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  1. Abstract

    Methane is a powerful greenhouse gas, more potent than carbon dioxide, and emitted from a variety of natural sources including wetlands, permafrost, mammalian guts and termites. As increases in global temperatures continue to break records, quantifying the magnitudes of key methane sources has never been more pertinent. Over the last 40 years, the contribution of termites to the global methane budget has been subject to much debate. The most recent estimates of termite emissions range between 9 and 15 Tg CH4year−1, approximately 4% of emissions from natural sources (excluding wetlands). However, we argue that the current approach for estimating termite contributions to the global methane budget is flawed. Key parameters, namely termite methane emissions from soil, deadwood, living tree stems, epigeal mounds and arboreal nests, are largely ignored in global estimates. This omission occurs because data are lacking and research objectives, crucially, neglect variation in termite ecology. Furthermore, inconsistencies in data collection methods prohibit the pooling of data required to compute global estimates. Here, we summarise the advances made over the last 40 years and illustrate how different aspects of termite ecology can influence the termite contribution to global methane emissions. Additionally, we highlight technological advances that may help researchers investigate termite methane emissions on a larger scale. Finally, we consider dynamic feedback mechanisms of climate warming and land‐use change on termite methane emissions. We conclude that ultimately the global contribution of termites to atmospheric methane remains unknown and thus present an alternative framework for estimating their emissions. To significantly improve estimates, we outline outstanding questions to guide future research efforts.

     
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  2. null (Ed.)
    Termites are important ecosystem engineers in tropical habitats, with different feeding groups able to decompose wood, grass, litter, and soil organic matter. In most tropical regions, termite abundance and species diversity are assumed to increase with rainfall, with highest levels found in rainforests. However, in the Australian tropics, this pattern is thought to be reversed, with lower species richness and termite abundance found in rainforest than drier habitats. The potential mechanisms underlying this pattern remain unclear. We compared termite assemblages (abundance, activity, diversity, and feeding group composition) across five sites along a precipitation gradient (ranging from ∼800 to 4,000 mm annual rainfall), spanning dry and wet savanna habitats, wet sclerophyll, and lowland and upland rainforests in tropical North Queensland. Moving from dry to wet habitats, we observed dramatic decreases in termite abundance in both mounds and dead wood occupancy, with greater abundance and activity at savanna sites (low precipitation) compared with rainforest or sclerophyll sites (high precipitation). We also observed a turnover in termite species and feeding group diversity across sites that were close together, but in different habitats. Termite species and feeding group richness were highest in savanna sites, with 13 termite species from wood-, litter-, grass-, dung-, and soil-feeding groups, while only five termite species were encountered in rainforest and wet sclerophyll sites—all wood feeders. These results suggest that the Australian termite diversity anomaly may be partly driven by how specific feeding groups colonized habitats across Australia. Consequently, termites in Australian rainforests may be less important in ecosystem processes, such as carbon and nutrient cycling during decomposition, compared with termites in other tropical rainforests. 
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  3. Summary

    Woody biomass is a large carbon store in terrestrial ecosystems. In calculating biomass, tree stems are assumed to be solid structures. However, decomposer agents such as microbes and insects target stem heartwood, causing internal wood decay which is poorly quantified.

    We investigated internal stem damage across five sites in tropical Australia along a precipitation gradient. We estimated the amount of internal aboveground biomass damaged in living trees and measured four potential stem damage predictors: wood density, stem diameter, annual precipitation, and termite pressure (measured as termite damage in downed deadwood).

    Stem damage increased with increasing diameter, wood density, and termite pressure and decreased with increasing precipitation. High wood density stems sustained less damage in wet sites and more damage in dry sites, likely a result of shifting decomposer communities and their differing responses to changes in tree species and wood traits across sites.

    Incorporating stem damage reduced aboveground biomass estimates by > 30% in Australian savannas, compared to only 3% in rainforests. Accurate estimates of carbon storage across woody plant communities are critical for understanding the global carbon budget. Future biomass estimates should consider stem damage in concert with the effects of changes in decomposer communities and abiotic conditions.

     
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  4. Abstract

    Models estimating decomposition rates of dead wood across space and time are mainly based on studies carried out in temperate zones where microbes are dominant drivers of decomposition. However, most dead wood biomass is found in tropical ecosystems, where termites are also important wood consumers. Given the dependence of microbial decomposition on moisture with termite decomposition thought to be more resilient to dry conditions, the relative importance of these decomposition agents is expected to shift along gradients in precipitation that affect wood moisture.

    Here, we investigated the relative roles of microbes and termites in wood decomposition across precipitation gradients in space, time and with a simulated drought experiment in tropical Australia. We deployed mesh bags with non‐native pine wood blocks, allowing termite access to half the bags. Bags were collected every 6 months (end of wet and dry seasons) over a 4‐year period across five sites along a rainfall gradient (ranging from savanna to wet sclerophyll to rainforest) and within a simulated drought experiment at the wettest site. We expected microbial decomposition to proceed faster in wet conditions with greater relative influence of termites in dry conditions.

    Consistent with expectations, microbial‐mediated wood decomposition was slowest in dry savanna sites, dry seasons and simulated drought conditions. Wood blocks discovered by termites decomposed 16–36% faster than blocks undiscovered by termites regardless of precipitation levels. Concurrently, termites were 10 times more likely to discover wood in dry savanna compared with wet rainforest sites, compensating for slow microbial decomposition in savannas. For wood discovered by termites, seasonality and drought did not significantly affect decomposition rates.

    Taken together, we found that spatial and seasonal variation in precipitation is important in shaping wood decomposition rates as driven by termites and microbes, although these different gradients do not equally impact decomposition agents. As we better understand how climate change will affect precipitation regimes across the tropics, our results can improve predictions of how wood decomposition agents will shift with potential for altering carbon fluxes.

    Read the freePlain Language Summaryfor this article on the Journal blog.

     
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  5. Abstract

    Variation in decay rates across woody species is a key uncertainty in predicting the fate of carbon stored in deadwood, especially in the tropics. Quantifying the relative contributions of biotic decay agents, particularly microbes and termites, under different climates and across species with diverse wood traits could help explain this variation.

    To fill this knowledge gap, we deployed woody stems from 16 plant species native to either rainforest (n = 10) or savanna (n = 6) in northeast Australia, with and without termite access. For comparison, we also deployed standardized, non‐native pine blocks at both sites. We hypothesized that termites would increase rates of deadwood decay under conditions that limit microbial activity. Specifically, termite contributions to wood decay should be greater under dry conditions and in wood species with traits that constrain microbial decomposers.

    Termite discovery of stems was surprisingly low with only 17.6% and 22.6% of accessible native stems discovered in the rainforest and savanna respectively. Contrary to our hypothesis, stems discovered by termites decomposed faster only in the rainforest. Termites discovered and decayed pine blocks at higher rates than native stems in both the rainforest and savanna.

    We found significant variation in termite discovery and microbial decay rates across native wood species within the same site. Although wood traits explained 85% of the variation in microbial decay, they did not explain termite‐driven decay. For stems undiscovered by termites, decay rates were greater in species with higher wood nutrient concentrations and syringyl:guiacyl lignin ratios but lower carbon concentrations and wood densities.

    Synthesis. Ecosystem‐scale predictions of deadwood turnover and carbon storage should account for the impact of wood traits on decomposer communities. In tropical Australia, termite‐driven decay was lower than expected for native wood on the ground. Even if termites are present, they may not always increase decomposition rates of fallen native wood in tropical forests. Our study shows how the drivers of wood decay differ between Australian tropical rainforest and savanna; further research should test whether such differences apply world‐wide.

     
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  6. A globally distributed field experiment shows that wood decay, particularly by termites, depends on temperature. 
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  7. Abstract

    Despite the importance of fine roots for the acquisition of soil resources such as nitrogen and water, the study of linkages between traits and both population and community dynamics remains focused on aboveground traits. We address this gap by investigating associations between belowground traits and metrics of species dynamics. Our analysis included 85 species from a long‐term data set on the transition from old field to forest in eastern North America (the Buell‐Small Succession Study) and the new Fine‐Root Ecology Database. Given the prominent roles of life form (woody vs. non‐woody) and species origin (native vs. exotic) in defining functional relationships, we also assessed whether traits or their relationships with species dynamics differed for these groups. Species that reached their peak abundance early in succession had fine‐root traits corresponding to resource acquisitive strategies (i.e., they were thinner, less dense, and had higher nitrogen concentrations) while species that peaked progressively later had increasingly conservative strategies. In addition to having more acquisitive root traits than native species, exotics diverged from the above successional trend, having consistently thinner fine roots regardless of the community context. Species with more acquisitive fine‐root morphologies typically had faster rates of abundance increase and achieved their maximal rates in fewer years. Decreasing soil nutrient availability and increasing belowground competition may become increasingly strong filters in successional communities, acting on root traits to promote a transition from acquisitive to conservative foraging. However, disturbances that increase light and soil resource availability at local scales may allow acquisitive species, especially invasive exotics, to continue colonizing late into the community transition to forest.

     
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  8. Abstract

    Transitioning across biological scales is a central challenge in land surface models. Processes that operate at the scale of individual leaves must be scaled to canopies, and this is done using dedicated submodels. Here, we focus on a submodel that prescribes how light and nitrogen are distributed through plant canopies. We found a mathematical inconsistency in a submodel implemented in the Community and Energy Land Models (CLM and ELM), which incorporates twigs, branches, stems, and dead leaves in nitrogen scaling from leaf to canopy. The inconsistency leads to unrealistic (physically impossible) values of the nitrogen scaling coefficient. The mathematical inconsistency is a general mistake, that is, would occur in any model adopting this particular submodel. We resolve the inconsistency by allowing distinct profiles of stems and branches versus living leaves. We implemented the updated scheme in the ELM and find that the correction reduces global mean gross primary production (GPP) by 3.9 Pg C (3%). Further, when stems and branches are removed from the canopy in the updated model (akin to models that ignore shading from stems), global GPP increases by 4.1 Pg C (3.2%), because of reduced shading. Hence, models that entirely ignore stem shading also introduce errors in the global spatial distribution of GPP estimates, with a strong signal in the tropics, increasing GPP there by over 200 g C m−2 yr−1. Appropriately incorporating stems and other nonphotosynthesizing material into the light and nitrogen scaling routines of global land models, will improve their biological realism and accuracy.

     
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