Chelsea J. Little

Meta-Ecosystems 2.0: Rooting the Theory into the Field

The meta-ecosystem framework demonstrates the significance of among-ecosystem spatial flows for ecosystem dynamics and has fostered a rich body of theory. The high level of abstraction of the models, however, impedes applications to empirical systems. We argue that further understanding of spatial dynamics in natural systems strongly depends on dense exchanges between field and theory. From empiricists, more and specific quantifications of spatial flows are needed, defined by the major categories of organismal movement (dispersal, foraging, life-cycle, and migration). In parallel, the theoretical framework must account for the distinct spatial scales at which these naturally common spatial flows occur. Integrating all levels of spatial connections among landscape elements will upgrade and unify landscape and meta-ecosystem ecology into a single framework for spatial ecology.

Worldwide cross-ecosystem carbon subsidies and their contribution to ecosystem functioning

Ecosystems are widely inter-connected by spatial flows of resources1,2, yet primarily studied in a local context. Meta-ecosystem models suggest that cross-ecosystem subsidies can play an essential role in ecosystem functioning, notably by controlling local availability of resources for biological communities3–6. The general contribution of these resource connections to ecosystem functioning, however, remains unclear in natural systems, due to the heterogeneity and dispersion of data across the ecological literature. Here we provide the first quantitative synthesis on spatial flows of carbon connecting ecosystems worldwide. These cross-ecosystem subsidies range over eight orders of magnitude, between 10−3 and 105 gC m−2 yr−1, and are highly diverse in their provenance. We found that spatial carbon flows and local carbon fluxes are of the same order of magnitudes in freshwater and benthic ecosystems, suggesting an underlying dependency of these systems on resources provided by connected terrestrial and pelagic ecosystems respectively. By contrast, in terrestrial systems, cross-ecosystem subsidies were two to three orders of magnitude lower than local production (grasslands and forests), indicating a weaker quantitative influence on functioning. Those subsidies may still be qualitatively important, however, as some have high nutrient content7,8. We also find important gaps in carbon flow quantification, notably of cross-ecosystem subsidies driven by animal movements, which likely leads to general underestimations of the magnitude and direction of cross-ecosystem linkages9. Overall, we demonstrate strong ecosystem couplings, suggesting that ecosystems can be vulnerable to alterations of these flows and pointing to an urgent need to re-think ecosystem functioning in a spatial perspective.

Do priority effects outweigh environmental filtering in a guild of dominant freshwater macroinvertebrates

Abiotic conditions have long been considered essential in structuring freshwater macroinvertebrate communities. Ecological drift, dispersal and biotic interactions also structure communities, and although these mechanisms are more difficult to detect, they may be of equal importance in natural communities. Here, we hypothesized that in 10 naturally replicated headwater streams in eastern Switzerland, locally dominant amphipod species would be associated with differences in environmental conditions. We conducted repeated surveys of amphipods and used a hierarchical joint species distribution model to assess the influence of different drivers on species co-occurrences. The species had unique environmental requirements, but a distinct spatial structure in their distributions was unrelated to habitat. Species co-occurred much less frequently than predicted by the model, which was surprising because laboratory and field evidence suggests they are capable of coexisting in equal densities. We suggest that niche preemption may limit their distribution and that a blocking effect related to the specific linear configuration of streams determines which species colonizes and dominates a given stream catchment, thus suggesting a new solution a long-standing conundrum in freshwater ecology.

Upstream trophic structure modulates downstream community dynamics via resource subsidies

Abstract In many natural systems, the physical structure of the landscape dictates the flow of resources. Despite mounting evidence that communities’ dynamics can be indirectly coupled by reciprocal among ecosystem resource flows, our understanding of how directional resource flows might indirectly link biological communities is limited. We here propose that differences in community structure upstream should lead to different downstream dynamics, even in the absence of dispersal of organisms. We report an experimental test of the effect of upstream community structure on downstream community dynamics in a simplified but highly controlled setting, using protist microcosms. We implemented directional flows of resources, without dispersal, from a standard resource pool into upstream communities of contrasting interaction structure and then to further downstream communities of either one or two trophic levels. Our results demonstrate that different types of species interactions in upstream habitats may lead to different population sizes and levels of biomass in these upstream habitats. This, in turn, leads to varying levels of detritus transfer (dead biomass) to the downstream communities, thus influencing their population densities and trophic interactions in predictable ways. Our results suggest that the structure of species interactions in directionally structured ecosystems can be a key mediator of alterations to downstream habitats. Alterations to upstream habitats can thus cascade down to downstream communities, even without dispersal.

On Embedding Meta-ecosystems into a Socioecological Framework: A Reply to Renaud et al.

had a limited appearance (but see [5,6]). Such limited uptake may reflect the formal nature of the scientific process, but Chapron et al. [1] demonstrate that there may be a place for satire in scientific journals after all. Conservation lends itself to satire because it is a value-laden topic full of social, political, and ethical obstacles [6]. We thus applaud Chapron et al. [1] for their use of satire and encourage others to do so too where appropriate, even if the views being expressed are sadly closer to reality than exaggeration. After all, the joke is on us. Nature has been around for a few billion years and will be around for a good while longer. Nature needs us a lot less than we need her. With that in mind, and understanding Earth’s new and potentially destructive climate, we have, of course, also booked our seats to the ‘second planet’ along with Chapron and his mates [1], leaving those unwilling to put up with the admittedly rather hefty price tag and terrible interstellar food to stew, roast, bake, or boil on Earth a little longer.

Bottom-up and top-down control of dispersal across major organismal groups

Ecology and evolution unfold in spatially structured communities, where dispersal links dynamics across scales. Because dispersal is multicausal, identifying general drivers remains challenging. In a coordinated distributed experiment spanning organisms from protozoa to vertebrates, we tested whether two fundamental determinants of local dynamics, top-down and bottom-up control, generally explain active dispersal. We show that both factors consistently increased emigration rates and use metacommunity modelling to highlight consequences on local and regional dynamics.In a coordinated distributed dispersal experiment involving seven laboratories, the authors show that both top-down predation risk and bottom-up resource limitation increase emigration rates across 21 species ranging from protozoa to vertebrates.

Landscape configuration alters spatial arrangement of terrestrial-aquatic subsidies in headwater streams

ContextFreshwater ecosystems depend on surrounding terrestrial landscape for resources. Most important are terrestrial leaf litter subsidies, which differ depending on land use. We lack a good understanding of the variation of these inputs across spatial scales.ObjectivesWe sought to determine: (1) the relative importance of local versus catchment-level forestation for benthic leaf litter biomass in streams, (2) how landscape configuration alters these relationships, and (3) how land use affects the quality and diversity of leaf litter subsidies.MethodsWe measured biomass and identity of benthic leaf litter in 121 reaches in 10 independent catchments seasonally over the course of a year. We assessed direct and indirect effects of forestation, reach position, and seasonality on leaf litter biomass using structural equation models, and assessed how leaf litter diversity varied with land use.ResultsIn catchments with forested headwaters, the degree of forestation and reach position in the catchment influenced benthic leaf litter biomass indirectly through local reach-scale forestation. In catchments where forest was only located downstream, or with minimal forest, none of these factors influenced reach-level benthic leaf litter. Leaf litter diversity peaked in fall in all land use types, but was generally lowest in forested reaches.ConclusionsNot only habitat amount, but its location relative to other habitats is important for ecosystem function in the context of cross-ecosystem material flows. Here, lack of upstream forest altered spatial patterns of leaf litter storage. Studies with high spatiotemporal resolution may further reveal effects of landscape configuration on other ecosystems.

Bottom-up and top-down control of dispersal across major organismal groups: a coordinated distributed experiment

Organisms rarely experience a homogeneous environment. Rather, ecological and evolutionary dynamics unfold in spatially structured and fragmented landscapes, with dispersal as the central process linking these dynamics across spatial scales. Because dispersal is a multi-causal and highly plastic life-history trait, finding general drivers that are of importance across species is challenging but highly relevant for ecological forecasting. We here tested whether two fundamental ecological forces and main determinants of local population dynamics, top-down and bottom-up control, generally explain dispersal in spatially structured communities. In a coordinated distributed experiment spanning a wide range of actively dispersing organisms, from protozoa to vertebrates, we show that bottom-up control, that is resource limitation, consistently increased dispersal. While top-down control, that is predation risk, was an equally important dispersal driver as bottom-up control, its effect depended on prey and predator space use and whether dispersal occurred on land, in water or in the air: species that routinely use more space than their predators showed increased dispersal in response to predation, specifically in aquatic environments. After establishing these general causes of dispersal, we used a metacommunity model to show that bottom-up and top-down control of dispersal has important consequences for local population fluctuations as well as cascading effects on regional metacommunity dynamics. Context-dependent dispersal reduced local population fluctuations and desynchronized dynamics between communities, two effects that increase population and community stability. Our study provides unprecedented insights into the generality of the positive resource dependency of dispersal as well as a robust experimental test of current theory predicting that predator-induced dispersal is modulated by prey and predator space use. Our experimental and theoretical work highlights the critical importance of the multi-causal nature of dispersal as well as its cascading effects on regional community dynamics, which are specifically relevant to ecological forecasting.

Exploring the relationship between tychoparthenogenesis and inbreeding depression in the Desert Locust, Schistocerca gregaria

Abstract Tychoparthenogenesis, a form of asexual reproduction in which a small proportion of unfertilized eggs can hatch spontaneously, could be an intermediate evolutionary link in the transition from sexual to parthenogenetic reproduction. The lower fitness of tychoparthenogenetic offspring could be due to either developmental constraints or to inbreeding depression in more homozygous individuals. We tested the hypothesis that in populations where inbreeding depression has been purged, tychoparthenogenesis may be less costly. To assess this hypothesis, we compared the impact of inbreeding and parthenogenetic treatments on eight life‐history traits (five measuring inbreeding depression and three measuring inbreeding avoidance) in four laboratory populations of the desert locust, Schistocerca gregaria, with contrasted demographic histories. Overall, we found no clear relationship between the population history (illustrated by the levels of genetic diversity or inbreeding) and inbreeding depression, or between inbreeding depression and parthenogenetic capacity. First, there was a general lack of inbreeding depression in every population, except in two populations for two traits. This pattern could not be explained by the purging of inbreeding load in the studied populations. Second, we observed large differences between populations in their capacity to reproduce through tychoparthenogenesis. Only the oldest laboratory population successfully produced parthenogenetic offspring. However, the level of inbreeding depression did not explain the differences in parthenogenetic success between all studied populations. Differences in development constraints may arise driven by random and selective processes between populations.