Global contribution of invertebrates to forest litter decomposition.

Forest litter decomposition is an essential component of global carbon and nutrient turnover. Invertebrates play important roles in litter decomposition, but the regional pattern of their effects is poorly understood. We examined 476 case studies across 93 sites and performed a meta-analysis to estimate regional effects of invertebrates on forest litter decomposition. We then assessed how invertebrate diversity, climate and soil pH drive regional variations in invertebrate-mediated decomposition. We found that (1) invertebrate contributions to litter decomposition are 1.4 times higher in tropical and subtropical forests than in forests elsewhere, with an overall contribution of 31% to global forest litter decomposition; and (2) termite diversity, together with warm, humid and acidic environments in the tropics and subtropics are positively associated with forest litter decomposition by invertebrates. Our results demonstrate the significant difference in invertebrate effects on mediating forest litter decomposition among regions. We demonstrate, also, the significance of termites in driving litter mass loss in the tropics and subtropics. These results are particularly pertinent in the tropics and subtropics where climate change and human disturbance threaten invertebrate biodiversity and the ecosystem services it provides.

Termite sensitivity to temperature affects global wood decay rates.

Deadwood is a large global carbon store with its store size partially determined by biotic decay. Microbial wood decay rates are known to respond to changing temperature and precipitation. Termites are also important decomposers in the tropics but are less well studied. An understanding of their climate sensitivities is needed to estimate climate change effects on wood carbon pools. Using data from 133 sites spanning six continents, we found that termite wood discovery and consumption were highly sensitive to temperature (with decay increasing >6.8 times per 10°C increase in temperature)-even more so than microbes. Termite decay effects were greatest in tropical seasonal forests, tropical savannas, and subtropical deserts. With tropicalization (i.e., warming shifts to tropical climates), termite wood decay will likely increase as termites access more of Earth's surface.

The impact of invertebrate decomposers on plants and soil.

Soil invertebrates make significant contributions to the recycling of dead plant material across the globe. However, studies focussed on the consequences of decomposition for plant communities largely ignore soil fauna across all ecosystems, because microbes are often considered the primary agents of decay. Here, we explore the role of invertebrates as not simply facilitators of microbial decomposition, but as true decomposers, able to break down dead organic matter with their own endogenic enzymes, with direct and indirect impacts on the soil environment and plants. We recommend a holistic view of decomposition, highlighting how invertebrates and microbes act in synergy to degrade organic matter, providing ecological services that underpin plant growth and survival.

Carbon flux and forest dynamics: Increased deadwood decomposition in tropical rainforest tree-fall canopy gaps.

Tree mortality rates are increasing within tropical rainforests as a result of global environmental change. When trees die, gaps are created in forest canopies and carbon is transferred from the living to deadwood pools. However, little is known about the effect of tree-fall canopy gaps on the activity of decomposer communities and the rate of deadwood decay in forests. This means that the accuracy of regional and global carbon budgets is uncertain, especially given ongoing changes to the structure of rainforest ecosystems. Therefore, to determine the effect of canopy openings on wood decay rates and regional carbon flux, we carried out the first assessment of deadwood mass loss within canopy gaps in old-growth rainforest. We used replicated canopy gaps paired with closed canopy sites in combination with macroinvertebrate accessible and inaccessible woodblocks to experimentally partition the relative contribution of microbes vs. termites to decomposition within contrasting understorey conditions. We show that over a 12 month period, wood mass loss increased by 63% in canopy gaps compared with closed canopy sites and that this increase was driven by termites. Using LiDAR data to quantify the proportion of canopy openings in the study region, we modelled the effect of observed changes in decomposition within gaps on regional carbon flux. Overall, we estimate that this accelerated decomposition increases regional wood decay rate by up to 18.2%, corresponding to a flux increase of 0.27 Mg C ha-1  year-1 that is not currently accounted for in regional carbon budgets. These results provide the first insights into how small-scale disturbances in rainforests can generate hotspots for decomposer activity and carbon fluxes. In doing so, we show that including canopy gap dynamics and their impacts on wood decomposition in forest ecosystems can help improve the predictive accuracy of the carbon cycle in land surface models.

Termites can decompose more than half of deadwood in tropical rainforest.

Termite-mediated decomposition is an important, but often overlooked, component of the carbon cycle. Using a large-scale suppression experiment in Borneo, Griffiths et al. found that termites contribute between 58 and 64% of mass loss from dead wood.

Ants are the major agents of resource removal from tropical rainforests.

Ants are diverse and abundant, especially in tropical ecosystems. They are often cited as the agents of key ecological processes, but their precise contributions compared with other organisms have rarely been quantified. Through the removal of food resources from the forest floor and subsequent transport to nests, ants play an important role in the redistribution of nutrients in rainforests. This is an essential ecosystem process and a key energetic link between higher trophic levels, decomposers and primary producers. We used the removal of carbohydrate, protein and seed baits as a proxy to quantify the contribution that ants, other invertebrates and vertebrates make to the redistribution of nutrients around the forest floor, and determined to what extent there is functional redundancy across ants, other invertebrate and vertebrate groups. Using a large-scale, field-based manipulation experiment, we suppressed ants from c. 1 ha plots in a lowland tropical rainforest in Sabah, Malaysia. Using a combination of treatment and control plots, and cages to exclude vertebrates, we made food resources available to: (i) the whole foraging community, (ii) only invertebrates and (iii) only non-ant invertebrates. This allowed us to partition bait removal into that taken by vertebrates, non-ant invertebrates and ants. Additionally, we examined how the non-ant invertebrate community responded to ant exclusion. When the whole foraging community had access to food resources, we found that ants were responsible for 52% of total bait removal whilst vertebrates and non-ant invertebrates removed the remaining 48%. Where vertebrates were excluded, ants carried out 61% of invertebrate-mediated bait removal, with all other invertebrates removing the remaining 39%. Vertebrates were responsible for just 24% of bait removal and invertebrates (including ants) collectively removed the remaining 76%. There was no compensation in bait removal rate when ants and vertebrates were excluded, indicating low functional redundancy between these groups. This study is the first to quantify the contribution of ants to the removal of food resources from rainforest floors and thus nutrient redistribution. We demonstrate that ants are functionally unique in this role because no other organisms compensated to maintain bait removal rate in their absence. As such, we strengthen a growing body of evidence establishing ants as ecosystem engineers, and provide new insights into the role of ants in maintaining key ecosystem processes. In this way, we further our basic understanding of the functioning of tropical rainforest ecosystems.

The value of trophic interactions for ecosystem function: dung beetle communities influence seed burial and seedling recruitment in tropical forests.

Anthropogenic activities are causing species extinctions, raising concerns about the consequences of changing biological communities for ecosystem functioning. To address this, we investigated how dung beetle communities influence seed burial and seedling recruitment in the Brazilian Amazon. First, we conducted a burial and retrieval experiment using seed mimics. We found that dung beetle biomass had a stronger positive effect on the burial of large than small beads, suggesting that anthropogenic reductions in large-bodied beetles will have the greatest effect on the secondary dispersal of large-seeded plant species. Second, we established mesocosm experiments in which dung beetle communities buried Myrciaria dubia seeds to examine plant emergence and survival. Contrary to expectations, we found that beetle diversity and biomass negatively influenced seedling emergence, but positively affected the survival of seedlings that emerged. Finally, we conducted germination trials to establish the optimum burial depth of experimental seeds, revealing a negative relationship between burial depth and seedling emergence success. Our results provide novel evidence that seed burial by dung beetles may be detrimental for the emergence of some seed species. However, we also detected positive impacts of beetle activity on seedling recruitment, which are probably because of their influence on soil properties. Overall, this study provides new evidence that anthropogenic impacts on dung beetle communities could influence the structure of tropical forests; in particular, their capacity to regenerate and continue to provide valuable functions and services.

Assessing the Importance of Intraspecific Variability in Dung Beetle Functional Traits.

Functional diversity indices are used to facilitate a mechanistic understanding of many theoretical and applied questions in current ecological research. The use of mean trait values in functional indices assumes that traits are robust, in that greater variability exists between than within species. While the assertion of robust traits has been explored in plants, there exists little information on the source and extent of variability in the functional traits of higher trophic level organisms. Here we investigated variability in two functionally relevant dung beetle traits, measured from individuals collected from three primary forest sites containing distinct beetle communities: body mass and back leg length. In doing so we too addressed the following questions: (i) what is the contribution of intra vs. interspecific differences in trait values; (ii) what sample size is needed to provide representative species mean trait values; and (iii) what impact does omission of intraspecific trait information have on the calculation of functional diversity (FD) indices from naturally assembled communities? At the population level, interspecific differences explained the majority of variability in measured traits (between 94% and 96%). In accordance with this, the error associated with calculating FD without inclusion of intraspecific variability was low, less than 20% in all cases. This suggests that complete sampling to capture intraspecific variance in traits is not necessary even when investigating the FD of small and/or naturally formed communities. To gain an accurate estimation of species mean trait values we encourage the measurement of 30-60 individuals and, where possible, these should be taken from specimens collected from the site of study.

Predatory functional response and prey choice identify predation differences between native/invasive and parasitised/unparasitised crayfish.

BACKGROUND: Invasive predators may change the structure of invaded communities through predation and competition with native species. In Europe, the invasive signal crayfish Pacifastacus leniusculus is excluding the native white clawed crayfish Austropotamobius pallipes. METHODOLOGY AND PRINCIPAL FINDINGS: This study compared the predatory functional responses and prey choice of native and invasive crayfish and measured impacts of parasitism on the predatory strength of the native species. Invasive crayfish showed a higher (>10%) prey (Gammarus pulex) intake rate than (size matched) natives, reflecting a shorter (16%) prey handling time. The native crayfish also showed greater selection for crustacean prey over molluscs and bloodworm, whereas the invasive species was a more generalist predator. A. pallipes parasitised by the microsporidian parasite Thelohania contejeani showed a 30% reduction in prey intake. We suggest that this results from parasite-induced muscle damage, and this is supported by a reduced (38%) attack rate and increased (30%) prey handling time. CONCLUSIONS AND SIGNIFICANCE: Our results indicate that the per capita (i.e., functional response) difference between the species may contribute to success of the invader and extinction of the native species, as well as decreased biodiversity and biomass in invaded rivers. In addition, the reduced predatory strength of parasitized natives may impair their competitive abilities, facilitating exclusion by the invader.

Climate change and evolutionary adaptations at species' range margins.

During recent climate warming, many insect species have shifted their ranges to higher latitudes and altitudes. These expansions mirror those that occurred after the Last Glacial Maximum when species expanded from their ice age refugia. Postglacial range expansions have resulted in clines in genetic diversity across present-day distributions, with a reduction in genetic diversity observed in a wide range of insect taxa as one moves from the historical distribution core to the current range margin. Evolutionary increases in dispersal at expanding range boundaries are commonly observed in virtually all insects that have been studied, suggesting a positive feedback between range expansion and the evolution of traits that accelerate range expansion. The ubiquity of this phenomenon suggests that it is likely to be an important determinant of range changes. A better understanding of the extent and speed of adaptation will be crucial to the responses of biodiversity and ecosystems to climate change.