elevated CO2 and temperature combinations, indicative of future climate change scenarios, is not known. We therefore investigated key physiological responses of velvet swimming crabs, Necora puber, kept for 14 days at one of four pCO(2)/temperature treatments (400 mu atm/10 degrees C, 1000 mu atm/10 degrees C, 400 mu atm/15 degrees C or 1000 mu atm/15 degrees C) to experimental emersion and recovery. Pre-exposure to elevated pCO(2) and temperature increased pre-emersion bicarbonate ion concentrations [HCO3-], increasing resistance to short periods of emersion (90 min). However, there was still a significant acidosis following 180 min emersion in all treatments. The recovery of extracellular acid-base via the removal of extracellular pCO(2) and lactate after emersion was significantly retarded by exposure to both elevated temperature and pCO(2). If elevated environmental pCO(2) and temperature lead to slower recovery after emersion, then some predominantly subtidal species that also inhabit the low to mid shore, such as N. puber, may have a reduced physiological capacity to retain their presence in the low intertidal zone, ultimately affecting their bathymetric range of distribution, as well as the structure and diversity of intertidal assemblages.

The Importance of Regional, System-Wide and Local Spatial Scales in Structuring Temperate Estuarine Fish Communities

An extensive literature base worldwide demonstrates how spatial differences in estuarine fish assemblages are related to those in the environment at (bio)regional, estuary-wide or local (within-estuary) scales. Few studies, however, have examined all three scales, and those including more than one have often focused at the level of individual environmental variables rather than scales as a whole. This study has identified those spatial scales of environmental differences, across regional, estuary-wide and local levels, that are most important in structuring ichthyofaunal composition throughout south-western Australian estuaries. It is the first to adopt this approach for temperate microtidal waters. To achieve this, we have employed a novel approach to the BIOENV routine in PRIMER v6 and a modified global BEST test in an alpha version of PRIMER v7. A combination of all three scales best matched the pattern of ichthyofaunal differences across the study area (rho = 0.59; P = 0.001), with estuary-wide and regional scales accounting for about twice the variability of local scales. A shade plot analysis showed these broader-scale ichthyofaunal differences were driven by a greater diversity of marine and estuarine species in the permanently-open west coast estuaries and higher numbers of several small estuarine species in the periodically-open south coast estuaries. When interaction effects were explored, strong but contrasting influences of local environmental scales were revealed within each region and estuary type. A quantitative decision tree for predicting the fish fauna at any nearshore estuarine site in south-western Australia has also been produced. The estuarine management implications of the above findings are highlighted.

Long-term iceshelf-covered meiobenthic communities of the Antarctic continental shelf resemble those of the deep sea

Since strong regional warming has led to the disintegration of huge parts of the Larsen A and B ice shelves east of the Antarctic Peninsula in 1995 and 2002, meiofaunal communities covered by ice shelves for thousands of years could be investigated for the first time. Based on a dataset of more than 230,000 individuals, meiobenthic higher taxa diversity and composition of Larsen continental shelf stations were compared to those of deep-sea stations in the Western Weddell Sea to see whether the food-limiting conditions in the deep sea and the food-poor shelf regime at times of iceshelf coverage has resulted in similar meiobenthic communities, on the premises that food availability is the main driver of meiobenthic assemblages. We show here that this is indeed the case; in terms of meiobenthic communities, there is greater similarity between the deep sea and the inner Larsen embayments than there is similarity between the deep sea and the former Larsen B iceshelf edge and the open continental shelf. We also show that resemblance to Antarctic deep-sea meiofaunal communities was indeed significantly higher for communities of the innermost Larsen B area than for those from intermediate parts of Larsen A and B. Similarity between communities from intermediate parts and the deep sea was again higher than between those of the ice-edge and the open shelf. Meiofaunal densities were low at the inner parts of Larsen A and B, and comparable to deep-sea densities, again likely owing to the low food supply at both habitats. We suggest that meiobenthic communities have not yet recovered from the food-limiting conditions present at the time of iceshelf coverage. Meiofaunal diversity on the other hand seemed driven by sediment structure, being higher in coarser sediments.

Effects of elevated CO2 and temperature on an intertidal meiobenthic community

In the near future, the marine environment is likely to be subjected to simultaneous increases in temperature and decreased pH. The potential effects of these changes on intertidal, meiofaunal assemblages were investigated using a mesocosm experiment. Artificial Substrate Units containing meiofauna from the extreme low intertidal zone were exposed for 60 days to eight experimental treatments (four replicates for each treatment) comprising four pH levels: 8.0 (ambient control), 7.7 & 7.3 (predicted changes associated with ocean acidification), and 6.7 (CO2 point-source leakage from geological storage), crossed with two temperatures: 12 °C (ambient control) and 16 °C (predicted). Community structure, measured using major meiofauna taxa was significantly affected by pH and temperature. Copepods and copepodites showed the greatest decline in abundance in response to low pH and elevated temperature. Nematodes increased in abundance in response to low pH and temperature rise, possibly caused by decreased predation and competition for food owing to the declining macrofauna density. Nematode species composition changed significantly between the different treatments, and was affected by both seawater acidification and warming. Estimated nematode species diversity, species evenness, and the maturity index, were substantially lower at 16 °C, whereas trophic diversity was slightly higher at 16 °C except at pH 6.7. This study has demonstrated that the combination of elevated levels of CO2 and ocean warming may have substantial effects on structural and functional characteristics of meiofaunal and nematode communities, and that single stressor experiments are unlikely to encompass the complexity of abiotic and biotic interactions. At the same time, ecological interactions may lead to complex community responses to pH and temperature changes in the interstitial environment