Insect outbreaks in forest and agriculture monocultures led Charles Elton to propose, a half-century ago, that higher plant diversity stabilized animal foodweb dynamics in natural ecosystems. We tested this hypothesis by studying arthropod community dynamics in a long-term experimental manipulation of grassland plant species diversity. Over the course of a decade, we found that higher plant diversity increased the stability (i.e. lowered year-to-year variability) of a diverse (>700 species) arthropod community across trophic levels. As the number of plant species increased, the stability of both herbivore and predator species richness and of total herbivore abundance increased. The underlying mechanisms driving these diversity-stability relationships were plant diversity, via effects on primary productivity and plant community stability, and portfolio effects. Taken together, our results show that higher plant diversity provides more temporally consistent food and habitat resources to arthropod foodwebs. Consequently, actively managing for high plant diversity may have stronger than expected benefits for increasing animal diversity and controlling pest outbreaks.

Plant diversity is predicted to be positively linked to the diversity of herbivores and predators in a foodweb. Yet, the relationship between plant and animal diversity is explained by a variety of competing hypotheses, with mixed empirical results for each hypothesis. We sampled arthropods for over a decade in an experiment that manipulated the number of grassland plant species. We found that herbivore and predator species richness were strongly, positively related to plant species richness, and that these relationships were caused by different mechanisms at herbivore and predator trophic levels. Even more dramatic was the threefold increase, from low- to high-plant species richness, in abundances of predatory and parasitoid arthropods relative to their herbivorous prey. Our results demonstrate that, over the long term, the loss of plant species propagates through food webs, greatly decreasing arthropod species richness, shifting a predator-dominated trophic structure to being herbivore dominated, and likely impacting ecosystem functioning and services.

We experimentally separated the effects of two components of plant diversity-plant species richness and plant functional group richness-on insect communities. Plant species richness and plant functional group richness had contrasting effects on insect abundances, a result we attributed to three factors. First, lower insect abundances at higher plant functional group richness were explained by a sampling effect, which was caused by the increasing likelihood that one low-quality group, C4 grasses, would be present and reduce average insect abundances by 25%. Second, plant biomass, which was positively related to plant functional group richness, had a strong, positive effect on insect abundances. Third, a positive effect of plant species richness on insect abundances may have been caused by greater availability of alternate plant resources or greater vegetational structure. In addition, a greater diversity of insect species, whose individual abundances were often unaffected by changes in plant species richness, may have generated higher total community abundances. After controlling for the strong, positive influence of insect abundance on insect diversity through rarefaction, insect species richness increased as plant species richness and plant functional group richness increased. Although these variables did not explain a high proportion of variation individually, plant species richness and plant functional group richness had similar effects on insect diversity and opposing effects on insect abundances, and both factors may explain how the loss of plant diversity influences higher trophic levels.

Characteristics used to categorize plant species into functional groups for their effects on ecosystem functioning may also be relevant to higher trophic levels. In addition, plant and consumer diversity should be positively related because more diverse plant communities offer a greater variety of resources for the consumers. Thus, the Functional group composition and richness of a plant community may affect the composition and diversity of the herbivores and even higher trophic levels associated with that community. We tested this hypothesis by sampling arthropods with a Vacuum sampler (34531 individuals of 494 species) from an experiment in which we manipulated plant Functional group richness and composition. Plant manipulations included all combinations of three functional groups (forbs, C-3 graminoids, and C-4 graminoids) removed zero, one, or two at a time from grassland plots at Cedar Creek Natural History Area. MN. Although total arthropod species richness was unrelated to plant Functional group richness or composition, the species richness of some arthropod orders was affected by plant Functional group composition.Two plant characteristics explained most of the effects of plant functional groups on arthropod species richness. Nutritional quality, a characteristic related to ecosystem Functioning, and taxonomic diversity, a characteristic not used to designate plant Functional groups, seemed to affect arthropod species richness both directly and indirectly. Thus, plant functional groups designated For their effects on ecosystem processes will only be partially relevant to consumer diversity and abundance.

Just as long-term nitrogen loading of grasslands decreases plant species richness and increases plant biomass, we have found that nitrogen loading decreases insect species richness and increases insect abundances. We sampled 54 plots that had been maintained at various rates of nitrogen addition for 14 years. Total insect species richness and effective insect diversity, as well as herbivore and predator species richness, were significantly, negatively related to the rate of nitrogen addition. However, there was variation in trophic responses to nitrogen. Detritivore species richness increased as nitrogen addition increased, and parasitoids showed no response. Insect abundances, measured as the number of insects and insect biovolume (an estimate of biomass), were significantly, positively related to the rate of nitrogen addition, as were the abundances of herbivores and detritivores. Parasitoid abundance was negatively related to the rate of nitrogen addition. Changes in the insect community were correlated with changes in the plant community. As rates of nitrogen addition increased, plant species richness decreased, plant productivity and plant tissue nitrogen increased, and plant composition shifted from C-4 to C-3 grass species. Along this gradient, total insect species richness and effective insect diversity were most strongly, positively correlated with plant species richness. Insect biovolume was negatively correlated with plant species richness. Responses of individual herbivores varied along the nitrogen gradient, but numbers of 13 of the 18 most abundant herbivores were positively correlated with their host plant biomass. Although insect communities did not respond as strongly as plant communities, insect species richness, abundance, and composition were impacted by nitrogen addition. This study demonstrates that long-term nitrogen loading affects the entire food chain, simplifying both plant and insect communities.

Declining biodiversity represents one of the most dramatic and irreversible aspects of anthropogenic global change, yet the ecological implications of this change are poorly understood. Recent studies have shown that biodiversity loss of basal species, such as autotrophs or plants, affects fundamental ecosystem processes such as nutrient dynamics and autotrophic production. Ecological theory predicts that changes induced by the loss of biodiversity at the base of an ecosystem should impact the entire system. Here we show that experimental reductions in grassland plant richness increase ecosystem vulnerability to invasions by plant species, enhance the spread of plant fungal diseases, and alter the richness and structure of insect communities. These results suggest that the loss of basal species may have profound effects on the integrity and functioning of ecosystems.

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.

1.The empirical relationships among body size, species richness and number of individuals may give insight into the factors controlling species diversity and the relative abundances of species. To determine these relationships, we sampled the arthropods of grasslands and savannahs at Cedar Creek, MN using sweep nets (90 525 individuals of 1225 species) and pitfall traps (12 721 individuals of 92 species). Specimens were identified, enumerated and measured to determine body size.2.Both overall and within abundant taxonomic orders, species richness and numbers of individuals peaked at body sizes intermediate for each group. Evolution could create unimodal diversity patterns by random diversification around an ancestral body size or from size‐dependent fitness differences. Local processes such as competition or predation could also create unimodal diversity distributions.3.The average body size of a species depended significantly on its taxonomic order, but on contemporary trophic role only within the context of taxonomic order.4.Species richness (Si) within size classes was related to the number of individuals (Ii) asSi = Ii0·5. This relationship held across a 100 000‐fold range of body sizes. Within size classes, abundance distributions of size classes were all similar power functions. A general rule of resource division, together with similar minimum population sizes, is sufficient to generate the relationship between species richness and number of individuals.5.Smaller bodied species had slightly shallower abundance distributions and may, in general, persist at lower densities than larger species.6.Our results suggest there may be fewer undescribed small arthropod species than previously thought and that most undescribed species will be smaller than arthropods.

Because a diversity of resources should support a diversity of consumers, most models predict that increasing plant diversity increases animal diversity. We report results of a direct experimental test of the dependence of animal diversity on plant diversity. We sampled arthropods in a well-replicated grassland experiment in which plant species richness and plant functional richness were directly manipulated. In simple regressions, both the number of species planted (log(2) transformed) and the number of functional groups planted significantly increased arthropod species richness but not arthropod abundance. However, the number of species planted was the only significant predictor of arthropod species richness when both predictor variables were included in ANOVAs or a MANOVA. Although highly significant, arthropod species richness regressions had low R-2 values, high intercepts (24 arthropod species in monocultures), and shallow slopes. Analyses of relations among plants and arthropod trophic groups indicated that herbivore diversity was influenced by plant, parasite, and predator diversity. Furthermore, herbivore diversity was more strongly correlated with parasite and predator diversity than with plant diversity. Together with regression results, this suggests that, although increasing plant diversity significantly increased arthropod diversity, local herbivore diversity is also maintained by, and in turn maintains, a diversity of parasites and predators.

-We investigated the effects of prescribed burning on the composition, abundance, species richness and diversity of oak savanna arthropod communities in a replicated, large-scale, 30-y experiment. We employed four sampling methods over 3 y and caught 11,215 arthropods of 551 species. Species had varied and often negatively correlated short vs. longterm responses to burning. In the years savannas were burned, species richness and abundance of arthropods, especially Homoptera and Lepidoptera, were reduced and species compositions were sometimes more similar. However, despite its major effect on vegetation, frequency of burning did not affect arthropod abundance and species richness, but sometimes caused savannas to be similar in species or taxonomic order composition. The Shannon diversity index was unaffected by burning. On the whole, prescribed burns necessary to maintain grasslands and savannas do not appear to be harmful to, or to greatly impact, the

BIOLOGICAL diversity, population size and body size are interdependent(1-8), but there is little consensus on the nature or causes of these relations. Here we analyse the most thoroughly sampled ecological community to date, a grassland insect community sample containing 89,596 individuals of 1,167 species. Each taxonomic order had a distinct body size at which both species richness and number of individuals were highest, but these peak sizes varied more than 100-fold among five major orders, These results suggest that there may be fewer undiscovered small insect species than previously thought. Moreover, we found a surprisingly strong, simple, but unreported, relation between species richness (S) and the number of individuals (I) within size classes, S =I-0.5. Because this held across numerous body types and a 100,000-fold body-size range, there may be a general rule that is independent of body size for the relations among interspecific resource division, abundance and diversity.