Two decades of research [1][2][3][4] have not established whether tropical insect herbivores are dominated by specialists or generalists. This impedes our understanding of species coexistence in diverse rainforest communities. Host specificity and species richness of tropical insects are also key parameters in mapping global patterns of biodiversity 1,4,5 . Here we analyse data for over 900 herbivorous species feeding on 51 plant species in New Guinea and show that most herbivorous species feed on several closely related plant species. Because species-rich genera are dominant in tropical floras, monophagous herbivores are probably rare in tropical forests. Furthermore, even between phylogenetically distant hosts, herbivore communities typically shared a third of their species. These results do not support the classical view that the coexistence of herbivorous species in the tropics is a consequence of finely divided plant resources; non-equilibrium models of tropical diversity 6 should instead be considered. Low host specificity of tropical herbivores reduces global estimates of arthropod diversity from 31 million (ref. 1) to 4-6 million species. This finding agrees with estimates based on taxonomic collections, reconciling an order of magnitude discrepancy between extrapolations of global diversity based on ecological samples of tropical communities with those based on sampling regional faunas 7,8 .Host specificity is difficult to measure, and the limitations of existing studies include sampling only certain taxonomic groups rather than entire guilds, or sampling limited numbers of host plant species and lineages. Studies are often of insufficient duration, producing samples too small for quantitative analysis, or insects are sampled destructively, which precludes feeding experiments and the study of immature stages. Further, previous studies 2 failed to consider the phylogenetic relationships of host plants by using measures of host specificity that relied on counts of higher plant taxa (for example, genera or families). This approach can be misleading when taxonomic ranks are not commensurate with plant lineages. We examined the impacts of sampling bias and phylogenetic effects on estimates of host specificity by analysing the largest available data set of its kind. The leaf-chewing insect community on 51 plant species was characterized by using a sample

Understanding variation in resource specialization is important for progress on issues that include coevolution, community assembly, ecosystem processes, and the latitudinal gradient of species richness. Herbivorous insects are useful models for studying resource specialization, and the interaction between plants and herbivorous insects is one of the most common and consequential ecological associations on the planet. However, uncertainty persists regarding fundamental features of herbivore diet breadth, including its relationship to latitude and plant species richness. Here, we use a global dataset to investigate host range for over 7,500 insect herbivore species covering a wide taxonomic breadth and interacting with more than 2,000 species of plants in 165 families. We ask whether relatively specialized and generalized herbivores represent a dichotomy rather than a continuum from few to many host families and species attacked and whether diet breadth changes with increasing plant species richness toward the tropics. Across geographic regions and taxonomic subsets of the data, we find that the distribution of diet breadth is fit well by a discrete, truncated Pareto power law characterized by the predominance of specialized herbivores and a long, thin tail of more generalized species. Both the taxonomic and phylogenetic distributions of diet breadth shift globally with latitude, consistent with a higher frequency of specialized insects in tropical regions. We also find that more diverse lineages of plants support assemblages of relatively more specialized herbivores and that the global distribution of plant diversity contributes to but does not fully explain the latitudinal gradient in insect herbivore specialization.

In the wake of widespread loss of old-growth forests throughout the tropics, secondary forests will likely play a growing role in the conservation of forest biodiversity. We considered a complex hierarchy of factors that interact in space and

Despite recent progress in understanding mechanisms of tree species coexistence in tropical forests, a simple explanation for the even more extensive diversity of insects feeding on these plants has been missing. We compared folivorous insects from temperate and tropical trees to test the hypothesis that herbivore species coexistence in more diverse communities could reflect narrow host specificity relative to less diverse communities. Temperate and tropical tree species of comparable phylogenetic distribution supported similar numbers of folivorous insect species, 29.0 ± 2.2 and 23.5 ± 1.8 per 100 square meters of foliage, respectively. Host specificity did not differ significantly between community samples, indicating that food resources are not more finely partitioned among folivorous insects in tropical than in temperate forests. These findings suggest that the latitudinal gradient in insect species richness could be a direct function of plant diversity, which increased sevenfold from our temperate to tropical study sites.

Most eukaryotic organisms are arthropods. Yet, their diversity in rich terrestrial ecosystems is still unknown. Here we produce tangible estimates of the total species richness of arthropods in a tropical rainforest. Using a comprehensive range of structured protocols, we sampled the phylogenetic breadth of arthropod taxa from the soil to the forest canopy in the San Lorenzo forest, Panama. We collected 6144 arthropod species from 0.48 hectare and extrapolated total species richness to larger areas on the basis of competing models. The whole 6000-hectare forest reserve most likely sustains 25,000 arthropod species. Notably, just 1 hectare of rainforest yields >60% of the arthropod biodiversity held in the wider landscape. Models based on plant diversity fitted the accumulated species richness of both herbivore and nonherbivore taxa exceptionally well. This lends credence to global estimates of arthropod biodiversity developed from plant models.M ost eukaryote species are terrestrial arthropods (1), and most terrestrial arthropods occur in tropical rainforests (2). However, considerably greater sampling effort is required in tropical arthropod surveys to yield realistic estimates of global species richness (3-7). A basic hindrance to estimating global biodiversity lies in a lack of empirical data that establish local biodiversity, which can be scaled up to achieve a global estimate.Although many studies reported species richness for selected groups of well-studied insect taxa, no satisfactory estimate of total arthropod species richness exists for a single tropical rainforest location to date.The unstructured collection and small-scale survey of tropical arthropods cannot yield convincing estimates of total species richness at a specific forest (7-9). Most studies either target few arthropod orders or trophic guilds, or use a limited array of sampling methods, or ignore the diverse upper canopy regions of tropical forests (10-15). Moreover, sampling protocols have rarely been structured in such a way that, with increased sampling, incomplete data on local diversity (7) can be extrapolated to estimate total species richness across multiple spatial scales (16). Where such structured estimates are made, it is invariably for insect herbivores on their host plants (5). However, species accumulation rates may differ markedly for nonherbivore guilds, which include more than half of all described arthropod species (1, 17). As the degree of host specificity (effective specialization) of other guilds can be much lower than that of insect herbivores, or may be driven by different factors (18,19), global estimates based on herbivores alone are questionable. Consequently, extensive cross-taxon surveys with structured protocols at reference sites may be the only effective approach toward estimating total arthropod species richness in tropical forests (3).To provide a comprehensive estimate of total arthropod species richness in a tropical rainforest, we established a collaboration involving 102 researchers with expertise encom...

Inventory of the caterpillars, their food plants and parasitoids began in 1978 for today's Area de Conservacion Guanacaste (ACG), in northwestern Costa Rica. This complex mosaic of 120 000 ha of conserved and regenerating dry, cloud and rain forest over 0-2000 m elevation contains at least 10 000 species of non-leaf-mining caterpillars used by more than 5000 species of parasitoids. Several hundred thousand specimens of ACG-reared adult Lepidoptera and parasitoids have been intensively and extensively studied morphologically by many taxonomists, including most of the co-authors. DNA barcoding -the use of a standardized short mitochondrial DNA sequence to identify specimens and flush out undisclosed species -was added to the taxonomic identification process in 2003.

Summary 1.The extent to which plant-herbivore feeding interactions are specialized is key to understand the processes maintaining the diversity of both tropical forest plants and their insect herbivores. However, studies documenting the full complexity of tropical plant-herbivore food webs are lacking. 2. We describe a complex, species-rich plant-herbivore food web for lowland rain forest in Papua New Guinea, resolving 6818 feeding links between 224 plant species and 1490 herbivore species drawn from 11 distinct feeding guilds. By standardizing sampling intensity and the phylogenetic diversity of focal plants, we are able to make the first rigorous and unbiased comparisons of specificity patterns across feeding guilds. 3. Specificity was highly variable among guilds, spanning almost the full range of theoretically possible values from extreme trophic generalization to monophagy. 4. We identify guilds of herbivores that are most likely to influence the composition of tropical forest vegetation through density-dependent herbivory or apparent competition. 5. We calculate that 251 herbivore species (48 of them unique) are associated with each rain forest tree species in our study site so that the 200 tree species coexisting in the lowland rain forest community are involved in 50 000 trophic interactions with 9600 herbivore species of insects. This is the first estimate of total herbivore and interaction number in a rain forest plant-herbivore food web. 6. A comprehensive classification of insect herbivores into 24 guilds is proposed, providing a framework for comparative analyses across ecosystems and geographical regions.

Recent advances in understanding insect communities in tropical forests have contributed little to our knowledge of large-scale patterns of insect diversity, because incomplete taxonomic knowledge of many tropical species hinders the mapping of their distribution records. This impedes an understanding of global biodiversity patterns and explains why tropical insects are under-represented in conservation biology. Our study of approximately 500 species from three herbivorous guilds feeding on foliage (caterpillars, Lepidoptera), wood (ambrosia beetles, Coleoptera) and fruit (fruitflies, Diptera) found a low rate of change in species composition (beta diversity) across 75,000 square kilometres of contiguous lowland rainforest in Papua New Guinea, as most species were widely distributed. For caterpillars feeding on large plant genera, most species fed on multiple host species, so that even locally restricted plant species did not support endemic herbivores. Large plant genera represented a continuously distributed resource easily colonized by moths and butterflies over hundreds of kilometres. Low beta diversity was also documented in groups with differing host specificity (fruitflies and ambrosia beetles), suggesting that dispersal limitation does not have a substantial role in shaping the distribution of insect species in New Guinea lowland rainforests. Similar patterns of low beta diversity can be expected in other tropical lowland rainforests, as they are typically situated in the extensive low basins of major tropical rivers similar to the Sepik-Ramu region of New Guinea studied here.