Dynamics of plant and arthropod diversity during old field succession

The successional dynamics of arthropod diversity in 18 abandoned agricultural fields (age 15-54 yr) at Cedar Creek. MN. USA were determined using sweep net sampling (44833 individuals of 618 species). Total arthropod species richness and equitability (J), but not abundance, increased significantly with field successional age. Herbivore and parasite species richness, but not detritivore and predator species richness, also increased significantly with field age. All of these arthropod variables were significantly positively correlated with plant species richness in the fields. When plant species richness was included as a covariate in regressions, there were no longer any significant effects of field age. These results supported the hypothesis that increases in arthropod diversity with field age are influenced by increases in plant diversity. The additional significant positive dependence of herbivore species richness on predator species richness suggests that predator-prey interactions may also influence the successional dynamics of arthropod diversity. Nine of the ten most common arthropod species decreased in abundance with field age, two of them significantly. The abundances of these two generalist forb-feeding species, Melanoplus femurrubrum (Orthoptera: Acrididae) and Scaphytopius acutus (Homoptera: Cicadellidae). each depended significantly on amount of forbs. The average body size of arthropod species (total and herbivores) decreased significantly with field age. An efficiency vs specialization hypothesis predicts such a decrease. Because plants in later secondary succession are generally less palatable, a diversity of smaller, potentially more specialized herbivores may have an advantage over larger and more efficient herbivores in later succession.     

Feedback effects between plant and flower-visiting insect communities along a primary succession gradient

Primary successions of glacier forelands are unique model systems to investigate community dynamics and assembly processes. However, successional changes of plant and insect communities have been mainly analysed separately. Therefore, changes in plant–insect interactions along successional gradients on glacier forelands remain unknown, despite their relevance to ecosystem functioning. This study assessed how successional changes of the vegetation influenced the composition of the flower-visiting insect assemblages of two plant species, Leucanthemopsis alpina (L.) Heyw. and Saxifraga bryoides L., selected as the only two insect-pollinated species occurring along the whole succession. In addition, we investigated the links between reproductive output of these plants and pollinator abundance through experimental exclusion of pollinators. Plant community structure changed along the succession, affecting the distribution and the abundance of insects via idiosyncratic responses of different insect functional groups. L. alpina interacted with ubiquitously distributed pollinators, while S. bryoides pollinators were positively associated with insect-pollinated plant species density and S. bryoides abundance. With succession proceeding, insect assemblages became more functionally diverse, with the abundance of parasitoids, predators and opportunists positively related to an increase in plant cover and diversity. The reproductive output of both plant species varied among successional stages. Contrary to our expectation, the obligate insect-pollinated L. alpina showed a reproductive output rather independent from pollinator abundance, while the reproductive output of the self-fertile S. bryoides seemed linked to pollinator abundance. Observing ecological interactions and using functional traits, we provided a mechanistic understanding of community assembly processes along a successional gradient. Plant community diversity and cover likely influenced insect community assembly through bottom-up effects. In turn, pollinators regulate plant reproductive output through top-down control. We emphasise that dynamics of alpine plant and insect communities may be structured by biotic interactions and feedback processes, rather than only be influenced by harsh abiotic conditions and stochastic events.     

The relationships of plant and insect diversities in succession

The basic features of an intensive study on the various stages of a secondary succession, from fallow Held to birch woodland, are described. The α-β diversities of the green plants, and two orders of insects, Hetcroptera and adult Coleoptera, are described. For the vegetation, in addition to taxonotnic diversity, structural diversity, with both spatial and architectural components, was recognized. It was found that up to a successional age of 16 months, the taxonomic diversities of plants and insects rose; thereafter the diversity of the plant species declined far more than the insect species diversity. It was concluded that in the later successional stages the maintenance of a high level of taxonomic diversity of these orders of insects is correlated with the rising structural diversity of the green plants, which virtually compensates for their falling taxonomic diversity. The larger fungi appear to show a similar trend to the insects.     

Core taxa underpin soil microbial community turnover during secondary succession

Understanding the processes that underpin the community assembly of bacteria is a key challenge in microbial ecology. We studied soil bacterial communities across a large-scale successional gradient of managed and abandoned grasslands paired with mature forest sites to disentangle drivers of community turnover and assembly. Diversity partitioning and phylogenetic null-modelling showed that bacterial communities in grasslands remain compositionally stable following abandonment and secondary succession but they differ markedly from fully afforested sites. Zeta diversity analyses revealed the persistence of core microbial taxa that both reflected and differed from whole-scale community turnover patterns. Differences in soil pH and C:N were the main drivers of community turnover between paired grassland and forest sites and the variability of pH within successional stages was a key factor related to the relative dominance of deterministic assembly processes. Our results indicate that grassland microbiomes could be compositionally resilient to abandonment and secondary succession and that the major changes in microbial communities between grasslands and forests occur fairly late in the succession when trees have established as the dominant vegetation. We also show that core taxa may show contrasting responses to management and abandonment in grasslands.     

Relationship between plant development, tannin concentration and insects associated with Copaifera langsdorffii (Fabaceae)

Plant development is the main factor that determines the insect-ontogeny interaction, since it leads to variations in resource quality and availability. The aim of this study was to test the hypothesis that plant development and varying tannin concentration leads to changes in species richness, abundance and composition of ants, free-feeding herbivores and galling insects associated with Copaifera langsdorffii (Fabaceae). The plant ontogeny and tannin concentration effects on insects were tested on 60 individuals with height varying from 0.9 to 11.0 m. A positive correlation was observed for tree height and species richness and abundance of ants, free-feeding and galling insects. In contrast, we did not find a significant relation between leaf tannin concentration and plant height, or richness and abundance of the different insect guilds. The assemblage of ants (composition of species) did not change between saplings and adults of C. langsdorffii. However, the assemblage of free-feeding herbivores and galling insects varied between the two development stages studied. The present study reveals an ontogenetic succession pattern for herbivore insects along the C. langsdorffii growth, probably due to both indirect and direct benefits from the host plant architecture and quality. Those plants with more complex architectures should support a wider diversity of insects, since they present higher number of sites for egg laying, housing, feeding and better environmental conditions. This is the first work to investigate the host plant ontogeny effect on insects in Cerrado “Savanna” vegetation. The pattern described, along with other previous studies, suggests a vast occurrence of ontogenetic succession in tropical areas.     

Non-native plants rarely provide suitable habitat for native gall-inducing species

For insect herbivores, a critical niche requirement—possibly the critical niche requirement—is the presence of suitable host plants. Current research suggests that non-native plants are not as suitable as native plants for native herbivores, resulting in decreases in insect abundance and richness on non-native plants. Like herbivores, gall-forming insects engage in complex, species-specific interactions with host plants. Galls are plant tissue tumors (including bulbous or spindle-shaped protrusions on leaves, stems and other plant organs) that are induced by insects through physical or chemical damage (prompting plants to grow a protective tissue shell around the insect eggs and larvae). As such, we hypothesized that gall-inducing insect species richness would be higher on native than non-native plants. We also predicted higher gall-inducing insect species richness on woody than herbaceous plants. We used an extensive literature review in which we compiled gall host plant species by genus, and we assigned native or non-native (or mixed) status to each genus. We found that native plants host far more gall-inducing insect species than non-native plants; woody plants host more gall-inducing species than herbaceous plants; and native woody plants host the most gall-inducing species of all. Gall-inducing species generally are a very cryptic group, even for experts, and hence do not elicit the conservation efforts of more charismatic insects such as plant pollinators. Our results suggest that non-native plants, particularly non-native woody species, diminish suitable habitat for gall-inducing species in parallel with similar results found for other herbivores, such as Lepidopterans. Hence, the landscape-level replacement of native with non-native species, particularly woody ones, degrades taxonomically diverse gall-inducing species (and their inquilines and parasitoids), removing multiple layers of diversity from forest ecosystems.

     

Top-down effects of insect herbivores during early succession: influence on biomass and plant dominance

We tested the hypothesis that phytophagous insects would have a strong top-down effect on early successional plant communities and would thus alter the course of succession. To test this hypothesis, we suppressed above-ground insects at regular intervals with a broad-spectrum insecticide through the first 3 years of old-field succession at three widely scattered locations in central New York State. Insect herbivory substantially reduced total plant biomass to a similar degree at all three sites by reducing the abundance of meadow goldenrod, Solidago altissima. As a result, Euthamia graminifolia dominated control plots whereas S. altissima dominated insecticide-treated plots by the third year of succession. S. altissima is the dominant old-field herbaceous species in this region but typically requires at least 5 years to become dominant. Past explanations for this delay have implicated colonization limitation whereas our data demonstrate that insect herbivory is a likely alternative explanation. A widespread, highly polyphagous insect, the xylem-tapping spittlebug, Philaenus spumarius, appeared to be the herbivore responsible for the reduction in standing crop biomass at all three sites. Insect herbivory typically caused little direct leaf tissue loss for the ten plant species we examined, including S. altissima. Consequently, the amount of leaf area removed was not a reliable indicator of the influence of insect herbivory on standing crop biomass or on early succession. Overall, we found a strong top-down effect of insect herbivores on biomass at several sites, so our results may be broadly applicable. These findings run counter to generalizations that top-down effects of herbivores, particularly insects, are weak in terrestrial systems. These generalizations may not apply to insects, such as spittlebugs, that can potentially mount an effective defense (i.e., spittle) against predators and subsequently reach relatively high abundance on common plant species. Our results suggest that insect herbivory may play an important but often overlooked role during early old-field succession. Received: 26 December 1998 / Accepted: 3 April 1999     

Gall-inducing insect species richness as indicators of forest age and health

The changes in the plant community that occur during the process of succession affect the availability of resources for the community of herbivores. In this study, the richness of galling insects was evaluated in restored stands of Amazonian tropical rain forest of several ages (0-21 yr), as well as in areas of primary forest in Brazil. The richness of gallers increased with the age of the restored stands. Fifty-eight percent of the variation in the richness of galling insects was explained by forest stand age, but an increase in richness was observed at intermediate stages of succession. The greatest similarity among groups was found between the initial successional stages and intermediate ones. The results indicate a recovery of both host plants and insect community and that succession directly affects the richness and composition of these herbivores.     

Dynamics of vesicular-arbuscular mycorrhizae during old field succession

Summary The species composition of vesicular-arbuscular mycorrhizal (VAM) fungal communities changed during secondary succession of abandoned fields based on a field to forest chronosequence. Twenty-five VAM fungal species were identified. Seven species were clearly early successional and five species were clearly late successional. The total number of VAM fungal species did not increase with successional time, but diversity as measured by the Shannon-Wiener index tended to increase, primarily because the community became more even as a single species, Glomus aggregatum, became less dominant in the older sites. Diversity of the VAM fungal community was positively correlated with soil C and N. The density of VAM fungi, as measured by infectivity and total spore count, first increased with time since abandonment and then decreased in the late successional forest sites. Within 12 abandoned fields, VAM fungal density increased with increasing soil pH, H₂O soluble soil C, and root biomass, but was inversely related to extractable soil P and percent cover of non-host plant species. The lower abundance of VAM fungi in the forest sites compared with the field sites agrees with the findings of other workers and corresponds with a shift in the dominant vegetation from herbaceous VAM hosts to woody ectomycorrhizal hosts.     

Plant and insect cuticular lipids serve as behavioral cues for insects

The roles of plant and insect cuticular lipids in insect and plant interactions are reviewed. Emphasis is given to the influence that the host plant and the surface lipids of the host plant have upon insect herbivores and the predators and parasitoids of these herbivores. Variations in cuticular lipids of herbivorous insects are dependent upon the host plant, and these variations may affect the behavior of predators and parasitoids. The cuticular lipids of species which interact on multiple trophic levels are compared. Similarities were found between the hydrocarbons of herbivorous insects, their host plants, and their predators or parasitoids.     

Effects of decomposers and herbivores on plant performance and aboveground plant-insect interactions

Most ecologists acknowledge that plants are subject to complex interactions between both below- and aboveground dwelling animals. However, these complex interactions are seldomly investigated simultaneously. In a factorial common garden experiment we tested single and combined effects of decomposers, root herbivores and leaf herbivores on the growth, flower visitation, and abundance of naturally colonizing aphids and parasitoids on wild mustard ( Sinapis arvensis ). We found that the individual presence of either root herbivores or decomposers resulted in increased aphid abundance, demonstrating that the same aboveground plant–insect interaction can be released by different belowground processes. Enhanced aphid densities caused higher numbers of parasitoids. Furthermore, decomposers increased plant growth and plant fitness (measured as the number of seeds produced), indicating that mustard may benefit from nutrients provided by decomposers, regardless whether plants are attacked by root herbivores or leaf herbivores, or both simultaneously. More flower visits were observed in plants attacked by root herbivores but without leaf herbivores than in plants with both herbivores, suggesting that root herbivory can modify flower attractivity to pollinators. Our results suggest that patterns in plant–insect interactions above the ground are not only affected by aboveground factors but also by a wealth of different belowground processes mediated by the plant.     

Secondary succession on sandy old‐fields in Hungary

Abstract. Development of semi‐natural vegetation has recently been a primary concern of restoration efforts. A primary management question is whether active intervention is required or spontaneous secondary succession could suffice. We studied 54 old‐fields in central Hungary, which differed in time since abandonment but which had similar environmental conditions and management histories. The sites were grouped into four age groups according to the time elapsed since cultivation abandonment: 1–5, 6–10, 11–23 and 24–33 yr. In each old‐field we recorded the species and estimated their abundances. We grouped species in two ways: according to life form (annuals, biennials, perennials, woody plants) and according to coenological behaviour (weeds, sand and steppe generalists, specialists). We analysed the changes in species number and abundance in these categories as a function of site age. Contrary to other successional studies, the total number of species did not change significantly among the four age groups. A significant change was detected between the first two age groups as to life‐form composition. Species number and abundance of annuals decreased, while the perennials and woody plants increased. As to coenological behaviour, species number changed only in the first two age groups, while abundance changed in the first three. Weeds quickly disappeared and specialists established and spread, while the species number and abundance of generalists did not change significantly. We concluded that the basic shifts in species composition are almost completed within 10 yr. Most of the late successional species colonized and weeds disappeared. We conclude that there was no need for active intervention in this system: the spontaneous secondary succession leads to semi‐natural vegetation.     

Plant species diversity, plant biomass and responses of the soil community on abandoned land across Europe: idiosyncracy or above-belowground time lags

We examined the relationship between plant species diversity, productivity and the development of the soil community during early secondary succession on former arable land across Europe. We tested the hypothesis that increasing the initial plant species diversity enhances the biomass production and consequently stimulates soil microbial biomass and abundance of soil invertebrates. We performed five identical field experiments on abandoned arable land in five European countries (CZ, NL, SE, SP and UK) which allowed us to test our hypothesis in a range of climate, soil and other environmental factors that varied between the experimental sites. The initial plant diversity was altered by sowing seed mixtures of mid-successional grassland species with two or five grass species, one or five legumes and one or five forbs. The results of low and high sown diversity treatments were compared with plots that were naturally colonized by species present in the seed bank. In three out of the five field sites, there was no correlation between plant species number and plant biomass production, one site had a positive and the other a negative relation. Treatments with a high diversity seed mixture had a higher biomass than the naturally colonized plots. However, there was no significant difference between high and low sown diversity plots at four out of five sites. The three-year study did not give any evidence of a general bottom-up effect from increased plant biomass on biomass of bacteria, saprophytic fungi or abundance of microarthropods. The biomass of arbuscular mycorrhizal was negatively related to plant biomass. The abundance of nematodes increased after abandonment and was related to plant biomass at four sites. Our results support the hypothesis that plant species diversity may have idiosyncratic effects on soil communities, even though studies on a longer term could reveal time lags in the response to changes in composition and biomass production of plant communities.     

Species richness and parasitism in a fragmented landscape: experiments and field studies with insects on<Emphasis Type="Italic"> Vicia sepium</Emphasis>

Effects of habitat fragmentation on species diversity and herbivore-parasitoid interactions were analyzed using the insect community of seed feeders and their parasitoids in the pods of the bush vetch (Vicia sepium L.). Field studies were carried out on 18 old meadows differing in area and isolation. The area of these meadows was found to be the major determinant of species diversity and population abundance of endophagous insects. Effects of isolation were further analyzed experimentally using 16 small plots with potted vetch plants isolated by 100–500 m from vetch populations on large old meadows. The results showed that colonization success greatly decreased with increasing isolation. In both cases, insect species were not equally affected. Parasitoids suffered more from habitat loss and isolation than their phytophagous hosts. Minimum area requirements, calculated from logistic regressions, were higher for parasitoids than for herbivores. In addition, percent parasitism of the herbivores significantly decreased with area loss and increasing isolation of Vicia sepium plots, supporting the trophic-level hypothesis of island biogeography. Species with high rates of absence on meadows and isolated plant plots were not only characterized by their high trophic level, but also by low abundance and high spatial population variability. Thus conservation of large and less isolated habitat remnants enhances species diversity and parasitism of potential pest insects, i.e., the stability of ecosystem functions. Received: 4 January 1999 / Accepted: 8 September 1999     

Insect communities on experimental mugwort (Artemisia vulgaris L.) plots along an urban gradient

We studied the ability of insect herbivores and their natural enemies to colonize exposed, potted mugwort plants (Artemisia vulgaris L.) along a rural-urban gradient in 1994 in Hamburg (northern Germany). Ectophagous insects, leafmines and galls were monitored weekly from mid-May to mid-September. Endophagous insects were counted by harvesting and dissecting the stems at the end of the growing season. The rural-urban gradient was characterized by a gradient of vegetation-free areas and increasing proportion of ground covered in concrete, tarmac, paving and other impermeable surfaces surrounding the Artemisia plots, i.e. six different zones of increasing isolation. Numbers of insect species (herbivores, parasitoids and predators) decreased along the gradient from 43 to 12. Monophagous herbivores were not more affected than polyphagous herbivores, but parasitoids, especially rare species, were more strongly affected by isolation than predators. Some dominant herbivorous species were very successful colonizers and occurred in inner city sites devoid of all natural vegetation. Sometimes their abundance increased in the inner city to significantly higher densities than in the urban fringe. Isolation appeared to be the main reason for the observed patterns, since area and soil conditions were held constant in the experiment. Microclimate and pollution were considered to play a minor role. Received: 16 January 1997 / Accepted: 26 August 1997     

Interacting effects of herbivory and fertility on a synthesized plant community

1 Outdoor microcosms were used to investigate the effects of invertebrate herbivory on plant community composition, and thereby infer possible effects on the rate of secondary succession, at differing levels of soil fertility.
2 A mixture containing 24 grassland plant species of widely different functional types was established, with 12 microcosms at each of three fertility levels. Four generalist herbivores ( Helix aspersa , Cepaea hortensis , Arianta arbustorum and Sitobion avenae ) were added to half of the microcosms. Above-ground biomass of each species was harvested after 2 years. Reproductive variables were also measured for one species, Poa annua .
3 At both moderate and high soil fertility generalist invertebrate herbivores fed selectively on early successional, fast-growing species, thus increasing the relative abundance of later successional, slow-growing species. This supports the hypothesis that herbivory increases the rate of secondary succession. Flowering and viable seed production of early successional ephemerals was also reduced by the invertebrate herbivores across a wide range of soil fertility. This would seriously reduce the ability of a species to persist in the community, thereby further hastening the rate of succession.     

Meta-analysis of tree diversity effects on the abundance,diversity and activity of herbivores' enemies

The natural enemies hypothesis predicts that the abundance and diversity of antagonists such as predators and parasitoids of herbivores increases with the diversity of plants, which can lead to more effective top-down control of insect herbivores. However, although the hypothesis has received large support in agricultural systems, fewer studies have been conducted in forest ecosystems and a comprehensive synthesis of previous research is still lacking.We conducted a meta-analysis of 65 publications comparing the diversity, abundance or activity of various groups of natural enemies (including birds, bats, spiders and insect parasitoids) in pure vs. mixed forest stands. We tested the effects of forest biome, natural enemy taxon and type of study (managed vs experimental forest).We found a significant positive effect of forest tree diversity on natural enemy abundance and diversity but not on their activity. The effect of tree diversity on natural enemies was stronger towards lower latitudes but was not contingent on the natural enemy taxon level.Overall, our study contributes toward a better understanding of the “natural enemies hypothesis” in forest systems and provides new insights about the mechanisms involved. Furthermore, we outline potential avenues for strengthening forest resistance to the growing threat of herbivorous insects.     

Feral horses dung piles as potential invasion windows for alien plant species in natural grasslands

Small scale disturbances could act as patches that provide sites for the colonization of competitively inferior species, promoting the establishment of non-native species in some cases. We analyzed the vegetation associated with feral horse dung piles in montane pampas grasslands in Mid-East Argentina and described the changes following their abandonment, evaluating whether dung piles act as invasion windows, allowing the entrance of alien plant species. We estimated the portion of the study area directly covered by horse manure and dung height was used to estimate the time elapsed after the abandonment of each pile. Vegetation replacement on dung piles of different ages was assessed and compared with grassland controls using discriminant analysis. We used regression analysis to look for changes in vegetation cover, species richness, species diversity and evenness in response to height (age) of the dung piles, and principal component analyses (PCA) to identify groups of plants associated with different successional stages. We compared cover of alien plant species on dung piles with grassland controls using one-way ANOVA. On average, 2.5% of the study area was covered by horse dung. Total vegetation cover, species richness, diversity and evenness increased after the piles were abandoned. Characteristic plant groups were associated with initial, middle and last phases of the studied succession. Vegetation on the dung piles significantly differed from that in grassland controls and two species were consistently associated with dung piles: the invasive Red Star Thistle, Centaurea calcitrapa, and a native grazing-intolerant grass, Nassella clarazii. Non-native species cover was also higher in dung piles than in control plots. Dung piles cover a significant portion of grassland area in our study site, produce significant changes in the vegetation and are associated with some invasive alien plants that could eventually colonize more pristine areas in the vicinity. On the other hand, they might represent refuges for palatable species, since horses seem to avoid them for grazing.     

Community structure and diversity of bryophytes and vascular plants in abandoned fen meadows

We examined effects of abandonment on species diversity and species composition by comparing 21 calcareous fen meadows in the pre-alpine zone of central and northeastern Switzerland. Meadows were divided into three classes of successional stages (mown: annually mown in late summer, young fallow: 4–15 years, and old fallow: >15 years of abandonment). In each fen, we measured litter mass in four 20 cm×20 cm plots, as well as (aboveground) biomass and species density (number of species per unit area) of bryophytes and vascular plants. Bryophyte biomass was reduced in abandoned fens, whereas litter mass and aboveground biomass of vascular plants increased. Species density of both taxonomic groups was lower in abandoned than in mown fens. Young and old successional stages were not different except for bryophytes, for which old successional stages had higher species density than young stages. We used litter mass and aboveground biomass of vascular plants as covariables in analyses of variance to reveal their effects on species density of both taxonomic groups. For bryophytes, litter mass was more important than vascular plant biomass in explaining variance of species density. This indicates severe effects of burying by litter on bryophyte species density. For species density of vascular plants, both vascular plant biomass and litter mass were of similar importance in explaining the decreased species density. Canonical correspondence analyses showed that abandonment also had an effect on species composition of both bryophytes and vascular plants. However, young and old successional stages were not different indicating fast initial changes after abandonment, but slow secondary succession afterwards. Furthermore, indicator species analysis showed that there was no establishment of new species after abandonment that might dramatically alter fen communities. Re-introduction of mowing as a nature conservation strategy may thus be very promising – even for old fallows.     

Numerical responses of different trophic groups of invertebrates to manipulations of plant diversity in grasslands

We studied the effects of plant diversity on abundance of invertebrate herbivores, parasitoids and predators in two grassland communities (one in Switzerland and one in Sweden) in which plant species richness and functional diversity have been experimentally manipulated. Among herbivores, the abundance of only the most sessile and specialised groups (leafhoppers and wingless aphids) was affected by plant diversity. At both sites, numbers of leafhoppers in sweep net samples showed a linear, negative relationship with plant species number whereas numbers of wingless aphids in suction samples increased with the number of plant functional groups (grasses, legumes, and non-legume forbs) present in the plot. Activity of carabid beetles and spiders (as revealed by pitfall catches) and the total number of predators in pitfalls at the Swiss site decreased linearly with increases in the number of plant species and plant functional groups. Abundance of more specialised enemies, hymenopteran parasitoids, was not affected by the manipulations of plant diversity. Path analysis and analysis of covariance indicated that plant diversity effects on invertebrate abundance were mostly indirect and mediated by changes in plant biomass and cover. At both sites, plant species composition (i.e. the identity of plant species in a mixture) affected numbers of most of the examined groups of invertebrates and was, therefore, a more important determinant of invertebrate abundance in grasslands than plant species richness per se or the number of plant functional groups. The presence of legumes in a mixture was especially important and led to higher numbers of most invertebrate groups. The similarity of invertebrate responses to plant diversity at the two study sites indicates that general patterns in abundance of different trophic groups can be detected across plant diversity gradients under different environmental conditions.