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On the Geographic Origin of the Medfly Ceratitis Capitata (Wiedemann) (Diptera: Tephritidae)

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On the Geographic Origin of the Medfly Ceratitis Capitata (Wiedemann) (Diptera: Tephritidae) Proceedings of 6th International Fruit Fly Symposium 6–10 May 2002, Stellenbosch, South Africa pp. 45–53 On the geographic origin of the Medfly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) M. De Meyer1*, R.S. Copeland2,3, R.A. Wharton2 & B.A. McPheron4 1Entomology Section, Koninklijk Museum voor Midden Afrika, Leuvensesteenweg 13, B-3080 Tervuren, Belgium 2Department of Entomology, Texas A&M University, College Station, TX 77843, U.S.A. 3International Centre for Insect Physiology and Ecology, P.O. Box 30772, Nairobi, Kenya 4College of Agricultural Sciences, Penn State University, University Park, PA , U.S.A. The Mediterranean fruit fly, Ceratitis capitata (Wiedemann), is a widespread species found on five continents. Evidence indicates that the species originated in the Afrotropical Region and may have spread worldwide, mainly through human activities. Historical accounts on adventive populations outside Africa date back at least 150 years. The exact geographic origin of the Medfly within Africa has been much debated. Recent research regarding phylogeny, biogeography, host plant range, and abundance of the Medfly and its congeners within the subgenus Ceratitis s.s., all support the view that the species originated in eastern Africa,possibly the Highlands,and dispersed from there. Only molecular evidence contradicts this view, with higher mitochondrial DNA diversity in western Africa, suggesting that the hypothesis of a West African origin should still be considered. INTRODUCTION dance records,and parasitoid data.There is consid- The Mediterranean fruit fly, or Medfly, Ceratitis erable evidence that the geographic origin of the capitata (Wiedemann),is among the most important Medfly must have been situated in southern or pests of cultivated fruits (White & Elson-Harris eastern Africa, although there are contradictory 1992). The genus to which it belongs, Ceratitis findings that still require us to consider the possi- MacLeay, comprises close to 90 species, all with an bility of a western African origin. Afrotropical distribution. The Medfly is the only species that readily colonizes tropical and mild MATERIALS AND METHODS temperate habitats far outside its home range and A large part of the data provided herein is is now reported from five different continents extracted from a relational database, compiled by (Gilstrap & Hart 1987). the first author, that comprises all collection and Although there is no doubt that the species literature data for the genus Ceratitis in Africa. This originated from the Afrotropical Region, the exact includes all available information associated either provenance is unknown and the source of much with particular specimens or published reports. debate (Balachowsky 1950; Maddison & Bartlett Individual database records include locality and 1989; Gasparich et al. 1997). Precise knowledge of date of collection, collector, sex of specimen, the geographic origin of the Medfly is, however, collection depository,associated literature records, deemed important, especially with regard to host plant (if known) and the type of relationship applied aspects such as pest management. A with the plant (reared, found on, or other associa- reconstruction of its geographic origin and subse- tions), as well as any additional information (e.g. quent dispersal throughout the region could pro- collecting reference numbers, lures, trap type, and vide essential information on the parasitoid habitat description when given). At the time of community, genetic diversity, and endemic host writing, this database comprised more than 4300 plant spectrum of the species as well as indicate block records,representing over 12 000 specimens environmental thresholds that delimit its distribu- (for further details, see De Meyer 2001). A large tion. part of the data is based on the results of recent This paper aims to synthesize current knowledge surveys in Kenya undertaken by R.S.C., as part of and evidence with regard to this debate.It presents USAID and USDA projects, funded through Texas an overview of published information, as well as A&M University (coordinated by R.A.W.). In total, recent findings of the authors regarding this 2004 samples (480 958 fruits) were collected matter, especially in the fields of phylogeny recon- from 518 plant species, representing 88 families struction, host plant data, distribution and abun- (Copeland et al. 2002). Twenty-seven per cent of *To whom correspondence should be addressed. the collections produced tephritids. These data E-mail: [email protected] were used for analysis regarding host plant 46 Proceedings of the 6th International Fruit Fly Symposium evidence, distribution and abundance data. Host Although a type locality does not necessarily plant distributions are based on existing floras for indicate the geographic origin of any species, it is the region (Agnew & Agnew 1994; Beentje 1994; still noteworthy in the fact that the type more likely Coates-Palgrave 1983) and data on the Tropicos originated from the Afrotropical Region, rather web site (http://mobot. mobot.org). than from, for example, the Mediterranean region. The taxonomic and phylogenetic information is based primarily on recent revisions of the subgen- PHYLOGENETIC EVIDENCE era of the genus Ceratitis by the first author (some- The Medfly belongs to the genus Ceratitis which times in collaboration with other specialists) (De comprises 87 species, grouped in six subgenera: Meyer 1996, 1998, 1999, 2000; De Meyer & Cope- Acropteromma Bezzi (1 species),Ceratalaspis Hancock land 2001, De Meyer & Freidberg, in press). (31 species), Ceratitis sensu stricto (eight species), Parasitoid evidence is based on recent research by Hoplolophomyia Bezzi (one species), Pardalaspis the third author, and on published reports, while Bezzi (10 species),and Pterandrus Bezzi (36 species). molecular evidence is based upon Gasparich et al. All these subgenera have been systematically (1997), and additional findings by the fourth reviewed recently (De Meyer 1996, 1998, 2000; De author. Meyer & Copeland 2001; De Meyer and Freidberg, in press) with redescriptions of all species. Unam- TAXONOMIC HISTORY biguous identification for all species is currently Ceratitis capitata was described by Wiedemann possible. The genus as a whole is restricted to the (1824) based on a single male specimen that is Afrotropical Region (including islands in the Indian deposited in the Zoological Museum of Copenha- and Atlantic Oceans), with only C. capitata having gen, Denmark (see De Meyer 2000 for description adventive populations in other biogeographical of present condition of the type specimen). The regions. original description (Wiedemann 1824) mentions From a preliminary cladistic analysis (De Meyer a dual type locality: ‘India orient.’ and ‘in mare 1999) based on 53 representatives of the genus, indico’,which is repeated in the German translation it was shown that the subgenus Ceratitis s.s. forms published a few years later (Wiedemann 1830): a distinct monophyletic clade. Currently, more ‘Aus Ostindien’and ‘im Indischen Ocean gefangen’. elaborate analyses, including all recognized spe- According to Pont (1995) ‘India orient.’ in Wiede- cies of the genus,confirmed the monophyly for the mann’s publications refer to east India or the East subgenus Ceratitis. The interrelationships within Indies (Indonesia), but usually the latter. However, this clade were analysed separately by De Meyer no historical records are known that indicate that (2000) through a cladistic analysis. This resulted in Ceratitis capitata was ever present or established in two completely resolved most parsimonious trees Indonesia or the Southeast Asian region in general. differing solely in the position of C. pinax Munro. A Even records for India are probably only based on strict consensus tree is shown in Fig. 1a. The same quarantine interception and there is no reliable consensus tree, with replacement of the species evidence that there was ever an established popu- by their current distribution in the Afrotropical lation on this subcontinent (Kapoor 1993, 2002). Region, is represented in Fig. 1b. This ‘area clado- The type specimen was collected by Dagobert Karl gram’shows a basal lineage that is found in south- Daldorff, who made a voyage from Denmark to ern Africa (C. cornuta (Bezzi), although at the Tranquebar (India) in 1790/91 (Henriksen 1923). It moment of preparing this article, this species is is therefore not unlikely that the specimen was probably newly recorded for Kenya as well), collected on board the ship in the Indian Ocean, followed by a number of species found in south- presumably emerging from fruits that were taken ern to eastern Africa, with the single exception of as provisions. The latter could have happened in C. brachychaeta Freidberg which is also reported the Cape region, which was a regular stop-over for from two localities in Central Africa (Democratic ships sailing to the East in those days. This could Republic of Congo and Cameroon). The closest place the original type locality in the Cape, relative of the Medfly, C. caetrata Munro, is limited South Africa. It cannot be confirmed, however, to Kenya. The sister clade of caetrata/capitata con- whether the ship that took Daldorff to the East sists of three species that are limited to the islands actually stopped in South Africa or any other place. of the Indian Ocean. Daldorff’s log does not give any indication in this respect. A discussion on the origin of the type Host plant data specimen can be found
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