Guild-specific patterns of species richness and host specialization in plant-herbivore food webs from a tropical forest.

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.

Low beta diversity of herbivorous insects in tropical forests.

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.

Multiple paternity in a natural population of a wild tobacco fly, Bactrocera cacuminata (Diptera: Tephritidae), assessed by microsatellite DNA markers.

Mating frequency has important implications for patterns of sexual selection and sexual conflict and hence for issues such as speciation and the maintenance of genetic diversity. Knowledge of natural mating patterns can also lead to more effective control of pest tephritid species, in which suppression programmes, such as the sterile insect technique (SIT) are employed. Multiple mating by females may compromise the success of SIT. We investigated the level of polyandry and sperm utilization in a Brisbane field population of the tropical fruit fly, Bactrocera cacuminata (Hering), using seven polymorphic microsatellite loci. The offspring of 22 wild-caught gravid females were genotyped to determine the number of males siring each brood and paternity skew, using the programs gerud and scare. Our data showed that 22.7% of females produced offspring sired by at least two males. The mean number of mates per female was 1.72. Paternal contributions of double-sired broods were skewed with the most successful male having sired between 76.9% and 87.5% of the offspring. These results have implications for SIT, because the level of remating we have identified would indicate that wild females could mate with one or more resident fertile males.

Attractiveness of various combinations of colors and shapes to females and males of Bactrocera minax (Diptera: Tephritidae) in a commercial mandarin grove in Bhutan.

Bactrocera minax (Enderlein) (Diptera: Tephritidae) is a major pest of citrus fruit in the region from Nepal through to southwestern China. In tests on wild adult populations of B. minax in a mandarin, Citrus reticulata Blanco, orchard in western Bhutan, both males and females were more attracted to 50-mm-diameter spheres than to 50-mm discs of the same color. Furthermore, they were more attracted to spheres colored with orange or green-yellow mixtures than to similar spheres colored red, yellow, green, blue, black, or white. The UV reflectance from the orange (600-650 nm) and yellow-green mixtures (530 nm) was similar to that proposed byProkopy (1977) as eliciting strong attractant responses in other species of Tephritidae.

Effect of physiological and experiential state ofBactrocera tryoni flies on intra-tree foraging behavior for food (bacteria) and host fruit.

Using caged host trees on which we manipulated food and oviposition sites, we investigated the foraging behavior of individually-releasedBactrocera tryoni (Diptera: Tephritidae) females in relation to state of fly hunger for protein, presence or absence of bacteria as a source of protein, degree of prior experience with host fruit, and quality of host fruit for oviposition. One aim was to evaluate whether it is immature or matureB. tryoni females that are responsible for initially inoculating host fruit surfaces with "fruit-fly-type" bacteria, the odor of which is known to attractB. tryoni females. We found that 3-week-old immature females provided with sucrose but deprived of protein from eclosion had a much greater propensity than 3-week-old protein-fed mature females to visit vials containing fruit-fly-type bacteria, irrespective of whether vials were associated with adjacent host fruit or not. In the absence of associated bacteria in vials, immature females had a much lower propensity than mature females to visit host fruit. In the presence of bacteria in vials, however, propensity of immature and mature females to visit fruit was about equal. Mature (but not immature) females were more inclined to visit fruit that ranked higher for oviposition (nectarines) than fruit that ranked lower (sweet oranges). Mature females that attempted oviposition during a single 3-min exposure period to a nectarine prior to release were much more likely to find a nectarine than were mature females naive to fruit or immature females with or without prior contact with fruit. Exposure to a nectarine before release did not affect the propensity of either mature or immature females to alight on an odorless visual model of a nectarine, however. As judged by numbers of leaves visited, protein-deprived immature females were more active than protein-fed mature females, irrespective of the sorts of resources on a tree. Together, our findings lead us to conclude that (1) the firstB. tryoni females to arrive on the fruit of a host tree and therefore inoculate the fruit with fruit-fly-type bacteria are unlikely to be sexually immature, but to be mature as a result of having earlier acquired protein elsewhere, (2) the odor of colonies of fruit-fly-type bacteria when associated with host fruit will attract protein-hungry but not protein-fed females, and (3) the odor of the fruit itself will attract mature females (especially experienced ones) but not immature females. These findings illustrate the value of considering jointly the state of a resource patch together with the physiological and experiential state of the individual when investigating the foraging behavior of an insect.