RESUMEN
Tephritid fruit flies are internationally significant pests of horticulture. Because they are also highly invasive and of major quarantine concern, significant effort is placed in developing full or partial pest risk assessments (PRAs) for fruit flies, while large investments can be made for their control. Competition between fruit fly species, driven by the need to access and utilise fruit for larval development, has long been recognised by researchers as a fundamental component of fruit fly biology, but is entirely absent from the fruit fly PRA literature and appears not be considered in major initiative planning. First presenting a summary of the research data which documents fruit fly competition, this paper then identifies four major effects of fruit fly competition that could impact a PRA or large-scale initiative: (i) numerical reduction of an existing fruit fly pest species following competitive displacement by an invasive fruit fly; (ii) displacement of a less competitive fruit fly pest species in space, time or host; (iii) ecological resistance to fruit fly invasion in regions already with competitively dominant fruit fly species; and (iv) lesser-pest fruit fly resurgence following control of a competitively superior species. From these four major topics, six more detailed issues are identified, with each of these illustrated by hypothetical, but realistic biosecurity scenarios from Australia/New Zealand and Europe. The scenarios identify that the effects of fruit fly competition might both positively or negatively affect the predicted impacts of an invasive fruit fly or targeted fruit fly control initiative. Competition as a modifier of fruit fly risk needs to be recognised by policy makers and incorporated into fruit fly PRAs and major investment initiatives.
RESUMEN
The genus Bactrocera (Diptera: Tephritidae) is endemic to the monsoonal rainforests of South-east Asia and the western Pacific where the larvae breed in ripe, fleshy fruits. While most Bactrocera remain rainforest restricted, species such as Bactrocera dorsalis, Bactrocera zonata and Bactrocera tryoni are internationally significant pests of horticulture, being both highly invasive and highly polyphagous. Almost universally in the literature it is assumed that Bactrocera breed continuously if temperature and hosts are not limiting. However, despite that, these flies show distinct seasonality. If discussed, seasonality is generally attributed to the fruiting of a particular breeding host (almost invariably mango or guava), but the question appears not to have been asked why flies do not breed at other times of the year despite other hosts being available. Focusing initially on B. tryoni, for which more literature is available, we demonstrate that the seasonality exhibited by that species is closely correlated with the seasons of its endemic rainforest environment as recognised by traditional Aboriginal owners. Evidence suggests the presence of a seasonal reproductive arrest which helps the fly survive the first two-thirds of the dry season, when ripe fruits are scarce, followed by a rapid increase in breeding at the end of the dry season as humidity and the availability of ripe fruit increases. This seasonal phenology continues to be expressed in human-modified landscapes and, while suppressed, it also partially expresses in long-term cultures. We subsequently demonstrate that B. dorsalis, across both its endemic and invasive ranges, shows a very similar seasonality although reversed in the northern hemisphere. While high variability in the timing of B. dorsalis population peaks is exhibited across sites, a four-month period when flies are rare in traps (Dec-Mar) is highly consistent, as is the fact that nearly all sites only have one, generally very sharp, population peak per year. While literature to support or deny a reproductive arrest in B. dorsalis is not available, available data is clear that continuous breeding does not occur in this species and that there are seasonal differences in reproductive investment. Throughout the paper we reinforce the point that our argument for a complex reproductive physiology in Bactrocera is based on inductive reasoning and requires specific, hypothesis-testing experiments to confirm or deny, but we do believe there is ample evidence to prioritise such research. If it is found that species in the genus undergo a true reproductive diapause then there are very significant implications for within-field management, market access, and biosecurity risk planning which are discussed. Arguably the most important of these is that insects in diapause have greater stress resistance and cold tolerance, which could explain how tropical Bactrocera species have managed to successfully invade cool temperate regions.
