Impact of the colonisation of Zaprionus (Diptera, Drosophilidae) in different ecosystems of the Neotropical Region: 2 years after the invasion
Introduction
Some of the most dramatic ecological events in recent decades have been species invasions. Nevertheless, according to Carey et al. (1996), research in invasion biology has tended to focus on the technology of control of invading organisms, rather than on attempting to understand why invasions succeed or fail, with the potential pay-off in the prevention of future invasion.
Rising evidence suggests that even the removal or the addition of species that strongly interact with disturbance regimes frequently results in discrete changes in ecosystem structure, and function. Thus, biological invasions offer model systems for understanding the mechanisms by which species or functional groups alter disturbance regimes (Mack and D'Antonio, 1998). These alterations occur in both disrupted and intact systems resulting in profound changes in many cases, including direct species replacements and changes in ecosystem processes that ultimately control plant and animal activity (Knops et al., 1999). In order to understand the changes in the receptor ecosystem, it is essential for us to study the impact of recently introduced species in natural environments.
We generally accept that flies belonging to the family Drosophilidae are excellent organisms for biological studies; according to Powell (1997), no other model has been so thoroughly studied. The genus Zaprionus is part of the family Drosophilidae and currently comprises 52 described species (Chassagnard and Kraaijeveld, 1991, Chassagnard and Tsacas, 1993). However, genetic data suggests that Zaprionus is included in the genus Drosophila (Kwiatowski & Ayala, 1999); in this case, Drosophila would be defined as a monophyletic taxon. Zaprionus inhabit the Australian, Afrotropical, Oriental and Palaearctic regions; among the typical genera of the afrotropical fauna, this taxon is of special significance in view of its abundance and the large body size of some of its species (Tsacas et al., 1981).
Zaprionus indianus constitutes one of the most successful colonising species of this genus (Chassagnard and Tsacas, 1993). According to Parkash and Yadav (1993), populations of this species have habitat-generalist or broad niche-width characteristics, i.e. it utilises diverse food resources and displays adaptation to variable climatic conditions. The first published record of this species for the American continent (Vilela, 1999) refers to individuals observed on fallen persimmon fruits (Diospyrus kaki L.; Ebenaceae) in the metropolitan area of the city of São Paulo, São Paulo state, Brazil, in March 1999. In that same period, during the fig crop (Ficus carica L.; Moraceae) in the area of Valinhos (São Paulo, Brazil), the presence of a large number of Z. indianus was found feeding on and ovipositing in fruits which were still ripening, making them inappropriate for human consumption. Up to where it is known, this species has not been considered a pest in its original area of occurrence. However, with the return of several fig lots and a loss on the order of 50% of that crop, this African insect seems to be reaching the pest status in the main fig growing area in the state of São Paulo (http://www.iac.br/∼cenfit/artigos/zaprionus).
During 1999 and 2000, several drosophilists collected Z. indianus in South America (Galinkin and Tidon, 2000, Moraes et al., 2000, Goni et al., 2001, Toni et al., 2001). These insects are conspicuous because they have a pair of white stripes along the sub median area of the dorsal surface of the head and thorax, an unusual feature for the neotropical drosophilids. According to Vilela (1999), collections made since 1943 in several regions of Brazil failed to detect any specimen of any species belonging to the genus Zaprionus, which seems to indicate that this afrotropical species has only very recently been introduced in Brazil. Vilela (op cit) still inquires whether this insect will be able to spread its distribution throughout the several ecosystems of the American continent in competition with local species, making this a possible introductory route for Z. indianus in South America.
In this study, data are presented from two years of collectionfrom cerrados (neotropical savanna) and gallery forests in central Brazil. Hypotheses on the introductory route of Z. indianus into South America, as well as aspects of this species adaptation to different environmental conditions in space and time, are also discussed.
Section snippets
Materials and methods
Monthly collections of drosophilids were accomplished in two areas close to the city of Brası́lia, capital of Brazil: the Ecological Reserve of IBGE (RECOR) and the National Park of Brası́lia (PN). Located 35km south of Brası́lia (15° 56′ S; 47° 53′ W), RECOR is part of a 10.000-ha environmental protection area . In addition, RECOR is part of the Reservation Nucleus for the Cerrado Biosphere, created in 1993 by UNESCO. The other site, PN, is a 30,000 ha park located 10 km
Results
We have collected drosophilids in central Brazil since 1998 (although without chronological regularity, see Table 1). In February 1999 we captured individuals of the genus Zaprionus for the first time. In addition to Z. indianus, we identified D. simulans, D. malerkotliana, D. hydei, D. busckii, D. immigrans, and Scaptodrosophila latifasciaeformis among the introduced species. The endemic species include representatives of the cardini, guarani, repleta, tripunctata, saltans and willistoni
Discussion
For millions of years, oceans and other natural barriers have restricted the distribution of the world's biota. During the last 100 years, human activities, especially international travel and trade, have circumvented these barriers, and species are invading new continents at an increasing rate. Biological invasions of insect, plant, and fungal pest species often cause substantial disturbance to ecosystems as well as severe socio-economic impacts (Liebhold et al., 1995).
Vilela (1999) asked: “Is
Conclusions
Invasions of non-indigenous species threaten native biodiversity, ecosystem functioning, animal and plant health, and human economies (Myers et al., 2000). According to Vermeij (1996), the invasion process is composed of three phases: arrival, establishment, and spread. Arrival occurs when a species is initially transported to the new area (e.g., transportation to a new continent). Establishment occurs when a population becomes abundant enough to prevent extinction, and spread is the process by
Acknowledgments
We are grateful to A. C. Franco and R. Henriques for the critical reading of the manuscript; R. Leão and R. Constantino for helping with the figures, Parque Nacional de Brası́lia and Reserva Ecológica do IBGE for the access to study sites. We wish to thank two anonymous reviewers and especially Mark Schwartz for his suggestions to the final version of the manuscript. This study was supported by the Conselho Nacional de Desenvolvimento Cientı́fico e Tecnológico (CNPq), Coordenação
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