Richard Arthur Ian Drew

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.

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.

Influence of host plant structure and microclimate on the abundance and behavior of a tephritid fly

Microclimate and host plant architecture significantly influence the abundance and behavior of insects. However, most research in this field has focused at the invertebrate assemblage level, with few studies at the single-species level. Using wild Solanum mauritianum plants, we evaluated the influence of plant structure (number of leaves and branches and height of plant) and microclimate (temperature, relative humidity, and light intensity) on the abundance and behavior of a single insect species, the monophagous tephritid fly Bactrocera cacuminata (Hering). Abundance and oviposition behavior were signficantly influenced by the host structure (density of foliage) and associated microclimate. Resting behavior of both sexes was influenced positively by foliage density, while temperature positively influenced the numbers of resting females. The number of ovipositing females was positively influenced by temperature and negatively by relative humidity. Feeding behavior was rare on the host plant, as was mating. The relatively low explanatory power of the measured variables suggests that, in addition to host plant architecture and associated microclimate, other cues (e.g., olfactory or visual) could affect visitation and use of the larval host plant by adult fruit flies. For 12 plants observed at dusk (the time of fly mating), mating pairs were observed on only one tree. Principal component analyses of the plant and microclimate factors associated with these plants revealed that the plant on which mating was observed had specific characteristics (intermediate light intensity, greater height, and greater quantity of fruit) that may have influenced its selection as a mating site.

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

SummaryUsing 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.

Alimentary tract bacteria isolated and identified with API-20E and molecular cloning techniques from Australian tropical fruit flies, Bactrocera cacuminata and B. Tryoni

Abstract Bacteria were isolated from the crop and midgut of field collected Bactrocera cacuminata (Hering) and Bactrocera tryoni (Froggatt) (Diptera: Tephritidae). Two methods were used, firstly isolation onto two types of bacteriological culture media (PYEA and TSA) and identification using the API-20E diagnostic kit, and secondly, analysis of samples using the 16S rRNA gene molecular diagnostic method. Using the API-20E method, 10 genera and 17 species of bacteria in the family Enterobacteriaceae were identified from cultures growing on the nutrient agar. The dominant species in both the crop and midgut were Citrobacter freundii, Enterobacter cloacae and Klebsiella oxytoca. Providencia rettgeri, Klebsiella pneumoniae ssp ozaenae and Serratia marcescens were isolated from B. tryoni only. Using the molecular cloning technique that is based on 16S rRNA gene sequences, five bacteria classes were dignosed — Alpha-, Beta-, Gamma- and Delta- Proteobacteria and Firmicutes — including five families, Leuconostocaceae, Enterococcaceae, Acetobacteriaceae, Comamonadaceae and Enterobacteriaceae. The bacteria affiliated with Firmicutes were found mainly in the crop while the Gammaproteobacteria, especially the family Enterobacteriaceae, was dominant in the midgut. This paper presents results from the first known application of molecular cloning techniques to study bacteria within tephritid species and the first record of Firmicutes bacteria in these flies.

Attraction of fruit flies of the genus Bactrocera to colored mimics of host fruit

In tests on feral populations of polyphagous Bactrocera tryoni (Froggatt) adults on host guava trees, both sexes were significantly more attracted to Tangletrap‐coated 50 mm diameter spheres colored blue or white than to similar spheres colored red, orange, yellow, green, or black or to Tangletrap‐coated 50 mm diameter yellow‐green guava fruit. In contrast, in tests on feral populations of oligophagous Bactrocera cacuminata (Hering) on host wild tobacco plants, both sexes were significantly more attracted to Tangletrap‐coated 15 mm diameter spheres colored orange or yellow than to other colors of spheres or to Tangletrap‐coated 15 mm diameter green wild tobacco fruit. Both sexes of both tephritid species were significantly more attracted to blue (in the case of B. tryoni) or orange (in the case of B. cacuminata) 50 mm spheres displayed singly than to blue or orange 15 mm spheres displayed in clusters, even though fruit of wild tobacco plants are borne in clusters. Finally, B. tryoni adults were significantly less attracted to non‐ultraviolet reflecting bluish fruit‐mimicking spheres than to bluish fruit‐mimicking spheres having a slightly enhanced level of ultraviolet reflectance, similar to the reflectance of possible native host fruit of B. tryoni, whose bluish skin color is overlayed with ultraviolet‐reflecting waxy bloom. Responses to fruit visual stimuli found here are discussed relative to responses found in other tephritid species.

Host specialization and species richness of fruit flies (Diptera: Tephritidae) in a New Guinea rain forest

Frugivorous dacine fruit flies were studied in a lowland tropical rain forest in Papua New Guinea to determine their host specificity, abundance, and the number of species attacking various plant species. Plant species hosted 0-3 fruit fly species at median (1-3 quartile) densities of 1 (0-17) fruit flies per 100 fruits. Fruit flies were mostly specialized to a single plant family (83% species) and within each family to a single genus (88% species), while most of the species (66%) were able to feed on >1 congeneric plant species. Only 30 from the 53 studied plant species were colonized by fruit flies. The plant-fruit fly food web, including these 30 plant species and the total of 29 fruit fly species feeding on them, was divided into 14 compartments, each including 1-8 plant species hosting mutually disjunct assemblages of fruit flies. This structure minimizes indirect interactions among plant species via shared herbivores. The local species pool was estimated at 152 ± 32 (± SE) fruit fly species. Forty per cent of all taxonomically described species known from Papua New Guinea were reared or trapped in our study area. Such a high proportion indicates low beta-diversity of fruit flies. Steiner traps were highly efficient in sampling the lure-responsive fruit fly species as they re-collected 84% of all species trapped in the same area 5 y before. Fruit fly monitoring by these traps is a cheap, simple and efficient method for the study of spatial and temporal changes in rain-forest communities.

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.

Seasonal phenology of Bactrocera minax (Diptera: Tephritidae) in western Bhutan

The Chinese citrus fruit fly, Bactrocera (Tetradacus) minax (Enderlein), is one of the major citrus pests in Bhutan and can cause >50% mandarin (Citrus reticulata Blanco) fruit drop. As part of the development of a management strategy for the fly in mandarin orchards, population monitoring and experimental manipulations were carried out to determine: (i) adult emergence period; (ii) adult phenology patterns; (iii) period of crop susceptibility; and (iv) period from fruit drop to pupation. In western Bhutan, adult flies emerge from the overwintering pupal stage in late April/early May. Most flies are mature by the end of May and it is inferred that mating occurs at this time: from the beginning of June males rapidly disappear from the population and by mid- to late June are rare or absent from traps. Mature females are present in the mandarin crop at the beginning of June, but very little oviposition occurs until mid-June, while most damage has occurred by mid-July. Initiation of oviposition into mandarins is almost certainly linked to crop phenology. Adult flies disappear from the orchard system during August. After fruit drop, larvae were recorded leaving the fruit to pupate within 13 days. The use of early to mid-season protein bait sprays and/or targeted use of systemic insecticides during the one month oviposition period, plus the removal of fallen fruit once every 10 days, are recommended as control strategies.

Facilitation in Ovipositional Behavior of Bactrocera tryoni Flies

Mature female Queensland fruit flies, Bactrocera tryoni Froggatt, released into canopies of field-caged trees containing host fruit, did not discriminate between fruit having or lacking two resident conspecific females when selecting fruit on which to alight. When released individually onto a fruit, mature females showed no greater or lesser propensity to initiate boring with the ovipositor into fruit having or lacking a resident female engaged in pre-ovipositional or post-ovipositional behavior. However, boring propensity of introduced females was significantly enhanced or facilitated when released onto fruit on which a conspecific female was in the act of ovipositing. Introduced females were significantly more prone to initiate boring into high-ranking fruit (nectarines) than low-ranking fruit (lemons) and to do so when having a high rather than a low egg load. Neither fruit type nor egg load of introduced females had a significant effect on the occurrence of encounters between introduced and resident females, on the occurrence of contests, or on the propensity of introduced females to initiate boring in the presence of a resident female. Among introduced females, there was a consistent trend to initiate boring earlier in the presence than in the absence of a resident female. We discuss circumstances, likely to be narrow, under which boring propensity of an intruding female might be facilitated in the presence of a resident female.