Ant exclusion in citrus over an 8-year period reveals a pervasive yet changing effect of ants on a Mediterranean spider assemblage

Ants and spiders are ubiquitous generalist predators that exert top-down control on herbivore populations. Research shows that intraguild interactions between ants and spiders can negatively affect spider populations, but there is a lack of long-term research documenting the strength of such interactions and the potentially different effects of ants on the diverse array of species in a spider assemblage. Similarly, the suitability of family-level surrogates for finding patterns revealed by species-level data (taxonomic sufficiency) has almost never been tested in spider assemblages. We present a long-term study in which we tested the impact of ants on the spider assemblage of a Mediterranean citrus grove by performing sequential 1-year experimental exclusions on tree canopies for 8 years. We found that ants had a widespread influence on the spider assemblage, although the effect was only evident in the last 5 years of the study. During those years, ants negatively affected many spiders, and effects were especially strong for sedentary spiders. Analyses at the family level also detected assemblage differences between treatments, but they concealed the different responses to ant exclusion shown by some related spider species. Our findings show that the effects of experimental manipulations in ecology can vary greatly over time and highlight the need for long-term studies to document species interactions.     

Tree Species Richness Strengthens Relationships between Ants and the Functional Composition of Spider Assemblages in a Highly Diverse Forest

In species‐rich ecosystems, such as subtropical and tropical forests, higher trophic level interactions are key mediators of ecosystem functioning. Plant species loss may alter these interactions, but the effects of plant diversity might be modified by intraguild interactions, particularly among predators. We analyzed the relationships between spiders and ants, two dominant predatory arthropod taxa, on tree saplings across a gradient from medium to high woody plant species richness in a subtropical forest in Southeast China. Neither ant nor spider total biomass was significantly related to plant species richness. By contrast, the biomass distribution of web‐building and hunting spiders changed and spider family richness increased in the presence of ants, resulting in more web builder‐dominated assemblages. However, these relationships depended on the plant communities, and were stronger in plots with higher plant species richness. Our results indicate that in addition to potential effects of ants on hunting spiders in particular, ants could indirectly influence intraguild interactions within spider assemblages. The observed shifts in the spider assemblages with increasing ant presence and plant species richness may have functional consequences, as web‐building and hunting spiders have distinct prey spectra. The relationships among ants, spiders, and plant species richness might contribute to explaining the non‐significant relationship between the overall effects of predators and plant diversity previously observed in the same forest plots. Our findings thus give insight into the complexity of biotic interactions in such species‐rich ecosystems.     

Intraguild interactions among generalist predator functional groups drive impact on herbivore and decomposer prey

Different functional groups of generalist predators may complement each other in controlling prey populations; but intraguild interactions, common among generalist predators, may also reduce the strength of top–down control. In natural communities greater alterations to ecosystem function are expected if a whole functional group declines in abundance or is lost. Therefore studying functional group diversity is important for predicting effects of predator loss. We studied the top–down impact of web‐building spiders, hunting spiders and ants, which are highly abundant generalist predators in most terrestrial ecosystems, on prey from the herbivore and decomposer system of a grassland food web. The density of the three predator groups was manipulated by continuous removal in a three‐factorial designed field experiment, which was carried out for two years. We found no positive effect of increasing predator functional group richness on prey control. However there was evidence for strong composition effects between the functional groups. The presence of ants in predator assemblages reduced the prey suppression through mostly trait‐mediated intraguild interactions, while hunting and web‐building spiders contributed additively to prey suppression and reduced the density of herbivore and decomposer prey by 50–60%. A trophic cascade on plant biomass triggered by web‐builders and hunting spiders was diminished at levels of higher predator group diversity. In conclusion, our experiments showed that intraguild interactions strongly influence the strength of top–down control by generalist predators. Among spiders there was evidence for a positive relation between functional group richness and prey suppression but the overall outcome strongly depended on the occurrence of interference, driven by trait‐mediated indirect interactions.     

Trophic structure of the spider community of a Mediterranean citrus grove: A stable isotope analysis

Spiders are dominant terrestrial predators that consume a large variety of prey and engage in intraguild predation. Although the feeding habits of certain species are well known, the trophic structure of spider assemblages still needs to be investigated. Stable isotope analysis enables characterisation of trophic relationships between organisms because it tracks the energy flow in food webs and indicates the average number of trophic transfers between a given species and the base of the web, thus being a useful tool to estimate the magnitude of intraguild predation in food webs. Using this technique, we studied the trophic groups of spiders and their links within the arthropod food web of a Mediterranean organic citrus grove. We assessed the trophic positions of the 25 most common spider species relative to other arthropod predators and potential prey in the four seasons of the year, both in the canopy and on the ground. The analyses showed great seasonal variation in the isotopic signatures of some arthropod species, as well as the existence of various trophic groups and a wide range of trophic levels among spiders, even in species belonging to the same family. Differences in δ¹⁵N between spiders and the most abundant prey in the grove usually spanned two trophic levels or more. Our findings provide field evidence of widespread intraguild predation in the food web and caution against using spider families or guilds instead of individual species when studying spider trophic interactions.     

Negative effects of ant foraging on spiders in Douglas-fir canopies

 Spiders and ants are potential competitors and mutual predators. Indirect evidence from previous research has suggested that ant foraging may significantly lower the abundance of arboreal spiders in young Douglas-fir plantations in western Oregon. This study tested the effect of foraging by ants, dominated by Camponotus spp., on spider assemblages in Douglas-fir canopies in a 5-month ant-exclusion experiment. The biomass of potential prey organisms on foliage, dominated by Psocoptera, increased significantly by 1.9- to 2.4-fold following ant exclusion. The removal of ants did not affect the abundance of flying arthropods in the vicinity of tree canopies as indicated by sticky trap catches. The abundance of hunting spiders, the majority being Salticidae, increased significantly by 1.5- to 1.8-fold in trees without ants in the late summer; neither the abundance of web-building spiders nor the average body size of hunting and web-building spiders were significantly affected by ant removal. Spider diversity and community structure did not differ significantly between control and ant-removal trees. The majority of prey captured by ants were Aphidoidea (48.1%) and Psocoptera (12.5%); spiders represented only 1.4% of the ants’ diet. About 40% of observed ants were tending Cinara spp. aphids. Our observations suggest that the lower abundance of hunting spiders in control canopies with ants may be due to interference competition with ants resulting from ant foraging and aphid-tending activities. Direct predation of spiders by ants appeared to be of minor importance in this study system. This study did not provide sufficient evidence for exploitative competition for prey between ants and spiders. Received: 21 February 1996 / Accepted: 14 August 1996     

Ant versus bird exclusion effects on the arthropod assemblage of an organic citrus grove

1. Predation‐exclusion experiments have highlighted that top‐down control is pervasive in terrestrial communities, but most of these experiments are simplistic in that they only excluded a single group of predators and the effect of removal was evaluated on a few species from the community. The main goal of our study was to experimentally establish the relative effects of ants and birds on the same arthropod assemblage of canopy trees. 2. We conducted 1‐year long manipulative experiments in an organic citrus grove intended to quantify the independent effects of bird and ant predators on the abundance of arthropods. Birds were excluded with plastic nets whereas ants were excluded with sticky barriers on the trunks. The sticky barrier also excluded other ground dwelling insects, like the European earwig Forficula auricularia L. 3. Both the exclusion of ants and birds affected the arthropod community of the citrus canopies, but the exclusion of ants was far more important than the exclusion of birds. Indeed, almost all groups of arthropods had higher abundance in ant‐excluded than in control trees, whereas only dermapterans were more abundant in bird‐excluded than in control trees. A more detailed analysis conducted on spiders also showed that the effect of ant exclusion was limited to a few families rather than being widespread over the entire diverse spectrum of spiders. 4. Our results suggest that the relative importance of vertebrate and invertebrate predators in regulating arthropod populations largely depends on the nature of the predator–prey system.     

Intraguild interactions between spiders and ants and top-down control in a grassland food web

In most terrestrial ecosystems ants (Formicidae) as eusocial insects and spiders (Araneida) as solitary trappers and hunters are key predators. To study the role of predation by these generalist predators in a dry grassland, we manipulated densities of ants and spiders (natural and low density) in a two-factorial field experiment using fenced plots. The experiment revealed strong intraguild interactions between ants and spiders. Higher densities of ants negatively affected the abundance and biomass of web-building spiders. The density of Linyphiidae was threefold higher in plots without ant colonies. The abundance of Formica cunicularia workers was significantly higher in spider-removal plots. Also, population size of springtails (Collembola) was negatively affected by the presence of wandering spiders. Ants reduced the density of Lepidoptera larvae. In contrast, the abundance of coccids (Ortheziidae) was positively correlated with densities of ants. To gain a better understanding of the position of spiders, ants and other dominant invertebrate groups in the studied food web and important trophic links, we used a stable isotope analysis (¹⁵N and ¹³C). Adult wandering spiders were more enriched in ¹⁵N relative to ¹⁴N than juveniles, indicating a shift to predatory prey groups. Juvenile wandering and web-building spiders showed δ¹⁵N ratios just one trophic level above those of Collembola, and they had similar δ¹³C values, indicating that Collembola are an important prey group for ground living spiders. The effects of spiders demonstrated in the field experiment support this result. We conclude that the food resource of spiders in our study system is largely based on the detrital food web and that their effects on herbivores are weak. The effects of ants are not clear-cut and include predation as well as mutualism with herbivores. Within this diverse predator guild, intraguild interactions are important structuring forces.     

Environmental and spatial drivers of spider diversity at contrasting microhabitats

The relative importance of environmental and spatial drivers of animal diversity varies across scales, but identifying these scales can be difficult if a sampling design does not match the scale of the target organisms' interaction with their habitat. In this study, we quantify and compare the effects of environmental variation and spatial proximity on ground‐dwelling spider assemblages sampled from three distinct microhabitat types (open grassland, logs, trees) that recur across structurally heterogeneous grassy woodlands. We used model selection and multivariate procedures to compare the effects of different environmental attributes and spatial proximity on spider assemblages at each microhabitat type. We found that species richness and assemblage composition differed among microhabitat types. Bare ground cover had a negative effect on spider richness under trees, but a positive effect on spider richness in open grassland. Turnover in spider assemblages from open grassland was correlated with environmental distance, but not geographic distance. By contrast, turnover in spiders at logs and trees was correlated with geographic distance, but not environmental distance. Our study suggests that spider assemblages from widespread and connected open grassland habitat were more affected by environmental than spatial gradients, whereas spiders at log and tree habitats were more affected by spatial distance among these discrete but recurring microhabitats. Deliberate selection and sampling of small‐scale habitat features can provide robust information about the drivers of arthropod diversity and turnover in landscapes.     

Negative Correlation between Ant and Spider Abundances in the Canopy of a Bornean Tropical Rain Forest

In tropical rain forests, high canopy trees have diverse and abundant populations of ants and spiders. However, accessing high trees and their fauna remains difficult; thus, how ants and spiders interact in the canopy remains unclear. To better understand the interspecific interactions between these two dominant arthropod groups, we investigated their spatial distributions at the canopy surface in a tropical rain forest in Borneo. We sampled ants and spiders six times between 2009 and 2011 by sweeping with an insect net at the tree crown surfaces of 190 emergent or tall (≥20 m in height) trees. We collected 438 ant individuals belonging to 94 species and 1850 spider individuals (1630 juveniles and 220 adults) belonging to 142 morphospecies (adults only) from a total of 976 samples. The fact that we collected four times more spider individuals than ant individuals suggests that fewer ants forage at the tree crown surface than previously thought. The number of spider individuals negatively correlated with the number of ant individuals and the number of ant species, indicating significant exclusivity between ant and spider spatial distributions at the tree crown surface. Niche‐overlap between the two taxa confirmed this observation. Although our data do not address the causes of these spatial distributions, antagonistic interspecific interactions such as interference behaviors and intra‐guild predation are ecological mechanisms that give rise to exclusive spatial distributions.     

Bird predation affects diurnal and nocturnal web-building spiders in a Mediterranean citrus grove

Spiders and birds can greatly decrease insect populations, but birds also limit spider densities in some habitats. Bird predation is thought to be one of the causes behind nocturnal activity in spiders, so night-active spiders that hide in retreats during the day should be less affected by bird foraging than day-active spiders. However, this hypothesis has not yet been tested. We investigated the importance of bird predation on the spider community of a Mediterranean organic citrus grove. We excluded birds by placing net cages over the trees and we conducted visual searches in the canopies to sample web-building spiders. As there are many nocturnal species in the family Araneidae, we conducted searches both by day and by night to compare the abundance of active araneids in these two time periods. We sampled the tree trunks with cardboard bands to collect hunting spiders. In bird-excluded canopies there were more spiders of the families Araneidae and Theridiidae. There were higher numbers of active Araneidae at night, but these were just as negatively affected by bird predation as day-active Araneidae, so there was no evidence of nocturnal activity serving as an anti-predator strategy. We did not find any negative impact of birds on hunting spiders. Our results contrast with other studies reporting a negative effect of birds on hunting but not on web-building spiders.     

Remote sensing improves prediction of tropical montane species diversity but performance differs among taxa

Texture information from passive remote sensing images provides surrogates for habitat structure, which is relevant for modeling biodiversity across space and time and for developing effective ecological indicators. However, the applicability of this information might differ among taxa and diversity measures. We compared the ability of indicators developed from texture analysis of remotely sensed images to predict species richness and species turnover of six taxa (trees, pyraloid moths, geometrid moths, arctiinae moths, ants, and birds) in a megadiverse Andean mountain rainforest ecosystem. Partial least-squares regression models were fitted using 12 predictors that characterize the habitat and included three topographical metrics derived from a high-resolution digital elevation model and nine texture metrics derived from very high-resolution multi-spectral orthophotos. We calculated image textures derived from mean, correlation, and entropy statistics within a relatively broad moving window (102 m × 102 m) of the near infra-red band and two vegetation indices. The model performances of species richness were taxon dependent, with the lowest predictive power for arctiinae moths (4%) and the highest for ants (78%). Topographical metrics sufficiently modeled species richness of pyraloid moths and ants, while models for species richness of trees, geometrid moths, and birds benefited from texture metrics. When more complexity was added to the model such as additional texture statistics calculated from a smaller moving window (18 m × 18 m), the predictive power for trees and birds increased significantly from 12% to 22% and 13% to 27%, respectively. Gradients of species turnover, assessed by non-metric two-dimensional scaling (NMDS) of Bray-Curtis dissimilarities, allowed the construction of models with far higher predictability than species richness across all taxonomic groups, with predictability for the first response variable of species turnover ranging from 64% (birds) to 98% (trees) of the explained change in species composition, and predictability for the second response variable of species turnover ranging from 33% (trees) to 74% (pyraloid moths). The two NMDS axes effectively separated compositional change along the elevational gradient, explained by a combination of elevation and texture metrics, from more subtle, local changes in habitat structure surrogated by varying combinations of texture metrics. The application of indicators arising from texture analysis of remote sensing images differed among taxa and diversity measures. However, these habitat indicators improved predictions of species diversity measures of most taxa, and therefore, we highly recommend their use in biodiversity research.     

Effects of ants on the foraging of birds in spruce trees

I experimentally excluded ants from randomly selected spruce trees Picea abies near colonies of the wood ant Formica aquilonia. Foraging activity of birds in these trees was then compared to the foraging activity of birds in neighboring spruce trees, where ants were allowed to continue foraging. Birds which foraged in the foliage showed the effects of competition with ants: they visited the trees without ants more frequently, and for longer periods. In addition, the insects and spiders that they utilized as food were more abundant in the foliage of trees without ants. Cone-foraging birds, however, which fed on seeds in cones at the tops of the trees, did not show a preference for trees without ants. The differences of tree usage between foliage-gleaning and coneforaging birds can be explained by alteration of the birds' food supply by wood ants: ants did not feed on seeds in cones, and so did not compete with cone-foraging birds. However, foraging wood ants did feed on arthropods living in the foliage, thus reducing the amount of food available to birds there.     

Host tree architecture mediates the effect of predators on herbivore survival

Abstract.  1. Vegetation structural complexity is an important factor influencing ecological interactions between different trophic levels. In order to investigate relationships between the architecture of trees, the presence of arthropod predators, and survival and parasitism of the autumnal moth Epirrita autumnata Borkhausen, two sets of experiments were conducted.
2. In one experiment, the architectural complexity of mountain birch was manipulated to separate the effects of plant structure and age. In the other experiment the trees were left intact, but chosen to represent varying degrees of natural complexity. Young autumnal moth larvae were placed on the trees and their survival was monitored during the larval period.
3. The larvae survived longer in more complex trees if predation by ants was prevented with a glue ring, whereas in control trees smaller canopy size improved survival times in one experiment. The density of ants observed in the trees was not affected by canopy size but spider density was higher on smaller trees. The effect of canopy structure on larval parasitism was weak; larger canopy size decreased parasitism only in one year. Until the fourth instar the larvae travelled shorter distances in trees with reduced branchiness than in trees with reduced foliage or control treatments. Canopy structure manipulation by pruning did not alter the quality of leaves as food for larvae.
4. The effect of canopy structure on herbivore survival may depend on natural enemy abundance and foraging strategy. In complex canopies herbivores are probably better able to escape predation by ambushing spiders but not by actively searching ants.     

Spider assemblages in bird burrows

The arachnofauna of bird burrows, which are excavated in the soil, is incompletely understood. The bird burrows have a potential to serve as refugia for cavity-adopting spider species particularly in anthropogenic habitats formed by sand mining, which otherwise suffer from a limited availability of other types of cavities. We hypothesized that the presence of abundant food resources and safe shelter in bird burrows excavated in the soil allow the existence of specialized assemblage of spiders, similarly to those known from cavities and nests made by other birds and mammals. Here we examined 353 burrows that had been previously occupied by Riparia riparia, Merops apiaster or Passer montanus. The burrows were examined in early May and early September in 24 Czech sand pits and construction sites. The burrows hosted a species-rich assemblage of spiders that, however, occurred at low abundances. The spiders were represented prevalently by species known to have distributions centered in anthropogenic and urban habitats, including species that were never observed outside of heated buildings in the study area and species that only were previously known from mammalian burrows and loess and karstic soil crevices. Spider assemblages strongly differed between bird burrows and cavities and nests made by other birds and mammals. The species composition and overall low abundance of spiders in bird burrows was unexpected and was in strong contrast to previous observations of spider assemblages in cavities and nests made by other birds and mammals. Concluded, we rejected the initial hypothesis and, instead, we have shown that bird burrows are rarely occupied by spiders and if occupied, the spider species that adopt old bird burrows and the nests inside of them predominantly consist of species known to have their distribution centered to anthropogenic and urban habitats.     

Contrasting cascades: insectivorous birds increase pine but not parasitic mistletoe growth

1. Intraguild predation occurs when top predators feed upon both intermediate predators and herbivores. Intraguild predators may thus have little net impact on herbivore abundance. Variation among communities in the strength of trophic cascades (the indirect effects of predators on plants) may be due to differing frequencies of intraguild predation. Less is known about the influence of variation within communities in predator-predator interactions upon trophic cascade strength. 2. We compared the effects of a single predator community between two sympatric plants and two herbivore guilds. We excluded insectivorous birds with cages from ponderosa pine Pinus ponderosa trees parasitized by dwarf mistletoe Arceuthobium vaginatum. For 3 years we monitored caged and control trees for predatory arthropods that moved between the two plants, foliage-feeding caterpillars and sap-feeding hemipterans that were host-specific, and plant damage and growth. 3. Excluding birds increased the abundance of ant-tended aphids on pine and resulted in an 11% reduction in pine woody growth. Mutualist ants protected pine-feeding aphids from predatory arthropods, allowing aphid populations to burgeon in cages even though predatory arthropods also increased in cages. By protecting pine-feeding aphids from predatory arthropods but not birds, mutualist ants created a three-tiered linear food chain where bird effects cascaded to pine growth via aphids. 4. In contrast to the results for tended aphids on pine, bird exclusion had no net effects on untended pine herbivores, the proportion of pine foliage damaged by pine-feeding caterpillars, or the proportion of mistletoe plants damaged by mistletoe-feeding caterpillars. These results suggest that arthropod predators, which were more abundant in cages as compared with control trees, compensated for bird predation of untended pine and mistletoe herbivores. 5. These contrasting effects of bird exclusion support food web theory: where birds were connected to pine by a linear food chain, a trophic cascade occurred. Where birds fed as intraguild predators, the reticulate food webs linking birds to pine and mistletoe resulted in no net effects on herbivores or plant biomass. Our study shows that this variation in food web structure occurred between sympatric plants and within plants between differing herbivore guilds.     

Emergent Impacts of Ant and Spider Interactions: Herbivory Reduction in a Tropical Savanna Tree

Multiple predators often have effects on their common prey populations that cannot be predicted by summing the effects of each predator at a time. When predators forage on the same vegetation substrate, intraguild interactions might cause emergent outcomes for the plants on which the predators co‐occur. We experimentally evaluated the effects of spiders and ants on herbivory and reproduction in the extrafloral nectary‐bearing tree Qualea multiflora (Vochysiaceae). Plants were divided in four experimental groups, depending on the presence or absence of ants and spiders. We compared the effects of each treatment on richness and abundance of chewing and sucking herbivores and on herbivory (leaf area loss). We also evaluated the impact of predators on the production of buds, fruits and seeds, and weight of the fruits. The presence of ants reduced the abundance and richness of spiders, but spiders did not affect the abundance and richness of ants. Only the removal of ants resulted in a significant increase in the abundance of herbivores and herbivore richness. Herbivory, however, was also affected by spiders. In addition, we found a significant interaction effect of ants and spiders on herbivory, indicating an emergent multiple predator effect. Neither ants nor spiders had an impact on the number of buds produced, number of fruits per bud, and seeds per fruits or fruit weight. This study highlights the importance of evaluating the effect of the predator fauna as a whole and not only one specific group on herbivory.     

Factors shaping latitudinal patterns in communities of arboreal spiders in northern Europe

Predator effects on herbivores are often referred to as examples of biotic interactions that weaken with latitude, but more studies are needed to test for the generality of this pattern. To further the understanding of large‐scale geographical patterns in abundance and diversity of predatory arthropods, from 2008–2011 we explored spider communities in the canopies of primary forest trees of the boreal zone (Pinus sylvestris, Picea abies, Betula pubescens and B. pendula) along five latitudinal gradients in northern Europe, from 59 to 70°N and from 10 to 60°E. The abundance of arboreal spiders in Norway and Finland was about a half of that in Russia, presumably due to more intensive forest management in Scandinavia. The abundance, taxonomic and functional diversity of arboreal spiders generally decreased with latitude; however, actual weather conditions during the study years had little effect on this pattern. Coniferous species supported higher abundance and diversity of arboreal spiders than birches, but the poleward decrease in either abundance or taxonomic diversity of arboreal spiders did not differ between coniferous and deciduous tree species. In contrast, functional diversity on birches decreased with latitude greater than on coniferous trees. Euryphagous spiders showed stronger decrease with latitude in terms of both abundance and taxonomic diversity than more specialized (steno‐ and oligophagous) spiders. We attribute the general decrease in density and diversity of spiders with latitude to an increased pressure from apex predators (birds) rather than to direct effects of climate or changes in prey abundance. The abundance of spiders declined with the latitude to the greater extent than the densities of their potential prey did, suggesting a decrease in the strength of predator–prey interactions towards the north.     

Spatial refuge from intraguild predation: implications for prey suppression and trophic cascades

The ability of predators to elicit a trophic cascade with positive impacts on primary productivity may depend on the complexity of the habitat where the players interact. In structurally-simple habitats, trophic interactions among predators, such as intraguild predation, can diminish the cascading effects of a predator community on herbivore suppression and plant biomass. However, complex habitats may provide a spatial refuge for predators from intraguild predation, enhance the collective ability of multiple predator species to limit herbivore populations, and thus increase the overall strength of a trophic cascade on plant productivity. Using the community of terrestrial arthropods inhabiting Atlantic coastal salt marshes, this study examined the impact of predation by an assemblage of predators containing Pardosa wolf spiders, Grammonota web-building spiders, and Tytthus mirid bugs on herbivore populations (Prokelisia planthoppers) and on the biomass of Spartina cordgrass in simple (thatch-free) and complex (thatch-rich) vegetation. We found that complex-structured habitats enhanced planthopper suppression by the predator assemblage because habitats with thatch provided a refuge for predators from intraguild predation including cannibalism. The ultimate result of reduced antagonistic interactions among predator species and increased prey suppression was enhanced conductance of predator effects through the food web to positively impact primary producers. Behavioral observations in the laboratory confirmed that intraguild predation occurred in the simple, thatch-free habitat, and that the encounter and capture rates of intraguild prey by intraguild predators was diminished in the presence of thatch. On the other hand, there was no effect of thatch on the encounter and capture rates of herbivores by predators. The differential impact of thatch on the susceptibility of intraguild and herbivorous prey resulted in enhanced top-down effects in the thatch-rich habitat. Therefore, changes in habitat complexity can enhance trophic cascades by predator communities and positively impact productivity by moderating negative interactions among predators.     

Pollination ecology of Acacia gerrardii Benth. (Fabaceae: Mimosoideae) under extremely hot-dry conditions

Talh trees (Acacia gerrardii Benth.) are acacias that are native to the arid and semiarid Africa and west Asia. We investigated the flowering biology, pod set and flower visitors of Talh and discussed the role of these visitors in pollen transfer. The Talh trees blossomed laterally on the nodes of one-year-old twigs. Each node produced 21 flower buds seasonally. Each flower bud opened to a flower head (FH) of 60 florets. The bagged FHs podded significantly (p ⩽ 0.05) less than did the unbagged FHs. The FHs were visited by 31 insect species (25 genera, 16 families and 5 orders). The major taxa were honeybees, megachilids, butterflies, ants, beetles and thrips. Each of honeybees, megachilids and beetles showed a significant (p ⩽ 0.05) hourly pattern, while each of butterflies, ants and thrips had no hourly pattern (p > 0.05). Furthermore, some birds and mammals touched the Talh FHs. Talh trees evolved a mass flowering behavior to face pre- and post-flowering obstacles. Megachilids seemed to play the major effort of zoophily because of their relatively high numbers of individuals and species and their effective movement behavior on the FH surface. Nevertheless, honeybees and other insects and vertebrate taxa also contributed to the pollen transfer. These results greatly contribute to our understanding of the pollination ecology of acacias, especially Arabian acacias.     

Spiders in mofette fields—Survival of the toughest in natural carbon dioxide springs?

In mofette fields, natural carbon dioxide springs, organisms have to stand extreme CO₂ concentrations up to 100%. These hostile conditions are spatially small-scaled and further influenced by earth tides, wind and temperature. The present project investigated the influence of increased atmospheric CO₂ concentration on spiders as representatives of above-ground organisms by means of pitfall traps in three mofette fields, differing in habitat conditions in the Plesná valley, eastern Cheb Basin, Czech Republic.Among the 71 recorded spider species four were rarely found in the Czech Republic. A canonical correspondence analysis revealed significant influences of environmental parameters on the spider assemblages. Two groups of spiders are clearly distinguishable, one being positively influenced by humidity and the second by temperature. A cluster analysis showed distinct and congruent results: spider assemblages of pitfall traps at spots with a mean CO₂ concentration above 7.6% grouped close together and this grouping was independent of site. At >7.6% CO₂ significantly fewer individuals and species were found in comparison to areas with lower CO₂ concentration. Between 2.5 and 10% CO₂, spiders indicated increased CO₂ concentrations much more sensitively than endogeic organisms (Nematoda, Collembola) in a nearby mofette field. Unlike in nematodes, collembolans and plants, no mofettovageous or mofettophilous spiders were detected. In contrast to humidity, CO₂ concentration and temperature, the vegetation cover was not among the factors, which significantly influenced spiders. This is explained by the fact that mofettophilous plants occurred at spots where almost no spiders could live. In a field experiment, most Pardosa pullata males tested passed a 30 cm long corridor with increased carbon dioxide concentration. These results and that of pitfall traps showed that relatively large and wandering specimens respectively were able to transit moderately hostile spots. Further experiments are necessary to find out if there is any active avoidance of high-CO₂ areas by spiders.