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...

By maintaining a forest-like structure, shaded cocoa plantations contribute to the conservation of ants that usually live in the soil, leaf litter or canopy of tropical forests.Here we synthesize the available information on the diversity and community structure of ants in shaded cocoa plantations in the Atlantic forest region of Brazil, compare ant assemblages in cocoa agroforests with forests and other forms of agriculture, and discuss how these shaded plantations contribute to the conservation of the ants in the Atlantic Forest region. We also discuss ants of economical importance and of special interest, including Camponotus, Dolichoderus, Gnamptogenys, Pachycondyla, Pseudomyrmex and other litter dwelling genera. We discuss the situation of the tramp ant Wasmannia auropunctata in the Bahian cocoa-producing region where it is considered as native, and that of the two cryptobiotic genera Thaumatomyrmex and Typhlomyrmex, as well as that of proven and possible endangered army ant and Ponerini species. A total of 192 ant species from four strata were found in extensive sampling of a cocoa plantation with a relatively simple shade canopy (comprised primarily of Erythrina). Species richness in the cocoa plantations corresponded roughly to that of low diversity native forests, and species composition of cocoa plantations was most similar to native habitats (forest and mangroves) while ant composition in other agricultural habitats was most similar to that of urban areas. Although occurrences of Wasmannia auropunctata were similar in cocoa plantations and forests, abundance of Thaumatomyrmex and Typhlomyrmex, generally thought to be rare ants, was relatively high in cocoa plantations. These results, from cocoa plantations with relatively simple shade, demonstrate the importance of cocoa for ant conservation in the Atlantic forest region of Brazil. It is likely that cocoa plantations with a greater number of vegetation strata and higher tree species richness (such as traditional cabruca plantations) provide even more important habitat for ants generally and for ant species of conservation concern.

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.

Evolution may improve the invasiveness of populations, but it often remains unclear whether key adaptation events occur after introduction into the recipient habitat (i.e. post-introduction adaptation scenario), or before introduction within the native range (i.e. prior-adaptation scenario) or at a primary site of invasion (i.e. bridgehead scenario). We used a multidisciplinary approach to determine which of these three scenarios underlies the invasion of the tropical ant Wasmannia auropunctata in a Mediterranean region (i.e. Israel). Species distribution models (SDM), phylogeographical analyses at a broad geographical scale and laboratory experiments on appropriate native and invasive populations indicated that Israeli populations followed an invasion scenario in which adaptation to cold occurred at the southern limit of the native range before dispersal to Israel. We discuss the usefulness of combining SDM, genetic and experimental approaches for unambiguous determination of eco-evolutionary invasion scenarios.

Many studies have focused on the impacts of climate change on biological assemblages, yet little is known about how climate interacts with other major anthropogenic influences on biodiversity, such as habitat disturbance. Using a unique global database of 1128 local ant assemblages, we examined whether climate mediates the effects of habitat disturbance on assemblage structure at a global scale. Species richness and evenness were associated positively with temperature, and negatively with disturbance. However, the interaction among temperature, precipitation and disturbance shaped species richness and evenness. The effect was manifested through a failure of species richness to increase substantially with temperature in transformed habitats at low precipitation. At low precipitation levels, evenness increased with temperature in undisturbed sites, peaked at medium temperatures in disturbed sites and remained low in transformed sites. In warmer climates with lower rainfall, the effects of increasing disturbance on species richness and evenness were akin to decreases in temperature of up to 98C. Anthropogenic disturbance and ongoing climate change may interact in complicated ways to shape the structure of assemblages, with hot, arid environments likely to be at greatest risk.

The importance of termites as decomposers in tropical forests has long been recognized. Studies on the richness and diversity of termite species and their ecological function have flourished in more recent times, but these have been mostly conducted in a thin stratum within a standing man's reach. Our aims were to evaluate the specific richness and composition of the termite assemblage in the canopy of a tropical rainforest and to determine its originality with respect to the sympatric ground-level fauna. We conducted systematic searches for canopy termites, together with conventional sampling of the sympatric ground-level fauna, in the San Lorenzo forest, Panama. We hypothesized that (1) the canopy accommodates two categories of wood-feeding termites (long-distance foragers and small-colony "one-piece" species) and possibly soil-feeders in suspended soil-like habitats; (2) due to the abundance of soil-feeders, the overall diversity of the ground fauna is higher than that of the canopy; (3) differences in microclimate and resource accessibility favour vertical stratification among wood-feeders. Sixty-three canopy samples yielded ten species of termites, all wood-feeders. Five of these were not found at ground level, although a total of 243 ground samples were collected, representing 29 species. In addition to long-distance foragers (Microcerotermes and Nasutitermes spp.) and small-colony termites (mostly Kalotermitidae), the canopy fauna included Termes hispaniolae, a wood-feeding Termitidae from an allegedly soil-feeding genus, living in large dead branches. Soil-feeders were absent from the canopy, probably because large epiphytes were scarce. As predicted, the ground fauna was much richer than that of the canopy, but the species richness of both habitats was similar when only wood-feeders were considered. Vertical stratification was strongly marked among wood-feeders, as all common species, apart from the arboreal-nesting Microcerotermes arboreus, could unequivocally be assigned to either a ground or a canopy group. The canopy, therefore, contributes significantly to the total species richness of the termite assemblage, and the diversity, abundance and ecological importance of canopy termites in tropical rainforests may be higher than previously recognized.

In this manuscript we present a focus stacking system, composed of commercial photographic equipment. The system is inexpensive compared to high-end commercial focus stacking solutions. We tested this system and compared the results with several different software packages (CombineZP, Auto-Montage, Helicon Focus and Zerene Stacker). We tested our final stacked picture with a picture obtained from two high-end focus stacking solutions: a Leica MZ16A with DFC500 and a Leica Z6APO with DFC290. Zerene Stacker and Helicon Focus both provided satisfactory results. However, Zerene Stacker gives the user more possibilities in terms of control of the software, batch processing and retouching. The outcome of the test on high-end solutions demonstrates that our approach performs better in several ways. The resolution of the tested extended focus pictures is much higher than those from the Leica systems. The flash lighting inside the Ikea closet creates an evenly illuminated picture, without struggling with filters, diffusers, etc. The largest benefit is the price of the set-up which is approximately € 3,000, which is 8 and 10 times less than the LeicaZ6APO and LeicaMZ16A set-up respectively. Overall, this enables institutions to purchase multiple solutions or to start digitising the type collection on a large scale even with a small budget.

A reliable characterization of community diversity and composition, necessary to allow inter-site comparisons and to monitor changes, is especially difficult to reach in speciose invertebrate communities. Spatial components of the sampling design (sampling interval, extent and grain) as well as temporal variations of species density affect the measures of diversity (species richness S, Buzas and Gibson's evenness E and Shannon's heterogeneity H). Our aim was to document the small-scale spatial distribution of leaf litter ants in a subtropical dry forest of the Argentinian Chaco and analyze how the community characterization was best achieved with a minimal sampling effort. The work was based on the recent standardized protocol for collecting ants of the leaf litter (''A.L.L.'': 20 samples at intervals of 10 m). To evaluate the consistency of the sampling method in time and space, the selected site was first subject to a preliminary transect, then submitted after a 9-month interval to an 8-fold oversampling campaign (160 samples at interval of 1.25 m). Leaf litter ants were extracted from elementary 1 m 2 quadrats with Winkler apparatus. An increase in the number of samples collected increased S and decreased E but did not affect much H. The sampling interval and extent did not affect S and H beyond a distance of 10 m between samples. An increase of the sampling grain had a similar effect on S than a corresponding increase of the number of samples collected, but caused a proportionaly greater increase of H. The density of species m (2 varied twofold after a 9-month interval; the effect on S could only be partially corrected by rarefaction. The measure of species numerical dominance was little affected by the season. A single standardized A.L.L. transect with Winkler samples collected B/45% of the species present in the assemblage. All frequent species were included but their relative frequency was not always representative. A log series distribution of species occurrences was oberved. Fisher's a and Shannon's H were the most appropriate diversity indexes. The former was useful to rarefy or abundify S and the latter was robust against sample size effects. Both parametric and Soberó n and Llorente extrapolation methods outperformed nonparametric methods and yielded a fair estimate of total species richness along the transect, a minimum value of S for the habitat sampled.Conservation biologists and environmental planners need reliable methods to evaluate the biological value of sites and to monitor changes over time. A major difficulty encountered when conducting diversity inventories is that species diversity cannot be recorded without reference to space, time and collection method. Components of species diversity include species richness (S, the number of species) and species evenness (E, equitability