The majority of living plant species are pollinated by insects, and this interaction is thought to have played a major role in driving the diversification of modern angiosperms. But while flower-insect interactions have been well studied from the perspective of plants in the form of pollination biology, few studies have been carried out from an entomological perspective, where flowers are resources to exploit. As a consequence, it remains unknown how many insect species actually utilise floral resources, especially since many flower-visitors do not carry out pollination and may therefore be widely ignored in pollination studies. In this review, I attempt to present an overview of the taxonomic range of flower-visiting invertebrates and estimate the proportion of described species that regularly utilise flowers. The flower-visiting habit has likely evolved independently hundreds of times across more than a dozen modern invertebrate orders. I speculate, based on reviewing the literature and discussions with experts, that *30 % of arthropod species ([350,000 described species) may regularly utilise flowers to feed, find a mate, or acquire other resources. When extrapolated to the estimated global diversity of the phylum Arthropoda, perhaps more than a million species regularly visit flowers. However, generating more accurate estimates will require much more work from the perspective of flower-visiting insects, including the often-ignored species that do not pollinate host plants. In particular, sampling techniques in addition to traditional observation protocols should be encouraged to ensure that all flower-visitors are recorded. Greater efforts to identify flower-visiting species beyond the level of order or family will also enhance our understanding of flower-visitor diversity.

Arguably the majority of species on Earth utilise tropical rainforest canopies, and much progress has been made in describing arboreal assemblages, especially for arthropods. The most commonly described patterns for tropical rainforest insect communities are host specificity, spatial specialisation (predominantly vertical stratification), and temporal changes in abundance (seasonality and circadian rhythms). Here I review the recurrent results with respect to each of these patterns and discuss the evolutionary selective forces that have generated them in an attempt to unite these patterns in a holistic evolutionary framework. I propose that species can be quantified along a generalist-specialist scale not only with respect to host specificity, but also other spatial and temporal distribution patterns, where specialisation is a function of the extent of activity across space and time for particular species. When all of these distribution patterns are viewed through the paradigm of specialisation, hypotheses that have been proposed to explain the evolution of host specificity can also be applied to explain the generation and maintenance of other spatial and temporal distribution patterns. The main driver for most spatial and temporal distribution patterns is resource availability. Generally, the distribution of insects follows that of the resources they exploit, which are spatially stratified and vary temporally in availability. Physiological adaptations are primarily important for host specificity, where nutritional and chemical variation among host plants in particular, but also certain prey species and fungi, influence host range. Physiological tolerances of abiotic conditions are also important for explaining the spatial and temporal distributions of some insect species, especially in drier forest environments where desiccation is an ever-present threat. However, it is likely that for most species in moist tropical rainforests, abiotic conditions are valuable indicators of resource availability, rather than physiologically limiting factors. Overall, each distribution pattern is influenced by the same evolutionary forces, but at differing intensities. Consequently, each pattern is linked and not mutually exclusive of the other distribution patterns. Most studies have examined each of these patterns in isolation. Future work should focus on examining the evolutionary drivers of these patterns in concert. Only then can the relative strength of resource availability and distribution, host defensive phenotypes, and biotic and abiotic interactions on insect distribution patterns be determined.

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.

Summary1. We tested the hypotheses that feeding guild structure of beetle assemblages changed with different arboreal microhabitats and that these differences were consistent across rainforest tree species. 2. Hand collection and beating techniques were used from the gondola of the Australian Canopy Crane to collect beetles from five microhabitats (mature leaves, flush leaves, flowers, fruit and suspended dead wood) within the rainforest canopy. A simple randomization procedure was implemented to test whether the abundances of each feeding guild on each microhabitat were different from that expected based on a null hypothesis of random distribution of individuals across microhabitats. 3. Beetles from different feeding guilds were not randomly distributed, but congregated on those microhabitats that are likely to provide the highest concentrations of their preferred food sources. Herbivorous beetles, in particular, were over-represented on flowers and flush foliage and underrepresented on mature leaves and dead wood. Proportional numbers of species within each feeding guild were remarkably uniform across tree species for each microhabitat, but proportional abundances of feeding guilds were all significantly non-uniformly distributed between host tree species, regardless of microhabitat, confirming patterns previously found for arthropods in trees in temperate and tropical forests. 4. These results show that the canopy beetle community is partitioned into discrete assemblages between microhabitats and that this partitioning arises because of differences in feeding guild structure as a function of the diversity and the temporal and spatial availability of resources found on each microhabitat.

Elevational gradients affect the production of plant secondary metabolites through changes in both biotic and abiotic conditions. Previous studies have suggested both elevational increases and decreases in host-plant chemical defences. We analysed the correlation of alkaloids and polyphenols with elevation in a community of nine Ficus species along a continuously forested elevational gradient in Papua New Guinea. We sampled 204 insect species feeding on the leaves of these hosts and correlated their community structure to the focal compounds. Additionally, we explored species richness of folivorous mammals along the gradient. When we accounted for Ficus species identity, we found a general increase in flavonoids and alkaloids. Elevational trends in non-flavonol polyphenols were less pronounced or showed non-linear correlations with elevation. The abundance of insect herbivores decreased with elevation, while the species richness of folivorous mammals showed an elevational increase. Insect community structure was affected mainly by alkaloid concentration and diversity. Although our results show an elevational increase in several groups of metabolites, the drivers behind these trends likely differ. Flavonoids probably provide figs with protection against abiotic stressors, such as UV-irradiation. In contrast, alkaloids affect insect herbivores and may provide protection against mammalian herbivores and pathogens. Concurrent analysis of multiple compound groups alongside ecological data is an important approach for understanding the selective landscape that shapes plant defences.

Abstract.  1. The degree of infestation by New Zealand sooty beech scale insects (Ultracoelostoma assimile, Homoptera: Margarodidae) varies dramatically among adjacent southern beech trees (Nothofagus spp., Fagaceae), but has previously been assumed to be uniformly or randomly distributed within individual host trees. In this study, a full‐census survey was conducted from ground level to canopy level on 14 naturally occurring, canopy‐dominant red beech (Nothofagus fusca) trees (size range 38.7–107.6 cm diameter at breast height) to determine the degree of within‐tree heterogeneity in herbivore density.2. The within‐tree distribution of the sooty beech scale was vertically stratified and highly heterogeneous, with the greatest densities occurring on bark surfaces in the canopy rather than on the trunk, and on the lower rather than upper sides of the branches. The spatial distribution was strongly negatively correlated with trunk and branch diameter, and increasing bark thickness (as a function of diameter) provides a plausible explanation for differences in the establishment and population density of sooty beech scale insects with trunk and branch size. Furthermore, there was a significant change in the spatial distribution of scale insect populations on trunks and branches of trees of increasing diameter at breast height. This indicates a strong temporal component to the spatial dynamics of the sooty beech scale insect driven by changing host phenology. Future studies on phytophagous insects infesting large host trees need to consider more explicitly changes in population dynamics through space and time.3. Because of the high degree of within‐tree heterogeneity in population density, the total population size of scale insects on an individual tree could not be predicted from any measure of population density low on the trunk. However, the dry weight biomass of sooty mould fungi growing on the ground beneath infested trees was a remarkably accurate predictor of the total population size of scale insects. The use of sooty mould fungi as a relative measure of population size could be incorporated into studies of other honeydew‐producing hemipterans, since the growth of sooty mould is a distinctive feature synonymous with high concentrations of honeydew production worldwide.

Altered abiotic conditions resulting from human-induced climate change are already driving changes in the spatial and temporal distributions of many organisms. For insects, how species are distributed across elevations is relatively well known, but data on their seasonality at different elevations are lacking. Here we show seasonal variation in beetle abundance and species richness along two spatially-distinct elevational transects (350–1000 m and 100–1000 m asl) in the rainforests of northern Australia. Temperature was the best predictor of temporal abundance and species richness patterns, while rainfall had little influence. Elevation had little effect on seasonal changes in abundance or diversity. Adults of most beetle species exhibited long season-lengths (>6 months of the year) with distinct peaks in abundance during the summer wet-season. We found evidence of phenotypic variation among the more widespread species, with seasonal peaks in abundance often not coinciding across elevations or transects. Due to the wide elevational range of most species, and the lack of consistency in the seasonality of wide-spread individual species, we suggest that many beetles inhabiting the low to mid-elevation mountains in the Wet Tropics, and potentially other tropical rainforests, are not as vulnerable to extinction due to climate change as many other organisms.

Accurate estimates of invertebrate biomass are essential for quantifying community structure, food web dynamics and energy flow in terrestrial ecosystems. In this paper, length‐mass and length × width‐mass regressions were carried out for 18 invertebrate taxonomic groups collected from the canopy of an Australian tropical rainforest. In an additional analysis, invertebrates were divided among seven body shape categories based on the ratio of body length to body width (from short and squat to long and thin) in an attempt to develop accurate equations for estimating biomass that can be applied to any taxonomic group in any locality. For most groups, the inclusion of body width to the predictor variable improved the model, confirming that body shape is an important factor in the accuracy of biomass estimations. The most accurate method for estimating invertebrate biomass was the use of taxon‐specific equations, followed by equations based on body shape. Single whole‐fauna equations were very inaccurate for estimating biomass, especially for insects that are either very squat or very long and thin. In accordance with previous studies, it was concluded that the most accurate method for estimating invertebrate biomass from proxy body measurements is the use of taxon‐specific regression equations, especially those that incorporate body width in the model. However, equations based on body shape categories may be useful for estimating the biomass of groups for which no length‐mass relationship has been determined, while single, whole‐fauna equations should be avoided.