Concurrent and legacy effects of sheep trampling on soil organic carbon stocks in a typical steppe, China.

Grazing plays a key role in ecosystem biogeochemistry, particularly soil carbon (C) pools. The non-trophic interactions between herbivores and soil processes through herbivore trampling have recently attracted extensive attention. However, their concurrent and legacy effects on the ecosystem properties and processes are still not clear, due to their effects being hard to separate via field experiments. In this study, we conducted a 2-year simulated-sheep-trampling experiment with four trampling intensity treatments (i.e., T0, T40, T80, and T120 for 0, 40, 80, and 120 hoofprints m-2, respectively) in a typical steppe to explore the concurrent and legacy effects of trampling on grassland ecosystem properties and processing. In 2017 (trampling treatment year), we found that trampling decreased aboveground biomass (AGB) of plant community and community-weighted mean shoot C concentration (CWM C), soil available nitrogen (N) and available phosphorus (P), but did not affect plant species diversity and belowground biomass (BGB). We show that compared with T0, trampling increased soil bulk density (BD) at T80, and decreased soil organic carbon (SOC) stocks. After the cessation of trampling for two years (i.e., in 2019), previous trampling increased plant diversity and BGB, reaching the highest values at T80, but decreased soil available N and available P. Compared with T0, previous trampling significantly increased soil BD at T120, while significantly decreased CWM C at T80 and T120, and reduced SOC stocks at T80. Compared with 2017, the trampling negative legacy effects amplified at T80 but weakened at T40 and T120. We also show that trampling-induced decreases in soil available N, AGB of Fabaceae and CWM C were the main predictors of decreasing SOC stocks in 2017, while previous trampling-induced legacy effects on soil available P, AGB of Poaceae and CWM C contributed to the variations of SOC stocks in 2019. Taken together, short-term trampling with low intensity could maintain most plant functions, while previous trampling with low intensity was beneficial to most plant and soil functions. The results of this study show that T40 caused by sheep managed at a stocking rate of 2.7 sheep ha-1 is most suitable for grassland adaptive management in the typical steppe. The ecosystem functions can be maintained under a high stocking rate through the process of providing enough time to rebuild sufficient vegetation cover and restore soil through measures such as regional rotational grazing and seasonal grazing.

Impact of three co-occurring physical ecosystem engineers on soil Collembola communities.

The interplay between organisms with their abiotic environment may have profound effects within ecological networks, but are still poorly understood. Soil physical ecosystem engineers (EEs) modify the abiotic environment, thereby potentially affecting the distribution of other species, such as microarthropods. We focus on three co-occurring physical EEs (i.e. cattle, vegetation, macrodetritivore) known for their profound effect on soil properties (e.g. pore volume, microclimate, litter thickness). We determined their effects on Collembola community composition and life-form strategy (a proxy for vertical distribution in soil) in a European salt marsh. Soil cores were collected in grazed (compacted soil, under short and tall vegetation) and non-grazed areas (decompacted soil, under short and tall vegetation), their pore structure analysed using X-ray computed tomography, after which Collembola were extracted. Collembola species richness was lower in grazed sites, but abundances were not affected by soil compaction or vegetation height. Community composition differed between ungrazed sites with short vegetation and the other treatments, due to a greater dominance of epigeic Collembola and lower abundance of euedaphic species in this treatment. We found that the three co-occurring EEs and their interactions modify the physical environment of soil fauna, particularly through changes in soil porosity and availability of litter. This alters the relative abundance of Collembola life-forms, and thus the community composition within the soil. As Collembola are known to play a crucial role in decomposition processes, these compositional changes in litter and soil layers are expected to affect ecosystem processes and functioning.

Species co-occurrence networks: Can they reveal trophic and non-trophic interactions in ecological communities?

Co-occurrence methods are increasingly utilized in ecology to infer networks of species interactions where detailed knowledge based on empirical studies is difficult to obtain. Their use is particularly common, but not restricted to, microbial networks constructed from metagenomic analyses. In this study, we test the efficacy of this procedure by comparing an inferred network constructed using spatially intensive co-occurrence data from the rocky intertidal zone in central Chile to a well-resolved, empirically based, species interaction network from the same region. We evaluated the overlap in the information provided by each network and the extent to which there is a bias for co-occurrence data to better detect known trophic or non-trophic, positive or negative interactions. We found a poor correspondence between the co-occurrence network and the known species interactions with overall sensitivity (probability of true link detection) equal to 0.469, and specificity (true non-interaction) equal to 0.527. The ability to detect interactions varied with interaction type. Positive non-trophic interactions such as commensalism and facilitation were detected at the highest rates. These results demonstrate that co-occurrence networks do not represent classical ecological networks in which interactions are defined by direct observations or experimental manipulations. Co-occurrence networks provide information about the joint spatial effects of environmental conditions, recruitment, and, to some extent, biotic interactions, and among the latter, they tend to better detect niche-expanding positive non-trophic interactions. Detection of links (sensitivity or specificity) was not higher for well-known intertidal keystone species than for the rest of consumers in the community. Thus, as observed in previous empirical and theoretical studies, patterns of interactions in co-occurrence networks must be interpreted with caution, especially when extending interaction-based ecological theory to interpret network variability and stability. Co-occurrence networks may be particularly valuable for analysis of community dynamics that blends interactions and environment, rather than pairwise interactions alone.

Experimentally reducing species abundance indirectly affects food web structure and robustness.

Studies on the robustness of ecological communities suggest that the loss or reduction in abundance of individual species can lead to secondary and cascading extinctions. However, most such studies have been simulation-based analyses of the effect of primary extinction on food web structure. In a field experiment we tested the direct and indirect effects of reducing the abundance of a common species, focusing on the diverse and self-contained assemblage of arthropods associated with an abundant Brazilian shrub, Baccharis dracunculifolia D.C. (Asteraceae). Over a 5-month period we experimentally reduced the abundance of Baccharopelma dracunculifoliae (Sternorrhyncha: Psyllidae), the commonest galling species associated with B. dracunculifolia, in 15 replicate plots paired with 15 control plots. We investigated direct effects of the manipulation on parasitoids attacking B. dracunculifoliae, as well as indirect effects (mediated via a third species or through the environment) on 10 other galler species and 50 associated parasitoid species. The experimental manipulation significantly increased parasitism on B. dracunculifoliae in the treatment plots, but did not significantly alter either the species richness or abundance of other galler species. Compared to control plots, food webs in manipulated plots had significantly lower values of weighted connectance, interaction evenness and robustness (measured as simulated tolerance to secondary extinction), even when B. dracunculifoliae was excluded from calculations. Parasitoid species were almost entirely specialized to individual galler species, so the observed effects of the manipulation on food web structure could not have propagated via the documented trophic links. Instead, they must have spread either through trophic links not included in the webs (e.g. shared predators) or non-trophically (e.g. through changes in habitat availability). Our results highlight that the inclusion of both trophic and non-trophic direct and indirect interactions is essential to understand the structure and dynamics of even apparently discrete ecological communities.

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.

Does biocide treatment for mosquito control alter carbon dynamics in floodplain ponds?

Shallow lentic aquatic ecosystems, such as ponds, are important repositories of carbon (C) and hotspots of C cycling and greenhouse gas emission. Tube-dwelling benthic invertebrates, such as chironomids, may be key players in C dynamics in these water bodies, yet their role in the C-budget at ecosystem level remains unclear. We tested whether a 41 % reduction in chironomid abundance after application of the mosquito control biocide Bacillus thuringiensis israelensis (Bti) had implications for the C-fluxes to the atmosphere, C-pools, and C-transformation (i.e. organic matter decomposition) in ponds. Data were collected over one year in the shallow, deep and riparian zones of 12 experimental floodplain pond mesocosms (FPMs), half of them treated with Bti. C-fluxes were measured as CO2 and CH4 emissions, atmospheric deposition, and emerging insects. C-pools were measured as dissolved inorganic and organic C in surface and porewater, sediment organic C, C in plant and in macroinvertebrate biomass. Despite seasonal variability, treated FPMs, for which higher CH4 emissions have been reported, showed a trend towards less dissolved organic C in porewater, while no effect was observed for all remaining components of the C-budget. We attribute the effect of Bti on the C-budget to the reduction in macroinvertebrates biomass, the increase in CH4 emissions, and the input of C from the Bti excipients. This finding suggests that changes in tube-dwelling macroinvertebrates have a weak influence on C cycling in ponds and confirms the existence of long-lasting effects of Bti on specific components of C-budgets.

More than a meal integrating non-feeding interactions into food webs.

Organisms eating each other are only one of many types of well documented and important interactions among species. Other such types include habitat modification, predator interference and facilitation. However, ecological network research has been typically limited to either pure food webs or to networks of only a few (<3) interaction types. The great diversity of non-trophic interactions observed in nature has been poorly addressed by ecologists and largely excluded from network theory. Herein, we propose a conceptual framework that organises this diversity into three main functional classes defined by how they modify specific parameters in a dynamic food web model. This approach provides a path forward for incorporating non-trophic interactions in traditional food web models and offers a new perspective on tackling ecological complexity that should stimulate both theoretical and empirical approaches to understanding the patterns and dynamics of diverse species interactions in nature.

Can large herbivores enhance ecosystem carbon persistence?

There is growing interest in aligning the wildlife conservation and restoration agenda with climate change mitigation goals. However, the presence of large herbivores tends to reduce aboveground biomass in some open-canopy ecosystems, leading to the possibility that large herbivore restoration may negatively influence ecosystem carbon storage. Belowground carbon storage is often ignored in these systems, despite the wide recognition of soils as the largest actively-cycling terrestrial carbon pool. Here, we suggest a shift away from a main focus on vegetation carbon stocks, towards inclusion of whole ecosystem carbon persistence, in future assessments of large herbivore effects on long-term carbon storage. Failure to do so may lead to counterproductive biodiversity and climate impacts of land management actions.

Converting Ecological Currencies: Energy, Material, and Information Flows.

Understanding how the three currencies of life - energy, material, and information - interact is a key step towards synthesis in ecology and evolution. However, current theory focuses on the role of matter as a resource and energy, and typically ignores how the same matter can have other important effects as a carrier of information or modifier of the environment. Here we present the hypothesis that the dynamic conversion of matter by organisms among its three currencies mediates the structure and function of ecosystems, and that these effects can even supersede the effects of matter as a resource. Humans are changing the information in the environment and this is altering species interactions and flows of matter within and among ecosystems.