Spatial and temporal hotspots of termite‐driven decomposition in the Serengeti

Ecosystem engineers are organisms that directly or indirectly control the availability of resources to other organisms by causing physical state changes in biotic or abiotic materials. Termites (Insecta, Isoptera) are among the most important ecosystem engineers in tropical ecosystems. We used a field experiment in the tall grasslands of Serengeti National Park, Tanzania, to investigate 1) the consumption by termites of grass litter and dung baits along the landscape gradient of catena position, and 2) seasonal variation in litter and dung removal. Our maps of termitaria and patterns of bait removal revealed clear spatial and temporal hotspots of termite activity. In the dry season termites removed more baits at the top‐catena positions than at the bottom positions, but there was no effect of catena position in the wet season. Spatial hotspots of termite activity overlapped with those of both mammalian herbivores and predators. Within the framework of ecosystem engineering, this study suggests that intraspecific aspects of spatial heterogeneity and temporal variability deserve much greater consideration.     

Termite mound cover and abundance respond to herbivore‐mediated biotic changes in a Kenyan savanna

Both termites and large mammalian herbivores (LMH) are savanna ecosystem engineers that have profound impacts on ecosystem structure and function. Both of these savanna engineers modulate many common and shared dietary resources such as woody and herbaceous plant biomass, yet few studies have addressed how they impact one another. In particular, it is unclear how herbivores may influence the abundance of long‐lived termite mounds via changes in termite dietary resources such as woody and herbaceous biomass. While it has long been assumed that abundance and areal cover of termite mounds in the landscape remain relatively stable, most data are observational, and few experiments have tested how termite mound patterns may respond to biotic factors such as changes in large herbivore communities. Here, we use a broad tree density gradient and two landscape‐scale experimental manipulations—the first a multi‐guild large herbivore exclosure experiment (20 years after establishment) and the second a tree removal experiment (8 years after establishment)—to demonstrate that patterns in Odontotermes termite mound abundance and cover are unexpectedly dynamic. Termite mound abundance, but areal cover not significantly, is positively associated with experimentally controlled presence of cattle, but not wild mesoherbivores (15–1,000 kg) or megaherbivores (elephants and giraffes). Herbaceous productivity and tree density, termite dietary resources that are significantly affected by different LMH treatments, are both positive predictors of termite mound abundance. Experimental reductions of tree densities are associated with lower abundances of termite mounds. These results reveal a richly interacting web of relationships among multiple savanna ecosystem engineers and suggest that termite mound abundance and areal cover are intimately tied to herbivore‐driven resource availability.     

Termite mounds differ in their importance for herbivores across savanna types,seasons and spatial scales

Herbivores do not forage uniformly across landscapes, but select for patches of higher nutrition and lower predation risk. Macrotermes mounds contain higher concentrations of soil nutrients and support grasses of higher nutritional value than the surrounding savanna matrix, attracting mammalian grazers that preferentially forage on termite mound vegetation. However, little is known about the spatial extent of such termite influence on grazing patterns and how it might differ in time and space. We measured grazing intensity in three African savanna types differing in rainfall and foliar nutrients and predicted that the functional importance of mounds for grazing herbivores would increase as the difference in foliar nutrient levels between mound and savanna matrix grasses increases and the mounds become more attractive. We expected this to occur in nutrient‐poor areas and during the dry season when savanna matrix grass nutrient levels are lower. Tuft use and grass N and P content were measured along transects away from termite mounds, enabling calculation of the spatial extent of termite influence on mammalian grazing. Using termite mound densities estimated from airborne light detection and ranging (LiDAR), we further upscaled field‐based results to determine the percentage of the landscape influenced by termite activity. Grasses in close proximity to termite mounds were preferentially grazed at all sites and in both seasons, but the strength of mound influence varied between savanna types and seasons. In the wet season, mounds had a relatively larger effect on grazers at the landscape scale in the nutrient‐poor, wetter savanna, whereas in the dry season the pattern was reversed with more of the landscape influenced at the nutrient‐rich, driest site. Our results reveal that termite mounds enhance the value of savanna landscapes for herbivores, but that their functional importance varies across savanna types and seasons.     

Termite‐induced heterogeneity in African savanna vegetation: mechanisms and patterns

Objectives: To (1) assess the strength of evidence for the role of termites in vegetation heterogeneity in African savannas, and (2) identify the mechanisms by which termites induce such heterogeneity. Location: African savannas. Methods: We conducted a review of the literature, a meta‐analysis and qualitative systems analysis to identify mechanisms to explain the observed patterns. Results: The review provided evidence for termite‐induced heterogeneity in floristic composition and vegetation patterning in savannas across Africa. Termites induced vegetation heterogeneity directly or indirectly through their nest‐building and foraging activities, associated nutrient cycling and their interaction with mammalian herbivores and fire. The literature reviewed indicated that termite mounds essentially act as islands of fertility, which are responsible for ecosystem‐level spatial heterogeneity in savannas. This was supported by the meta‐analysis, which demonstrated that mounds of Ancistrotermes, Macrotermes, Odontotermes (family Macrotermitinae), Cubitermes (family Termitinae) and Trinervitermes (Nasutitermitinae) are significantly enriched in clay (75%), carbon (16%), total nitrogen (42%), calcium (232%), potassium (306%) and magnesium (154%) compared to the surrounding savanna soil. Conclusions: Termite activity is one of the major factors that induce vegetation patterning in African savannas. The implications of this are discussed and research questions for future studies and modelling efforts are indicated.     

Relative roles of termites and saprotrophic microbes as drivers of wood decay: A wood block test

Deadwood in tropical ecosystems represents an important but poorly studied carbon (C) pool. Biologically mediated decay of this pool occurs by both saprotrophic microbes and macro‐invertebrates, such as termites. The activity of these decay agents is influenced by abiotic conditions, especially water availability in tropical systems. While saprotrophic microbial activity is directly controlled by moisture, termites employ various morphological and behavioural modifications that should allow for continued activity in dry conditions. We therefore hypothesized that the relative role of termites would be enhanced in the dry season and a dry compared to a wet site. We deployed a novel wood bait (Pinus radiata) at two sites (rainforest and savanna), with or without access holes cut into termite‐excluding mesh. Mass loss from wood baits was measured after a dry season and after a full dry/wet annual cycle. Mass loss was higher at the rainforest site, demonstrating the overall role of moisture in driving wood decay. Counter to expectations, we found no evidence that the relative role of termites was higher at the dry site, nor during the dry season. However, the prevalence of termites was higher in the savanna compared to the rainforest. While termites clearly impact wood decay, these findings indicate that the relative importance of termites in the fate of deadwood may not reflect their mere presence within and across ecosystems. If moisture availability shifts under climate change, our results suggest similar functional responses between termites and saprotrophic microbes in driving C loss from deadwood.     

Nectar quality affects ant aggressiveness and biotic defense provided to plants

Ant–plant mutualisms are useful models for investigating how plant traits mediate interspecific interactions. As plant‐derived resources are essential components of ant diets, plants that offer more nutritious food to ants should be better defended in return, as a result of more aggressive behavior toward natural enemies. We tested this hypothesis in a field experiment by adding artificial nectaries to individuals of the species Vochysia elliptica (Vochysiaceae). Ants were offered one of four liquid foods of different nutritional quality: amino acids, sugar, sugar + amino acids, and water (control). We used live termites (Nasutitermes coxipoensis) as herbivore competitors and observed ant behavior toward them. In 88 hr of observations, we recorded 1,009 interactions with artificial nectaries involving 1,923 individual ants of 26 species. We recorded 381 encounters between ants and termites, of which 38% led to attack. Sixty‐one percent of these attacks led to termite exclusion from the plants. Recruitment and patrolling were highest when ants fed upon nectaries providing sugar + amino acids, the most nutritious food. This increase in recruitment and patrolling led to higher encounter rates between ants and termites, more frequent attacks, and faster and more complete termite removal. Our results are consistent with the hypothesis that plant biotic defense is mediated by resource quality. We highlight the importance of qualitative differences in nectar composition for the outcome of ant–plant interactions. Abstract in Portuguese is available with online material.     

Infestation of African savanna ecosystems by subterranean termites

This review is an analysis of the dimensions of termite infestation in African savannas. The aim of this work is to draw the attention of ecologists, conservationists, policy makers and farmers to the current and future threats of subterranean termites to the functioning and sustainability of such ecosystem habitats. This study analyzes and describes termite problem (questionable changes in density and assemblage structure) in selected African savannah ecosystems, synthesizes information on the effects of various human induced habitat disturbance regimes on termites’ assemblage structure, predators, nests and feed resources to generate hypotheses relating termite infestation with anthropogenic activities; it describes and critiques existing termite management practices. The review is suggestive that the infestation and resultant undesirable effects of subterranean termites in African savannahs are largely a consequence of the inappropriate savannah management practices (overgrazing, indiscriminate tree cutting and overhunting) undertaken by humans in pursuit of various livelihood options. Based on the evidence presented herein, we hypothesized that (1) human induced habitat disturbance in savanna ecosystems alters the feeding group composition of termites’ assemblages, favoring grass harvesters and polyphagus termite feeders that forage on more abundant food items, paying little attention to rarer food items and (2) habitat disturbance through activities like heavy grazing and overhunting results in decline in the populations of both macro and microscopic termite predators, which eventually enhances the proliferation of termite populations, escalates the density of termite nests particularly epigeal mounds and intensifies consumption of herbaceous savannah vegetation. The review calls for dedicated efforts to develop ecological thresholds of savannah biotic and abiotic ecosystem components in which human induced disturbance regimes trigger the destructive behavior of termites. This would provide information that will act as a precautionary savannah habitat monitoring and decision support tool to prevent future infestation of savannah habitats with termites. Also, the review shows that majority of the termite control practices are ineffective, ecologically unsustainable and above all, do not address the root cause of termite infestation and thus merely provide temporary relief to the problem. As such, termite control methods that attempt to enhance proliferation of termite predators need to be studied, developed and emphasized. This review reveals that human induced habitat disturbance depletes termites’ predator populations, leading to proliferation of termite populations particularly grass harvesters that intensify their consumption on grass biomass and eventually contribute to denudation of herbaceous vegetation cover in savannah ecosystems.     

Termites facilitate and ungulates limit savanna tree regeneration

Both large herbivores and termites are key functional groups in savanna ecosystems, and in many savanna areas, large termite mounds (termitaria) are associated with distinct woody clusters. Studies on the effect of large mammals on tree regeneration are few, and the results are conflicting. Large herbivores have been found to be important seedling predators in some areas, but facilitate tree regeneration by outcompeting small mammals and reducing grass cover in other areas. Through the use of the experimental fencing of termite mounds and adjacent savanna areas in this study, we investigated how termites and large herbivores influence tree regeneration. Termite mounds had a higher number of seedlings, more species richness, more alpha diversity (OD) and lower evenness (E) than savanna plots. Large herbivores did not significantly affect overall seedling density, species richness, OD or E. Beta diversity was higher in savanna areas than on termitaria, and beta diversity decreased in savanna areas when herbivores were excluded. Herbivore exclusion increased the density of the 12 (40 %) most common seedling species, representing 79 % of all seedlings, and fenced plots had relatively taller seedlings than open plots. Thus, termites were the main determinants of tree regeneration in our study area, but large mammals regulated the most common species. Although our study confirms previous work suggesting that large herbivores affect tree regeneration, we found that termites were an even more important determinant. Termite impacts on tree regeneration deserve increased attention by savanna ecologists.     

Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature – herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local‐scale herbivory, and its interaction with nutrient enrichment and climate, within global‐scale models to better predict land–atmosphere interactions under future climate change.     

Herbivory and nutrient limitation protect warming tundra from lowland species’ invasion and diversity loss

Herbivory and nutrient limitation can increase the resistance of temperature‐limited systems to invasions under climate warming. We imported seeds of lowland species to tundra under factorial treatments of warming, fertilization, herbivore exclusion and biomass removal. We show that warming alone had little impact on lowland species, while exclusion of native herbivores and relaxation of nutrient limitation greatly benefitted them. In contrast, warming alone benefitted resident tundra species and increased species richness; however, these were canceled by negative effects of herbivore exclusion and fertilization. Dominance of lowland species was associated with low cover of tundra species and resulted in decreased species richness. Our results highlight the critical role of biotic and abiotic filters unrelated to temperature in protecting tundra under warmer climate. While scarcity of soil nutrients and native herbivores act as important agents of resistance to invasions by lowland species, they concurrently promote overall species coexistence. However, when these biotic and abiotic resistances are relaxed, invasion of lowland species can lead to decreased abundance of resident tundra species and diminished diversity.     

Strong positive effects of termites on savanna bird abundance and diversity are amplified by large herbivore exclusion

Vast areas of the African savanna landscapes are characterized by tree‐covered Macrotermes termite mounds embedded within a relatively open savanna matrix. In concert with termites, large herbivores are important determinants of savanna woody vegetation cover. The relative cover of woody species has considerable effects on savanna function. Despite the potentially important ecological relationships between termite mounds, woody plants, large herbivores, and birds, these associations have previously received surprisingly little attention. We experimentally studied the effects of termites and large herbivores on the avian community in Lake Mburo National Park, Uganda, where woody vegetation is essentially limited to termite mounds. Our experiment comprised of four treatments in nine replicates; unfenced termite mounds, fenced mounds (excluding large mammals), unfenced adjacent savanna, and fenced savanna. We recorded species identity, abundance, and behavior of all birds observed on these plots over a two‐month period, from late dry until wet season. Birds used termite mounds almost exclusively, with only 3.5% of observations occurring in the treeless intermound savanna matrix. Mean abundance and species richness of birds doubled on fenced (large herbivores excluded) compared to unfenced mounds. Feeding behavior increased when large mammals were excluded from mounds, both in absolute number of observed individuals, and relative to other behaviors. This study documents the fundamental positive impact of Macrotermes termites on bird abundance and diversity in an African savanna. Birds play crucial functional roles in savanna ecosystems, for example, by dispersing fruits or regulating herbivorous insect populations. Thus, the role of birds in savanna dynamics depends on the distribution and abundance of termite mounds.     

Plant composition,not richness,drives occurrence of specialist herbivores

1. How herbivore plant diversity relationships are shaped by the interplay of biotic and abiotic environmental variables is only partly understood. For instance, plant diversity is commonly assumed to determine abundance and richness of associated specialist herbivores. However, this relationship can be altered when environmental variables such as temperature covary with plant diversity. 2. Using gall‐inducing arthropods as focal organisms, biotic and abiotic environmental variables were tested for their relevance to specialist herbivores and their relationship to host plants. In particular, the hypothesis that abundance and richness of gall‐inducing arthropods increase with plant richness was addressed. Additionally, the study asked whether communities of gall‐inducing arthropods match the communities of their host plants. 3. Neither abundance nor species richness of gall‐inducing arthropods was correlated with plant richness or any other of the tested environmental variables. Instead, the number of gall species found per plant decreased with plant richness. This indicates that processes of associational resistance may explain the specialised plant herbivore relationship in our study. 4. Community composition of gall‐inducing arthropods matched host plant communities. In specialised plant herbivore relationships, the presence of obligate host plant species is a prerequisite for the occurrence of its herbivores. 5. It is concluded that the abiotic environment may only play an indirect role in shaping specialist herbivore communities. Instead, the occurrence of specialist herbivore communities might be best explained by plant species composition. Thus, plant species identity should be considered when aiming to understand the processes that shape diversity patterns of specialist herbivores.     

Does shrub invasion indirectly limit grass establishment via seedling herbivory? A test at grassland‐shrubland ecotones

Question: Does shrub invasion at ecotones indirectly limit grass establishment by increasing mammalian seedling herbivory? Location: Chihuahuan Desert, New Mexico, USA. Methods: We tested the hypothesis that herbivore‐related mortality of seedlings of the dominant perennial grass Bouteloua eriopoda would be highest in shrub‐dominated portions of grassland‐shrubland ecotones. We tested the hypothesis in two Chihuahuan Desert sites featuring similar shrub encroachment patterns but different shrub species, grass cover, and different abundances of small mammals. Within each site we transplanted B. eriopoda seedlings to grass‐dominated, middle, and shrub‐dominated positions of replicate ecotones during the time of year (mid‐summer) when they would naturally appear and monitored seedling fates. We estimated population size/activity of putative small mammal herbivores. Results: Seedlings were killed by mammals in greater numbers in shrubland than in grassland or middle ecotone positions at the site with large herbivore numbers. At the site with low herbivore numbers, most seedlings were killed in middle ecotone positions. The abundance patterns of herbivores did not parallel patterns of seedling herbivory across the ecotones or between sites. Conclusions: Seedling herbivory is an important process and is related to vegetation composition, but the mechanisms underlying the relationship are not clear. We speculate that variation in small mammal foraging behavior may contribute to seedling herbivory patterns. Restoration strategies in the Chihuahuan Desert need to account for the abundance and/or behavior of native herbivores.     

Ant colonization and coarse woody debris decomposition in temperate forests

Ants are ubiquitous, abundant and have widespread impacts on ecological communities and ecosystem processes. However, ant effects on coarse woody debris decomposition are unexplored. Several ant species colonize coarse woody debris for nesting, and this puts them in contact with fauna and microbes that utilize coarse woody debris as habitat and food, potentially influencing nutrient cycling and, ultimately, forest productivity. We report results from a field experiment employing 138 artificial ant nests (routed pine blocks) across five locations in southeastern US deciduous forests. We examine the correspondence between ant, termite and wood-eating fungi colonization and variation in coarse woody debris decomposition. After 1 year, nests colonized by ants had 5% more mass than those not colonized. Ant colonization corresponded with significantly less termite- and fungal-mediated decomposition of the nests. Without ants, termites removed 11.5% and fungi removed 4% more wood biomass. Ants, termites and wood-eating fungi all colonized pine nests where temperatures were highest, and ants also preferred higher soil moisture whereas termites and fungi responded negatively to high soil moisture when temperatures were higher. Ants reduce termite colonies through predation, and may inhibit fungi through the secretion of antimicrobial compounds. Our results indicate that interactions between forest understory ants, termites and fungi may influence the rate of coarse woody debris decomposition—biotic interactions that potentially influence forest structure and function.     

Plant water stress and previous herbivore damage affect insect performance

1. Short‐term changes in plant resistance traits can be affected by abiotic factors or damage by herbivores, although how the combined effects of abiotic factors and previous damage affect subsequent insect larval development is not well understood. 2. Complementary glasshouse and field experiments were conducted to evaluate whether plant water stress and previous herbivore damage influenced monarch (Danaus plexippus) larval development on common milkweed, Asclepias syriaca. 3. In the glasshouse, water stress altered a suite of A. syriaca functional traits but did not affect nutrient content, whereas herbivore damage increased leaf nitrogen (N) and reduced the carbon:nitrogen (C:N) ratio. A bioassay experiment showed that monarch larval survival was lower on well‐watered plants that were previously damaged by monarch larva than on damaged and drought‐stressed plants. Bioassay larvae consumed less leaf tissue of previously damaged plants, whereas monarch larval mass was affected additively by water stress and previous damage, after correcting for the amount of leaf tissue consumed. 4. In a 2‐year field experiment, monarch larval performance was higher on previously damaged A. syriaca plants that received experimentally reduced rainfall, relative to plants receiving ambient rainfall. 5. Collectively, these results from glasshouse and field experiments suggest that insect performance was highest on previously damaged plants under water stress and highlight the additive and interactive roles of abiotic and biotic factors on herbivore performance.     

Disproportionate effects of non‐colonial small herbivores on structure and diversity of grassland dominated by large herbivores

The response of semiarid grasslands to small, non‐colonial herbivores has received little attention, focusing primarily on the effects of granivore assemblages on annual plant communities. We studied the long‐term effects of both small and large herbivores on vegetation structure and species diversity of shortgrass steppe, a perennial semiarid grassland considered marginal habitat for small mammalian herbivores. We hypothesized that 1) large generalist herbivores would affect more abundant species and proportions of litter‐bare ground‐vegetation cover through non‐selective herbivory, 2) small herbivores would affect less common species through selective but limited consumption, and 3) herbivore effects on plant richness would increase with increasing aboveground net primary production (ANPP). Plant community composition was assessed over a 14‐year period in pastures grazed at moderate intensities by cattle and in exclosures for large (cattle) and large‐plus‐small herbivores (additional exclusion of rabbits and rodents). Exclusion of large herbivores affected litter and bare ground and basal cover of abundant, common and uncommon species. Additional exclusion of small herbivores did not affect uncommon components of the plant community, but had indirect effects on abundant species, decreased the cover of the dominant grass Bouteloua gracilis and total vegetation, and increased litter and species diversity. There was no relationship between ANPP and the intensity of effects of either herbivore body size on richness. Exclusion of herbivores of both body sizes had complementary and additive effects which promoted changes in vegetation composition and physiognomy that were linked to increased abundance of tall and decreased abundance of short species. Our findings show that small mammalian herbivores had disproportionately large effects on plant communities relative to their small consumption of biomass. Even in small‐seeded perennial grasslands with a long history of intensive grazing by large herbivores, non‐colonial small mammalian herbivores should be recognized as an important driver of grassland structure and diversity.     

Temperate forest termites: ecology,biogeography, and ecosystem impacts

1. Wood decomposition in temperate forests is dominated by termites, fungi, and some species of ants and beetles. Outside of urban areas, temperate termite ecology is largely unknown, particularly when compared to tropical termites and other temperate organisms in the functional guild of wood‐decomposing animals. 2. This review combines climate habitat modelling with knowledge of species physiology, behaviour, and community interactions to identify and prioritise future research on temperate termite ecology and biogeography. 3. Using a correlative climate model, the regional distributions of three common temperate forest termite species are shown to correlate with different aspects of climate (e.g. mean versus minimum monthly temperature), but that overall their distributions within temperate systems correlate more strongly with temperature variables than with precipitation variables. 4. Existing data are synthesised to outline how the subterranean, wood‐nesting behaviour of most temperate forest termite species links their activity to an additional set of non‐climate controls: wood type and tree species, soil depth, fungal activity, ant abundances and phenology, and competitive asymmetries among termite species. 5. Although fine‐scale estimates of temperate termite abundances are rare, we provide upper bounds on their ecosystem impacts and illustrate how their regional abundances may influence forest structure and habitat availability for other organisms. 6. This review highlights that rigorous ecological studies in non‐urban, intact ecosystems – with a particular focus on community interactions – are critically needed to accurately project future abundances, economic impacts, and ecosystem effects of temperate forest termites.     

Associational refuge in practice: can existing vegetation facilitate woodland restoration?

Herbivores can dramatically diminish revegetation success, but associational refuge theory predicts that neighbouring plants could hinder browsing of planted seedlings. The key to strategic restoration using associational refuge is to define which patch variables are effective against the appropriate herbivores, at multiple scales, and to understand which stages of the foraging process these variables disrupt. Our study aimed to test the capacity of existing vegetation to act as associational refuge for planted seedlings by affecting search, detection and consumption decisions, and more generally influence herbivore foraging patterns. We conducted a field trial with free‐ranging, mammalian herbivores and nursery‐raised, native tree seedlings. We quantified seedling browsing damage over time in relation to a suite of existing patch variables at two spatial scales (100 m² and 4 m²). After two months, 78% of seedlings were browsed, suffering mean foliage loss of 90.5%. Focal seedlings were almost exclusively consumed by swamp wallabies Wallabia bicolor, an abundant generalist browser. Once a swamp wallaby investigated a seedling, the probability of consumption was high (86%). At the large scale, browsing of seedlings was delayed in patches with lower canopy cover and fewer browsed plant species. Seedlings in fern‐dominated patches escaped browsing for longer than those in grass‐dominated patches. At the small scale, browsing was delayed with higher cover of understorey vegetation. Associational refuge was provided by vegetation with characteristics, and at spatial scales, consistent with disrupted search and detection of focal seedlings by herbivores. Thus strategic placement of seedlings in existing vegetation – based on understanding which herbivore species is responsible and how it responds to vegetation – can take advantage of associational refuge during restoration. However, given rapid seedling detection by herbivores, associational refuge may be inadequate in the long‐term under high browsing pressure unless high absolute numbers of seedlings are planted among refuge.     

Litter type and termites regulate root decomposition across contrasting savanna land‐uses

Decomposition is a vital ecosystem process, increasingly modified by human activity. Theoretical frameworks and empirical studies that aim to understand the interplay between human land‐use, macro‐fauna and decomposition processes have primarily focused on leaf and wood litter. For a whole‐plant understanding of how land‐use and macro‐fauna influence decomposition, investigating root litter is required. Using litterbags, we quantified rates of root decomposition across contrasting tropical savanna land‐uses, namely wildlife and fire‐dominated protected areas and livestock pastureland without fire. By scanning litterbags for termite intrusion, we differentiated termite and microbial driven decomposition. Root litter was buried underneath different tree canopies (leguminous and non‐leguminous trees) and outside canopies to account for savanna landscape effects. Additionally, we established a termite cafeteria‐style experiment and common garden to explore termite selectivity of root litter and root trait relationships, respectively. After one year, we found no significant differences in root litter mass loss between wildlife dominated areas and pastureland. Instead, we found consistent species differences in root litter mass loss across land‐uses and additive and non‐additive effects of termites on root decomposition across plant species. Termite selectivity for root litter species occurred for both root and leaf litter buried near termite mounds, but was not explained by root traits measured in the common garden. Termite foraging was greater under leguminous tree canopies than other canopies; however, this did not influence rates of root decomposition. Our study suggests that land‐use has a weak direct effect on belowground processes in savannas. Instead, changes in herbaceous species composition and termite foraging have stronger impacts on belowground decomposition. Moreover, termites were not generalist decomposers of root litter, but their impact varies depending on plant species identity and likely associated root traits. This root litter selectivity by termites is likely to be an important contributor to spatial heterogeneity in savanna nutrient cycling.     

Effects of two locust control methods on wood-eating termites in arid Australia

Termites are ubiquitous detritivores and are a key influence on soil function and nutrient cycles, particularly in arid and semi-arid ecosystems. Locust control presents a unique hazard to termites and the effective functioning of ecosystems as a consequence of the overlap between pesticide applications and termite populations in grassland and desert landscapes. We monitored the effects of locust control methods using ultra-low-volume (ULV) barrier application of a chemical pesticide, fipronil, and a blanket application of a fungal biopesticide, Metarhizium acridum, on wood-eating termites in arid western New South Wales, Australia. We tested the hypothesis that spray applications decrease termite activity at wood baits using a BACI designed field experiment over 2 years. Our replicated control and treatment sites represented the spatial scale of Australian locust control activities. There was no detectable impact of either locust control treatment on termite activity, bait mass loss or termite community composition measures. Non-significant differences in termite survey measures among sites suggested that climate and environmental conditions were stronger drivers of our termite measures than the single, localized and unreplicated application of pesticides more commonly used in locust control operations in arid Australia. A lack of evidence for an impact of our fipronil or Metarhizium application methods supports their use as low hazard locust control options with minimal large scale and longer-term effects on termites in Australian arid rangelands. Future research would be necessary to determine the probable short-term impacts of treatments on individual termite colonies and the possible impacts on non-wood eating termite species in the arid-zone.