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...

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

A severe outbreak of Lepidoptera followed the 1997–98 El Niño Southern Oscillation event, during which the climate in central Panama was unusually dry. The outbreak involved the larvae of at least 12 species of Lepidoptera and occurred at a seasonally dry, deciduous forest site, where extensive background data were available regarding climate, tree species and non-outbreak herbivory levels. Most Lepidoptera were associated with only one or two larval host plant species belonging to the same family, and the majority were monophagous during this study. During the outbreak, caterpillar densities for the major outbreak species averaged 1.6 larvae per young leaf and 0.18 larvae per leaf for leaves of all ages. For canopy trees and lianas, the mean level of leaf damage was 13.8%, ranging from 1–100%. Seven out of 20 tree species sustained most of the damage, with 21–37% of the leaf area consumed. Relative to non-outbreak years, damage levels increased by more than 250% during the outbreak. Single-species outbreaks were observed in other areas with a similar drought, but wetter forests in central Panama did not experience outbreaks during this period. Historically, fewer outbreaks have been reported from tropical forests than from temperate forests, however, similar El Niño-related outbreaks have occurred recently in several other locations throughout the tropics.

Abstract. 1. Networks of feeding interactions among insect herbivores and natural enemies such as parasitoids, describe the structure of these assemblages and may be critically linked to their dynamics and stability. The present paper describes the first quantitative study of parasitoids associated with gall-inducing insect assemblages in the tropics, and the first investigation of vertical stratification in quantitative food web structure.2. Galls and associated parasitoids were sampled in the understorey and canopy of Parque Natural Metropolitano in the Pacific forest, and in the understorey of San Lorenzo Protected Area in the Caribbean forest of Panama. Quantitative host-parasitoid food webs were constructed for each assemblage, including 34 gall maker species, 28 host plants, and 57 parasitoid species.3. Species richness was higher in the understorey for parasitoids, but higher in the canopy for gall makers. There was an almost complete turnover in gall maker and parasitoid assemblage composition between strata, and the few parasitoid species shared between strata were associated with the same host species.4. Most parasitoid species were host specific, and the few polyphagous parasitoid species were restricted to the understorey.5. These results suggest that, in contrast to better-studied leaf miner-parasitoid assemblages, the influence of apparent competition mediated by shared parasitoids as a structuring factor is likely to be minimal in the understorey and practically absent in the canopy, increasing the potential for coexistence of parasitoid species.6. High parasitoid beta diversity and high host specificity, particularly in the poorly studied canopy, indicate that tropical forests may be even more species rich in hymenopteran parasitoids than previously suspected.