Effects of large herbivores on grassland arthropod diversity

Both arthropods and large grazing herbivores are important components and drivers of biodiversity in grassland ecosystems, but a synthesis of how arthropod diversity is affected by large herbivores has been largely missing. To fill this gap, we conducted a literature search, which yielded 141 studies on this topic of which 24 simultaneously investigated plant and arthropod diversity. Using the data from these 24 studies, we compared the responses of plant and arthropod diversity to an increase in grazing intensity. This quantitative assessment showed no overall significant effect of increasing grazing intensity on plant diversity, while arthropod diversity was generally negatively affected. To understand these negative effects, we explored the mechanisms by which large herbivores affect arthropod communities: direct effects, changes in vegetation structure, changes in plant community composition, changes in soil conditions, and cascading effects within the arthropod interaction web. We identify three main factors determining the effects of large herbivores on arthropod diversity: (i) unintentional predation and increased disturbance, (ii) decreases in total resource abundance for arthropods (biomass) and (iii) changes in plant diversity, vegetation structure and abiotic conditions. In general, heterogeneity in vegetation structure and abiotic conditions increases at intermediate grazing intensity, but declines at both low and high grazing intensity. We conclude that large herbivores can only increase arthropod diversity if they cause an increase in (a)biotic heterogeneity, and then only if this increase is large enough to compensate for the loss of total resource abundance and the increased mortality rate. This is expected to occur only at low herbivore densities or with spatio‐temporal variation in herbivore densities. As we demonstrate that arthropod diversity is often more negatively affected by grazing than plant diversity, we strongly recommend considering the specific requirements of arthropods when applying grazing management and to include arthropods in monitoring schemes. Conservation strategies aiming at maximizing heterogeneity, including regulation of herbivore densities (through human interventions or top‐down control), maintenance of different types of management in close proximity and rotational grazing regimes, are the most promising options to conserve arthropod diversity.     

Heterogeneous patterns of abundance of epigeic arthropod taxa along a major elevation gradient

Species diversity is the variable most commonly studied in recent ecological research. Ecological processes, however, are driven by individuals and affected by their abundances. Understanding the variation in animal abundances along climatic gradients is important for predicting changes in ecosystem processes under global warming. High abundances make arthropods, despite their small body sizes, important actors in food webs, yet abundance distributions of major arthropod taxa along climatic gradients remain poorly documented. We sampled arthropod assemblages in disturbed and undisturbed vegetation types along an elevational gradient of 860–4550 m asl on the southern slopes of Mt. Kilimanjaro, Tanzania. In our analysis, we focused on 13 taxa of arthropods that represented three major functional groups: predators, herbivores, and decomposers. Abundance patterns were unimodal for most of the taxa and functional groups, including decomposer arthropods, and most of them peaked at low elevations in lower montane forest. When we assigned beetles to functional groups, however, decomposer beetle abundances declined almost linearly, and abundances of predator beetles (ca. 2400 m asl) and herbivore beetles (ca. 3000 m asl, undisturbed vegetation) peaked at higher elevations and exhibited unimodal patterns. Temperature, not primary productivity, was the best predictor of abundance for most of the taxa and groups. Disturbance was only of minor importance. Our results revealed different trends in the response of arthropod abundance along the elevational gradient that depended on the level of taxonomic and functional resolution. This highlights the need for more comparisons of different taxa along the same climatic gradients.     

Biotic filtering and mass effects in small shrub patches: is arthropod community structure predictable based on the quality of the vegetation?

1. Community assembly is affected by four processes: dispersal, filtering effects (selection), ecological drift and evolution. The role of filtering relative to dispersal and drift should decline with patch size, hampering possibilities to predict which organisms will be observed within small‐sized patches. However, vegetation structure is known to have a marked impact on species assemblages, and plant quality may act as a biotic filter. This challenges the assumption of unpredictable species assemblages in small‐sized vegetation patches. 2. Using 32 stands of five shrub species in south‐west Finland, this study investigated whether biotic filtering effects caused by patch‐forming plants are strong enough to overcome the mixing of mobile arthropod assemblages across small patches. 3. Stochastic variation did not hide the signals of biotic filtering and dispersal in the small shrub patches. Habitat richness around the patches explained a three times larger share of variation in the species composition than did the identity of the patch‐forming plant, but it had less effect on the abundance of arthropods. A radius of 50–100 m around a patch explained the species composition best. 4. Abundance patterns varied between the feeding guilds; the patch‐forming shrub influenced the abundances of detritivores and leaf‐feeding herbivores, whereas the abundances of flower‐visiting herbivores appeared to track the flowering phenology of the plants. Shrub identity had little effect on omnivores or predators. Predator abundances were correlated with the abundance of potential prey. 5. The results of this study suggest that community composition within a vegetation patch may be predictable even if dispersal overrides local filtering effects, as suggested by the mass‐effects paradigm.     

Mammalian herbivores,grass height and rainfall drive termite activity at different spatial scales in an African savanna

Termites have a large influence on ecosystem functioning. Understanding what drives termite activity patterns improves understanding of nutrient cycling, productivity, and heterogeneity in savannas. We present a mechanistic framework that relates the interactive effects of rainfall, grassland structure, large herbivore presence, and soil factors to termite activity. To test this framework, we used grass litterbags to monitor termite activity at ten sites across Hluhluwe‐iMfolozi Park, South Africa. We assessed the effects of abiotic and biotic factors on termite activity at two scales: the large (landscape) scale, variation in bait removal among 300 m² plots that were distributed across the park and at the small (within‐plot) scale (1–300 m²). Half of our sites were located inside large herbivore exclosures to test for the effect of mammalian herbivore presence. At the landscape scale, termite grass removal declined towards higher rainfall and in the presence of mammalian herbivores. Removal did not depend on soil factors. At the small scale, removal declined with increasing grass height, particularly in the 1 m surrounding the bait bag. Resource quality did not affect bait removal. We suggest that competition for forage drives the negative effect of mammalian herbivores on termites, whereas lower bait removal in taller swards may be due to direct negative effects from rainfall, fire and/or competition with free‐living microbes. Ultimately, we suggest that the impact of termites on nutrient cycling is most pronounced when abiotic (rainfall) and biotic conditions (mammalian herbivory) limit grass removal by fire and decomposition by free‐living microbes.     

Context‐dependent biotic interactions control plant abundance across altitudinal environmental gradients

Many biotic interactions influence community structure, yet most distribution models for plants have focused on plant competition or used only abiotic variables to predict plant abundance. Furthermore, biotic interactions are commonly context‐dependent across abiotic gradients. For example, plant–plant interactions can grade from competition to facilitation over temperature gradients. We used a hierarchical Bayesian framework to predict the abundances of 12 plant species across a mountain landscape and test hypotheses on the context‐dependency of biotic interactions over abiotic gradients. We combined field‐based estimates of six biotic interactions (foliar herbivory and pathogen damage, fungal root colonization, fossorial mammal disturbance, plant cover and plant diversity) with abiotic data on climate and soil depth, nutrients and moisture. All biotic interactions were significantly context‐dependent along temperature gradients. Results supported the stress gradient hypothesis: as abiotic stress increased, the strength or direction of the relationship between biotic variables and plant abundance generally switched from negative (suggesting suppressed plant abundance) to positive (suggesting facilitation/mutualism). For half of the species, plant cover was the best predictor of abundance, suggesting that the prior focus on plant–plant interactions is well‐justified. Explicitly incorporating the context‐dependency of biotic interactions generated novel hypotheses about drivers of plant abundance across abiotic gradients and may improve the accuracy of niche models.     

Biogeography of parasitism in freshwater fish: spatial patterns in hot spots of infection

Contrary to species occurrence, little is known about the determinants of spatial patterns of intraspecific variation in abundance, particularly for parasitic organisms. In this study, we provide a multi‐faceted overview of spatial patterns in parasite abundance and examine several potential underlying processes. We first tested for a latitudinal gradient in local abundance of the regionally most common parasite species and whether these species achieve higher abundances at the same localities (shared hot spots of infection). Secondly, we tested whether intraspecific similarity in local abundance between sites follows a spatial distance decay pattern or is better explained by variation in extrinsic biotic and abiotic factors between localities related to local parasite transmission success. We examined the infection landscape of a model fish host system (common and upland bullies, genus Gobiomorphus: Eleotridae) across its entire distributional range. We applied general linear models to test the effect of latitude on each species local abundance independently, including the abundance of each co‐infecting species as another predictor. We computed multiple regressions on distance matrices among localities based on abundance of each of the four most common trematode species, as well as for geographic distance, biotic and abiotic distinctness of the localities. Our results showed that the most widely distributed parasites of bullies also achieve the highest mean local abundances, following the abundance – occupancy relationship. Variation in local abundance of any focal parasite species was independent of latitude, the abundance of co‐occurring species and spatial distance or disparity in biotic attributes between localities. For only one parasite species, similarity of abundance between sites covaried with the extent of abiotic differences between sites. The lack of association between hot spots of infection for co‐occurring species reinforces the geographic mosaic scenario in which hosts and parasites coevolve by suggesting non‐deterministic, species‐specific variation in parasite abundance across space.     

Salinity cascading and predator effects on consumer and producer community structure: a mesocosm experiment

1. An observed community structure is often shaped by a combination of bottom-top and top-bottom processes, affected by biotic and abiotic factors. These factors affect community structure either directly (each species separately) or indirectly, via other species. Few studies have observed the combined effects of biotic and abiotic factors on primary producers and consumer communities composed of numerous species. 2. This study investigated the combined effects of a predatory hemipteran and water salinity on abundance and species richness of the periphyton and arthropod communities in an artificial pool experiment. 3. The abundance and species richness of both periphyton and arthropod communities were affected by the combination of salinity and predator. 4. It is suggested that the observed consumer community is composed of arthropod species that vary in salinity tolerance and response to the predator. In addition, the abundance of euryhaline species or species that are not consumed by this particular predator increases indirectly in specific treatment combinations that eventually result in reduced interspecific competition. Periphyton variables were strongly correlated to densities of larval Ochlerotatus caspius (Diptera: Culicidae), suggesting that this species may be largely responsible for structuring the producer community. 5. It is suggested that O. caspius distribution is the result of female oviposition habitat selection that is manipulated by chemical signals. Therefore, chemical signals that inform about habitat suitability also play an important role in shaping both consumer and primary producer communities.     

Effects of bird predation on canopy arthropods in wandoo Eucalyptus wandoo woodland

Abstract: As top predators, birds may have significant effects on arthropod abundances and affect the trophic structure of arthropod communities through predation of lower order predators (e.g. spiders) and by competition for prey. We investigated the effects of bird predation on canopy arthropods in south‐western Australia by using plastic bird mesh to exclude insectivorous birds from the foliage of wandoo Eucalyptus wandoo saplings. Exclosure resulted in an increase in the number of herbivorous and predatory arthropods. Total arthropods (with and without ants), spiders, adult Coleoptera, and larval Lepidoptera were significantly more abundant on meshed than unmeshed saplings. All size‐classes of arthropods, taxa grouped, were more abundant on meshed than unmeshed saplings, but with no evidence of a disproportionate increase of the largest arthropods on meshed saplings. All size‐classes of spiders increased in abundance on saplings from which birds were excluded. There were significant differences in the total abundance of arthropods (with and without ants), spiders (Araneae), sucking bugs (Homoptera), adult beetles (Coleoptera), larval moths (Lepidoptera), and wasps and ants (Hymenoptera) for both unmeshed and meshed saplings between sample periods. These seasonal patterns of abundance and differences between sample periods appeared to be determined by seasonal weather patterns, with the lowest numbers associated with drier and hotter conditions in summer and autumn than in winter and spring. The conclusion reached is that eucalypt forest birds have limited effects on temporal variation in canopy arthropod abundances, but depress abundances, and affect the size and trophic composition of the fauna. Given the cascading effects of birds as predators on arthropods, successful conservation management of eucalypt ecosystems, including plantations and revegetation, should be planned to maximize bird numbers and diversity.     

Multiple Environmental Controls on Cockroach Assemblage Structure in a Tropical Rain Forest

Arthropod abundance and diversity are remarkable in tropical forests, but are also spatially patchy. This has been attributed either to resources, predators, abiotic conditions or disturbances, but whether such factors may simultaneously shape arthropod assemblage structure is little known. We used cockroaches to test for multiple environmental controls on assemblage structure in 25 km² of Amazonian forest. We performed nocturnal, direct searches for cockroaches in 30 plots (250 m × 2 m) during two seasons, and gathered data on biotic and abiotic factors from previous studies. Cockroach abundance increased with dry litter mass, a measure of resource amount, while species richness increased with litter phosphorus content, a measure of resource availability. Cockroach abundance and species richness decreased with ant relative abundance. Cockroach species composition changed along the gradient of: (1) soil clay content, which correlates with a broad differentiation between flood‐prone and non‐flooded forest; (2) soil relative moisture, consistent with known interspecific variation in desiccation tolerance; and (3) according to the abundance of ants, a potential predator. Turnover in species composition was correlated with abiotic conditions—sorting species according to physiological requirements and to disturbance‐related life history traits—and to ants' selective pressure. Cockroach abundance, diversity, and composition seem to be controlled by distinct sets of environmental factors, but predators which were represented by ants, emerged as a common factor underlying cockroach distribution. Such patterns of community structure may have been previously overlooked by undue focus on single or a few factors, and may be common to tropical forest arthropods.     

The Multiple Impacts of Tropical Forest Fragmentation on Arthropod Biodiversity and on their Patterns of Interactions with Host Plants

Tropical rain forest fragmentation affects biotic interactions in distinct ways. Little is known, however, about how fragmentation affects animal trophic guilds and their patterns of interactions with host plants. In this study, we analyzed changes in biotic interactions in forest fragments by using a multitrophic approach. For this, we classified arthropods associated with Heliconia aurantiaca herbs into broad trophic guilds (omnivores, herbivores and predators) and assessed the topological structure of intrapopulation plant-arthropod networks in fragments and continuous forests. Habitat type influenced arthropod species abundance, diversity and composition with greater abundance in fragments but greater diversity in continuous forest. According to trophic guilds, coleopteran herbivores were more abundant in continuous forest and overall omnivores in fragments. Continuous forest showed a greater diversity of interactions than fragments. Only in fragments, however, did the arthropod community associated with H aurantiaca show a nested structure, suggesting novel and/or opportunistic host-arthropod associations. Plants, omnivores and predators contributed more to nestedness than herbivores. Therefore, Heliconia-arthropod network properties do not appear to be maintained in fragments mainly caused by the decrease of herbivores. Our study contributes to the understanding of the impact of fragmentation on the structure and dynamics of multitrophic arthropod communities associated with a particular plant species of the highly biodiverse tropical forests. Nevertheless, further replication of study sites is needed to strengthen the conclusion that forest fragmentation negatively affects arthropod assemblages.     

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.     

Flow‐mediated predator–prey dynamics influence fish populations in a tropical river

相似文献   

An invasive plant-fungal mutualism reduces arthropod diversity

Ecological theory holds that competition and predation are the most important biotic forces affecting the composition of communities. Here, we expand this framework by demonstrating that mutualism can fundamentally alter community and food web structure. In large, replicated field plots, we manipulated the mutualism between a dominant plant ( Lolium arundinaceum ) and symbiotic fungal endophyte ( Neotyphodium coenophialum ). The presence of the mutualism reduced arthropod abundance up to 70%, reduced arthropod diversity nearly 20%, shifted arthropod species composition relative to endophyte-free plots and suppressed the biomass and richness of other plant species in the community. Herbivorous arthropods were more strongly affected than carnivores, and for both herbivores and carnivores, effects of the mutualism appeared to propagate indirectly via organisms occurring more basally in the food web. The influence of the mutualism was as great or greater than previously documented effects of competition and predation on arthropod communities. Our work demonstrates that a keystone mutualism can significantly reduce arthropod biodiversity at a broad community scale.     

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.     

Local-scale drivers of tree survival in a temperate forest

Tree survival plays a central role in forest ecosystems. Although many factors such as tree size, abiotic and biotic neighborhoods have been proposed as being important in explaining patterns of tree survival, their contributions are still subject to debate. We used generalized linear mixed models to examine the relative importance of tree size, local abiotic conditions and the density and identity of neighbors on tree survival in an old-growth temperate forest in northeastern China at three levels (community, guild and species). Tree size and both abiotic and biotic neighborhood variables influenced tree survival under current forest conditions, but their relative importance varied dramatically within and among the community, guild and species levels. Of the variables tested, tree size was typically the most important predictor of tree survival, followed by biotic and then abiotic variables. The effect of tree size on survival varied from strongly positive for small trees (1-20 cm dbh) and medium trees (20-40 cm dbh), to slightly negative for large trees (>40 cm dbh). Among the biotic factors, we found strong evidence for negative density and frequency dependence in this temperate forest, as indicated by negative effects of both total basal area of neighbors and the frequency of conspecific neighbors. Among the abiotic factors tested, soil nutrients tended to be more important in affecting tree survival than topographic variables. Abiotic factors generally influenced survival for species with relatively high abundance, for individuals in smaller size classes and for shade-tolerant species. Our study demonstrates that the relative importance of variables driving patterns of tree survival differs greatly among size classes, species guilds and abundance classes in temperate forest, which can further understanding of forest dynamics and offer important insights into forest management.     

Asexual endophytes and associated alkaloids alter arthropod community structure and increase herbivore abundances on a native grass

Despite their minute biomass, microbial symbionts of plants potentially alter herbivory, diversity and community structure. Infection of grasses by asexual endophytic fungi often decreases herbivore loads and alters arthropod diversity. However, most studies to date have involved agronomic grasses and often consider only infection status (infected vs. uninfected), without explicitly measuring endophyte-produced alkaloids, which vary among endophyte isolates and may impact consumers. We combined field experiments and population surveys to investigate how endophyte infection and associated alkaloids influence abundances, species richness, evenness and guild structure of arthropod communities on a native grass, Achnatherum robustum (sleepygrass). Surprisingly, we found that endophyte-produced alkaloids were associated with increased herbivore abundances and species richness. Our results suggest that, unlike what has been found in agronomic grass systems, high alkaloid levels in native grasses may not protect host grasses from arthropod herbivores, and may instead more negatively affect natural enemies of herbivores.
Ecology Letters (2010) 13: 106–117     

Abundance and diversity of birch‐feeding leafminers along latitudinal gradients in northern Europe

Latitudinal patterns in biotic interactions, including herbivory, have been widely debated during the past years. In particular, recent meta‐analysis questioned the hypothesis that herbivory increases from the poles towards the equator. Our study was designed to verify this hypothesis by exploring latitudinal patterns in abundance and diversity of birch‐feeding insect herbivores belonging to the leafminer guild in northern Europe, from 59° to 69°N. We collected branches from five mature trees of two birch species (Betula pendula and B. pubescens) at each study site (ten sites for each of five latitudinal gradients) twice per season (in early and late summer of 2008–2011) and attributed all mines found on leaves of these branches to a certain taxon of insects. Latitudinal patterns were quantified by calculating Spearman rank correlation coefficients between both abundance and diversity of leafmining taxa and latitudes of sampling sites. In general, both abundance and diversity of leafminers significantly decreased with latitude. However, we discovered pronounced variation in patterns of latitudinal changes among study years and leafminer taxa. Variation among study years was best explained by mean temperatures in July at the northern ends of our gradients. During cold years, abundance of leafminers significantly decreased with latitude, while during warm years the abundance was either independent of latitude or even increased towards the pole. In the northern boreal forests (66° to 69°N), herbivores demonstrated larger changes in densities in response to temperature variations than in the boreo‐nemoral forests (59° to 62°N). Our data suggest that climate warming will result in a stronger increase in herbivory at higher latitudes than at lower latitudes.     

Disturbance,fish, and variation in the predatory insect guild of coastal streams

Species distributions, and thus the composition of communities, are determined by many interacting biotic and abiotic factors. We analyzed the variation in composition of the invertebrate predator guild across eight small, steep coastal streams in eastern Canada, in relation to variation in several broad categories of environmental variables: disturbance-related physical characteristics, top-down effects (fish), and bottom-up effects (prey composition and productivity-related physical/chemical variables). Similarity in composition (relative and absolute abundances of the 19 species belonging to the Rhyacophilidae, Perlodidae, and Chloroperlidae) declined significantly with distance. Streams that were most similar in predator composition were also most similar in physical factors related to disturbance, but were not more similar in prey abundance/composition or in environmental factors associated with productivity (chemistry, canopy cover, and riparian forest type). Similarity in the relative abundance of species within the Rhyacophilidae was linked to variation in the presence/abundance of brook trout. These results suggest that the invertebrate predator guilds of these small coastal streams are structured first and foremost by factors associated with the magnitude and variability of the flow regime, and second by response to top-down factors (predation by brook trout). There was no evidence that bottom-up effects altered the composition or abundance of the predator guild. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Sonja Stendera     

Nonequilibrial community structure of sawflies on arroyo willow

Summary We use Wiens' (1984) nonequilibrium-equilibrium continuum concept to evaluate aspects of community structure for sawfly herbivores (Hymenoptera: Tenthredinidae) that attack arroyo willow, Salix lasiolepis. The sawfly community on arroyo willow shares many characteristics of nonequilibrium communities: unsaturation, biotic decoupling, lack of density dependence, and loose emergent patterns. Species abundances exhibit highly significant differences among clones (P<0.001) and among years (P<0.001) and exhibit significant and complex clone-by-year interactions both for absolute and relative abundances. Organizing forces, such as interspecific competition and regulation by natural enemies, are largely absent from the arroyo willow system. Sawfly densities exhibit significant positive covariation among willow clones and appear to be affected by largely stochastic abiotic forces, primarily winter precipitation.