Ecological Investigations with the Continuous Plankton Recorder: The Zooplankton (other than Copepoda and Young Fish) in the Southern North Sea, 1932-37.

I. The report describes the main monthly changes in the distribution and abundance of the zooplankton, other than Copepoda and young fish (dealt with in separate reports), over the southern part of the North Sea from 1932 to 1937. The work is part of the survey carried out by Continuous Plankton Recorders towed at a depth of 10 metres on regular steamship lines between England and the Continent. 2. The limitations to the sampling method are discussed, and it is shown to be unsuitable for recording Mysidacea and Euphausiacea on account of their marked diurnal variation due presumably to vertical migration; they are omitted from the report. 3. The changing distribution of Sagitta, Limacina, Clione, Lamellibranch larvae, Cladocera, Caprellid Amphipoda, Decapod larvae, Echinoderm larvae and Oikopleura are shown in a series of monthly charts while their seasonal fluctuations are compared in time-chart histograms. 4. The Alima larvae of Squilla are recorded on a few occasions in the regions where the Channel opens into the North Sea. 5. The distributional characteristics of the different forms, i.e. their tendencies to even or " patchy " production, are compared.

Contents, Preface and Explanation

THE Hull Bulletins of Marine Ecology are reports on investigations carried out by the Department of Oceanography in University College Hull. The researches were carried out in close co-operation with the Ministry of Agriculture and Fisheries and the Fisheries Division, Scottish Home Department (previously Fishery Board for Scotland). The papers of the present volume are just a beginning, a first attempt at showing how the changing plankton can be studied over wide areas, just as the changing weather conditions can be studied, with the passage of time.

Continuous Plankton Records: General Introduction to the 1938-39 Survey

The Bulletins in this volume, except the last, deal entirely with the results of this expanded survey of 1938 and 1939. They show for the first time, month by month, the main changes in the plankton over practically the whole of the North Sea for a year and eight months. They form an important basis for comparison with the results of the post-war survey, revived on an even more extensive scale and to be described in later volumes.

Picoeukaryote distribution in relation to nitrate uptake in the oceanic nitracline

We investigated the relationship between picoeukaryote phytoplankton (< 2 mu m) and the deep layer of new production (NO3- uptake) in the nitracline of the eastern subtropical North Atlantic Ocean. Indices of NO3- uptake kinetics obtained within the lower 15 % of the euphotic zone demonstrate that subsurface NO3- uptake maxima are coincident with localised peaks in maximum uptake rates (V-max) and, crucially, with maximum picoeukaryote abundance. The mean rate of NO3- utilization at the nitracline is typically 10-fold higher than in surface waters despite much lower in situ irradiance. These observations confirm a high affinity for NO3-, most likely by the resident picoeukaryote community, and we conservatively estimate mean cellular uptake rates of between 0.27 and 1.96 fmol NO3- cell(-1) h(-1). Greater scrutiny of the taxonomic composition of the picoeukaryote group is required to further understand this deep layer of new production and its importance for nitrogen cycling and export production, given longstanding assumptions that picoplankton do not contribute directly to export fluxes.

Mesoscale fronts as foraging habitats: composite front mapping reveals oceanographic drivers of habitat use for a pelagic seabird

The oceanographic drivers of marine vertebrate habitat use are poorly understood yet fundamental to our knowledge of marine ecosystem functioning. Here, we use composite front mapping and high-resolution GPS tracking to determine the significance of mesoscale oceanographic fronts as physical drivers of foraging habitat selection in northern gannets Morus bassanus. We tracked 66 breeding gannets from a Celtic Sea colony over 2 years and used residence time to identify area-restricted search (ARS) behaviour. Composite front maps identified thermal and chlorophyll-a mesoscale fronts at two different temporal scales—(i) contemporaneous fronts and (ii) seasonally persistent frontal zones. Using generalized additive models (GAMs), with generalized estimating equations (GEE-GAMs) to account for serial autocorrelation in tracking data, we found that gannets do not adjust their behaviour in response to contemporaneous fronts. However, ARS was more likely to occur within spatially predictable, seasonally persistent frontal zones (GAMs). Our results provide proof of concept that composite front mapping is a useful tool for studying the influence of oceanographic features on animal movements. Moreover, we highlight that frontal persistence is a crucial element of the formation of pelagic foraging hotspots for mobile marine vertebrates.

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.

Environmental hypoxia but not minor shell damage affects scope for growth and body condition in the blue mussel Mytilus edulis (L.)

The effects of short-term (7 d) exposure to environmental hypoxia (2.11 mg O-2 L-1; control: 6.96 mg O-2 L-1) and varying degrees of shell damage (1 or 2, 1 mm diameter holes; control: no holes) on respiration rate, clearance rate, ammonia excretion rate, scope for growth (SFG) and body condition index were investigated in adult blue mussels (Mytilus edulis). There was a significant hypoxia-related reduction in SFG (>6.70 to 0.92J g(-1) h(-1)) primarily due to a reduction in energy acquisition as a result of reduced clearance rates during hypoxia. Shell damage had no significant affect on any of the physiological processes measured or the SFG calculated. Body condition was unaffected by hypoxia or shell damage. In conclusion, minor physical damage to mussels had no effect on physiological energetics but environmental hypoxia compromised growth, respiration and energy acquisition presumably by reducing feeding rates.

Feeding and overwintering of Antarctic krill across its major habitats: The role of sea ice cover, water depth, and phytoplankton abundance

Antarctic krill (Euphausia superba) were sampled in contrasting habitats: a seasonally ice-covered deep ocean (Lazarev Sea), ice-free shelves at their northern range (South Georgia) and the Antarctic Peninsula (Bransfield Strait), and shelf and oceanic sites in the Scotia Sea. Across 92 stations, representing a year-round average, the food volume in krill stomachs comprised 71 +/- 29% algae, 17 +/- 21% protozoans, and 12 +/- 25% metazoans. Fatty acid trophic markers showed that copepods were consistently part of krill diet, not a switch food. In open waters, both diatom and copepod consumption increased with phytoplankton abundance. Under sea ice, ingestion of diatoms became rare, whereas feeding on copepods remained constant. During winter, larvae contained high but variable proportions of diatom markers, whereas in postlarvae the role of copepods increased with krill body length. Overwintering differed according to habitat. Krill from South Georgia had lower lipid stores than those from the Bransfield Strait or Lazarev Sea. Feeding effort was much reduced in Lazarev Sea krill, whereas most individuals from the Bransfield Strait and South Georgia contained phytoplankton and seabed detritus in their stomachs. Their retention of essential body reserves indicates that krill experienced most winter hardship in the Lazarev Sea, followed by South Georgia and then Bransfield Strait. This was reflected in the delayed development from juveniles to adults in the Lazarev Sea. Circumpolar comparisons of length frequencies suggest that krill growth conditions are more favorable in the southwest Atlantic than in the Lazarev Sea or off East Antarctica because of longer phytoplankton bloom periods and rewarding access to benthic food.

Ocean Net Heat Flux Influences Seasonal to Interannual Patterns of Plankton Abundance

Changes in the net heat flux (NHF) into the ocean have profound impacts on global climate. We analyse a long-term plankton time-series and show that the NHF is a critical indicator of ecosystem dynamics. We show that phytoplankton abundance and diversity patterns are tightly bounded by the switches between negative and positive NHF over an annual cycle. Zooplankton increase before the transition to positive NHF in the spring but are constrained by the negative NHF switch in autumn. By contrast bacterial diversity is decoupled from either NHF switch, but is inversely correlated (r=-0.920) with the magnitude of the NHF. We show that the NHF is a robust mechanistic tool for predicting climate change indicators such as spring phytoplankton bloom timing and length of the growing season.

Response of a floodplain fish community to river-floodplain connectivity: natural versus managed reconnection

To restore lateral connectivity in highly regulated river-floodplain systems, it has become necessary to implement localized, "managed" connection flows, made possible using floodplain irrigation infrastructure. These managed flows contrast with "natural", large-scale, overbank flood pulses. We compared the effects of a managed and a natural connection event on (i) the composition of the large-bodied fish community and (ii) the structure of an endangered catfish population of a large floodplain lake. The change in community composition following the managed connection was not greater than that exhibited between seasons or years during disconnection. By contrast, the change in fish community structure following the natural connection was much larger than that attributed to background, within-and between-year variability during disconnection. Catfish population structure only changed significantly following the natural flood. While the natural flood increased various population rates of native fishes, it also increased those of non-native carp, a pest species. To have a positive influence on native biodiversity, environmental flows may need to be delivered to floodplains in a way that simulates the properties of natural flood pulses. A challenge, however, will be managing river-floodplain connectivity to benefit native more than non-native species.

Family-level AMBI is valid for use in the north-eastern Atlantic but not for assessing the health of microtidal Australian estuaries

Analysis of benthic macroinvertebrate samples at a higher taxonomic level than species, e.g. family, potentially provides a more cost-effective protocol for environmental impact assessments and monitoring as it requires less time, funds and taxonomic expertise. Using the AMBI database, species ecological group scores are shown to be coherent within families. Faunal data from a wide range of environmental impact scenarios in the north-eastern Atlantic demonstrate that AMBI, calculated from mean values for families, exhibits a strong linear relationship with species-level AMBI, the correlation improving by using square-root transformed rather than untransformed abundances. In many regions of the world, however, the sensitivity of benthic macroinvertebrates to environmental perturbations is unknown, precluding the use of AMBI for environmental assessments. Yet the families are essentially the same as in the AMBI database. The utility of family-level AMBI is tested using data for four south-western Australian estuaries previously subjected to environmental quality assessments, but where only 17 species of the 144 taxa are included in the AMBI database. Although family-level AMBI scores reflect differences in environmental quality spatially and temporally within an estuary, they do not follow variations in environmental quality among estuaries. Indeed, south-western Australia estuaries are numerically dominated by families with high AMBI scores, probably due to the detrimental effects of natural accumulations of organic material in estuaries with long residence times. As taxonomic distinctness follows trends in environmental quality among estuaries, as well as temporally and spatially within a system, it provides an appropriate substitute for assessing the 'heath' of microtidal estuaries.

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.

Fishery Exploitation and Stock Assessment of the Endangered Nassau Grouper, Epinephelus Striatus (Actinopterygii: Perciformes: Serranidae), in the Turks and Caicos Islands

The Nassau grouper, Epinephelus striatus (Bloch, 1792), is an endangered species that has been historically overexploited in numerous fisheries throughout its range in the Caribbean and tropical West Atlantic. Data relating fishery exploitation levels to stock abundance of the species are deficient, and protective regulations for the Nassau grouper are yet to be implemented in the Turks and Caicos Islands (TCI). The goal of this study was to conduct a stock assessment and evaluate the exploitation status of the Nassau grouper in the TCI. Materials and methods. Calibrated length cohort analysis was applied to published fisheries data on Nassau grouper landings in the TCI. The total lengths of Nassau groupers among the catches of spearfishers, lobster trappers, and deep sea fishers on the island of South Caicos during 2006 and 2008 were used with estimates of growth, natural mortality, and total annual landings to derive exploitation benchmarks. Results. The TCI stock experienced low to moderate fishing mortality (0.28, 0.18) and exploitation rates (0.49, 0.38) during the period of the study (2006, 2008). However, 21.2%-64.4% of all landings were reproductively immature. Spearfishing appeared to contribute most to fishing mortality relative to the use of lobster traps or hydraulic reels along bank drop-offs. Conclusion. In comparison with available fisheries data for the wider Caribbean, the results reveal the TCI as one of the remaining sites, in addition to the Bahamas, with a substantial Nassau grouper stock. In light of increasing development and tourism in the TCI, continued monitoring is essential to maintain sustainable harvesting practices.