Relating plankton assemblages to environmental variables using instruments towed by ships-of-opportunity

Undulating Oceanographic Recorders (UORs) and Continuous Plankton Recorders (CPRs) equipped with a suite of sensors were towed by merchant vessels in the North Sea between 1988 and 1991, recording a range of environmental variables. These were used to interpret the results of analyses of the plankton taken on CPR tows off the northeast coast of the UK in 1989 and in the Skagerrak and Kattegat in July 1988 and through 1989. Correlations were found between the biota and the environmental variables. The tidal front off the northeast coast of the UK and the front between the low salinity water in the Kattegat and the higher salinity water in the Skagerrak were dominant factors correlating with the distribution of the plankton assemblages. Discontinuities, defining the positions of the fronts, in the values of physical variables (temperature and, where measured, salinity and turbidity) were closely identified with geographical divisions between plankton assemblages. Measures of irradiance were found to be important on several occasions, presumably due to diel migrations of the zooplankton.

Quantifying the impact of environmental variables upon catch-per-unit-effort of the Blue Shark Prionace glauca in the Western English Channel

The effect of environmental variables on blue shark Prionace glauca catch per unit effort (CPUE) in a recreational fishery in the western English Channel, between June and September 1998–2011, was quantified using generalized additive models (GAMs). Sea surface temperature (SST) explained 1·4% of GAM deviance, and highest CPUE occurred at 16·7° C, reflecting the optimal thermal preferences of this species. Surface chlorophyll a concentration (CHL) significantly affected CPUE and caused 27·5% of GAM deviance. Additionally, increasing CHL led to rising CPUE, probably due to higher productivity supporting greater prey biomass. The density of shelf-sea tidal mixing fronts explained 5% of GAM deviance, but was non-significant, with increasing front density negatively affecting CPUE. Time-lagged frontal density significantly affected CPUE, however, causing 12·6% of the deviance in a second GAM and displayed a positive correlation. This outcome suggested a delay between the evolution of frontal features and the subsequent accumulation of productivity and attraction of higher trophic level predators, such as P. glauca.

Quantifying the impact of environmental variables upon catch-per-unit-effort of the Blue Shark Prionace glauca in the Western English Channel

The effect of environmental variables on blue shark Prionace glauca catch per unit effort (CPUE) in a recreational fishery in the western English Channel, between June and September 1998–2011, was quantified using generalized additive models (GAMs). Sea surface temperature (SST) explained 1·4% of GAM deviance, and highest CPUE occurred at 16·7° C, reflecting the optimal thermal preferences of this species. Surface chlorophyll a concentration (CHL) significantly affected CPUE and caused 27·5% of GAM deviance. Additionally, increasing CHL led to rising CPUE, probably due to higher productivity supporting greater prey biomass. The density of shelf-sea tidal mixing fronts explained 5% of GAM deviance, but was non-significant, with increasing front density negatively affecting CPUE. Time-lagged frontal density significantly affected CPUE, however, causing 12·6% of the deviance in a second GAM and displayed a positive correlation. This outcome suggested a delay between the evolution of frontal features and the subsequent accumulation of productivity and attraction of higher trophic level predators, such as P. glauca.

Linking environmental variables with regional- scale variability in ecological structure and standing stock of carbon within UK kelp forests

Kelp forests represent some of the most productive and diverse habitats on Earth. Understanding drivers of ecological patterns at large spatial scales is critical for effective management and conservation of marine habitats. We surveyed kelp forests dominated by Laminaria hyperborea (Gunnerus) Foslie 1884 across 9° latitude and >1000 km of coastline and measured a number of physical parameters at multiple scales to link ecological structure and standing stock of carbon with environmental variables. Kelp density, biomass, morphology and age were generally greater in exposed sites within regions, highlighting the importance of wave exposure in structuring L. hyperborea populations. At the regional scale, wave-exposed kelp canopies in the cooler regions (the north and west of Scotland) were greater in biomass, height and age than in warmer regions (southwest Wales and England). The range and maximal values of estimated standing stock of carbon contained within kelp forests was greater than in historical studies, suggesting that this ecosystem property may have been previously undervalued. Kelp canopy density was positively correlated with large-scale wave fetch and fine-scale water motion, whereas kelp canopy biomass and the standing stock of carbon were positively correlated with large-scale wave fetch and light levels and negatively correlated with temperature. As light availability and summer temperature were important drivers of kelp forest biomass, effective management of human activities that may affect coastal water quality is necessary to maintain ecosystem functioning, while increased temperatures related to anthropogenic climate change may impact the structure of kelp forests and the ecosystem services they provide.

Linking environmental variables with regional- scale variability in ecological structure and standing stock of carbon within UK kelp forests

Kelp forests represent some of the most productive and diverse habitats on Earth. Understanding drivers of ecological patterns at large spatial scales is critical for effective management and conservation of marine habitats. We surveyed kelp forests dominated by Laminaria hyperborea (Gunnerus) Foslie 1884 across 9° latitude and >1000 km of coastline and measured a number of physical parameters at multiple scales to link ecological structure and standing stock of carbon with environmental variables. Kelp density, biomass, morphology and age were generally greater in exposed sites within regions, highlighting the importance of wave exposure in structuring L. hyperborea populations. At the regional scale, wave-exposed kelp canopies in the cooler regions (the north and west of Scotland) were greater in biomass, height and age than in warmer regions (southwest Wales and England). The range and maximal values of estimated standing stock of carbon contained within kelp forests was greater than in historical studies, suggesting that this ecosystem property may have been previously undervalued. Kelp canopy density was positively correlated with large-scale wave fetch and fine-scale water motion, whereas kelp canopy biomass and the standing stock of carbon were positively correlated with large-scale wave fetch and light levels and negatively correlated with temperature. As light availability and summer temperature were important drivers of kelp forest biomass, effective management of human activities that may affect coastal water quality is necessary to maintain ecosystem functioning, while increased temperatures related to anthropogenic climate change may impact the structure of kelp forests and the ecosystem services they provide.

Global latitudinal variations in marine copepod diversity and environmental factors

Latitudinal gradients in diversity are among the most striking features in ecology. For terrestrial species, climate (i.e. temperature and precipitation) is believed to exert a strong influence on the geographical distributions of diversity through its effects on energy availability. Here, we provide the first global description of geographical variation in the diversity of marine copepods, a key trophic link between phytoplankton and fish, in relation to environmental variables. We found a polar-tropical difference in copepod diversity in the Northern Hemisphere where diversity peaked at subtropical latitudes. In the Southern Hemisphere, diversity showed a tropical plateau into the temperate regions. This asymmetry around the Equator may be explained by climatic conditions, in particular the influence of the Inter-Tropical Convergence Zone, prevailing mainly in the northern tropical region. Ocean temperature was the most important explanatory factor among all environmental variables tested, accounting for 54 per cent of the variation in diversity. Given the strong positive correlation between diversity and temperature, local copepod diversity, especially in extra-tropical regions, is likely to increase with climate change as their large-scale distributions respond to climate warming.

Aquatic Distribution And Heterotrophic Degradation Of Polycyclic Aromatic-Hydrocarbons (PAH) In The Tamar Estuary

Variations in the concentrations and microheterotrophic degradation rates of selected Polycyclic Aromatic Hydrocarbons (PAH) in the water column of the Tamar Estuary were investigated in relation to the major environmental variables. Concentrations of individual PAH varied typically between i and 50 ng l−1 Based on their observed environmental behaviour the PAH appeared divisible into two groupings: (1) low molecular weight PAH incorporating naphthalene, phenanthrene and anthracence and (a) the larger molecular weight homologues (fluoranthene, pyrene, chrysene, benz(a)anthracene, benzo(b)fluoranthene, benzo(k)fluoranthene, benzo(a)-pyrene). Group 1 PAH showed a complex distribution throughout the estuary with no significant correlations with either salinity or suspended particulates. Based on their relatively low particle affinity and high water solubilities and vapour pressures, volatilization is proposed as an important process in determining their fate. Microheterotrophic turnover times of naphthalene varied between x and 30 days, and were independent of suspended solids with maximum degradation rates located in the central and urban regions of the Estuary. When compared with the flushing times for the Tamar (3–5 days), it is probable that heterotrophic activity is important in the removal of naphthalene (and possibly the other Group 1 PAH) from the estuarine environment. In contrast Group 2 PAH concentrations exhibited highly significant correlations with suspended particulates. Highest concentrations occurred at the turbidity maximum, with a secondary concentration maximum localized to the industrialized portion of the estuary and associated with anthropogenic inputs. Laboratory degradation studies of benzo(a)pyrene in water samples taken from the estuary showed turnover times for the compound of between 2000 and 9000 days. Degradation rates correlated positively with suspended solids. The high particulate affinity and microbial refractivity of Group 2 PAH indicate sediment burial as the principal tate of these PAH in the Tamar Estuary. Estuarine sediments contained typically 50–1500 ng g−1 dry weight of individual PAH which were comparable to the levels of Group 2 PAH associated with the suspended particulates. Highest concentrations occurred at the riverine end of the estuary resulting from unresolved inputs in the catchment. Subsequent dilution by less polluted marine sediments together with slow degradation results in a seaward trend of decreasing concentrations. However, there is a secondary maximum of PAH superimposed on this trend which is associated with urban Plymouth.

Defining seasonal marine microbial community dynamics

Here we describe, the longest microbial time-series analyzed to date using high-resolution 16S rRNA tag pyrosequencing of samples taken monthly over 6 years at a temperate marine coastal site off Plymouth, UK. Data treatment effected the estimation of community richness over a 6-year period, whereby 8794 operational taxonomic units (OTUs) were identified using single-linkage preclustering and 21 130 OTUs were identified by denoising the data. The Alphaproteobacteria were the most abundant Class, and the most frequently recorded OTUs were members of the Rickettsiales (SAR 11) and Rhodobacteriales. This near-surface ocean bacterial community showed strong repeatable seasonal patterns, which were defined by winter peaks in diversity across all years. Environmental variables explained far more variation in seasonally predictable bacteria than did data on protists or metazoan biomass. Change in day length alone explains >65% of the variance in community diversity. The results suggested that seasonal changes in environmental variables are more important than trophic interactions. Interestingly, microbial association network analysis showed that correlations in abundance were stronger within bacterial taxa rather than between bacteria and eukaryotes, or between bacteria and environmental variables.

Structure and diversity of intertidal benthic diatom assemblages in contrasting shores: a case study from the Tagus estuary1

The structure of intertidal benthic diatoms assemblages in the Tagus estuary was investigated during a 2-year survey, carried out in six stations with different sediment texture. Nonparametric multivariate analyses were used to characterize spatial and temporal patterns of the assemblages and to link them to the measured environmental variables. In addition, diversity and other features related to community physiognomy, such as size-class or life-form distributions, were used to describe the diatom assemblages. A total of 183 diatom taxa were identified during cell counts and their biovolume was determined. Differences between stations (analysis of similarity (ANOSIM), R=0.932) were more evident than temporal patterns (R=0.308) and mud content alone was the environmental variable most correlated to the biotic data (BEST, rho=0.863). Mudflat stations were typically colonized by low diversity diatom assemblages (H' similar to 1.9), mainly composed of medium-sized motile epipelic species (250-1,000 mu m(3)), that showed species-specific seasonal blooms (e.g., Navicula gregaria Donkin). Sandy stations had more complex and diverse diatom assemblages (H' similar to 3.2). They were mostly composed by a large set of minute epipsammic species (<250 mu m(3)) that, generally, did not show temporal patterns. The structure of intertidal diatom assemblages was largely defined by the interplay between epipelon and epipsammon, and its diversity was explained within the framework of the Intermediate Disturbance Hypothesis. However, the spatial distribution of epipelic and epipsammic life-forms showed that the definition of both functional groups should not be over-simplified.

Reply to Haddock, S. H. D. Reconsidering evidence for potential climate-related increases in jellyfish

Following the publication of our paper (Attrill et al. 2007), we became quickly aware of a couple of errors. We have subsequently been collaborating with Dr. Chris Lynam (Lynam et al. 2004, 2005) to bring together our two datasets, explore the common patterns within our data, and attempt to provide a consensus on how climate is affecting gelatinous plankton in the North Sea. During this reanalysis, two errors within the data were discovered, one involving a transcription error of a column of residuals during de-trended analysis, the other a major data entry error deep in the Continuous Plankton Recorder (CPR) database for sector B2. Here we present a revised version of table 1 from Attrill et al. (2007) to incorporate corrections to these transcription and data entry errors. These corrections alter some of the results in our original data table, mainly to increase and strengthen the number of significant relations we found (e.g., for sector B2 and whole sea area); all previous main results remain robustly significant. Following discussions with Dr. Lynam, two clarifications of statements made in Attrill et al. (2007) are also required. Page 482, Results, last line of first column: ‘‘There were no...robust, consistent relations between jellyfish frequency and any environmental variables for B and D… contrary to the findings of previous shorter time series (Lynam et al. 2005).’’ The Lynam et al. (2004, 2005) papers presented no data for the D sector and found no link in the B sector, contrary to our revised results. Page 482, Discussion, paragraph 1, last sentence: ‘‘… positive association … North of Scotland (Lynam et al. 2005) … does not appear to be maintained.’’ Our paper did not report on any data that covered Lynam et al.’s (2005) North of Scotland area so the statement is not directly supported, although their positive relation North of Scotland, when considered in conjunction with inflow, may agree with the C2 and B2 results of Attrill et al. (2007).

Patterns of jellyfish abundance in the North Atlantic

A number of explanations have been advanced to account for the increased frequency and intensity at which jellyfish (pelagic cnidarians and ctenophores) blooms are being observed, most of which have been locally directed. Here, we investigate seasonal and inter-annual patterns in abundance and distribution of jellyfish in the North Atlantic Ocean to determine if there have been any system-wide changes over the period 1946–2005, by analysing records of the presence of coelenterates from the Continuous Plankton Recorder (CPR) survey. Peaks in jellyfish abundance are strongly seasonal in both oceanic and shelf areas: oceanic populations have a mid-year peak that is more closely related to peaks in phyto- and zooplankton, whilst the later peak of shelf populations mirrors changes in SST and reflects processes of advection and aggregation. There have been large amplitude cycles in the abundance of oceanic and shelf jellyfish (although not synchronous) over the last 60 years, with a pronounced synchronous increase in abundance in both areas over the last 10 years. Inter-annual variations in jellyfish abundance in oceanic areas are related to zooplankton abundance and temperature changes, but not to the North Atlantic Oscillation or to a chlorophyll index. The long-term inter-annual abundance of jellyfish on the shelf could not be explained by any environmental variables investigated. As multi-decadal cycles and more recent increase in jellyfish were obvious in both oceanic and shelf areas, we conclude that these are likely to reflect an underlying climatic signal (and bottom-up control) rather than any change in fishing pressure (top-down control). Our results also highlight the role of the CPR data in investigating long-term changes in jellyfish, and suggest that the cnidarians sampled by the CPR are more likely to be holoplanktic hydrozoans and not the much larger meroplanktic scyphozoans as has been suggested previously.

Long-term effects of ocean warming on the prokaryotic community: evidence from the vibrios

The long-term effects of ocean warming on prokaryotic communities are unknown because of lack of historical data. We overcame this gap by applying a retrospective molecular analysis to the bacterial community on formalin-fixed samples from the historical Continuous Plankton Recorder archive, which is one of the longest and most geographically extensive collections of marine biological samples in the world. We showed that during the last half century, ubiquitous marine bacteria of the Vibrio genus, including Vibrio cholerae, increased in dominance within the plankton-associated bacterial community of the North Sea, where an unprecedented increase in bathing infections related to these bacteria was recently reported. Among environmental variables, increased sea surface temperature explained 45% of the variance in Vibrio data, supporting the view that ocean warming is favouring the spread of vibrios and may be the cause of the globally increasing trend in their associated diseases.