From plankton to top predators: bottom-up control of a marine food web across four trophic levels

1. Abundant mid-trophic pelagic fish often play a central role in marine ecosystems, both as links between zooplankton and top predators and as important fishery targets. In the North Sea, the lesser sandeel occupies this position, being the main prey of many bird, mammal and fish predators and the target of a major industrial fishery. However, since 2003, sandeel landings have decreased by > 50%, and many sandeel-dependent seabirds experienced breeding failures in 2004. 2. Despite the major economic implications, current understanding of the regulation of key constituents of this ecosystem is poor. Sandeel abundance may be regulated 'bottom-up' by food abundance, often thought to be under climatic control, or 'top-down' by natural or fishery predation. We tested predictions from these two hypotheses by combining unique long-term data sets (1973–2003) on seabird breeding productivity from the Isle of May, SE Scotland, and plankton and fish larvae from the Continuous Plankton Recorder survey. We also tested whether seabird breeding productivity was more tightly linked to sandeel biomass or quality (size) of individual fish. 3. The biomass of larval sandeels increased two- to threefold over the study period and was positively associated with proxies of the abundance of their plankton prey. Breeding productivity of four seabirds bringing multiple prey items to their offspring was positively related to sandeel larval biomass with a 1-year lag, indicating dependence on 1-year-old fish, but in one species bringing individual fish it was strongly associated with the size of adult sandeels. 4. These links are consistent with bottom-up ecosystem regulation and, with evidence from previous studies, indicate how climate-driven changes in plankton communities can affect top predators and potentially human fisheries through the dynamics of key mid-trophic fish. However, the failing recruitment to adult sandeel stocks and the exceptionally low seabird breeding productivity in 2004 were not associated with low sandeel larval biomass in 2003, so other mechanisms (e.g. predation, lack of suitable food after metamorphosis) must have been important in this case. Understanding ecosystem regulation is extremely important for predicting the fate of keystone species, such as sandeels, and their predators.

Hormesis - The Stimulation Of Growth By Low-Levels Of Inhibitors

Hormesis is the name given to the stimulatory effects caused by low levels of potentially toxic agents. When this phenomenon was first identified it was called the Arndt-Schulz Law or Hueppe's Rule, because it was thought to occur generally. Although this generalisation is not accepted today, there has never been more evidence in its support, justifying a re-examination of the phenomenon. Evidence from the literature shows that not only has growth hormesis been observed in a range of taxa after exposure to a variety of agents, but also that the dose-response data have a consistent form. While there are a number of separate hypotheses to explain specific instances of hormesis, the evidence presented here suggests that different examples might have a common explanation, and the possibility of a general theory is considered.

Shore height and differentials between macrobenthic assemblages in vegetated and unvegetated areas of an intertidal sandflat

Intertidal macrobenthic faunal assemblages of a dual seagrass/callianassid-structured sandflat system were investigated in subtropical Moreton Bay, Queensland. Consistently across all 20 stations, the gastropod-dominated seagrass supported greater abundance (2.5×) and species richness (2×) than the amphipod-dominated sandflat. There was no evidence of along-shore or up-shore variation in the overall assemblage properties such as total abundance, species richness or diversity within either habitat type, except for variation in sandflat abundance between sites. But seagrass and sandflat assemblages both varied significantly in composition from site to site, and seagrass assemblage composition also varied with shore height. Shore height and site, however, only accounted for ≤41% of total variation. The two faunal assemblages showed a Bray–Curtis dissimilarity of 97.7% and within-habitat similarities of <20%. There was no consistency in distribution of greater diversity, dominance or evenness. No differential between any assemblage features in adjacent sandflat and seagrass samples changed with shore height, supporting hypotheses that such differentials are not maintained by predation. Macrofaunal species richness and diversity were closely coupled within sandflat stations but were uncoupled within seagrass ones, questioning the value of diversity as a comparative measure.

Macroecological study of Centropages typicus in the North Atlantic Ocean

Centropages typicus is a temperate neritic-coastal species of the North Atlantic Oceans, generally found between the latitudes of the Mediterranean and the Norwegian Sea. Therefore, the species experiences a large number of environments and adjusts its life cycle in response to changes in key abiotic parameters such as temperature. Using data from the Continuous Plankton Recorder (CPR) Survey, we review the macroecology of C. typicus and factors that influence its spatial distribution, phenology and year-to-year to decadal variability. The ecological preferences are identified and quantified. Mechanisms that allow the species to occur in such different environments are discussed and hypotheses are proposed as to how the species adapts to its environment. We show that temperature and both quantity and quality of phytoplankton are important factors explaining the space and time variability of C. typicus. These results show that C. typicus will not respond only to temperature increase in the region but also to changes in phytoplankton abundance, structure and composition and timing of occurrence. Methods such as a decision tree can help to forecast expected changes in the distribution of this species with hydro-climatic forcing. (C) 2007 Elsevier Ltd. All rights reserved.

Recruitment of lesser sandeel Ammodytes marinus in relation to density dependence and zooplankton composition

Recent recruitment failure of lesser sandeel Ammodytes marinus, a key prey fish in the North Sea, followed by several years of low spawning stock biomass, prompted us to investigate factors influencing the recruitment of this species. We tested 2 hypotheses that relate to ecological mechanisms of recruitment regulation in lesser sandeel: (1) a positive spawning stock–recruitment relationship is decoupled in years associated with high abundances of age-1 sandeels and (2) the survival success of early larvae depends specifically on the abundance of Calanus finmarchicus and not C. helgolandicus. The findings of the present study supported both hypotheses and resulted in a multiple linear recruitment model with pronounced predictive capabilities. The model includes interactions between age-1 abundance and spawning stock biomass, plus the effect of C. finmarchicus abundance, and it explained around 65% of the inter-annual variation in recruitment in contrast to only 12% by a traditional Ricker curve. We argue that early egg production in C. finmarchicus supports the survival of larvae, and that climate-generated shifts in the Calanus species composition lead to a mismatch in timing between food availability and the early life history of lesser sandeels.

Using the Continuous Plankton Recorder to investigate the absence of North Atlantic right whales (Eubalaena glacialis) from the Roseway Basin foraging ground

North Atlantic right whales (Eubalaena glacialis) were absent from Roseway Basin, located off southeastern Nova Scotia, for a 7-year period (1993–1999). The objective of this study was to examine the availability of the right whale's main prey, Calanus finmarchicus, in Roseway Basin during those 7 years to determine if the whales’ absence was due to inadequate prey resources. Since we had no historical data on zooplankton abundances at depth on the Scotian Shelf, near-surface zooplankton abundance data from the Continuous Plankton Recorder were used to infer water-column abundances. In addition, environmental parameters that are often correlated with high zooplankton concentrations were examined. The hypotheses tested were that changes in these parameters would be detectable between three time periods: pre-1993, 1993–1999 and post-1999. Calanus finmarchicus abundance was found to be lowest during 1993–1999, suggesting that right whales were not foraging in Roseway Basin because of the near-absence of their main prey species. Decreased in situ salinity and density proved to be indicators of the changes in circulation in the 1990s that may have affected the advection of C. finmarchicus onto the Scotian Shelf.

Macro-ecology of plankton–seabird associations in the North Pacific Ocean

In conjunction with the North Pacific Continuous Plankton Recorder program, we conducted surveys of seabirds from June 2002 to June 2007. Here, we tested the hypotheses of (i) east–west variations in coupled plankton and seabird abundance, and (ii) that surface-feeding and diving seabirds vary in their relationships to primary productivity and mesozooplankton species abundance and diversity. To test these hypotheses, we developed statistical models for 20 species of seabirds and 12 zooplankton taxonomic groups. Seabird density was highly variable between seasons, but was consistently higher in the western than eastern North Pacific. Seabird diversity was greater in the east. Zooplankton abundance did not differ between regions. We found associations at the “bulk” level between seabird density and net primary productivity, but only one association between seabirds and total zooplankton abundance or diversity. However, we found many relationships between seabird species and the abundance of different zooplankton summarized at the genus or family level. Some of these taxonomic relationships reflect direct predator–prey interactions, while others may reflect zooplankton that serve as ecological indicators of other prey, such as micronekton, upon which the birds may feed. Surface or near-surface feeding, mostly piscivorous seabirds, did not differ systematically from diving, mainly planktivorous seabirds in their zooplankton associations. Seabirds apparently respond to zooplankton taxonomic groupings more so than bulk zooplankton characteristics, such as abundance or diversity. Macro-ecological studies of remote marine ecosystems using zooplankton and seabirds as ecological indicators provide a framework for understanding and assessing spatial and temporal variations in these difficult-to-study pelagic environments.

Long-term individual foraging site fidelity – why some gannets don’t change their spots

Many established models of animal foraging assume that individuals are ecologically equivalent. However, it is increasingly recognized that populations may comprise individuals who differ consistently in their diets and foraging behaviors. For example, recent studies have shown that individual foraging site fidelity (IFSF, when individuals consistently forage in only a small part of their population's home range) occurs in some colonial breeders. Short‐term IFSF could result from animals using a win–stay, lose–shift foraging strategy. Alternatively, it may be a consequence of individual specialization. Pelagic seabirds are colonial central‐place foragers, classically assumed to use flexible foraging strategies to target widely dispersed, spatiotemporally patchy prey. However, tracking has shown that IFSF occurs in many seabirds, although it is not known whether this persists across years. To test for long‐term IFSF and to examine alternative hypotheses concerning its cause, we repeatedly tracked 55 Northern Gannets (Morus bassanus) from a large colony in the North Sea within and across three successive breeding seasons. Gannets foraged in neritic waters, predictably structured by tidal mixing and thermal stratification, but subject to stochastic, wind‐induced overturning. Both within and across years, coarse to mesoscale (tens of kilometers) IFSF was significant but not absolute, and foraging birds departed the colony in individually consistent directions. Carbon stable isotope ratios in gannet blood tissues were repeatable within years and nitrogen ratios were also repeatable across years, suggesting long‐term individual dietary specialization. Individuals were also consistent across years in habitat use with respect to relative sea surface temperature and in some dive metrics, yet none of these factors accounted for IFSF. Moreover, at the scale of weeks, IFSF did not decay over time and the magnitude of IFSF across years was similar to that within years, suggesting that IFSF is not primarily the result of win–stay, lose–shift foraging. Rather, we hypothesize that site familiarity, accrued early in life, causes IFSF by canalizing subsequent foraging decisions. Evidence from this and other studies suggests that IFSF may be common in colonial central‐place foragers, with far‐reaching consequences for our attempts to understand and conserve these animals in a rapidly changing environment.

Looking backwards to look forwards: the role of natural history in temperate reef ecology

Temperate reefs are superb tractable systems for testing hypotheses in ecology and evolutionary biology. Accordingly there is a rich history of research stretching back over 100 years, which has made major contributions to general ecological and evolutionary theory as well as providing better understanding of how littoral systems work by linking pattern with process. A brief resumé of the history of temperate reef ecology is provided to celebrate this rich heritage. As a community, temperate reef ecologists generally do well designed experiments and test well formulated hypotheses. Increasingly large datasets are being collected, collated and subjected to complex meta-analyses and used for modelling. These datasets do not happen spontaneously – the burgeoning subject of macroecology is possible only because of the efforts of dedicated natural historians whether it be observing birds, butterflies, or barnacles. High-quality natural history and old-fashioned field craft enable surveys or experiments to be stratified (i.e. replicates are replicates and not a random bit of rock) and lead to the generation of more insightful hypotheses. Modern molecular approaches have led to the discovery of cryptic species and provided phylogeographical insights, but natural history is still required to identify species in the field. We advocate a blend of modern approaches with old school skills and a fondness for temperate reefs in all their splendour.

Microplastics Alter the Properties and Sinking Rates of Zooplankton Faecal Pellets

Plastic debris is a widespread contaminant, prevalent in aquatic ecosystems across the globe. Zooplankton readily ingest microscopic plastic (microplastic, < 1 mm), which are later egested within their faecal pellets. These pellets are a source of food for marine organisms, and contribute to the oceanic vertical flux of particulate organic matter as part of the biological pump. The effects of microplastics on faecal pellet properties are currently unknown. Here we test the hypotheses that (1) faecal pellets are a vector for transport of microplastics, (2) polystyrene microplastics can alter the properties and sinking rates of zooplankton egests and, (3) faecal pellets can facilitate the transfer of plastics to coprophagous biota. Following exposure to 20.6 μm polystyrene microplastics (1000 microplastics mL–1) and natural prey (∼1650 algae mL–1) the copepod Calanus helgolandicus egested faecal pellets with significantly (P < 0.001) reduced densities, a 2.25-fold reduction in sinking rates, and a higher propensity for fragmentation. We further show that microplastics, encapsulated within egests of the copepod Centropages typicus, could be transferred to C. helgolandicus via coprophagy. Our results support the proposal that sinking faecal matter represents a mechanism by which floating plastics can be vertically transported away from surface waters.