Insects on flowers: The unexpectedly high biodiversity of flower-visiting beetles in a tropical rainforest canopy.

Insect biodiversity peaks in tropical rainforest environments where a large but as yet unknown proportion of species are found in the canopy. While there has been a proliferation of insect biodiversity research undertaken in the rainforest canopy, most studies focus solely on insects that inhabit the foliage. In a recent paper, we examined the distribution of canopy insects across five microhabitats (mature leaves, new leaves, flowers, fruit and suspended dead wood) in an Australian tropical rainforest, showing that the density (per dry weight gram of microhabitat) of insects on flowers were ten to ten thousand times higher than on the leaves. Flowers also supported a much higher number of species than expected based on their contribution to total forest biomass. Elsewhere we show that most of these beetle species were specialized to flowers with little overlap in species composition between different canopy microhabitats. Here we expand our discussion of the implications of our results with respect to specialization and the generation of insect biodiversity in the rainforest canopy. Lastly, we identify future directions for research into the biodiversity and specialization of flower-visitors in complex tropical rainforests.

Estimating global arthropod species richness: refining probabilistic models using probability bounds analysis.

A key challenge in the estimation of tropical arthropod species richness is the appropriate management of the large uncertainties associated with any model. Such uncertainties had largely been ignored until recently, when we attempted to account for uncertainty associated with model variables, using Monte Carlo analysis. This model is restricted by various assumptions. Here, we use a technique known as probability bounds analysis to assess the influence of assumptions about (1) distributional form and (2) dependencies between variables, and to construct probability bounds around the original model prediction distribution. The original Monte Carlo model yielded a median estimate of 6.1 million species, with a 90 % confidence interval of [3.6, 11.4]. Here we found that the probability bounds (p-bounds) surrounding this cumulative distribution were very broad, owing to uncertainties in distributional form and dependencies between variables. Replacing the implicit assumption of pure statistical independence between variables in the model with no dependency assumptions resulted in lower and upper p-bounds at 0.5 cumulative probability (i.e., at the median estimate) of 2.9-12.7 million. From here, replacing probability distributions with probability boxes, which represent classes of distributions, led to even wider bounds (2.4-20.0 million at 0.5 cumulative probability). Even the 100th percentile of the uppermost bound produced (i.e., the absolutely most conservative scenario) did not encompass the well-known hyper-estimate of 30 million species of tropical arthropods. This supports the lower estimates made by several authors over the last two decades.

The overlooked biodiversity of flower-visiting invertebrates.

Estimates suggest that perhaps 40% of all invertebrate species are found in tropical rainforest canopies. Extrapolations of total diversity and food web analyses have been based almost exclusively on species inhabiting the foliage, under the assumption that foliage samples are representative of the entire canopy. We examined the validity of this assumption by comparing the density of invertebrates and the species richness of beetles across three canopy microhabitats (mature leaves, new leaves and flowers) on a one hectare plot in an Australian tropical rainforest. Specifically, we tested two hypotheses: 1) canopy invertebrate density and species richness are directly proportional to the amount of resource available; and 2) canopy microhabitats represent discrete resources that are utilised by their own specialised invertebrate communities. We show that flowers in the canopy support invertebrate densities that are ten to ten thousand times greater than on the nearby foliage when expressed on a per-unit resource biomass basis. Furthermore, species-level analyses of the beetle fauna revealed that flowers support a unique and remarkably rich fauna compared to foliage, with very little species overlap between microhabitats. We reject the hypothesis that the insect fauna on mature foliage is representative of the greater canopy community even though mature foliage comprises a very large proportion of canopy plant biomass. Although the significance of the evolutionary relationship between flowers and insects is well known with respect to plant reproduction, less is known about the importance of flowers as resources for tropical insects. Consequently, we suggest that this constitutes a more important piece of the 'diversity jigsaw puzzle' than has been previously recognised and could alter our understanding of the evolution of plant-herbivore interactions and food web dynamics, and provide a better foundation for accurately estimating global species richness.

Quantifying uncertainty in estimation of tropical arthropod species richness.

There is a bewildering range of estimates for the number of arthropods on Earth. Several measures are based on extrapolation from species specialized to tropical rain forest, each using specific assumptions and justifications. These approaches have not provided any sound measure of uncertainty associated with richness estimates. We present two models that account for parameter uncertainty by replacing point estimates with probability distributions. The models predict medians of 3.7 million and 2.5 million tropical arthropod species globally, with 90% confidence intervals of [2.0, 7.4] million and [1.1, 5.4] million, respectively. Estimates of 30 million or greater are predicted to have <0.00001 probability. Sensitivity analyses identified uncertainty in the proportion of canopy arthropod species that are beetles as the most influential parameter, although uncertainties associated with three other parameters were also important. Using the median estimates suggests that in spite of 250 years of taxonomy and around 855,000 species of arthropods already described, approximately 70% await description.

How do beetle assemblages respond to cyclonic disturbance of a fragmented tropical rainforest landscape?

There are surprisingly few studies documenting effects of tropical cyclones (including hurricanes and typhoons) on rainforest animals, and especially insects, considering that many tropical forests are frequently affected by cyclonic disturbance. Consequently, we sampled a beetle assemblage inhabiting 18 upland rainforest sites in a fragmented landscape in north-eastern Queensland, Australia, using a standardised sampling protocol in 2002 and again 12 months after the passage of Severe Tropical Cyclone Larry (March 2006). The spatial configuration of sites allowed us to test if the effects of a cyclone and those from fragmentation interact. From all insect samples we extracted 12,568 beetles of 382 species from ten families. Beetle species composition was significantly different pre-and post-cyclone although the magnitude of faunal change was not large with 205 species, representing 96% of all individuals, present in both sampling events. Sites with the greatest changes to structure had the greatest changes in species composition. At the site level, increases in woody debris and wood-feeding beetle (Scolytinae) counts were significantly correlated but changes in the percent of ground vegetation were not mirrored by changes in the abundance of foliage-feeding beetles (Chrysomelidae). The overall direction of beetle assemblage change was consistent with increasing aridity, presumably caused by the loss of canopy cover. Sites with the greatest canopy loss had the strongest changes in the proportion of species previously identified in the pre-cyclone study as preferring arid or moist rainforest environments. The magnitude of fragmentation effects was virtually unaltered by the passage of Cyclone Larry. We postulate that in the short-term the effects of cyclonic disturbance and forest fragmentation both reduce the extent of moist, interior habitat.

Beetle assemblages from an Australian tropical rainforest show that the canopy and the ground strata contribute equally to biodiversity.

There remains great uncertainty about how much tropical forest canopies contribute to global species richness estimates and the relative specialization of insect species to vertical zones. To investigate these issues, we conducted a four-year sampling program in lowland tropical rainforest in North Queensland, Australia. Beetles were sampled using a trap that combines Malaise and flight interception trap (FIT) functions. Pairs of this trap, one on the ground and a second suspended 15-20 m above in the canopy were located at five sites, spaced 50 m or more apart. These traps produced 29986 beetles of 1473 species and 77 families. There were similar numbers of individuals (canopy 14473; ground 15513) and species (canopy 1158; ground 895) in each stratum, but significantly more rare species in the canopy (canopy 509; ground 283). Seventy two percent of the species (excluding rare species) were found in both strata. Using IndVal, we found 24 and 27% of the abundant species (n>or=20 individuals) to be specialized to the canopy and the ground strata, respectively, and equivalent analyses at the family level showed figures of 30 and 22%, respectively. These results show that the canopy and the ground strata both provide important contributions to rainforest biodiversity.