Effects of food diversity on diatom selection by harpacticoid copepods

The diversity of species assemblages which occupy a basal position in the trophic pyramid (typically unicellular algae in aquatic environments) is known to influence the interaction with organisms of higher trophic levels. A laboratory feeding experiment was conducted with cultures of three benthic diatom species (Navicula phyllepta, Grammatophora marina and Cylindrotheca closterium) as primary producers and three harpacticoid copepod species (Harpacticus obscurus, Paramphiascella fulvofasciata and Tigriopus brevicornis) as grazers to evaluate the effects of food diversity (and concomitant food availability) on grazing selectivity. This kind of selectivity experiments is singular for benthic harpacticoid copepods as so far, information on food selection of harpacticoids is scarce.Uptake of a unispecific food source by a single copepod species decreased as food diversity (and concomitant overall food concentration) increased. All three consumers reacted similarly to changing food diversity, but exhibited strong species-specific responses to food identity i.e. which diatom was added was crucial. Irrespective of level of food diversity, H. obscurus took up high amounts of G. marina, whereas both P. fulvofasciata and T. brevicornis preferred C. closterium when given the choice between different diatoms. As for zooplanktonic taxa, this experiment showed that in lower benthic marine food webs both prey organisms (primary producers) and grazers play a very specific role. Diversity of food and its identity are of critical importance at the base of the trophic pyramid, influencing trophic transfer from primary producers over grazers to higher trophic levels.     

A method for rearing sand-dwelling harpacticoid copepods in experimental conditions

A floating culture tube was developed for rearing the sand-dwelling harpacticoid Asellopsis intermedia (T. Scott) in experimental conditions. The culture tube was adapted from a polythene specimen tube with attached lid, the base of the tube and the centre of the lid being replaced by nylon mesh of 75 μm aperture. A thin layer of sand placed on the basal net provided ballast for the culture tube which then floated vertically at the surface of the water. Cultured harpacticoids remained in a healthy condition for several months, apparently grazing on the bacterial epigrowth of the sand grains.     

The role of phototrophic sulfur bacteria as food for meiobenthic harpacticoid copepods inhabiting eutrophic coastal lagoons

Laboratory experiments were undertaken using Amonardia normani and Schizopera cf. compacta, two meiobenthic harpacticoid copepods commonly found in coastal lagoons. The first experiments were designed to determine if the phototrophic sulfur bacteria Chromatium gracile can be ingested by these copepods and at what concentrations. Egestion rate was used as an index of feeding rate. The response of the egestion rate, expressed in numbers of faecal pellets produced by copepod per day, as a function of bacterial concentration followed the functional model. A. normani attained constant feeding rates from the bacterial concentration of 1 × 10⁷ cells ml^–1 (5 µg C ml^–1) onwards, S. cf. compacta attained constant feeding rates from 2.6 × 10⁷ cells ml^–1 (13 µg C ml^–1) onwards. The faecal pellet volume changed significantly (p<0.05) between food concentrations for A. normani but not for S. cf. compacta (p>0.05). In order to investigate the effect of the phototrophic bacterial diet on the population dynamics of A. normani three groups of nauplii were maintained at 2 × 10⁷ cells ml^–1 and observed every day. The mortality of these nauplii was very high compared to those maintained on a diatom diet (Nitzschia constricta); only in one of the groups did some copepodites develop but no adults were ever observed. Adults fed on bacteria did not have different (p>0.05) survival rates compared to those fed on diatoms, nevertheless, the number of nauplii produced was significantly less (p<0.05) on the bacterial diet. These results lead us to suggest that although the phototrophic sulfur bacteria (Chromatium gracile) can be ingested by both copepod species it cannot sustain the full development of the A. normani population. Thus, a bloom of phototrophic sulfur bacteria does not seem to be a favourable situation for opportunistic benthic copepods to colonize eutrophic coastal lagoons after a dystrophic crisis.     

Preliminary observations on the metabolic and reproductive strategies of harpacticoid copepods from an intertidal sandflat

The metabolic responses to temperature changes and the reproductive strategies of four species of harpacticoid copepod are examined in relation to their depth distribution, susceptibility to predation and assumptions about food supply. Two species, one of which is heavily preyed upon, are confined to the surface layers and two species occur deeper in the sediment through most of the year. These latter species appear to have adopted different metabolic strategies in response to lowered food supply but both have a lower Q₁₀ than surface dwelling species. In addition the surface species have the ability to regulate their respiratory demands at times of high, short-term, temperature fluctuations. Preliminary data suggests that generally, the surface species have shorter development times than those dwelling deeper in the sediment but until more information is available on the energetic costs of reproduction it cannot be shown that deeper dwelling species are more conservative in their reproductive strategies.     

How habitat-modifying organisms structure the food web of two coastal ecosystems

The diversity and structure of ecosystems has been found to depend both on trophic interactions in food webs and on other species interactions such as habitat modification and mutualism that form non-trophic interaction networks. However, quantification of the dependencies between these two main interaction networks has remained elusive. In this study, we assessed how habitat-modifying organisms affect basic food web properties by conducting in-depth empirical investigations of two ecosystems: North American temperate fringing marshes and West African tropical seagrass meadows. Results reveal that habitat-modifying species, through non-trophic facilitation rather than their trophic role, enhance species richness across multiple trophic levels, increase the number of interactions per species (link density), but decrease the realized fraction of all possible links within the food web (connectance). Compared to the trophic role of the most highly connected species, we found this non-trophic effects to be more important for species richness and of more or similar importance for link density and connectance. Our findings demonstrate that food webs can be fundamentally shaped by interactions outside the trophic network, yet intrinsic to the species participating in it. Better integration of non-trophic interactions in food web analyses may therefore strongly contribute to their explanatory and predictive capacity.     

Cascading predation effects of Daphnia and copepods on microbial food web components

1. We performed a mesocosm experiment to investigate the structuring and cascading effects of two predominant crustacean mesozooplankton groups on microbial food web components. The natural summer plankton community of a mesotrophic lake was exposed to density gradients of Daphnia and copepods. Regression analysis was used to reveal top–down impacts of mesozooplankton on protists and bacteria after days 9 and 15. 2. Selective grazing by copepods caused a clear trophic cascade via ciliates to nanoplankton. Medium‐sized (20–40 μm) ciliates (mainly Oligotrichida) were particularly negatively affected by copepods whereas nanociliates (mainly Prostomatida) became more abundant. Phototrophic and heterotrophic nanoflagellates increased significantly with increasing copepod biomass, which we interpret as an indirect response to reduced grazing pressure from the medium‐sized ciliates. 3. In Daphnia‐treatments, ciliates of all size classes as well as nanoflagellates were reduced directly but the overall predation effect became most strongly visible after 15 days at higher Daphnia biomass. 4. The response of bacterioplankton involved only modest changes in bacterial biomass and cell‐size distribution along the zooplankton gradients. Increasing zooplankton biomass resulted either in a reduction (with Daphnia) or in an increase (with copepods) of bacterial biovolume, activity and production. Patterns of bacterial diversity, as measured by polymerase chain reaction–denaturing gradient gel electrophoresis (PCR–DGGE), showed no distinct grouping after 9 days, whereas a clear treatment‐coupled similarity clustering occurred after 15 days. 5. The experiment demonstrated that zooplankton‐mediated predatory interactions cascade down to the bacterial level, but also revealed that changes occurred rather slowly in this summer plankton community and were most pronounced with respect to bacterial activity and composition.     

Some experimental approaches to the study of nutrition and parasitic infection

Two experimental models to examine the relationship between host nutrition and parasitic infection are considered. Moniliformis dubius, an acanthocephalan parasite of the laboratory rat, is dependent on the type and amount of host dietary carbohydrate for growth and reproduction. Starch, glucose, fructose, and mannose in host diets support parasite growth and development, but much lower dietary amounts of starch, fructose, and mannose are needed compared to free glucose. The pig-Ascaris suum relationship is considered as a model for human ascariasis. Experimental Ascaris infections in pigs resulted in reduced growth, reduced food intake, lowered biological value of dietary protein, and reduced intestinal lactase activity and lactose tolerance. These results are similar in some respects to reported effects of Ascaris infection in humans.     

Tracing carbon flow in an arctic marine food web using fatty acid-stable isotope analysis

Global warming and the loss of sea ice threaten to alter patterns of productivity in arctic marine ecosystems because of a likely decline in primary productivity by sea ice algae. Estimates of the contribution of ice algae to total primary production range widely, from just 3 to >50%, and the importance of ice algae to higher trophic levels remains unknown. To help answer this question, we investigated a novel approach to food web studies by combining the two established methods of stable isotope analysis and fatty acid (FA) analysis--we determined the C isotopic composition of individual diatom FA and traced these biomarkers in consumers. Samples were collected near Barrow, Alaska and included ice algae, pelagic phytoplankton, zooplankton, fish, seabirds, pinnipeds and cetaceans. Ice algae and pelagic phytoplankton had distinctive overall FA signatures and clear differences in delta(13)C for two specific diatom FA biomarkers: 16:4n-1 (-24.0+/-2.4 and -30.7+/-0.8 per thousand, respectively) and 20:5n-3 (-18.3+/-2.0 and -26.9+/-0.7 per thousand, respectively). Nearly all delta(13)C values of these two FA in consumers fell between the two stable isotopic end members. A mass balance equation indicated that FA material derived from ice algae, compared to pelagic diatoms, averaged 71% (44-107%) in consumers based on delta(13)C values of 16:4n-1, but only 24% (0-61%) based on 20:5n-3. Our estimates derived from 16:4n-1, which is produced only by diatoms, probably best represented the contribution of ice algae relative to pelagic diatoms. However, many types of algae produce 20:5n-3, so the lower value derived from it likely represented a more realistic estimate of the proportion of ice algae material relative to all other types of phytoplankton. These preliminary results demonstrate the potential value of compound-specific isotope analysis of marine lipids to trace C flow through marine food webs and provide a foundation for future work.     

Influence of pollution on the harpacticoid copepods of two North Sea estuaries

Seasonal monitoring of the meiobenthos in the Dutch estuaries revealed an anomaly in density and diversity of harpacticoid copepods in the Westerschelde.Another Dutch estuary, the Eems Dollard, has comparable hydrodynamical, physical and sedimentological, characteristics and a similar fauna, but even in the severely organically polluted oligohaline mudflats of this estuary, annual average density and diversity of endo-epibenthic harpacticoid communities are higher than at similar less enriched meso- to polyhaline biotopes of the Westerschelde.Besides the concentrations of inorganic pollutants, such as phosphates and nitrates, the concentrations of pesticides, cyanide, detergents, phenoles, oils, polychlorobenzenes, polycyclic aromatic hydrocarbons and heavy metals were compared in both estuaries and compared to suggested permissive levels when available. From this it appears that these pollutants are present in the Westerschelde either in too low concentrations to be considered dangerous or at concentrations comparable to those occurring in the Eems Dollard, except for heavy metals.The load of a.o. Zn, Cu and Pb is distinctly and persistently higher in sediments and suspensions of the Westerschelde than in the Eems Dollard and copper is continuously present in a concentration at which, according to bioassays, egg production and larval development of planktonic copepods are severely affected. The remarkable scarcity of harpacticoid life on nutrient rich mudflats of the Westerschelde is thus probably due to heavy metal pollution.Since no other hardbodied meio- and macrobenthic taxa nor the plankton of this estuary show such a marked impoverishment, benthic harpacticoids prove to be suitable as indicators for the first stages of ecosystem-breakdown in estuarine and coastal zones polluted by trace-metals.     

Adult-infant food transfer in common marmosets: an experimental study

Infant marmosets and tamarins obtain solid food items from adults during and after the time of weaning. In addition to providing nutrients, food transfers may provide infants with the opportunity to learn about diet. The aim of this study was to investigate patterns of begging and food transfer in captive groups of common marmosets during tests with 1) palatable novel food, 2) unpalatable novel food, and 3) familiar food. Infants may gain an opportunity to learn about diet breadth by preferentially begging for novel rather than familiar items, while adults may facilitate infant learning by differing in their willingness to transfer food depending upon the food's novelty or palatability. In this study, infants exhibited more interest and begging with novel food items than with familiar ones. However, food transfers were not more likely to occur with novel foods than with familiar ones, and in fact adults were more likely to refuse infant begging attempts with novel foods. The palatability of the food also did not influence the rates of refusal and transfer: adults transferred palatable and unpalatable novel foods with similar frequencies. These results suggest that infant marmosets gain an opportunity to learn about diet breadth by begging for novel foods, but adults do not preferentially transfer novel or palatable food items.     

Fatty acid profiles of algae mark the development and composition of harpacticoid copepods

1. The value of algal fatty acids (FA) as diet biomarkers for benthic harpacticoid copepods was investigated. A high proportion of 18:1ω9 and 18:2ω6 FA was observed in the lipid reserve fraction of copepods fed with cyanobacteria. In contrast, a high proportion of 16:1ω7 and ω3 FA (including eicosapentaenoic) was present in the lipid reserve fraction of copepods grown on diatoms. 2. Copepods that were grown on cyanobacteria showed reduced survival and took 26% more time to develop from the first copepodid stage to adult than copepods that were grown on diatoms. Copepods feeding on the cyanobacteria showed reduced FA content when compared with animals fed with diatoms. This reduction in FA content was more pronounced in the apolar lipid fraction (mainly reserve lipids) than in the polar (mainly structural) lipid fraction. 3. The FA profiles of algae were used to calculate a function discriminating between diatoms and cyanobacteria. This function was applied to the FA profiles in the reserve lipid fraction of copepods and correctly classified copepod diet. 16:1ω7, 18:2ω6 and 20:5ω3 were the most important FA in the discriminant function. The suitability of this chemometric method to infer copepod diet was further tested by using algal class FA data from literature to derive the discriminant functions. The correct classification of the diet when the functions were applied to FA composition of the copepod reserve lipids suggests that this method may be employed in trophic web studies. 18:3ω3, 18:1ω9 and 16:1ω7 were the most important FA in the functions discriminating diatoms, cyanobacteria and green algae. The identification and quantification of the whole suit of 16:1ω7, 18:1ω9, 18:2ω6, 18:3ω3 and 20:5ω3 in trophic web studies is therefore of paramount importance to infer diet origin of aquatic herbivores. 4. The FA profile of copepod polar lipids did not reflect that of the diet. The presence of long chain polyunsaturated FAs in the polar lipid fraction of copepods feeding on the cyanobacterium suggests that C18 FAs from the diet may be elongated and desaturated by the copepod. The ability to elongate and desaturated FAs may reduce the importance of some FAs as diet biomarkers while it may turn the copepods into valuable trophic intermediaries in transferring organic matter from microorganisms to higher trophic levels.     

Assessing the trophic impacts of marine mammals: From metabolism to food web indices

Marine mammals are an important part of ecosystems, and their trophic role and potential impact have been increasingly studied. One key question is how these large animals interact with fisheries or compete for similar resources. Consequently, some models once used only for fisheries management are now including pinnipeds and cetaceans. However, fish and marine mammals do not share the same ecology and bioenergetics, and complex ecosystem models may not be the best way to assess the impact of pinnipeds or cetaceans in food webs. Indeed, simpler methods based on thermodynamics might give us reasonable answers with limited amounts of data. Here, we present an assessment of two different approaches to assess the trophic role of marine mammals in the northern Gulf of St. Lawrence (Canada): mixed trophic impacts (MTI) based on ecosystem modeling and surface index (SI) impact based on bioenergetics. Our results show that while modeling represents a good way of getting a holistic view of the role of marine mammals in ecosystems, trophic impact estimates based on fundamental thermodynamics principles can also give us answers requiring less data. The body surface area approach presented here might provide a practical tool for ecologists, who are not necessarily ecosystem modelers, to study this issue.     

Novocriniidae, a new family of harpacticoid copepods from anchihaline caves in Belize

Huys, R. & Iliffe, T. M. 1998. Novocriniidae, a new family of harpacticoid copepods from anchihaline caves in Belize.
A new family of fusiform, semi-planktonic harpacticoid copepods is described from two anchihaline caves, Caye Chapel Cave and Columbus Caye Blue Hole, on the Belize Barrier Reef. The Novocriniidae fam. n. is proposed to accommodate Novocrinia trifida gen. et sp. n. and is placed in the tisbidimorph complex of families (Tisboidea) by virtue of its maxilliped morphology. The new genus shares several characters with other tisbidimorph families but cannot be placed in any of them without an unnatural extension of their diagnoses. The Novocriniidae display a suite of plesiomorphic character states in most appendages from the antennules to the PS, indicating their basal position on the tisbidimorph phylogenetic tree. They exhibit a unique sexually dimorphic modification of the antennae, involving the transformation of two setal elements into filamentous sensory tufts. The unusual presence of supernumerary elements on the male PS baseoendopod is briefly discussed as well as the presence of a genuine but incipient oral cone derived from the labrum and medially fused paragnaths.     

Relative impacts of copepods, cladocerans and nutrients on the microbial food web of a mesotrophic lake

Calanoid copepods, rather than cladocerans, frequently dominatethe zooplankton of lakes in New Zealand. The potential consequencesof this domination for the microbial community of mesotrophicLake Mahinerangi, New Zealand, were determined by field experimentsin which Boeckella and Daphnia were added to in situ enclosuresin the presence and absence of added nutrients. Boeckella hamataat ambient densities (2 and 81^–1) rapidly and severelysuppressed ciliate population growth over 4 days, even whenmicrobial growth was enhanced by added nutrients, but effectsof copepods on other components of the microbial community (bacteria,photosynthetic picoplankton, heterotrophic nano-flagellates,algae) were slight. In contrast, Daphnia carinata at the samedensities (but 3-fold higher biomasses per litre) had a relativelyweak effect on ciliates, suppressing ciliate abundance onlyafter 4 days at 8 Daphnia 1^–1 (330 µg 1^–1);this daphniid density also depressed abundances of large bacterialrods, some photosynthetic picoplankton and the dominant alga,Cyclotella. These results highlight the relative importanceof specific trophic linkages in a microbial food web; they alsosuggest that the dominance of Boeckella in many southern hemispherelakes may account for relatively low ciliate abundances in theselakes.     

Macroecological patterns and niche structure in a new marine food web

The integration of detailed information on feeding interactions with measures of abundance and body mass of individuals provides a powerful platform for understanding ecosystem organisation. Metabolism and, by proxy, body mass constrain the flux, turnover and storage of energy and biomass in food webs. Here, we present the first food web data for Lough Hyne, a species rich Irish Sea Lough. Through the application of individual-and size-based analysis of the abundance-body mass relationship, we tested predictions derived from the metabolic theory of ecology. We found that individual body mass constrained the flux of biomass and determined its distribution within the food web. Body mass was also an important determinant of diet width and niche overlap, and predator diets were nested hierarchically, such that diet width increased with body mass. We applied a novel measure of predator-prey biomass flux which revealed that most interactions in Lough Hyne were weak, whereas only a few were strong. Further, the patterning of interaction strength between prey sharing a common predator revealed that strong interactions were nearly always coupled with weak interactions. Our findings illustrate that important insights into the organisation, structure and stability of ecosystems can be achieved through the theoretical exploration of detailed empirical data.     

Planktonic food web in marine mesocosms in the Eastern Mediterranean: bottom-up or top-down regulation?

A mesocosm experiment was conducted in order to studythe structure of the planktonic food web. The dynamicsof pico-, nano- and microplankton populations werefollowed during 40 days in four large (40 m³)enclosures. In three tanks a gradient of addednutrients (nitrogen and phosphorus) was applied, whilea fourth tank was used as a control. On day 14, thetop predator (sea bream Sparus aurata larvae)was introduced into the tanks and part of the watercolumn in each tank was isolated in a plastic bagwithout fish larvae, to act as a control forpredation. Physical parameters, chlorophyll aand nutrient concentrations, as well as planktonconcentrations were monitored. A diatom bloom wasobserved in all four tanks, in the first phase endingwith silicate depletion. Flagellate and dinoflagellateabundance subsequently increased, these organismsbeing limited by zooplankton grazing. The zooplanktonpopulations were controlled by both resources (mostlyflagellates) and predation (by fish larvae) asindicated by the results of the control experiments. This revised version was published online in July 2006 with corrections to the Cover Date.     

Regime shifts in marine communities: a complex systems perspective on food web dynamics

Species composition and habitats are changing at unprecedented rates in the world''s oceans, potentially causing entire food webs to shift to structurally and functionally different regimes. Despite the severity of these regime shifts, elucidating the precise nature of their underlying processes has remained difficult. We address this challenge with a new analytic approach to detect and assess the relative strength of different driving processes in food webs. Our study draws on complexity theory, and integrates the network-centric exponential random graph modelling (ERGM) framework developed within the social sciences with community ecology. In contrast to previous research, this approach makes clear assumptions of direction of causality and accommodates a dynamic perspective on the emergence of food webs. We apply our approach to analysing food webs of the Baltic Sea before and after a previously reported regime shift. Our results show that the dominant food web processes have remained largely the same, although we detect changes in their magnitudes. The results indicate that the reported regime shift may not be a system-wide shift, but instead involve a limited number of species. Our study emphasizes the importance of community-wide analysis on marine regime shifts and introduces a novel approach to examine food webs.     

The food web in the lower part of the Seine estuary: a synthesis of existing knowledge

The Seine estuary illustrates the alterations to estuaries due to human activities heavy releases of pollutants of various origins and significant morphological changes beginning in the middle of the 19th century. The intertidal mudflat surface has been seriously reduced (< 30km²) since the channels of the Seine River came under management. While the role of the Seine estuary in the dynamics of the eastern English Channel ecosystem is recognized as important, the biological characteristics of the estuary remained relatively unknown until the 1990s. Biological diversity was progressively impoverished from the polyhaline zone to the oligohaline zone. In spite of a heavily contaminated environment, the macrobenthic and planktonic fauna of the Seine estuary remains similar to those of other northeastern Atlantic estuaries. The fauna exhibit clear contrasts between areas with very high abundance and others with very low abundance. The pelagic fauna, especially the copepod Eurytemora affinisand the shrimp Palaemon longirostris, are more abundant in the Seine estuary than in other estuaries. Diversified and abundant, Abra alba-Pectinaria koreni and Macoma balthica benthic communities occur, respectively, in the outer and inner parts of the estuary. In subtidal flats, benthic fauna is especially poor in terms of specific richness, abundance and biomass. Paradoxically, considering the high abundance of prey, fish are particularly scarce. Two food webs have been identified. In the oligohaline zone, where turbidity is maximum, the food web is exclusively planktonic, due to dredging that prevented benthic fauna from settling. In the polyhaline zone, fish populations that feed particularly on benthic fauna benefit from low turbidity and high oxygen concentrations. So, in spite of heavy organic and metallic contamination and human activities, the Seine estuary remains a highly productive ecosystem, which provides a nursery for marine fish and feeding grounds for migratory birds. A global management plan appears to be necessary in order to guarantee that the Seine estuary continues to function as it currently does.