Towards a multi‐trophic extension of metacommunity ecology

Metacommunity theory provides an understanding of how spatial processes determine the structure and function of communities at local and regional scales. Although metacommunity theory has considered trophic dynamics in the past, it has been performed idiosyncratically with a wide selection of possible dynamics. Trophic metacommunity theory needs a synthesis of a few influential axis to simplify future predictions and tests. We propose an extension of metacommunity ecology that addresses these shortcomings by incorporating variability among trophic levels in ‘spatial use properties’. We define ‘spatial use properties’ as a set of traits (dispersal, migration, foraging and spatial information processing) that set the spatial and temporal scales of organismal movement, and thus scales of interspecific interactions. Progress towards a synthetic predictive framework can be made by (1) documenting patterns of spatial use properties in natural food webs and (2) using theory and experiments to test how trophic structure in spatial use properties affects metacommunity dynamics.     

Environmental heterogeneity mediates a cross‐ecosystem trophic cascade

1. The flow of energy and nutrients across ecosystem boundaries can have significant community‐wide effects, but the role of productivity of the recipient habitat in mediating these effects remains unclear. This is especially true when organisms moving across ecosystem boundaries serve simultaneously as predators and prey. 2. In this study, the effects of odonates, primarily Enallagma civile (Hagen), on a salt marsh system were examined. Cages were used to exclude odonate predators, but not other arthropods, from experimental plots of the sea oxeye daisy, Borrichia frutescens (L.) in high and low productivity areas. The effects were assessed on the in situ arthropod community and the host plant. 3. There were strong direct effects of predation on the herbivores Pissonotus quadripustulatus Van Duzee and Asphondylia borrichiae Rossi and Strong, with higher densities where damselflies were excluded. Damselflies also served as prey for web‐building spiders. This resulted in lower spider densities inside cages, and a positive indirect effect on grasshopper densities. 4. Direct and indirect effects of odonates were greater in the high productivity area, resulting in a trophic cascade, with greater damage and reduced flowering and density of the host plant inside cages. 5. The results of this study support the subsidy hypothesis and show that theoretical models of trophic dynamics, which were developed to explain exchanges within ecosystems, may have predictive and explanatory value for exchanges across ecosystems as well.     

The effects of livestock grazing on biodiversity are multi‐trophic: a meta‐analysis

Anthropogenic disturbance has generated a significant loss of biodiversity worldwide and grazing by domestic herbivores is a contributing disturbance. Although the effects of grazing on plants are commonly explored, here we address the potential multi‐trophic effects on animal biodiversity (e.g. herbivores, pollinators and predators). We conducted a meta‐analysis on 109 independent studies that tested the response of animals or plants to livestock grazing relative to livestock excluded. Across all animals, livestock exclusion increased abundance and diversity, but these effects were greatest for trophic levels directly dependent on plants, such as herbivores and pollinators. Detritivores were the only trophic level whose abundance decreased with livestock exclusion. We also found that the number of years since livestock was excluded influenced the community and that the effects of grazer exclusion on animal diversity were strongest in temperate climates. These findings synthesise the effects of livestock grazing beyond plants and demonstrate the indirect impacts of livestock grazing on multiple trophic levels in the animal community. We identified the potentially long‐term impacts that livestock grazing can have on lower trophic levels and consequences for biological conservation. We also highlight the potentially inevitable cost to global biodiversity from livestock grazing that must be balanced against socio‐economic benefits.     

Spatial insurance in multi‐trophic metacommunities

Metacommunity theory suggests that dispersal is a key driver of diversity and ecosystem functioning in changing environments. The capacity of dispersal to mitigate effects of environmental change might vary among trophic groups, potentially resulting in changes in trophic interactions and food web structure. In a mesocosm experiment, we compared the compositional response of bacteria, phyto‐ and zooplankton to a factorial manipulation of acidification and dispersal. We found that the buffering capacity of dispersal varied among trophic groups: dispersal alleviated the negative effect of acidification on phytoplankton diversity mid‐experiment, but had no effect on the diversity of zooplankton and bacteria. Likewise, trophic groups differed in whether dispersal facilitated compositional change. Dispersal accelerated changes in phytoplankton composition under acidification, possibly mediated by changes in trophic interactions, but had no effect on the composition of zooplankton and bacteria. Overall, our results suggest that the potential for spatial insurance can vary among trophic groups.     

Fish trophic divergence along a lake productivity gradient revealed by historic patterns of invasion and eutrophication

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Carbon sources and trophic structure in a macrophyte‐dominated polyculture pond assessed by stable‐isotope analysis

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Tri‐trophic interactions: bridging species,communities and ecosystems

A vast body of research demonstrates that many ecological and evolutionary processes can only be understood from a tri‐trophic viewpoint, that is, one that moves beyond the pairwise interactions of neighbouring trophic levels to consider the emergent features of interactions among multiple trophic levels. Despite its unifying potential, tri‐trophic research has been fragmented, following two distinct paths. One has focused on the population biology and evolutionary ecology of simple food chains of interacting species. The other has focused on bottom‐up and top‐down controls over the distribution of biomass across trophic levels and other ecosystem‐level variables. Here, we propose pathways to bridge these two long‐standing perspectives. We argue that an expanded theory of tri‐trophic interactions (TTIs) can unify our understanding of biological processes across scales and levels of organisation, ranging from species evolution and pairwise interactions to community structure and ecosystem function. To do so requires addressing how community structure and ecosystem function arise as emergent properties of component TTIs, and, in turn, how species traits and TTIs are shaped by the ecosystem processes and the abiotic environment in which they are embedded. We conclude that novel insights will come from applying tri‐trophic theory systematically across all levels of biological organisation.     

Refining trophic dynamics through multi‐factor Bayesian mixing models: A case study of subterranean beetles

Food web dynamics are vital in shaping the functional ecology of ecosystems. However, trophic ecology is still in its infancy in groundwater ecosystems due to the cryptic nature of these environments. To unravel trophic interactions between subterranean biota, we applied an interdisciplinary Bayesian mixing model design (multi‐factor BMM) based on the integration of faunal C and N bulk tissue stable isotope data (δ¹³C and δ¹⁵N) with radiocarbon data (Δ¹⁴C), and prior information from metagenomic analyses. We further compared outcomes from multi‐factor BMM with a conventional isotope double proxy mixing model (SIA BMM), triple proxy (δ¹³C, δ¹⁵N, and Δ¹⁴C, multi‐proxy BMM), and double proxy combined with DNA prior information (SIA + DNA BMM) designs. Three species of subterranean beetles (Paroster macrosturtensis, Paroster mesosturtensis, and Paroster microsturtensis) and their main prey items Chiltoniidae amphipods (AM1: Scutachiltonia axfordi and AM2: Yilgarniella sturtensis), cyclopoids and harpacticoids from a calcrete in Western Australia were targeted. Diet estimations from stable isotope only models (SIA BMM) indicated homogeneous patterns with modest preferences for amphipods as prey items. Multi‐proxy BMM suggested increased—and species‐specific—predatory pressures on amphipods coupled with high rates of scavenging/predation on sister species. SIA + DNA BMM showed marked preferences for amphipods AM1 and AM2, and reduced interspecific scavenging/predation on Paroster species. Multi‐factorial BMM revealed the most precise estimations (lower overall SD and very marginal beetles' interspecific interactions), indicating consistent preferences for amphipods AM1 in all the beetles' diets. Incorporation of genetic priors allowed crucial refining of the feeding preferences, while integration of more expensive radiocarbon data as a third proxy (when combined with genetic data) produced more precise outcomes but close dietary reconstruction to that from SIA + DNA BMM. Further multidisciplinary modeling from other groundwater environments will help elucidate the potential behind these designs and bring light to the feeding ecology of one the most vital ecosystems worldwide.     

Combined stable‐isotope and fatty‐acid analyses demonstrate that large wood increases the autochthonous trophic base of a macroinvertebrate assemblage

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Management implications of the Macquarie Island trophic cascade revisited: a reply to Dowding et al. (2009)

1. The management of non-indigenous species is not without its complications. In Bergstrom et al. 's (2009) study, we demonstrated that feral cats Felis catus on sub-Antarctic Macquarie Island were exerting top-down control on the feral rabbit Oryctolagus cuniculus population, and that the eradication of the cats led to a substantial increase in rabbit numbers and an associated trophic cascade.
2. Dowding et al. (2009) claim our modelling was flawed for various reasons, but primarily that a reduction in the application of the rabbit control agent, Myxoma virus, coinciding with cat removal, was a major driver of rabbit population release.
3. We explore this proposition (as well as others) by examining rates of Myxoma viral release between 1991 and 2006 (with an attenuation factor for the years, 2003–2006) in association with presence/absence of cats against two estimates of rabbit population size. Myxoma viral release was a significant factor in the lower estimates of rabbit population, but the effect was small, and was not significant for higher rabbit population estimates. By contrast, the presence or absence of cats remained highly significant for both estimates.
4. Synthesis and applications. We re-affirm our position that top-down control of rabbit numbers by cats, prior to their eradication, was occurring on Macquarie Island. Nonetheless, we agree with Dowding et al. (2009) that systems with multiple invasive species represent complex situations that require careful scrutiny. Such scrutiny should occur in advance of, during, and following management interventions.     

Food web structure and biocontrol in a four-trophic level system across a landscape complexity gradient

Decline in landscape complexity owing to agricultural intensification may affect biodiversity, food web complexity and associated ecological processes such as biological control, but such relationships are poorly understood. Here, we analysed food webs of cereal aphids, their primary parasitoids and hyperparasitoids in 18 agricultural landscapes differing in structural complexity (42-93% arable land). Despite little variation in the richness of each trophic group, we found considerable changes in trophic link properties across the landscape complexity gradient. Unexpectedly, aphid-parasitoid food webs exhibited a lower complexity (lower linkage density, interaction diversity and generality) in structurally complex landscapes, which was related to the dominance of one aphid species in complex landscapes. Nevertheless, primary parasitism, as well as hyperparasitism, was higher in complex landscapes, with primary parasitism reaching levels for potentially successful biological control. In conclusion, landscape complexity appeared to foster higher parasitism rates, but simpler food webs, thereby casting doubt on the general importance of food web complexity for ecosystem functioning.     

Top‐Down Control of Reed Detritus Processing in a Lake Littoral Zone: Experimental Evidence of a Seasonal Compensation between Fish and Invertebrate Predation

We investigated whether predatory fish exert a top‐down control on reed leaf packs processing in a lake littoral zone through a trophic cascade. Exclosure experiments were repeated in summer and winter, under high and low natural fish abundance, respectively. Fish exclusion effects on detritus processing and fungal conditioning were consistent with trophic cascade predictions only in summer. In winter, however, results indicated that a trophic cascade was induced by predatory invertebrates. In both seasons, variations in detritivores abundance generally supported a cascade scenario, whereas several taxon‐specific departures occurred during the experimental periods. We conclude suggesting that predators may continuously regulate leaf detritus processing in lake littoral zones, through a seasonal shift in the relative contribution of fish and invertebrate predation. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Unpacking brown food‐webs: Animal trophic identity reflects rampant microbivory

Detritivory is the dominant trophic paradigm in most terrestrial, aquatic, and marine ecosystems, yet accurate measurement of consumer trophic position within detrital (=“brown”) food webs has remained unresolved. Measurement of detritivore trophic position is complicated by the fact that detritus is suffused with microbes, creating a detrital complex of living and nonliving biomass. Given that microbes and metazoans are trophic analogues of each other, animals feeding on detrital complexes are ingesting other detritivores (microbes), which should elevate metazoan trophic position and should be rampant within brown food webs. We tested these hypotheses using isotopic (¹⁵N) analyses of amino acids extracted from wild and laboratory‐cultured consumers. Vertebrate (fish) and invertebrate detritivores (beetles and moths) were reared on detritus, with and without microbial colonization. In the field, detritivorous animal specimens were collected and analyzed to compare trophic identities among laboratory‐reared and free‐roaming detritivores. When colonized by bacteria or fungi, the trophic positions of detrital complexes increased significantly over time. The magnitude of trophic inflation was mediated by the extent of microbial consumption of detrital substrates. When detrital complexes were fed to vertebrate and invertebrate animals, the consumers registered similar degrees of trophic inflation, albeit one trophic level higher than their diets. The wild‐collected detritivore fauna in our study exhibited significantly elevated trophic positions. Our findings suggest that the trophic positions of detrital complexes rise predictably as microbes convert nonliving organic matter into living microbial biomass. Animals consuming such detrital complexes exhibit similar trophic inflation, directly attributable to the assimilation of microbe‐derived amino acids. Our data demonstrate that detritivorous microbes elevate metazoan trophic position, suggesting that detritivory among animals is, functionally, omnivory. By quantifying the impacts of microbivory on the trophic positions of detritivorous animals and then tracking how these effects propagate “up” food chains, we reveal the degree to which microbes influence consumer groups within trophic hierarchies. The trophic inflation observed among our field‐collected fauna further suggests that microbial proteins represent an immense contribution to metazoan biomass. Collectively, these findings provide an empirical basis to interpret detritivore trophic identity, and further illuminate the magnitude of microbial contributions to food webs.     

Effects of diversity loss on ecosystem function across trophic levels and ecosystems: A test in a detritus‐based tropical food web

Abstract We investigated the effects of biodiversity loss across trophic levels and across ecosystems (terrestrial to aquatic) on ecosystem function, in a detritus‐based tropical food web. Diversities of consumers (stream shredders) and resources (leaf litter) were experimentally manipulated by varying the number of species from 3 to 1, using different species combinations, and the effects on leaf breakdown rates were examined. In single‐species shredder treatments, leaf diversity loss affected breakdown rates, but the effect depended on the identity of the leaves remaining in the system: they increased when the most preferred leaf species remained, but decreased when this species was lost (leaf preferences were the same for all shredders). In multi‐species shredder assemblages, breakdown rates were lower than expected from single‐species treatments, suggesting an important role of interspecific competition. This pattern was also evident when oneleaf species was available but not with higher leaf diversity, suggesting that lowered leaf diversity promotes competitive interactions among shredders. The influence of diversity and identity of species across trophic levels and ecosystems on stream functioning points to complex interactions that may well be reflected in other types of ecosystem.     

Plant and insect diversity along a pollution gradient: understanding species richness across trophic levels

We analysed species richness of plants and true bugs (Insecta, Heteroptera) along a pollution gradient in Scots pine stands in Central Germany. As a consequence of particulate deposition, pH-values of soils increased in the vicinity of the emission source. Therefore, emission increased productivity. Species richness of plants increased with decreasing distance from emission source, and thus with increasing productivity. Similarly, species richness of herbivorous Heteroptera increased with decreasing distance from emission source, whereas, surprisingly, abundance decreased. The proportion of specialised herbivorous bug species is largest in the vicinity of the emission source. Thus, the diversity pattern of herbivores may be explained by the specialisation hypothesis and not the consumer rarity hypothesis. Species richness and abundance of carnivorous Heteroptera showed no significant trend along the gradient. Overall our data favour the bottom-up control of species diversity in the analysed system.     

Unexpected consequences of control: competitive vs. predator release in a four-species assemblage of invasive mammals

Invasive species are frequently the target of eradication or control programmes to mitigate their impacts. However, manipulating single species in isolation can lead to unexpected consequences for other species, with outcomes such as mesopredator release demonstrated both theoretically and empirically in vertebrate assemblages with at least two trophic levels. Less is known about the consequences of species removal in more complex assemblages where a greater number of interacting invaders increases the potential for selective species removal to result in unexpected changes in community structure. Using a replicated Before-After Control-Impact field experiment with a four-species assemblage of invasive mammals we show that species interactions in the community are dominated by competition rather than predation. There was no measurable response of two mesopredators (rats and mice) following control of the top predator (stoats), but there was competitive release of rats following removal of a herbivore (possums), and competitive release of mice following removal of rats.