Trait-dependent declines of species following conversion of rain forest to oil palm plantations

Conversion of natural habitats to agriculture reduces species richness, particularly in highly diverse tropical regions, but its effects on species composition are less well-studied. The conversion of rain forest to oil palm is of particular conservation concern globally, and we examined how it affects the abundance of birds, beetles, and ants according to their local population size, body size, geographical range size, and feeding guild or trophic position. We re-analysed data from six published studies representing 487 species/genera to assess the relative importance of these traits in explaining changes in abundance following forest conversion. We found consistent patterns across all three taxa, with large-bodied, abundant forest species from higher trophic levels, declining most in abundance following conversion of forest to oil palm. Best-fitting models explained 39–66 % of the variation in abundance changes for the three taxa, and included all ecological traits that we considered. Across the three taxa, those few species found in oil palm tended to be small-bodied species, from lower trophic levels, that had low local abundances in forest. These species were often hyper-abundant in oil palm plantations. These results provide empirical evidence of consistent responses to land-use change among taxonomic groups in relation to ecological traits.     

Aerial arthropod communities of native and invaded forests, Robinson Crusoe Island, Chile

Invasive species significantly contribute to biological change and threaten biodiversity, with a growing body of evidence that plant invasions affect higher trophic levels. We explored the relative importance of plant invasion and forest structure on aerial arthropod abundance, diversity, and composition on Robinson Crusoe Island, Chile. We used flight intercept traps to sample aerial arthropods within distinct canopy strata of native and invaded forests over 3-mo periods in 2006 and 2007. Arthropod abundance and diversity were higher in native than invaded forest, and arthropod communities were distinct between forest types. In both forest types, arthropod abundance was highest in the lower canopy, and canopy strata exhibited some differences in arthropod community composition. Several morphospecies were distinctly associated with each forest type. The strong differences in aerial arthropod communities associated with the invasion of native forest by non-native plants may affect other trophic levels, such as insectivorous birds. Steps to stop invasive plant spread and to restore native forest composition and structure are needed to safeguard the integrity of native communities, from plants to higher-level consumers.     

Trees as templates for tropical litter arthropod diversity

Increased tree species diversity in the tropics is associated with even greater herbivore diversity, but few tests of tree effects on litter arthropod diversity exist. We studied whether tree species influence patchiness in diversity and abundance of three common soil arthropod taxa (ants, gamasid mites, and oribatid mites) in a Panama forest. The tree specialization hypothesis proposes that tree-driven habitat heterogeneity maintains litter arthropod diversity. We tested whether tree species differed in resource quality and quantity of their leaf litter and whether more heterogeneous litter supports more arthropod species. Alternatively, the abundance–extinction hypothesis states that arthropod diversity increases with arthropod abundance, which in turn tracks resource quantity (e.g., litter depth). We found little support for the hypothesis that tropical trees are templates for litter arthropod diversity. Ten tree species differed in litter depth, chemistry, and structural variability. However, the extent of specialization of invertebrates on particular tree taxa was low and the more heterogeneous litter between trees failed to support higher arthropod diversity. Furthermore, arthropod diversity did not track abundance or litter depth. The lack of association between tree species and litter arthropods suggests that factors other than tree species diversity may better explain the high arthropod diversity in tropical forests.     

Spiders and Other Arthropods as Indicators in Old-Growth Versus Logged Redwood Stands

This study, conducted in the Santa Cruz Mountains of California, provides data on the abundance and diversity of litter spiders and other arthropods in three redwood forest conditions: old growth, second growth, and tree farm. Litter spiders are linked to and reflect habitat structure and prey abundance and can act as indicators for redwood forest restoration and monitoring. There were significant declines in spider and other arthropod diversity and abundance with increased logging and decreased herb cover. The absolute and relative increase in nocturnal spiders and detritivores in unlogged sites suggests that guild structures of spiders and other arthropods can indicate forest recovery from logging disturbance. Furthermore, selectively harvested stands do not retain old‐growth levels of litter arthropod diversity or abundance. This study identifies potential indicator redwood litter spiders that show higher abundances in old‐growth areas, Zelotes sp. (Gnaphosidae), Xysticus sp. (Thomisidae), and Ceratinops inflata (Linyphiidae) and a possible old‐growth specialist, Phrurotimpus sp. (Liocranidae). These findings strengthen the case for including soil arthropods in redwood forest monitoring and assessment and for the preservation of undisturbed forest areas.     

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.     

Comparative patterns in taxonomic and functional spider diversities between tropical vs. temperate forests

High diversity in tropical compared to temperate regions has long intrigued ecologists, especially for highly speciose taxa like terrestrial arthropods in tropical rainforests. Previous studies showed that arthropod herbivores account for much tropical diversity, yet differences in the diversity of predatory arthropods between tropical and temperate systems have not been properly quantified. Here, we present the first standardized tropical–temperate forest quantification of spider diversities, a dominant and mega‐diverse taxon of generalist predators. Spider assemblages were collected using a spatially replicated protocol including two standardized sampling methods (vegetation sweep netting and beating). Fieldwork took place between 2010 and 2015 in metropolitan (Brittany) and overseas (French Guiana) French territories. We found no significant difference in functional diversity based on hunting guilds between temperate and tropical forests, while species richness was 13–82 times higher in tropical versus temperate forests. Evenness was also higher, with tropical assemblages up to 55 times more even than assemblages in temperate forests. These differences in diversity far surpass previous estimates and exceed tropical–temperate ratios for herbivorous taxa.     

Arthropods in epiphytes: a diversity component that is not effectively sampled by canopy fogging

Insecticide fogging is often used to document arthropod species richnessin forest canopies, but this technique may not effectively sample invertebratesthat are concealed within a variety of microhabitats. We quantified the effectsof fogging on invertebrates in canopy epiphyte mats of a Costa Rican cloudforest by extracting arthropods from 18 paired pre- and post-fogging samples.Mean abundance and morphospecies richness of living arthropods were respectivelyreduced by 33 and 30% in epiphyte material after fogging, but most organismssurvived the treatment. Relative abundances of major taxa were unaffected byfogging. Herbivores were less abundant after fogging than other trophic groups,and the median body length of non-mite arthropods present in epiphytes wassignificantly smaller after fogging. Examination of seven post-fogging samplesshowed that many arthropods killed by insecticide remained trapped within theepiphyte material. These results provide the first quantitative assessment of aspecific component of arboreal arthropod biodiversity that is missed by thefogging technique.     

Arthropod Distribution in a Tropical Rainforest: Tackling a Four Dimensional Puzzle

Quantifying the spatio-temporal distribution of arthropods in tropical rainforests represents a first step towards scrutinizing the global distribution of biodiversity on Earth. To date most studies have focused on narrow taxonomic groups or lack a design that allows partitioning of the components of diversity. Here, we consider an exceptionally large dataset (113,952 individuals representing 5,858 species), obtained from the San Lorenzo forest in Panama, where the phylogenetic breadth of arthropod taxa was surveyed using 14 protocols targeting the soil, litter, understory, lower and upper canopy habitats, replicated across seasons in 2003 and 2004. This dataset is used to explore the relative influence of horizontal, vertical and seasonal drivers of arthropod distribution in this forest. We considered arthropod abundance, observed and estimated species richness, additive decomposition of species richness, multiplicative partitioning of species diversity, variation in species composition, species turnover and guild structure as components of diversity. At the scale of our study (2km of distance, 40m in height and 400 days), the effects related to the vertical and seasonal dimensions were most important. Most adult arthropods were collected from the soil/litter or the upper canopy and species richness was highest in the canopy. We compared the distribution of arthropods and trees within our study system. Effects related to the seasonal dimension were stronger for arthropods than for trees. We conclude that: (1) models of beta diversity developed for tropical trees are unlikely to be applicable to tropical arthropods; (2) it is imperative that estimates of global biodiversity derived from mass collecting of arthropods in tropical rainforests embrace the strong vertical and seasonal partitioning observed here; and (3) given the high species turnover observed between seasons, global climate change may have severe consequences for rainforest arthropods.     

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.     

Impacts of the invader giant reed (<Emphasis Type="Italic">Arundo donax</Emphasis>) on riparian habitats and ground arthropod communities

Riparian areas have experienced long-term anthropogenic impacts including the effects of plant introductions. In this study, 27 plots were surveyed across three Mediterranean rivers in north-eastern Spain to explore the effects of the invader giant reed (Arundo donax) on riparian habitat features and the diversity, trophic structure, body size, and abundances of epigeal and hypogeal arthropods in riparian areas. Using pitfall traps and Berlese funnels, this study detected a significant increase in collembola abundance and a decrease in the abundance, body size and diversity of macro-arthropods at order and family levels in invaded plots compared to native stands. Invaded and un-invaded areas also differed in the taxonomical structure of arthropod assemblies but not in trophic guild proportions. However, the fact that arthropods were smaller in A. donax soils, together with the absence of particular taxa within each trophic guild or even an entire trophic group (parasitoids), suggests that food-web alterations in invaded areas cannot be discarded. Habitat features also differed between invaded and un-invaded areas with the poorest herbaceous understory and the largest leaf litter deposition and soil carbon stock observed in A. donax plots. The type of vegetation in riparian areas followed by the total native plant species richness were identified as major causal factors to changes in the abundance, diversity and composition of macro-arthropods. However, our analyses also showed that some alterations related to A. donax invasion were inconsistent across rivers, suggesting that A. donax effects may be context dependent. In conclusion, this study highlights an impoverishment of native flora and arthropod fauna in A. donax soils, and suggests major changes in riparian food webs if A. donax displaces native riparian vegetation.     

Tree diversity drives diversity of arthropod herbivores,but successional stage mediates detritivores

The high tree diversity of subtropical forests is linked to the biodiversity of other trophic levels. Disentangling the effects of tree species richness and composition, forest age, and stand structure on higher trophic levels in a forest landscape is important for understanding the factors that promote biodiversity and ecosystem functioning. Using a plot network spanning gradients of tree diversity and secondary succession in subtropical forest, we tested the effects of tree community characteristics (species richness and composition) and forest succession (stand age) on arthropod community characteristics (morphotype diversity, abundance and composition) of four arthropod functional groups. We posit that these gradients differentially affect the arthropod functional groups, which mediates the diversity, composition, and abundance of arthropods in subtropical forests. We found that herbivore richness was positively related to tree species richness. Furthermore, the composition of herbivore communities was associated with tree species composition. In contrast, detritivore richness and composition was associated with stand age instead of tree diversity. Predator and pollinator richness and abundance were not strongly related to either gradient, although positive trends with tree species richness were found for predators. The weaker effect of tree diversity on predators suggests a cascading diversity effect from trees to herbivores to predators. Our results suggest that arthropod diversity in a subtropical forest reflects the net outcome of complex interactions among variables associated with tree diversity and stand age. Despite this complexity, there are clear linkages between the overall richness and composition of tree and arthropod communities, in particular herbivores, demonstrating that these trophic levels directly impact each other.     

Invertebrate responses to the management of genetically modified herbicide-tolerant and conventional spring crops. II. Within-field epigeal and aerial arthropods

The effects of the management of genetically modified herbicide-tolerant (GMHT) crops on the abundances of aerial and epigeal arthropods were assessed in 66 beet, 68 maize and 67 spring oilseed rape sites as part of the Farm Scale Evaluations of GMHT crops. Most higher taxa were insensitive to differences between GMHT and conventional weed management, but significant effects were found on the abundance of at least one group within each taxon studied. Numbers of butterflies in beet and spring oilseed rape and of Heteroptera and bees in beet were smaller under the relevant GMHT crop management, whereas the abundance of Collembola was consistently greater in all GMHT crops. Generally, these effects were specific to each crop type, reflected the phenology and ecology of the arthropod taxa, were indirect and related to herbicide management. These results apply generally to agriculture across Britain, and could be used in mathematical models to predict the possible long-term effects of the widespread adoption of GMHT technology. The results for bees and butterflies relate to foraging preferences and might or might not translate into effects on population densities, depending on whether adoption leads to forage reductions over large areas. These species, and the detritivore Collembola, may be useful indicator species for future studies of GMHT management.     

The influence of habitat structure on arthropod diversity in Argentine semi-arid Chaco forest

Abstract. The Argentine Chaco is a mosaic of grassland and open forest habitats maintained by natural disturbance activities such as fire. Since the introduction of domestic livestock and other human activities, the balance of this mosaic has been significantly altered, both in plant species and structural composition. This study focuses on the impact of such changes on the diversity of ground-dwelling arthropods within semi-arid Chaco forest. Quantitative measures of habitat structure and arthropod diversity were taken in forest areas previously subjected to grazing, logging and ploughing. Results indicated that arthropod diversity was smaller on sites with reduced structural complexity, with marked changes in arthropod family composition. The habitat components relating to plant architectural and vertical diversity were particularly influential on arthropod diversity. The guild size ratio of predatory to non-predatory arthropods also differed significantly between habitats suggesting a change in the resource base available to some groups. The latter suggests a shift in the functional organisation of the forest ecosystem which could have important repercussions for the diversity of other trophic levels.     

The impact of high-altitude ski pistes on ground-dwelling arthropods in the Alps

The development of winter sport resorts above the timberline may affect every ecosystem component. We analyzed the effect of ski-pistes on the abundance and species richness of arthropods (namely carabids, spiders, opilionids, and grasshoppers) trapped in grasslands adjacent to the ski-run, on ski-pistes and at the edge between these two habitat types. Our results showed that diversity of brachypterous carabids, spiders, and grasshoppers decreased significantly from natural grasslands to ski-pistes. This was not true for the macropterous carabid guild, which included species with contrasting ecological requirements. The analysis of indicator species (IndVal) showed that most of the species (some of them precinctive to restricted areas in the north-western Alps) had clear preferences for natural grassland and few taxa were limited to ski-pistes. Generalized linear models suggested that the local extent of grass and rock cover can significantly affect assemblages: the low grass cover of ski-pistes, in particular, was a serious hindrance to colonization by spider, grasshopper, brachypterous, and some macropterous carabid species. The results obtained, support concerns over the possible disruption of local ecosystem functionality and over the conservation of arthropod species which are endemic to restricted alpine areas. In order to retain arthropod ground-dwelling fauna we suggest that: (i) new, environmentally friendly ways of constructing pistes should be developed to preserve as much soil and grass cover as possible; (ii) existing ski-pistes should be restored through management to promote the recovery of local vegetation.     

Invertebrates in the canopy of tropical rain forests How much do we really know?

The current state of knowledge of canopy invertebrates in tropical rain forests is reviewed using data drawn, without bias toward taxon, collecting method or biogeographical region, from 89 studies concerned with mass-collecting (>1000 individuals). The review is intended to identify the most serious gaps and biases in the distribution of higher taxa among forest types and biogeographical regions. With respect to knowledge, biogeographical regions can be ranked as Neotropical > Australian > Oriental > Afrotropical. The canopy of lowland wet and subtropical forests has been studied in greater detail, whereas the canopy of lowland dry and montane forests is much less well known. Collecting techniques influence greatly the present knowledge of canopy invertebrates. Invertebrates other than arthropods, often abundant in epiphytic habitats, phytotelmata and perched litter, are virtually unknown. The abundance of several groups, such as Acari, Collembola and Isoptera, is almost certainly seriously underestimated. Densities of invertebrate individuals in the canopy of tropical rain forests appear to be lower than in temperate forests, although invertebrate abundance is dissipated by the high standing-biomass of rain forests. Coleoptera, particularly Staphylinidae, Curculionidae and Chrysomelidae, along with Hymenoptera, Lepidoptera and Araneae appear to be the most speciose taxa in the canopy, and it is probable that this reflects their range of feeding habits and exploitation of rain forests habitats. The distribution of individuals among the major arthropod orders and across the studies examined is complex and depends on many factors. The amount of variance that can be directly explained by biogeography, forest types (subtropical, wet, dry or montane), or collecting methods appears to be about 11%. The explained variance increases when considering major families of Coleoptera (28%) or subfamilies of Chrysomelidae (40%). In all cases, the variance explained by the type of forest is much higher than by that explained by biogeography. These conclusions are similar when considering various prey-predator relationships in the canopy. This suggests that, at the higher taxa level, the composition of the invertebrate fauna in the canopy may vary comparatively more across forest types than across biogeographical regions and this is discussed briefly from a conservation viewpoint.     

Effects of barrier application of granular deltamethrin on subadult Ixodes scapularis (Acari: Ixodidae) and nontarget forest floor arthropods

We evaluated the effects of a single application of granular deltamethrin made against nymphal Ixodes scapularis Say on the diversity and abundance of forest arthropods taken in pitfall traps in oak forest sites for 16 wk after treatment in central New Jersey. Control of I. scapularis subadults on treated plots ranged between 97 and 100% and continued at least 12 wk postapplication. Significant short-term changes in arthropod assemblages were detected at one of three study sites within 4 wk posttreatment. Effects were not distributed equally across taxa. Seasonal changes in numbers and diversity of forest arthropods in the study areas may have affected the impact of the acaricide in the treatment area. Comparison with control areas indicated that reductions in abundance of some arthropod taxa in the treatment area were detectable 12 wk after treatment. Total arthropod species diversity was not significantly affected by the application, and no treatment effects were detected 16 wk postapplication, suggesting that the arthropod community had recovered from the effects of the application. The merits of barrier applications in integrated tick control programs are discussed.     

Tree Species Composition and Harvest Intensity Affect Herbivore Density and Leaf Damage on Beech,Fagus sylvatica,in Different Landscape Contexts

Most forests are exposed to anthropogenic management activities that affect tree species composition and natural ecosystem processes. Changes in ecosystem processes such as herbivory depend on management intensity, and on regional environmental conditions and species pools. Whereas influences of specific forest management measures have already been addressed for different herbivore taxa on a local scale, studies considering effects of different aspects of forest management across different regions are rare. We assessed the influence of tree species composition and intensity of harvesting activities on arthropod herbivores and herbivore-related damage to beech trees, Fagus sylvatica, in 48 forest plots in three regions of Germany. We found that herbivore abundance and damage to beech trees differed between regions and that – despite the regional differences - density of tree-associated arthropod taxa and herbivore damage were consistently affected by tree species composition and harvest intensity. Specifically, overall herbivore damage to beech trees increased with increasing dominance of beech trees – suggesting the action of associational resistance processes – and decreased with harvest intensity. The density of leaf chewers and mines was positively related to leaf damage, and several arthropod groups responded to beech dominance and harvest intensity. The distribution of damage patterns was consistent with a vertical shift of herbivores to higher crown layers during the season and with higher beech dominance. By linking quantitative data on arthropod herbivore abundance and herbivory with tree species composition and harvesting activity in a wide variety of beech forests, our study helps to better understand the influence of forest management on interactions between a naturally dominant deciduous forest tree and arthropod herbivores.     

River drying lowers the diversity and alters the composition of an assemblage of desert riparian arthropods

Summary 1. Many studies have shown negative effects of river drying on in‐stream animals. However, the influence of river drying on riparian animals remains poorly studied. We examined ground‐dwelling riparian arthropod assemblages along a drying section of the semi‐arid San Pedro River in southeastern Arizona, U.S.A. 2. We found strong differences in assemblage composition, taxon diversity and the abundance of key taxa between dry and flowing sites, with higher diversity and abundance of most taxa at flowing sites. 3. Changes in assemblage composition, taxon diversity and abundance of representative taxa were associated with a combined measure of water availability that included distance to water and type of water. Other environmental variables showed a weaker association with changes in these arthropod assemblages. 4. Thus, we found evidence that desert riparian arthropods are sensitive to river drying and to reduction in water resources. Increases in drying along this river may reduce the diversity and the abundance of many groups of ground‐dwelling arthropods, leading to marked shifts in community composition.     

Effects of an application of granular carbaryl on nontarget forest floor arthropods

We evaluated the effects of a single application of granular carbaryl made against nymphal Ixodes scapularis Say on the diversity and abundance of forest arthropods taken in pitfall traps in oak and mixed oak-pine forest sites for 12 wk after treatment in central New Jersey. Significant short-term changes in arthropod assemblages were detected immediately posttreatment. Effects were not distributed equally across taxa. Seasonal changes in numbers and diversity of forest arthropods in the study areas may have affected the impact of the acaricide in the treatment area. Comparison with control areas indicated that reductions in abundance of some arthropod taxa in the treatment area were detectable 12 wk after treatment.     

Structure of ground-dwelling arthropod assemblages in vegetation units of Área Natural Protegida Península Valdés,Patagonia, Argentina

Knowing the spatial variation of insect and arachnid assemblages and their relationship with habitat variables is critical to understand the structure and dynamics of these communities in arid environments. The aim of this paper was to analyze the variation in ground-dwelling arthropod assemblages across three representative vegetation units of the Área Natural Protegida Península Valdés (Patagonia, Argentina). We asked whether environmental differences among representative vegetation units were associated to distinct arthropod assemblages. We selected three plant communities: grass, dwarf-shrub, and shrub steppes, and established three sampling sites within each of them. We measured variables of vegetation structure and soil characteristics and collected the arthropods using 10 pitfall traps per site. We analyzed the structure of arthropod assemblages at both family and ant species taxonomic levels. Each plant community displayed a distinctive assemblage, with differences in diversity, taxa abundance, trophic structure and functional groups of ants. Vegetation variables explained a higher proportion of the variation in the structure of the ground-dwelling arthropod assemblages than the soil variables. This work highlights the importance of the different vegetation units for the conservation of ground-dwelling arthropod biodiversity in Península Valdés.