Integrating Ecosystem Engineering and Food Web Ecology: Testing the Effect of Biogenic Reefs on the Food Web of a Soft-Bottom Intertidal Area.
ABSTRACT: The potential of ecosystem engineers to modify the structure and dynamics of food webs has recently been hypothesised from a conceptual point of view. Empirical data on the integration of ecosystem engineers and food webs is however largely lacking. This paper investigates the hypothesised link based on a field sampling approach of intertidal biogenic aggregations created by the ecosystem engineer Lanice conchilega (Polychaeta, Terebellidae). The aggregations are known to have a considerable impact on the physical and biogeochemical characteristics of their environment and subsequently on the abundance and biomass of primary food sources and the macrofaunal (i.e. the macro-, hyper- and epibenthos) community. Therefore, we hypothesise that L. conchilega aggregations affect the structure, stability and isotopic niche of the consumer assemblage of a soft-bottom intertidal food web. Primary food sources and the bentho-pelagic consumer assemblage of a L. conchilega aggregation and a control area were sampled on two soft-bottom intertidal areas along the French coast and analysed for their stable isotopes. Despite the structural impacts of the ecosystem engineer on the associated macrofaunal community, the presence of L. conchilega aggregations only has a minor effect on the food web structure of soft-bottom intertidal areas. The isotopic niche width of the consumer communities of the L. conchilega aggregations and control areas are highly similar, implying that consumer taxa do not shift their diet when feeding in a L. conchilega aggregation. Besides, species packing and hence trophic redundancy were not affected, pointing to an unaltered stability of the food web in the presence of L. conchilega.
Project description:Detrital subsidies from marine macrophytes are prevalent in temperate estuaries, and their role in structuring benthic macrofaunal communities is well documented, but the resulting impact on ecosystem function is not understood. We conducted a field experiment to test the effects of detrital decay on soft-sediment primary production, community metabolism and nutrient regeneration (measures of ecosystem function). Twenty four (2 m(2)) plots were established on an intertidal sandflat, to which we added 0 or 220 g DW m(-2) of detritus from either mangroves (Avicennia marina), seagrass (Zostera muelleri), or kelp (Ecklonia radiata) (n = 6 plots per treatment). Then, after 4, 17 and 46 d we measured ecosystem function, macrofaunal community structure and sediment properties. We hypothesized that (1) detrital decay would stimulate benthic primary production either by supplying nutrients to the benthic macrophytes, or by altering the macrofaunal community; and (2) ecosystem responses would depend on the stage and rate of macrophyte decay (a function of source). Avicennia detritus decayed the slowest with a half-life (t50) of 46 d, while Zostera and Ecklonia had t50 values of 28 and 2.6 d, respectively. However, ecosystem responses were not related to these differences. Instead, we found transient effects (up to 17 d) of Avicennia and Ecklonia detritus on benthic primary production, where initially (4 d) these detrital sources suppressed primary production, but after 17 d, primary production was stimulated in Avicennia plots relative to controls. Other ecosystem function response variables and the macrofaunal community composition were not altered by the addition of detritus, but did vary with time. By sampling ecosystem function temporally, we were able to capture the in situ transient effects of detrital subsidies on important benthic ecosystem functions.
Project description:The escalating spread of invasive species increases the risk of disrupting the pathways of energy flow through native ecosystems, modify the relative importance of resource ('bottom-up') and consumer ('top-down') control in food webs and thereby govern biomass production at different trophic levels. The current lack of understanding of interaction cascades triggered by non-indigenous species underscores the need for more basic exploratory research to assess the degree to which novel species regulate bottom-up and/or top down control. Novel predators are expected to produce the strongest effects by decimating consumers, and leading to the blooms of primary producers. Here we show how the arrival of the invasive crab Rhithropanopeus harrisii into the Baltic Sea - a bottom-up controlled ecosystem where no equivalent predators ever existed - appeared to trigger not only strong top-down control resulting in a decline in richness and biomass of benthic invertebrates, but also an increase in pelagic nutrients and phytoplankton biomass. Thus, the addition of a novel interaction - crab predation - to an ecosystem has a potential to reduce the relative importance of bottom-up regulation, relax benthic-pelagic coupling and reallocate large amounts of nutrients from benthic to pelagic processes, resulting in a regime shift to a degraded ecosystem state.
Project description:Consumer effects on prey are well known for cascading through food webs and producing dramatic top-down effects on community structure and ecosystem function. Bottom-up effects of prey (primary producer) biodiversity are also well known. However, the role of consumer diversity in affecting community structure or ecosystem function is not well understood. Here, we show that herbivore species richness can be critical for maintaining the structure and function of coral reefs. In two experiments over 2 years, we constructed large cages enclosing single herbivore species, equal densities of mixed species of herbivores, or excluding herbivores and assessed effects on both seaweeds and corals. When compared with single-herbivore treatments, mixed-herbivore treatments lowered macroalgal abundance by 54-76%, enhanced cover of crustose coralline algae (preferred recruitment sites for corals) by 52-64%, increased coral cover by 22%, and prevented coral mortality. Complementary feeding by herbivorous fishes drove the herbivore richness effects, because macroalgae were unable to effectively deter fishes with different feeding strategies. Maintaining herbivore species richness appears critical for preserving coral reefs, because complementary feeding by diverse herbivores produces positive, but indirect, effects on corals, the foundation species for the ecosystem.
Project description:Understanding the fundamental laws that govern complex food web networks over large ecosystems presents high costs and oftentimes unsurmountable logistical challenges. This way, it is crucial to find smaller systems that can be used as proxy food webs. Intertidal rock pool environments harbour particularly high biodiversity over small areas. This study aimed to analyse their food web networks to investigate their potential as proxies of larger ecosystems for food web networks research. Highly resolved food webs were compiled for 116 intertidal rock pools from cold, temperate, subtropical and tropical regions, to ensure a wide representation of environmental variability. The network properties of these food webs were compared to that of estuaries, lakes and rivers, as well as marine and terrestrial ecosystems (46 previously published complex food webs). The intertidal rock pool food webs analysed presented properties that were in the same range as the previously published food webs. The niche model predictive success was remarkably high (73-88%) and similar to that previously found for much larger marine and terrestrial food webs. By using a large-scale sampling effort covering 116 intertidal rock pools in several biogeographic regions, this study showed, for the first time, that intertidal rock pools encompass food webs that share fundamental organizational characteristics with food webs from markedly different, larger, open and abiotically stable ecosystems. As small, self-contained habitats, intertidal rock pools are particularly tractable systems and therefore a large number of food webs can be examined with relatively low sampling effort. This study shows, for the first time that they can be useful models for the understanding of universal processes that regulate the complex network organization of food webs, which are harder or impossible to investigate in larger, open ecosystems, due to high costs and logistical difficulties.
Project description:Widespread overharvesting of top consumers of the world's ecosystems has "skewed" food webs, in terms of biomass and species richness, towards a generally greater domination at lower trophic levels. This skewing is exacerbated in locations where exotic species are predominantly low-trophic level consumers such as benthic macrophytes, detritivores, and filter feeders. However, in some systems where numerous exotic predators have been added, sometimes purposefully as in many freshwater systems, food webs are skewed in the opposite direction toward consumer dominance. Little is known about how such modifications to food web topology, e.g., changes in the ratio of predator to prey species richness, affect ecosystem functioning. We experimentally measured the effects of trophic skew on production in an estuarine food web by manipulating ratios of species richness across three trophic levels in experimental mesocosms. After 24 days, increasing macroalgal richness promoted both plant biomass and grazer abundance, although the positive effect on plant biomass disappeared in the presence of grazers. The strongest trophic cascade on the experimentally stocked macroalgae emerged in communities with a greater ratio of prey to predator richness (bottom-rich food webs), while stronger cascades on the accumulation of naturally colonizing algae (primarily microalgae with some early successional macroalgae that recruited and grew in the mesocosms) generally emerged in communities with greater predator to prey richness (the more top-rich food webs). These results suggest that trophic skewing of species richness and overall changes in food web topology can influence marine community structure and food web dynamics in complex ways, emphasizing the need for multitrophic approaches to understand the consequences of marine extinctions and invasions.
Project description:Impacts of invasive species are context dependent and linked to the ecosystem they occur within. To broaden the understanding of the impact of a globally widespread invasive oyster, Crassostrea (Magallana) gigas, intertidal surveys were carried out at 15 different sites in Europe. The impact of C. gigas on macro- (taxa surrounding oyster >?1 cm) and epifaunal (taxa on oyster <?1 cm) benthic communities and ? and ?-diversity was assessed and compared to those associated with native ecosystem engineers, including the flat oyster Ostrea edulis. Whilst the effect of C. gigas on benthic community structures was dependent on habitat type, epifaunal communities associated with low densities of O. edulis and C. gigas did not differ and changes in benthic assemblage structure owing to the abundance of C. gigas were therefore attributed to the presence of oyster shells. Macrofaunal ?-diversity increased with C. gigas cover in muddy habitats, while epifaunal ?-diversity decreased at greater oyster densities. Macrofaunal ?-diversity was greatest at low densities of C. gigas; however, it did not differ between samples without and increased densities of oysters. In contrast, epifaunal ?-diversity decreased with increasing oyster cover. Different environmental contexts enabled more independent predictions of the effect of C. gigas on native communities. These were found to be low and more importantly not differing from O. edulis. This indicates that, at low densities, C. gigas may be functionally equivalent to the declining native oyster in terms of biodiversity facilitation and aid in re-establishing benthic communities on shores where O. edulis has become extinct.
Project description:Quantitative description of food webs provides fundamental information for the understanding of population, community, and ecosystem dynamics. Recently, stable isotope mixing models have been widely used to quantify dietary proportions of different food resources to a focal consumer. Here we propose a novel mixing model (IsoWeb) that estimates diet proportions of all consumers in a food web based on stable isotope information. IsoWeb requires a topological description of a food web, and stable isotope signatures of all consumers and resources in the web. A merit of IsoWeb is that it takes into account variation in trophic enrichment factors among different consumer-resource links. Sensitivity analysis using realistic hypothetical food webs suggests that IsoWeb is applicable to a wide variety of food webs differing in the number of species, connectance, sample size, and data variability. Sensitivity analysis based on real topological webs showed that IsoWeb can allow for a certain level of topological uncertainty in target food webs, including erroneously assuming false links, omission of existent links and species, and trophic aggregation into trophospecies. Moreover, using an illustrative application to a real food web, we demonstrated that IsoWeb can compare the plausibility of different candidate topologies for a focal web. These results suggest that IsoWeb provides a powerful tool to analyze food-web structure from stable isotope data. We provide R and BUGS codes to aid efficient applications of IsoWeb.
Project description:The framework of ecological stoichiometry was developed primarily within the context of "green" autotroph-based food webs. While stoichiometric principles also apply in "brown" detritus-based systems, these systems have been historically understudied and differ from green ones in several important aspects including carbon (C) quality and the nutrient [nitrogen (N) and phosphorus (P)] contents of food resources for consumers. In this paper, we review work over the last decade that has advanced the application of ecological stoichiometry from green to brown food webs, focusing on freshwater ecosystems. We first review three focal areas where green and brown food webs differ: (1) bottom-up controls by light and nutrient availability, (2) stoichiometric constraints on consumer growth and nutritional regulation, and (3) patterns in consumer-driven nutrient dynamics. Our review highlights the need for further study of how light and nutrient availability affect autotroph-heterotroph interactions on detritus and the subsequent effects on consumer feeding and growth. To complement this conceptual review, we formally quantified differences in stoichiometric principles between green and brown food webs using a meta-analysis across feeding studies of freshwater benthic invertebrates. From 257 datasets collated across 46 publications and several unpublished studies, we compared effect sizes (Pearson's r) of resource N:C and P:C on growth, consumption, excretion, and egestion between herbivorous and detritivorous consumers. The meta-analysis revealed that both herbivore and detritivore growth are limited by resource N:C and P:C contents, but effect sizes only among detritivores were significantly above zero. Consumption effect sizes were negative among herbivores but positive for detritivores in the case of both N:C and P:C, indicating distinct compensatory feeding responses across resource stoichiometry gradients. Herbivore P excretion rates responded significantly positively to resource P:C, whereas detritivore N and P excretion did not respond; detritivore N and P egestion responded positively to resource N:C and P:C, respectively. Our meta-analysis highlights resource N and P contents as broadly limiting in brown and green benthic food webs, but indicates contrasting mechanisms of limitation owing to differing consumer regulation. We suggest that green and brown food webs share fundamental stoichiometric principles, while identifying specific differences toward applying ecological stoichiometry across ecosystems.
Project description:A central issue in ecology is understanding how complex and biodiverse food webs persist in the face of disturbance, and which structural properties affect disturbance propagation among species. However, our comprehension of assemblage mechanisms and disturbance propagation in food webs is limited by the multitude of stressors affecting ecosystems, impairing ecosystem management. By analysing directional food web components connecting species along food chains, we show that increasing species richness and constant feeding linkage density promote the establishment of predictable food web structures, in which the proportion of species co-present in one or more food chains is lower than what would be expected by chance. This reduces the intrinsic vulnerability of real food webs to disturbance propagation in comparison to random webs, and suggests that biodiversity conservation efforts should also increase the potential of ecological communities to buffer top-down and bottom-up disturbance in ecosystems. The food web patterns observed here have not been noticed before, and could also be explored in non-natural networks.
Project description:Agricultural intensification (AI) is currently a major driver of biodiversity loss and related ecosystem functioning decline. However, spatio-temporal changes in community structure induced by AI, and their relation to ecosystem functioning, remain largely unexplored. Here, we analysed 16 quantitative cereal aphid-parasitoid and parasitoid-hyperparasitoid food webs, replicated four times during the season, under contrasting AI regimes (organic farming in complex landscapes vs. conventional farming in simple landscapes). High AI increased food web complexity but also temporal variability in aphid-parasitoid food webs and in the dominant parasitoid species identity. Enhanced complexity and variability appeared to be controlled bottom-up by changes in aphid dominance structure and evenness. Contrary to the common expectations of positive biodiversity-ecosystem functioning relationships, community complexity (food-web complexity, species richness and evenness) was negatively related to primary parasitism rates. However, this relationship was positive for secondary parasitoids. Despite differences in community structures among different trophic levels, ecosystem services (parasitism rates) and disservices (aphid abundances and hyperparasitism rates) were always higher in fields with low AI. Hence, community structure and ecosystem functioning appear to be differently influenced by AI, and change differently over time and among trophic levels. In conclusion, intensified agriculture can support diverse albeit highly variable parasitoid-host communities, but ecosystem functioning might not be easy to predict from observed changes in community structure and composition.