Recovery of a biodiversity action plans species in northwest England: possible role of climate change, artificial habitat and water quality amelioration. Occasional Publication of the Marine Biological Association 16

The honeycomb reef worm Sabellaria alveolata is recognised as being an important component of intertidal communities. It is a priority habitat within the UK Biodiversity Action Plan and as a biogenic reef forming species is covered by Annex 1 of the EC habitats directive. S. alveolata has a lusitanean (southern) distribution, being largely restricted to the south and west coasts of England. A broad-scale survey of S. alveolata distribution along the north-west coasts was undertaken in 2003/2004. These records were then compared with previous distribution records, mainly those collected by Cunningham in 1984. More detailed mapping was carried out at Hilbre Island at the mouth of the River Dee, due to recent reports that S. alveolata had become re-established there after a long absence.

Long-Term Oceanographic and Ecological Research in the Western English Channel

Long-term research in the western English Channel, undertaken by the marine laboratories in Plymouth, is described and details of survey methods, sites, and time series given in this chapter. Major findings are summarized and their limitations outlined. Current research, with recent reestablishment and expansion of many sampling programmes, is presented, and possible future approaches are indicated. These unique long-term data sets provide an environmental baseline for predicting complex ecological responses to local, regional, and global environmental change. Between 1888 and the present, investigations have been carried out into the physical, chemical, and biological components (ranging from plankton and fish to benthic and intertidal assemblages) of the western English Channel ecosystem. The Marine Biological Association of the United Kingdom has performed the main body of these observations. More recent contributions come from the Continuous Plankton Recorder Survey, now the Sir Alister Hardy Foundation for Ocean Science, dating from 1957; the Institute for Marine Environmental Research, from 1974 to 1987; and the Plymouth Marine Laboratory, which was formed by amalgamation of the Institute for Marine Environmental Research and part of the Marine Biological Association, from 1988. Together, these contributions constitute a unique data series; one of the longest and most comprehensive samplings of environmental and marine biological variables in the world. Since the termination of many of these time series in 1987-1988 during a reorganisation of UK marine research, there has been a resurgence of interest in long-term environmental change. Many programmes have been restarted and expanded with support from several agencies. The observations span significant periods of warming (1921-1961; 1985-present) and cooling (1962-1980). During these periods of change, the abundance of key species underwent dramatic shifts. The first period of warming saw changes in zooplankton, pelagic fish, and larval fish, including the collapse of an important herring fishery. During later periods of change, shifts in species abundances have been reflected in other assemblages, such as the intertidal zone and the benthic fauna. Many of these changes appear to be related to climate, manifested as temperature changes, acting directly or indirectly. The hypothesis that climate is a forcing factor is widely supported today and has been reinforced by recent studies that show responses of marine organisms to climatic attributes such as the strength of the North Atlantic Oscillation. The long-term data also yield important insights into the effects of anthropogenic disturbances such as fisheries exploitation and pollution. Comparison of demersal fish hauls over time highlights fisheries effects not only on commercially important species but also on the entire demersal community. The effects of acute ("Torrey Canyon" oil spill) and chronic (tributyltin [TBT] antifoulants) pollution are clearly seen in the intertidal records. Significant advances in diverse scientific disciplines have been generated from research undertaken alongside the long-term data series.

A review of some common Indo-Malayan and western Pacific species of Chthamalus barnacles (Crustacea: Cirripedia)

The type specimens of the common tropical intertidal barnacles Chthamalus malayensis and C. moro, were re-investigated and compared with other specimens of Chthamalus from the Indian Ocean, Indo-Malaya, northern Australia, Vietnam, China and the western Pacific, using ‘arthropodal’ as well as shell characters. Chthamalus malayensis occurs widely in Indo-Malayan and tropical Australian waters. It ranges westwards in the Indian Ocean to East Africa and northwards in the Pacific to Vietnam, China and the Ryukyu Islands. Chthamalus malayensis has the arthropodal characters attributed to it by Pope (1965); conical spines on cirrus 1 and serrate setae with basal guards on cirrus 2. Chthamalus moro is currently fully validated only for the Philippines, Indonesia, Taiwan, the Xisha (Paracel) Islands, the Ryukyu Islands, the Mariana Islands, the Caroline Islands, Fiji and Samoa. It is a small species of the ‘challengeri’ subgroup, lacking conical spines on cirrus 1 and bearing pectinate setae without basal guards on cirrus 2. It may be a ‘relict’ insular species. Chthamalus challengeri also lacks conical spines on cirrus 1 and has pectinate setae without basal guards on cirrus 2. Records of C. challengeri south of Japan are probably erroneous. However, there is an undescribed species of the ‘challengeri’ subgroup in the Indian Ocean, Indo-Malaya, Vietnam and southern China and yet more may occur in the western Pacific. The subgroups ‘malayensis’ and ‘challengeri’ require genetic investigation. Some comments are included on the arthropodal characters and geographical distributions of Chthamalus antennatus, C. dalli and C. stellatus

Structural and functional vulnerability to elevated pCO2 in marine benthic communities

The effect of elevated pCO(2)/low pH on marine invertebrate benthic biodiversity, community structure and selected functional responses which underpin ecosystem services (such as community production and calcification) was tested in a medium-term (30 days) mesocosm experiment in June 2010. Standardised intertidal macrobenthic communities, collected (50.3567A degrees N, 4.1277A degrees W) using artificial substrate units (ASUs), were exposed to one of seven pH treatments (8.05, 7.8. 7.6, 7.4, 7.2, 6.8 and 6.0). Community net calcification/dissolution rates, as well as changes in biomass, community structure and diversity, were measured at the end of the experimental period. Communities showed significant changes in structure and reduced diversity in response to reduced pH: shifting from a community dominated by calcareous organisms to one dominated by non-calcareous organisms around either pH 7.2 (number of individuals and species) or pH 7.8 (biomass). These results were supported by a reduced total weight of CaCO3 structures in all major taxa at lowered pH and a switch from net calcification to net dissolution around pH 7.4 (a"broken vertical bar(calc) = 0.78, a"broken vertical bar(ara) = 0.5). Overall community soft tissue biomass did not change with pH and high mortality was observed only at pH 6.0, although molluscs and arthropods showed significant decreases in soft tissue. This study supports and refines previous findings on how elevated pCO(2) can induce changes in marine biodiversity, underlined by differential vulnerability of different phyla. In addition, it shows significant elevated pCO(2)-/low pH-dependent changes in fundamental community functional responses underpinning changes in ecosystem services.

The Nematode-Copepod Ratio And Its Use In Pollution Ecology

The nematode/copepod ratio is critically examined with a view to adding some precision to its proposed use in pollution ecology. At two unpolluted intertidal sites, differing markedly in sediment grade, the metabolic requirements of copepods are shown to be equivalent to the requirements of that fraction of the nematode population which feeds in the same way. The partitioning of this total energy requirement among individuals depends on the distribution of individual metabolic body sizes and the relative rates of metabolism. The distribution of body sizes is constrained by the sediment granulometry, which affects nematodes and copepods differently. These considerations enable precise predictions of the nematode/copepod ratios expected in unpolluted situations, against which observed ratios can be compared.

Effect Of Temperature On The Respiration Rate Of Meiofauna

The effect of temperature on respiration rate has been established, using Cartesian divers, for the meiofaunal sabellid polychaeteManayunkia aestuarina, the free-living nematodeSphaerolaimus hirsutus and the harpacticoid copepodTachidius discipes from a mudflat in the Lynher estuary, Cornwall, U.K. Over the temperature range normally experienced in the field, i.e. 5–20° C the size-compensated respiration rate (R c) was related to the temperature (T) in °C by the equation Log10 R c=-0.635+0.0339T forManayunkia, Log10 R c=0.180+0.0069T forSphaerolaimus and Log10 R c=-0.428+0.0337T forTachidius, being equivalent toQ 10 values of 2.19, 1.17 and 2.17 respectively. In order to derive the temperature response forManayunkia a relationship was first established between respiration rate and body size: Log10 R=0.05+0.75 Log10 V whereR=respiration in nl·O2·ind-1·h-1 andV=body volume in nl. TheQ 10 values are compared with values for other species derived from the literature. From these limited data a dichotomy emerges: species with aQ 10≏2 which apparently feed on diatoms and bacteria, the abundance of which are subject to large short term variability, and species withQ 10≏1 apparently dependent on more stable food sources.

Predicted levels of future ocean acidification and temperature rise could alter community structure and biodiversity in marine benthic communities

A mesocosm experiment was conducted to quantify the effects of reduced pH and elevated temperature on an intact marine invertebrate community. Standardised faunal communities, collected from the extreme low intertidal zone using artificial substrate units, were exposed to one of eight nominal treatments (four pH levels: 8.0, 7.7, 7.3 and 6.7, crossed with two temperature levels: 12 and 16°C). After 60 days exposure communities showed significant changes in structure and lower diversity in response to reduced pH. The response to temperature was more complex. At higher pH levels (8.0 and 7.7) elevated temperature treatments contained higher species abundances and diversity than the lower temperature treatments. In contrast, at lower pH levels (7.3 and 6.7), elevated temperature treatments had lower species abundances and diversity than lower temperature treatments. The species losses responsible for these changes in community structure and diversity were not randomly distributed across the different phyla examined. Molluscs showed the greatest reduction in abundance and diversity in response to low pH and elevated temperature, whilst annelid abundance and diversity was mostly unaffected by low pH and was higher at the elevated temperature. The arthropod response was between these two extremes with moderately reduced abundance and diversity at low pH and elevated temperature. Nematode abundance increased in response to low pH and elevated temperature, probably due to the reduction of ecological constraints, such as predation and competition, caused by a decrease in macrofaunal abundance. This community-based mesocosm study supports previous suggestions, based on observations of direct physiological impacts, that ocean acidification induced changes in marine biodiversity will be driven by differential vulnerability within and between different taxonomical groups. This study also illustrates the importance of considering indirect effects that occur within multispecies assemblages when attempting to predict the consequences of ocean acidification and global warming on marine communities.

Data rescue and re-use: Recycling old information to address new policy concerns

Information on past trends is essential to inform future predictions and underpin attribution needed to drive policy responses. It has long been recognised that sustained observations are essential for disentangling climate-driven change from other regional and local-scale anthropogenic impacts and environmental fluctuations or cycles in natural systems. This paper highlights how data rescue and re-use have contributed to the debate on climate change responses of marine biodiversity and ecosystems. It also illustrates via two case studies the re-use of old data to address new policy concerns. The case studies focus on (1) plankton, fish and benthos from the Western English Channel and (2) broad-scale and long-term studies of intertidal species around the British Isles. Case study 1 using the Marine Biological Association of the UK's English Channel data has shown the influence of climatic fluctuations on phenology (migration and breeding patterns) and has also helped to disentangle responses to fishing pressure from those driven by climate, and provided insights into ecosystem-level change in the English Channel. Case study 2 has shown recent range extensions, increases of abundance and changes in phenology (breeding patterns) of southern, warm-water intertidal species in relation to recent rapid climate change and fluctuations in northern and southern barnacle species, enabling modelling and prediction of future states. The case is made for continuing targeted sustained observations and their importance for marine management and policy development.

Diverse profiles of N‐acyl‐homoserine lactone molecules found in cnidarians

Many marine habitats, such as the surface and tissues of marine invertebrates, including corals, harbour diverse populations of microorganisms, which are thought to play a role in the health of their hosts and influence mutualistic and competitive interactions. Investigating the presence and stability of quorum sensing (QS) in these ecosystems may shed light on the roles and control of these bacterial communities. Samples of 13 cnidarian species were screened for the presence and diversity of N-acyl-homoserine lactones (AHLs; a prevalent type of QS molecule) using thin-layer chromatography and an Agrobacterium tumefaciens NTL4 biosensor. Ten of 13 were found to harbour species-specific, conserved AHL profiles. AHLs were confirmed in Anemonia viridis using liquid chromatography tandem mass spectrometry. To assess temporal role and stability, AHLs were investigated in A. viridis from intertidal pools over 16 h. Patterns of AHLs showed conserved profiles except for two mid-chain length AHLs, which increased significantly over the day, peaking at 20:00, but had no correlation with pool chemistry. Denaturing gel electrophoresis of RT-PCR-amplified bacterial 16S rRNA showed the presence of an active bacterial community that changed in composition alongside AHL profiles and contained a number of bands that affiliate with known AHL-producing bacteria. Investigations into the quorum sensing-controlled, species-specific roles of these bacterial communities and how these regulatory circuits are influenced by the coral host and members of the bacterial community are imperative to expand our knowledge of these interactions with respect to the maintenance of coral health.

Macroalgal blooms alter community structure and primary productivity in marine ecosystems

Eutrophication, coupled with loss of herbivory due to habitat degradation and overharvesting, has increased the frequency and severity of macroalgal blooms worldwide. Macroalgal blooms interfere with human activities in coastal areas, and sometimes necessitate costly algal removal programs. They also have many detrimental effects on marine and estuarine ecosystems, including induction of hypoxia, release of toxic hydrogen sulfide into the sediments and atmosphere, and the loss of ecologically and economically important species. However, macroalgal blooms can also increase habitat complexity, provide organisms with food and shelter, and reduce other problems associated with eutrophication. These contrasting effects make their overall ecological impacts unclear. We conducted a systematic review and meta-analysis to estimate the overall effects of macroalgal blooms on several key measures of ecosystem structure and functioning in marine ecosystems. We also evaluated some of the ecological and methodological factors that might explain the highly variable effects observed in different studies. Averaged across all studies, macroalgal blooms had negative effects on the abundance and species richness of marine organisms, but blooms by different algal taxa had different consequences, ranging from strong negative to strong positive effects. Blooms' effects on species richness also depended on the habitat where they occurred, with the strongest negative effects seen in sandy or muddy subtidal habitats and in the rocky intertidal. Invertebrate communities also appeared to be particularly sensitive to blooms, suffering reductions in their abundance, species richness, and diversity. The total net primary productivity, gross primary productivity, and respiration of benthic ecosystems were higher during macroalgal blooms, but blooms had negative effects on the productivity and respiration of other organisms. These results suggest that, in addition to their direct social and economic costs, macroalgal blooms have ecological effects that may alter their capacity to deliver important ecosystem services.

Macrofauna along the Sudanese Red Sea coast: potential effect of mangrove clearance on community and trophic structure

Mangroves along the Sudanese Red Sea coast are under constant anthropogenic pressure. To better understand the influence of mangrove clearance on the intertidal benthic community, we investigated the composition, biodiversity and standing stock of the macrofauna communities at high-, mid- and low-water levels in three contrasting habitats: a bare sand flat, a cleared mangrove and an intact mangrove. In addition, a community-wide metric approach based on taxon-specific carbon and nitrogen isotope values was used to compare the trophic structure between the three habitats. The habitats differed significantly in terms of macrofaunal standing stock, community composition and trophic structure. The high- and mid-water levels of the intact mangroves showed a distinct macrofaunal community characterized by elevated densities and biomass, largely governed by higher decapod and gastropod abundances. Diversity was similar for cleared and intact mangroves, but much lower for the bare sand flat. Community-wide metrics indicated highest trophic diversity and community niche breadth in the intact mangroves. Differences between the cleared and intact mangroves can be partly attributed to differences in sediment characteristics resulting from mangrove clearance. These results suggest a significant impact of mangrove clearance on the macrofaunal community and trophic structure. This study calls for further investigations and management actions to protect and restore these habitats, and ensure the survival of this ecologically valuable coastal ecosystem.

Coastal Zone Ecosystem Services: From science to values and decision making; a case study

This research is concerned with the following environmental research questions: socio-ecological system complexity, especially when valuing ecosystem services; ecosystems stock and services flow sustainability and valuation; the incorporation of scale issues when valuing ecosystem services; and the integration of knowledge from diverse disciplines for governance and decision making. In this case study, we focused on ecosystem services that can be jointly supplied but independently valued in economic terms: healthy climate (via carbon sequestration and storage), food (via fisheries production in nursery grounds), and nature recreation (nature watching and enjoyment). We also explored the issue of ecosystem stock and services flow, and we provide recommendations on how to value stock and flows of ecosystem services via accounting and economic values respectively. We considered broadly comparable estuarine systems located on the English North Sea coast: the Blackwater estuary and the Humber estuary. In the past, these two estuaries have undergone major land-claim. Managed realignment is a policy through which previously claimed intertidal habitats are recreated allowing the enhancement of the ecosystem services provided by saltmarshes. In this context, we investigated ecosystem service values, through biophysical estimates and welfare value estimates. Using an optimistic (extended conservation of coastal ecosystems) and a pessimistic (loss of coastal ecosystems because of, for example, European policy reversal) scenario, we find that context dependency, and hence value transfer possibilities, vary among ecosystem services and benefits. As a result, careful consideration in the use and application of value transfer, both in biophysical estimates and welfare value estimates, is advocated to supply reliable information for policy making.

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 impact of ocean acidification and warming on the skeletal mechanical properties of the sea urchin Paracentrotus lividus from laboratory and field observations

Increased atmospheric CO2 concentration is leading to changes in the carbonate chemistry and the temperature of the ocean. The impact of these processes on marine organisms will depend on their ability to cope with those changes, particularly the maintenance of calcium carbonate structures. Both a laboratory experiment (long-term exposure to decreased pH and increased temperature) and collections of individuals from natural environments characterized by low pH levels (individuals from intertidal pools and around a CO2 seep) were here coupled to comprehensively study the impact of near-future conditions of pH and temperature on the mechanical properties of the skeleton of the euechinoid sea urchin Paracentrotus lividus. To assess skeletal mechanical properties, we characterized the fracture force, Young's modulus, second moment of area, material nanohardness, and specific Young's modulus of sea urchin test plates. None of these parameters were significantly affected by low pH and/or increased temperature in the laboratory experiment and by low pH only in the individuals chronically exposed to lowered pH from the CO2 seeps. In tidal pools, the fracture force was higher and the Young's modulus lower in ambital plates of individuals from the rock pool characterized by the largest pH variations but also a dominance of calcifying algae, which might explain some of the variation. Thus, decreases of pH to levels expected for 2100 did not directly alter the mechanical properties of the test of P. lividus. Since the maintenance of test integrity is a question of survival for sea urchins and since weakened tests would increase the sea urchins' risk of predation, our findings indicate that the decreasing seawater pH and increasing seawater temperature expected for the end of the century should not represent an immediate threat to sea urchins vulnerability.

Intersex in the clam Scrobicularia plana (Da Costa): Widespread occurrence in English Channel estuaries and surrounding areas

Estuarine clams Scrobicularia plana were sampled from 108 intertidal locations around the English Channel and adjacent areas. Although S. plana is believed to be a strict gonochorist, 58% of the populations sampled included intersexed individuals (described as male clams exhibiting ovotestis). Over the entire region, on average, 8.6% of male clams exhibited intersex, although proportions of affected males ranged from 0% to 53% depending on location. The severity of intersex was assessed using a simple classification scale, with the majority of individuals showing low levels of impact. Sex ratios were significantly skewed at some sites. There were no significant relationships between the incidence and severity of intersex; or of associations with size or parasitism of individual clams. Intersex in S. plana is a useful tool to assess endocrine disruptive effects in estuaries, although mechanisms of impact and causative agents remain uncertain.