Communities of microscopic plant life, or phytoplankton, dominate the Earth's aquatic ecosystems. This important new book by Colin Reynolds covers the adaptations, physiology and population dynamics of phytoplankton communities in lakes and rivers and oceans. It provides basic information on composition, morphology and physiology of the main phyletic groups represented in marine and freshwater systems and in addition reviews recent advances in community ecology, developing an appreciation of assembly processes, co-existence and competition, disturbance and diversity. Although focussed on one group of organisms, the book develops many concepts relevant to ecology in the broadest sense, and as such will appeal to graduate students and researchers in ecology, limnology and oceanography.

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The Ecology of Phytoplankton

Transparent exopolymer particles (TEP) in aquatic environments.

Although recent articles state that jellyfish populations are increasing, most available evidence shows that jellyfish abundances fluctuate with climatic cycles. Reports of increasing prob- lems with jellyfish, especially in East Asia, are too recent to exclude decadal climate cycles. Jellyfish are infamous for their direct negative effects on human enterprise; specifically, they interfere with tourism by stinging swimmers, fishing by clogging nets, aquaculture by killing fish in net-pens and power plants by clogging cooling-water intake screens. They also have indirect effects on fisheries by feeding on zooplankton and ichthyoplankton, and, therefore, are predators and potential competitors of fish. Ironically, many human activities may contribute to increases in jellyfish populations in coastal waters. Increased jellyfish and ctenophore populations often are associated with warming caused by climate changes and possibly power plant thermal effluents. Jellyfish may benefit from eutrophication, which can increase small-zooplankton abundance, turbidity and hypoxia, all conditions that may favor jellyfish over fish. Fishing activities can remove predators of jellyfish and zooplanktivorous fish com- petitors as well as cause large-scale ecosystem changes that improve conditions for jellyfish. Aquacul- ture releases millions of jellyfish into Asian coastal waters yearly to enhance the jellyfish fishery. Aquaculture and other marine structures provide favorable habitat for the benthic stages of jellyfish. Changes in the hydrological regime due to dams and other construction can change the salinity to favor jellyfish. Accidental introductions of non-native gelatinous species into disturbed ecosystems have led to blooms with serious consequences. In many coastal areas, most of these environmental changes occur simultaneously. We summarize cases of problem jellyfish blooms and the evidence for anthropogenic habitat disruptions that may have caused them. Rapid development in East Asia makes that region especially vulnerable to escalating problems. We conclude that human effects on coastal environments are certain to increase, and jellyfish blooms may increase as a consequence.

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Anthropogenic causes of jellyfish blooms and their direct consequences for humans: a review

▪ Abstract It has often been argued that conserving biodiversity is necessary for maintaining ecosystem functioning. We critically evaluate the current evidence for this argument. Although there is substantial evidence that diversity is able to affect function, particularly for plant communities, it is unclear if these patterns will hold for realistic scenarios of extinctions, multitrophic communities, or larger spatial scales. Experiments are conducted at small spatial scales, the very scales at which diversity tends to increase owing to exotics. Stressors may affect function by many pathways, and diversity-mediated effects on function may be a minor pathway, except in the case of multiple-stressor insurance effects. In general, the conservation case is stronger for stability measures of function than stock and flux measures, in part because it is easier to attribute value unambiguously to stability and in part because stock and flux measures of functions are anticipated to be more affected by multitroph...

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Biodiversity-Ecosystem Function Research: Is It Relevant to Conservation?

Proposed links between biodiversity and ecosystem processes have generated intense interest and controversy in recent years. With few exceptions, however, empirical studies have focused on grassland plants and laboratory aquatic microbial systems, whereas there has been little attention to how changing animal diversity may influence ecosystem processes. Meanwhile, a separate research tradition has demonstrated strong top-down forcing in many systems, but has considered the role of diversity in these processes only tangentially. Integration of these research directions is necessary for more complete understanding in both areas. Several considerations suggest that changing diversity in multi-level food webs can have important ecosystem effects that can be qualitatively different than those mediated by plants. First, extinctions tend to be biased by trophic level: higher-level consumers are less diverse, less abundant, and under stronger anthropogenic pressure on average than wild plants, and thus face greater risk of extinction. Second, unlike plants, consumers often have impacts on ecosystems disproportionate to their abundance. Thus, an early consequence of declining diversity will often be skewed trophic structure, potentially reducing top-down influence. Third, where predators remain abundant, declining diversity at lower trophic levels may change effectiveness of predation and penetrance of trophic cascades by reducing trait diversity and the potential for compensation among species within a level. The mostly indirect evidence available provides some support for this prediction. Yet effects of changing animal diversity on functional processes have rarely been tested experimentally. Evaluating impacts of biodiversity loss on ecosystem function requires expanding the scope of current experimental research to multi-level food webs. A central challenge to doing so, and to evaluating the importance of trophic cascades specifically, is understanding the distribution of interaction strengths within natural communities and how they change with community composition. Although topology of most real food webs is extremely complex, it is not at all clear how much of this complexity translates to strong dynamic linkages that influence aggregate biomass and community composition. Finally, there is a need for more detailed data on patterns of species loss from real ecosystems (community “disassembly” rules).

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Biodiversity and ecosystem function: the consumer connection

Based on existing knowledge about phytoplankton responses to nutrients and food size spectra of herbivorous zooplankton, three different configurations of pelagic food webs are proposed for three different types of marine nutrient regimes: (1) upwelling systems, (2) oligotrophic oceanic systems, (3) eutrophicated coastal systems. Up-welling systems are characterised by high levels of plant nutrients and high ratios of Si to N and R. Phytoplankton consists mainly of diatoms together with a subdominant contribution of flagellates. Most phytoplankton falls into the food spectrum of herbivorous, crustacean zooplankton. Therefore, herbivorous crustaceans occupy trophic level 2 and zooplanktivorous fish occupy trophic level 3. Phytoplankton in oligotrophic, oceanic systems is dominated by picoplankton, which are too small to be ingested by copepods. Most primary production is channelled through the ‘microbial loop’ (picoplankton — heterotrophic nanoflagellates — ciliates). Sporadically, pelagic tunicates also consume a substantial proportion of primary production. Herbivorous crustaceans feed on heterotrophic nanoflagellates and ciliates, thus occupying a food chain position between 3 and 4, which leads to a food chain position between 4 and 5 for zooplanktivorous fish. By cultural eutrophication, N and P availability are elevated while Si remains unaffected or even declines. Diatoms decrease in relative importance while summer blooms of inedible algae (Phaeocystis, toxic dinoflagellates, toxic prymnesiophyceae, etc.) prevail. The spring bloom may still contain a substantial contribution of diatoms. The production of the inedible algae enters the pelagic energy flow via the detritus food chain: DOC release by cell lysis — bacteria — heterotrophic nanoflagellates — ciliates. Accordingly, crustacean zooplankton occupy food chain position 4 to 5 during the non-diatom seasons. Ecological efficiency considerations lead to the conclusion that fish production:primary production ratios should be highest in upwelling systems and substantially lower in oligotrophic and in culturally eutrophicated systems. Further losses of fish production may occur when carnivorous, gelatinous zooplankton (jellyfish) replace fish.

Pelagic food web configurations at different levels of nutrient richness and their implications for the ratio fish production:primary production

Top–down control of phytoplankton by crustacean mesozooplankton is a cornerstone of freshwater ecology. Apparently, trophic cascades are more frequently reported from freshwater than from marine plankton. We argue that this difference is real and mainly caused by biological differences at the zooplankton–phytoplankton link: cladocerans (particularly Daphnia) in the lakes and copepods in the sea. We derive these conclusions from recent literature and a number of own, similarly designed mesocosm experiments conducted in a lake, a brackish water and a marine site. In all experiments, phytoplankton were exposed to gradients of experimentally manipulated densities of zooplankton, including freshwater copepods and cladocerans, and marine copepods and appendicularians. The suggested reasons for the difference between lake and marine trophic cascades are: (1) Both copepods and cladocerans suppress only part of the phytoplankton size spectrum: cladocerans the small and copepods the large phytoplankton. (2) If not controlled by grazing, small phytoplankton may increase their biomass faster than large phytoplankton. (3) Copepods additionally release small phytoplankton from grazing pressure by intermediate consumers (protozoa) and competitors (predation on appendicularian eggs), while cladocerans do not release large phytoplankton from grazing pressure by any functional group. (4) Cladocerans sequester more of the limiting nutrient than copepods, leaving fewer nutrients available for compensatory growth of ungrazed phytoplankton.

Cladocerans versus copepods: the cause of contrasting top–down controls on freshwater and marine phytoplankton

The differences in the impact of two major groups of herbivorous zooplankton (Cladocera and Copepoda) on summer phytoplankton in a mesotrophic lake were studied. Field experiments were performed in which phytoplankton were exposed to different densities of two major types of herbivorous zooplankton, cladocerans and copepods. Contrary to expectation, neither of the two zooplankton groups significantly reduced phytoplankton biomass. However, there were strong and contrasting impacts on phytoplankton size structure and on individual taxa. Cladocerans suppressed small phytoplankton, while copepods suppressed large phytoplankton. The unaffected size classes compensated for the loss of those affected by enhanced growth. After contamination of the copepod mesocosms with the cladoceran Daphnia, the combined impact of both zooplankton groups caused a decline in total phytoplankton biomass.

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Complementary impact of copepods and cladocerans on phytoplankton

Here we report on a mesocom study performed to compare the top-down impact of microphagous and macrophagous zooplankton on phytoplankton. We exposed a species-rich, summer phytoplankton assemblage from the mesotrophic Lake Schohsee (Germany) to logarithmically scaled abundance gradients of the microphagous cladoceran Daphnia hyalina×galeata and of a macrophagous copepod assemblage. Total phytoplankton biomass, chlorophyll a and primary production showed only a weak or even insignificant response to zooplankton density in both gradients. In contrast to the weak responses of bulk parameters, both zooplankton groups exerted a strong and contrasting influence on the phytoplankton species composition. The copepods suppressed large phytoplankton, while nanoplanktonic algae increased with increasing copepod density. Daphnia suppressed small algae, while larger species compensated in terms of biomass for the losses. Autotrophic picoplankton declined with zooplankton density in both gradients. Gelatinous, colonial algae were fostered by both zooplankton functional groups, while medium-sized (ca. 3,000 µm3), non-gelatinous algae were suppressed by both. The impact of a functionally mixed zooplankton assemblage became evident when Daphnia began to invade and grow in copepod mesocosms after ca. 10 days. Contrary to the impact of a single functional group, the combined impact of both zooplankton groups led to a substantial decline in total phytoplankton biomass.

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Daphnia versus copepod impact on summer phytoplankton: functional compensation at both trophic levels

Grazing experiments were conducted with natural mesozooplankton from Kiel Bight, Germany, using radioactive labelled phytoplankton cultures and seston size fractions. The results of experiments using phytoplankton cultures indicated that bivalve veligers performed highest clearance of particles within a size range of 4.7 to 6.3 pm, whereas optimum particle size for copepods was 15 pm. The results of experiments using labelled natural seston size fractions identified bivalve veligers and appendicularians as those responsible for the removal of particles within the smallest size class (<2 μm) Seston size fractions larger than 5 pm were mainly cleared by copepods and nauplii. As particle size increased, the contribution of copepod clearance to total zooplankton clearance within size classes increased from 57 % (<5 pm size class) to more than 81 % (30 to 100 pm size class). When the nauplii clearance rates were included, the total copepod clearance accounted for 90 to 97.6% of the total volume cleared of particles bigger than 10 pm. Despite low abundances of bivalve veligers and appendicularians in Kiel Bight at the time of the experiment, we calculated that approximately 10 and 8.5 %, respectively, of the carbon ingested by total mesozooplankton was due to veliger and appendicularian grazing. The importance of bivalve veligers might be seen in their grazing on seston particles that escape predation by copepods and on the amount of energy that is therefore directed from the water column to the benthos when larvae settle.

https://www.int-res.com/articles/meps/199/m199p043.pdf

Frank Sommer

Papers

Pelagic food web configurations at different levels of nutrient richness and their implications for the ratio fish production:primary production

Cladocerans versus copepods: the cause of contrasting top–down controls on freshwater and marine phytoplankton

Complementary impact of copepods and cladocerans on phytoplankton

Daphnia versus copepod impact on summer phytoplankton: functional compensation at both trophic levels

Grazing by mesozooplankton from Kiel Bight, Baltic Sea, on different sized algae and natural seston size fractions