Synonymization of key pest species within the Bactrocera dorsalis species complex (Diptera: Tephritidae): Taxonomic changes based on a review of 20 years of integrative morphological, molecular, cytogenetic, behavioural and chemoecological data

Bactrocera papayae Drew & Hancock, Bactrocera philippinensis Drew & Hancock, Bactrocera carambolae Drew & Hancock, and Bactrocera invadens Drew, Tsuruta & White are four horticultural pest tephritid fruit fly species that are highly similar, morphologically and genetically, to the destructive pest, the Oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae). This similarity has rendered the discovery of reliable diagnostic characters problematic, which, in view of the economic importance of these taxa and the international trade implications, has resulted in ongoing difficulties for many areas of plant protection and food security. Consequently, a major international collaborative and integrated multidisciplinary research effort was initiated in 2009 to build upon existing literature with the specific aim of resolving biological species limits among B. papayae, B. philippinensis, B. carambolae, B. invadens and B. dorsalis to overcome constraints to pest management and international trade. Bactrocera philippinensis has recently been synonymized with B. papayae as a result of this initiative and this review corroborates that finding; however, the other names remain in use. While consistent characters have been found to reliably distinguish B. carambolae from B. dorsalis, B. invadens and B. papayae, no such characters have been found to differentiate the latter three putative species. We conclude that B. carambolae is a valid species and that the remaining taxa, B. dorsalis, B. invadens and B. papayae, represent the same species. Thus, we consider B. dorsalis (Hendel) as the senior synonym of B. papayae Drew and Hancock syn.n. and B. invadens Drew, Tsuruta & White syn.n. A redescription of B. dorsalis is provided. Given the agricultural importance of B. dorsalis, this taxonomic decision will have significant global plant biosecurity implications, affecting pest management, quarantine, international trade, postharvest treatment and basic research. Throughout the paper, we emphasize the value of independent and multidisciplinary tools in delimiting species, particularly in complicated cases involving morphologically cryptic taxa.

Paropsine beetles (Coleoptera: Chrysomelidae) in south-eastern Queensland hardwood plantations: identifying potential pest species

Paropsine chrysomelid beetles are significant defoliators of Australian eucalypts. In Queensland, the relatively recent expansion of hardwood plantations has resulted in the emergence of new pest species. Here I identify paropsine beetles collected from Eucalyptus cloeziana Muell. and E. dunnii Maiden, two of the major Eucalyptus species grown in plantations in south-eastern Queensland, and estimate the relative abundance of each paropsine species. Although I was unable to identify all taxa to species level, at least 17 paropsine species were collected, about one-third of which have not been previously associated with hardwood plantations. Paropsis atomaria Olivier and P. charybdis Stål were the most abundant species collected in E. cloeziana plantations, while Chrysophtharta cloelia (Stål) and P. atomaria were most commonly collected from E. dunnii. Occasional collections from Corymbia citriodora (Hook.) Hill and Johns, ssp. variegata revealed an additional four species implicated in plantation damage. Abundance and voltinism varied between species and sites. I predict which paropsine species are likely to threaten plantation productivity.

Responses of fruit flies (Tephritidae: Dacinae) to novel male attractants in north Queensland, Australia, and improved lures for some pest species

Male fruit fly attractants, cue-lure and methyl eugenol (ME), have been successfully used for the last 50 years in the monitoring and control of Dacini fruit flies (Bactrocera and Dacus species). However, over 50% of Dacini are non-responsive to either lure, including some pest species. A new lure, zingerone, has been found to weakly attract cue- and ME-responsive species in Malaysia. In Australia it attracted a weakly cue-responsive minor pest Bactrocera jarvisi (Tryon) and three non-responsive' species. Similar compounds were tested in Queensland and attracted cue- and ME-responsive species and two non-responsive' species. In this study, 14 novel compounds, including raspberry ketone formate (RKF) (Melolure) and zingerone, were field tested in comparison with cue-lure and ME at 17 sites in north Queensland. The most attractive novel lures were isoeugenol, methyl-isoeugenol, dihydroeugenol and zingerone. Several non-responsive' species responded to the new lures: Bactrocera halfordiae (Tryon), a species of some market access concern, was most attracted to isoeugenol; B.barringtoniae (Tryon), B.bidentata (May) and B.murrayi (Perkins) responded to isoeugenol, methyl-isoeugenol and dihydroeugenol; two new species of Dacus responded to zingerone. Bactrocera kraussi (Hardy), a cue-responsive minor pest in north Queensland, was significantly more attracted to isoeugenol than cue-lure. The cue-responsive D.absonifacies (May) and D.secamoneaeDrew were significantly more attracted to zingerone than cue-lure. Bactrocera yorkensisDrew & Hancock, a ME-responsive species was significantly more attracted to isoeugenol, methyl-isoeugenol and dihydroeugenol than ME. The preferential response to RKF or cue-lure was species specific. Six species were significantly more attracted to RKF, including the pests B.tryoni (Froggatt), B.frauenfeldi (Schiner) and minor pest B.bryoniae (Tryon); eight species were significantly more attracted to cue-lure including the pest B.neohumeralis (Hardy). These findings have significance in the search for optimal male lures for pest species elsewhere in the world.

The taxonomic status of pest species of Coptotermes in Southeast Asia: Resolving the paradox in the pest status of the termites, Coptotermes gestroi, C-havilandi and C-travians (Isoptera : Rhinotermitidae)

The taxonomic status of Coptotermes gestroi (Wasmann), C. havilandi Holmgren, C. travians (Haviland) and C. borneensis Oshima (Isoptera: Rhinotermitidae) is revised. The apparent discrepancy between the reported importance of C. havitandi in countries to which it has been introduced and the region from which it originated is shown to be due to misidentification and taxonomic confusion between C. travians, C. havilandi and C. gestroi. Based on an examination of specimens from Southeast Asia, two species are recognized, namely C. gestroi and C. travians. Coptotermes havilandi, described from imagos, is shown to be the same species as C. gestro described earlier from the soldier caste, and is designated a junior synonym. Coptotermes gestroi occurs from Assam through Burma and Thailand to Malaysia and the Indonesian archipelago, and has been introduced into other geographic regions, including parts of North and South America and the Caribbean. It is frequently found damaging wood in buildings, and is often intercepted outside its range in cargo onboard ships and sailing vessels, which is a likely mechanism for its spread to new geographical areas. Coptotermes gestroi has been misidentified in much literature as C. travians. Conversely, C. travians has been misidentified in recent literature in Peninsular Malaysia as C. havilandi and was redescribed from Borneo as C. borneensis, which is here designated a junior synonym of C. travians. It has a known distribution from Peninsular Malaysia to Borneo, and has not been found infesting wood in buildings. It is envisaged that the resolution of this taxonomic problem will enable the deployment of common pest management strategies for C. gestro the primary pest species of Coptotermes originating from Southeast Asia.

Managing maize under pest species competition: is Bt (Bacillus thuringiensis) maize the solution?

Transgenic crops that contain Cry genes from Bacillus thuringiensis (Bt) have been adopted by farmers over the last 17 years. Unlike traditional broad spectrum chemical insecticides, Bt's toxicity spectrum is relatively narrow and selective, which may indirectly benefit secondary insects that may become important pests. The economic damage caused by the rise of secondary pests could offset some or all of the benefits associated with the use of Bt varieties. We develop a bioeconomic model to analyze the interactions between primary and secondary insect populations and the impact of different management options on insecticide use and economic impact over time. Results indicate that some of the benefits associated with the adoption of genetically engineered insect resistant crops may be eroded when taking into account ecological dynamics. It is suggested that secondary pests could easily become key insect pests requiring additional measures - such as insecticide applications or stacked traits – to keep their populations under the economic threshold.

Multi-gene phylogenetic analysis of south-east Asian pest members of the Bactrocera dorsalis species complex (Diptera: Tephritidae) does not support current taxonomy

Bactrocera dorsalis sensu stricto, B. papayae, B. philippinensis and B. carambolae are serious pest fruit fly species of the B. dorsalis complex that predominantly occur in south-east Asia and the Pacific. Identifying molecular diagnostics has proven problematic for these four taxa, a situation that cofounds biosecurity and quarantine efforts and which may be the result of at least some of these taxa representing the same biological species. We therefore conducted a phylogenetic study of these four species (and closely related outgroup taxa) based on the individuals collected from a wide geographic range; sequencing six loci (cox1, nad4-3′, CAD, period, ITS1, ITS2) for approximately 20 individuals from each of 16 sample sites. Data were analysed within maximum likelihood and Bayesian phylogenetic frameworks for individual loci and concatenated data sets for which we applied multiple monophyly and species delimitation tests. Species monophyly was measured by clade support, posterior probability or bootstrap resampling for Bayesian and likelihood analyses respectively, Rosenberg's reciprocal monophyly measure, P(AB), Rodrigo's (P(RD)) and the genealogical sorting index, gsi. We specifically tested whether there was phylogenetic support for the four 'ingroup' pest species using a data set of multiple individuals sampled from a number of populations. Based on our combined data set, Bactrocera carambolae emerges as a distinct monophyletic clade, whereas B. dorsalis s.s., B. papayae and B. philippinensis are unresolved. These data add to the growing body of evidence that B. dorsalis s.s., B. papayae and B. philippinensis are the same biological species, which poses consequences for quarantine, trade and pest management.

Learning in Helicoverpa armigera (Lepidoptera: Noctuidae): a new look at the behaviour and control of a polyphagous pest

Recent experimental evidence has shown that learning occurs in the host selection behaviour of Helicoverpa armigera (Hübner), one of the world‘s most important agricultural pests. This paper discusses how the occurrence of learning changes our understanding of the host selection behaviour of this polyphagous moth. Host preferences determined from previous laboratory studies may be vastly different from preferences exhibited by moths in the field, where the abundance of particular hosts may be more likely to determine host preference. In support of this prediction, a number of field studies have shown that the ‘attractiveness’ of different hosts for H. armigera oviposition may depend on the relative abundance of these host species. Insect learning may play a fundamental role in the design and application of present and future integrated pest management strategies such as the use of host volatiles, trap crops and resistant crop varieties for monitoring and controlling this important pest species

Resolution of Cryptic Species Complexes of Tephritid Pests to Enhance SIT Application and Facilitate International Trade

Tephritid fruit flies (Diptera: Tephritidae) are considered by far the most important group of horticultural pests worldwide. Female fruit flies lay eggs directly into ripening fruit, where the maggots feed causing fruit loss. Each and every continent is plagued by a number of fruit fly pests, both indigenous as well as invasive ones, causing tremendous economic losses. In addition to the direct losses through damage, they can negatively impact commodity trade through restrictions to market access. The quarantine and regulatory controls put in place to manage them are expensive, while the on-farm control costs and loss of crop affect the general well-being of growers. These constraints can have huge implications on loss in revenues and limitations to developing fruit and vegetable-based agroindustries in developing, emergent and developed nations. Because fruit flies are a global problem, the study of their biology and management requires significant international attention to overcome the hurdles they pose. The Joint Food and Agriculture Organisation / International Atomic Energy Agency (FAO/IAEA) Programme on Nuclear Techniques in Food and Agriculture has been on the foreground in assisting Member States in developing and validating environment-friendly fruit fly suppression systems to support viable fresh fruit and vegetable production and export industries. Such international attention has resulted in the successful development and validation of a Sterile Insect Technique (SIT) package for the Mediterranean fruit fly. Although demands for R&D support with respect to Mediterranean fruit fly are diminishing due to successful integration of this package into sustainable control programmes against this pest in many countries, there were increasing demands from Member States in Africa, Asia and Latin America, to address other major fruit fly pests and a related, but sometimes neglected issue of tephritid species complexes of economic importance. Any research, whether it is basic or applied, requires a taxonomic framework that provides reliable and universally recognized entities and names. Among the currently recognized major fruit fly pests, there are groups of species whose morphology is very similar or identical, but biologically they are distinct species. As such, some insect populations that are grouped taxonomically within the same pest species, display different biological and genetic traits and show reproductive isolation which suggest that they are different species. On the other hand, different species may have been taxonomically described, but there may be doubt as to whether they actually represent distinct biological species or merely geographical variants of the same species. This uncertain taxonomic status has practical implications on the effective development and use of the SIT against such complexes, particularly at the time of determining which species to mass-rear, and significantly affects international movement of fruit and vegetables through the establishment of trade barriers to important agricultural commodities which are hosts to these pest tephritid species...

Using Trichogramma Westwood (Hymenoptera: Trichogrammatidae) for insect pest biological control in cotton crops: an Australian perspective.

Trichogramma Westwood egg parasitoids alone generally fail to suppress heliothine pests when released in established cotton-growing regions. Factors hindering their success include indiscriminate use of detrimental insecticides, compensation for minimal pest larval hatch due to their activity via reduced larval cannibalism or mortality in general, singly laid heliothine eggs avoiding detection and asynchronous development benefiting host over parasitoid. Yet, despite these limitations, relatively large Trichogramma pretiosum Riley populations pervade and effectively suppress Helicoverpa (Hardwick) pests in Australian Bt (Bacillus thuringiensis Berliner)-transgenic cotton, Gossypium hirsutum L., crops, especially in the Ord River Irrigation Area (ORIA) of tropical northern Australia, where their impact on the potentially resistant pest species, Helicoverpa armigera (Hubner), is considered integral to the local insecticide resistance management (IRM) strategy for continued, sustainable Bt-transgenic cotton production. When devoid of conventional insecticides, relatively warm and stable conditions of the early dry season in winter grown ORIA Bt-transgenic cotton crops are conducive to Trichogramma proliferation and biological control appears effective. Further, there is considerable scope to improve Trichogramma's biological control potential, in both the ORIA and established cotton-growing regions, via habitat manipulation. It is proposed that Trichogramma may prove equally effective in developing agricultural regions of monsoonal northern Australia, and that environmental constraints on Trichogramma survival, and those of other natural enemies, require due consideration prior to their successful application in biological control programs.

Early-season movement dynamics of phytophagous pest and natural enemies across a native vegetation-crop ecotone

There is limited understanding about how insect movement patterns are influenced by landscape features, and how landscapes can be managed to suppress pest phytophage populations in crops. Theory suggests that the relative timing of pest and natural enemy arrival in crops may influence pest suppression. However, there is a lack of data to substantiate this claim. We investigate the movement patterns of insects from native vegetation (NV) and discuss the implications of these patterns for pest control services. Using bi-directional interception traps we quantified the number of insects crossing an NV/crop ecotone relative to a control crop/crop interface in two agricultural regions early in the growing season. We used these data to infer patterns of movement and net flux. At the community-level, insect movement patterns were influenced by ecotone in two out of three years by region combinations. At the functional-group level, pests and parasitoids showed similar movement patterns from NV very soon after crop emergence. However, movement across the control interface increased towards the end of the early-season sampling period. Predators consistently moved more often from NV into crops than vice versa, even after crop emergence. Not all species showed a significant response to ecotone, however when a response was detected, these species showed similar patterns between the two regions. Our results highlight the importance of NV for the recruitment of natural enemies for early season crop immigration that may be potentially important for pest suppression. However, NV was also associated with crop immigration by some pest species. Hence, NV offers both opportunities and risks for pest management. The development of targeted NV management may reduce the risk of crop immigration by pests, but not of natural enemies.

Biodiversity, biosecurity and integrated pest management

Breaches of biosecurity, leading to incursions by invasive species, have the potential to cause substantial economic, social and environmental losses, including drastic reduction in biodiversity. It is argued that improving biosecurity reduces risk to biodiversity, while maintaining stable ecosystems through biodiversity can be a safeguard against biosecurity breaches. The global costs of invasive alien species (IAS) have been estimated at around US$350 billion, while alien invertebrate and vertebrate pests and weeds are estimated to cost Australia at least $7 billion a year. A striking, current, example is the incursion by Myrtle Rust (Puccinia psidii) an organism which can infect all members of the Myrtaceae, the most important family in the Australian flora. Myrtle rust was first detected on a property on the central coast of New South Wales in late April 2010. Two years later the disease has been detected in numerous locations in Queensland and New South Wales ranging from commercial plant nurseries and public amenities to large areas of bushland. This particular breach of biosecurity will, inevitably, diminish biodiversity of flora and fauna over large areas of the continent. Integrated pest management (IPM), an enrichment of diversity in managing invasive and other pest species, offers the best opportunity to address problems such as these. Australia's response to increasing biosecurity risk is comprehensive and includes national networking of scientists engaged in a complex program of biosecurity research and development, including studies of IPM. This network is being enhanced by the development of international linkages.

Plant composition modulates arthropod pest and predator abundance: Evidence for culling exotics and planting natives

We investigate the role of plant species in crops, pasture and native vegetation remnants in supporting agronomic pests and their predators. The study was conducted in three Australian States and across 290 sites sampled monthly for two years. Pastures played a key role in harbouring pest species consistent across States, while native vegetation hosted relatively more predators than other habitat types within each State. Furthermore, native plant species supported the lowest pest density and more predators than pests; in contrast, 75 of the exotic weed species surveyed hosted more pests than predators. Despite the role of pasture in harbouring pests, we found in NSW that pasture also supported the highest proportion of juvenile predators, while native vegetation remnants had the lowest. Our results indicate that non-crop habitat (native remnants or pasture) with few exotic weeds supports high predator and low pest arthropod densities, and that weeds are associated with high pest densities. By linking broad response variables such as ‘all pests’ with specific predictors such as ‘plant species’, our study will inform on-farm management actions of which weeds to control and which natives to plant or regenerate. This study shows the importance of knowing the function of habitats and plants species in supporting pests and predators in agricultural landscapes across multiple regions.

Functional benefits of predator species diversity depend on prey identity

1. Determining the functional significance of species diversity in natural enemy assemblages is a key step towards prediction of the likely impact of biodiversity loss on natural pest control processes. While the biological control literature contains examples in which increased natural enemy diversity hinders pest control, other studies have highlighted mechanisms where pest suppression is promoted by increased enemy diversity. 2. This study aimed to test whether increased predator species diversity results in higher rates of predation on two key, but contrasting, insect pest species commonly found in the rice ecosystems of south-east Asia. 3. Glasshouse experiments were undertaken in which four life stages of a planthopper (Nilaparvata lugens) and a moth (Marasmia patnalis) were caged with single or three-species combinations of generalist predators. 4. Generally, predation rates of the three-species assemblages exceeded expectation when attacking M. patnalis, but not when attacking N. lugens. In addition, a positive effect of increased predator species richness on overall predation rate was found with M. patnalis but not with N. lugens. 5. The results are consistent with theoretical predictions that morphological and behavioural differentiation among prey life stages promotes functional complementarity among predator species. This indicates that emergent species diversity effects in natural enemy assemblages are context dependent; they depend not only on the characteristics of the predators species, but on the identity of the species on which they prey.

The rodent species of the Ifugao Rice Terraces, Philippines - Target or non-target species for management?

We investigated the species diversity and habitat use of rodents in the Ifugao Rice Terraces (IRT), Luzon, Philippines, as a first step in their assessment either as pest species of rice or as potential non-target species of rodent control practice. Trapping was carried out in caneland and forest habitats adjacent to rice cropland using trap lines of 10 - 15 cage-traps. Four trapping rounds, each consisting of 5 nights trapping, were replicated at two sites during the months of May and June. A diverse rodent fauna was recorded, including the non-native pest species, Rattus tanezumi, and the native species, Rattus everetti and Chrotomys mindorensis. Results from trapping and spool-and-line tracking suggested that these native species do not contribute to rice damage and that several may actually be beneficial in the ricefield ecosystem as vermivores that feed on invertebrate pests. Control should therefore be directed at the pest species, R. tanezumi, minimising non-target effects on the non-pest rodent species.