Proceedings of 6th International Fruit Symposium 6–10 May 2002, Stellenbosch, South pp. 45–53

On the geographic origin of the Medfly capitata (Wiedemann) (Diptera: )

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, A&M University, College Station, TX 77843, U.S.A. 3International Centre for 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, (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 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 in De Meyer (2000). Ceratitis capitata is an extremely polyphagous De Meyer et al.: Geographic origin of the Medfly Ceratitis capitata 47

a

b

Fig. 1. a, Cladogram for the subgenus Ceratitis s.s.; b, area-cladogram for the subgenus Ceratitis s.s. (SA = southern Africa, EA = eastern Africa, CA = central Africa, COSM = cosmopolitan, IO = Indian Ocean islands). species with close to 400 different host plants If we consider, however,those hosts that C.capitata known worldwide (Liquido et al. 1998). In Africa and C. caetrata have in common, and from which more than 150 host plants are reported with only these two species have been reared (hence certainty (De Meyer et al. 2002). The close rela- hosts unique for the clade capitata/caetrata), only tives of C.capitata,within the subgenus Ceratitiss.s., Table 1. Host plants of Ceratitis caetrata. have a more restricted host spectrum, confined to a single host genus or a few genera at most. Ebenaceae: Euclea divinorum Hiern Ceratitis caetrata, which is considered the closest Rutaceae: Vepris trichocarpa (Engl.) Mziray relative of the Medfly, has six known hosts belong- ing to four different plant families (Table 1). All six Sapotaceae: Manilkara butugi Chiov. Mimusops bagshawei S. Moore hosts have largely a southern to eastern African Simaroubaceae: Brucea antidysenterica J.F. Mill. distribution. Only Harrisonia abyssinica Oliv. (Sima- Harrisonia abyssinica Oliv. roubaceae) is also reported from western Africa. 48 Proceedings of the 6th International Fruit Fly Symposium

Table 2. Number of hosts known per country and per region for three polyphagous and widespread Ceratitis species (W = western Africa, C = central Africa, E = eastern Africa, S = southern Africa).

Country C. capitata Regional C. cosyra Regional C. punctata Regional

Senegal W:7 2 W:11 W:8 Guinea 1 2 2 Liberia 1 Sierra Leone 3 1 Mali 2 Ivory Coast 1 2 3 Ghana 3 1 1 Benin 1 Togo 1 1 1 Nigeria 1 3 3 Cameroon 2 C:6 1 C:12 3 C:4 Congo (DR) 4 11 2 Uganda E:73 2 E:15 4 E:8 Kenya 67 13 4 Tanzania 8 3 4 Malawi 1 Zambia 1 1 1 Zimbabwe 4 S:55 8 S:12 S:2 South Africa 52 6 2 Namibia 1 3 three species remain: Manilkara butugi Chiov., (Table 2, columns ‘Regional’), the same pattern is Mimusops bagshawei S. Moore (both Sapotaceae) observed. and Brucea antidysenterica J.F.Mill. (Simarouba- ceae). These host species are restricted to eastern Distribution and abundance Africa(KenyaandTanzania).AtonelocalityinKenya Ceratitis capitata has been considered wide- that was intensively sampled (Kakamega Forest, spread throughout the Afrotropical Region. It is Western Province), and where both species were indeed reported from at least 39 countries in reared from Manilkara butugi and Mimusops that biogeographical entity. However,considering bagshawei, several other polyphagous species like records per country, it is clear that those from C. fasciventris and C. anonae were present but were western and central Africa are relatively scarce, never reared from these hosts. while those for eastern and southern Africa are Table 2 summarizes the number of different host abundant.This scarcity was already reported in the plants reported per country in the above-men- literature. Noteworthy in this respect are the tioned database for C. capitata, as well as two reports of the collecting trips organized by the other polyphagous and widely distributed Ceratitis Hawaiian Board of Agriculture and Forestry, and species (C. cosyra (Walker) and C. punctata the United States Department of Agriculture in (Wiedemann)),throughout their ranges from West search of biological control agents for Medfly on Africa, to central, eastern and eventually southern their continent of origin. Both Silvestri (1914) and Africa. This gives an indication of the relative Van Zwaluwenburg (1936), who collected inten- abundance of host plant records reported for sively in western Africa,pointed out that C.capitata eastern and southern regions compared to central are hard to come by in the region, despite and western regions. For the Medfly, the number repeated rearing efforts from a wide array of host of host records from West and Central African plants. More recently, Steck et al. (1986) con- countries is fairly limited,compared to records from ducted intensive sampling in Ghana and Camer- eastern or southern African countries. Ceratitis oon of coffee and other suitable hosts for C. cosyra and C. punctata have a less skewed list. capitata. A full year cycle of collecting coffee only When the data are grouped according to different yielded approximately 500 individuals, which is host plants reported per region for western, very limited compared to similar sampling efforts central, eastern and southern African countries in East Africa. De Meyer et al.: Geographic origin of the Medfly Ceratitis capitata 49

Fig. 2. Distribution of Ceratitis capitata in Africa (black triangles) versus total distribution of Ceratitis (open circles) (see text for explanation).

When we analyse the data in the above-men- 1937; Clausen et al. 1965; Steck et al. 1986; Whar- tioned database, a similar pattern is observed. For ton et al. 2000). Approximately 34 hymenopteran all Ceratitis species, the total records for Africa parasitoids have thus far been reared from fruit-in- are shown in Fig. 2 (open circles). Overall these festing tephritids in Africa (including three from records cover all regions relatively well, despite Mauritius and Réunion). The vast majority of larger concentrations in certain areas. If the re- these have come from batches with more than one cords for C. capitata are superimposed on the host fly, and are apparently not species-specific overall generic distribution (Fig. 2, black triangles), (though they may be confined to a narrow range there is an uneven distribution, with the majority of tephritid hosts). About 13 have been reared of records from the eastern and southern regions. from samples containing Medfly, and a few addi- For comparison, a similar superimposition for tional species have been laboratory-reared on C. cosyra shows a more even distribution (Fig. 3). Medfly following emergence from other hosts. Six of these are suspected to regularly attack Parasitoids Medfly. Only two parasitoids are apparently lim- Within the Afrotropical Region,Ceratitis fruit flies ited to Medfly: Fopius ceratitivorus Wharton and are readily attacked by parasitic Hymenoptera, Psyttalia humilis (Silvestri). Fopius ceratitivorus is particularly of the families Braconidae,Eulophidae, so far only reported from East Africa, with one un- Diapriidae, Chalcididae and Pteromalidae. Over published record from northeastern South Africa. the last 90 years, several collecting efforts have Psyttalia humilis is only known from South Africa been made to gather these parasitoids within the (but could be a host race of the more widespread framework of biological control of Medfly outside species P. concolor). The four other major Medfly its natural range (Silvestri 1913; Bianchi & Krauss parasitoids (Diachasmimorpha fullawayi (Silvestri), 50 Proceedings of the 6th International Fruit Fly Symposium

Fig. 3. Distribution of Ceratitis cosyra in Africa (black triangles) versus total distribution of Ceratitis (open circles) (see text for explanation).

Tetrastichus giffardianus Silvestri, Coptera robustior and southern Africa, and three known only from (Silvestri), Dirhinus giffardii Silvestri) have also been Kenya and Tanzania. Although it is not a prerequi- reared from other fruit-infesting tephritids and site that ancestral hosts should be unique to the occur in both East and West Africa. capitata/caetrata clade, the limited geographic distribution of the shared hosts could be seen as an DISCUSSION additional argument for eastern Africa being the Both the phylogenetic, host plant, abundance, geographic origin. In addition, the spectrum of distribution as well as parasitoid evidence seem host plants for Medfly in eastern and southern to indicate a southern to eastern origin of the Africa seems to be much wider than in other parts Mediterranean fruit fly,C.capitata. From the phylo- of its geographic range.There is certainly a collect- genetic analysis, the closest relatives of C. capitata, ing bias towards these regions (because of more those species grouped within the same clade as active sampling programmes) but other Ceratitis C. capitata within the subgenus Ceratitis s.s., are all species with a general distribution throughout mainly restricted to southern to eastern Africa or the continent do not show this tendency as is Indian Ocean islands. The Medfly’s sister taxon, illustratedwithrecordsforC.cosyraandC.punctata. C. caetrata, is only reported from several localities The same applies to distribution and abun- in Kenya. dance data. Again we can expect a bias towards The common but exclusive host plant spectrum eastern and southern African data because of for the clade capitata/caetrata is equally restricted larger sampling effort, but the tendency for to East Africa (Kenya and Tanzania). All of the six capitata is not repeated in other widely distrib- host plants of C. caetrata are shared by C. capitata. uted species (cf. Figs 2 & 3). Ceratitis capitata Five of these six hosts are found only in eastern seems truly to be more abundant and more De Meyer et al.: Geographic origin of the Medfly Ceratitis capitata 51 widely distributed in the eastern and southern Africa. In the same region, high infestation rates areas than in western areas (although it can be were also observed in Solanum seaforthianum present in large numbers in the latter in a particular Andrews, Mimusops obtusifolia Lam., Guettarda host at a particular time as observed in Ghana in speciosa L., Ekebergia capensis Sparrm. and Capsicum frutescens L.). A lower abundance could Harrisonia abyssinica Oliv. (Copeland et al. 2002). be interpreted as proof of original range, since The main argument against Balachowsky’s sugges- more natural enemies would be expected to tion is that the southern part of Morocco belongs suppress the population. The parasitoid evidence, to an entirely different biogeographical region (the however, does not seem to support this.Therefore, Palaearctic) which is clearly separated from the while abundance by itself is not a good criterion, Afrotropical Region, where all other relatives of C. it can be seen in this context as supporting capitata are found,by the Sahara Desert. Through- evidence, especially since in certain areas of west- out geological times the present day arid condi- ern Africa Medfly does not seem to occur at all. tions of the Sahara region were not always present. The parasitoid evidence is limited at this stage. For example, during the Early Holocene, there is Nevertheless, there is an indication that the only an indication that vegetation was denser than at hymenopteran parasitoids that are species-spe- present (Gowdie 1983), so there could have been cific regarding Medfly, are only found in eastern a connection in the past. But the fact that all and southern Africa. One would expect such close relatives of C.capitata are found in eastern to specialists in the centre of origin, rather than in southern Africa, without any connection to south- regions were the species was introduced or in- ern Morocco, cannot be explained by a founder vaded at a later stage. species in that part of the continent. Moreover, There seems to be, therefore, some convincing there is some historical evidence regarding when evidence to hypothesize the geographic origin of the Medfly was first observed in the Mediterranean the Medfly as somewhere within the region region (Fimiani 1989), with the first records from stretching from South Africa to East Africa. Such the Iberian Peninsula in 1842, Italy in 1863, and a distribution pattern along the southeastern France in 1885. A species that originated from stretch of the African continent is a common obser- the Mediterranean region, as suggested by vation in Ceratitis fruit flies (De Meyer 2001). For Balachowsky, would be expected to have a much C. capitata, the evidence pointing to an origin in older record of presence in that region. One could or near Kenya is particularly strong. In this area, even argue that the type locality of the species Medfly co-occurs in the same plant host with its would be expected to have been in that region, nearest relative, attacks a wide range of indige- given the entomological activity in Europe in the nous fruit species, reaching high densities in late 18th and early 19th centuries. regarding the several of them, and is attacked by an appar- argument that Medfly is the only Ceratitis species ently specific parasitoid. capable of surviving in a temperate climate, this Nevertheless there are some contradicting is only true in the sense that it is the only species hypotheses that suggest alternative regions of established in predominantly temperate countries. origin. Balachowsky (1950) argued that the south- Developmental studies in Réunion have indicated western part of Morocco might have been the that, for example, , which is also a origin of the Medfly, based on the following polyphagous species with adventive populations observations. The main host of C. capitata in that on the Indian Ocean islands, is better adapted to region is Argania spinosa (L.) Skeels (Sapotaceae) low temperatures than C. capitata (Duyck et al. which is endemic to the southern part of Morocco. 2002). There seems, therefore, little or no support Balachowsky suggested that, because of the for the hypothesis of a North African origin of the abundance of the fly in this fruit, this could be the Medfly. original host. An additional argument of that A second, and more convincing hypothesis is author is that C. capitata is the only species of the that of Gasparich et al. (1997), who studied mito- genus that is capable of surviving in a temperate chondrial DNA variation in more than 100 Medfly climate, hence it could have originated from the populations from the New and Old Worlds. They Mediterranean region. Although these are correct found the highest diversity of mitochondrial observations, it must be said that C. capitata is haplotypes in samples from western Africa (i.e. equally abundant in other hosts, given suitable Ghana and Nigeria). Moreover, this diversity was conditions. It maintains, for example, high num- found in relatively small samples from single host bers in coffee in the Central Highlands of eastern plants (Capsicum frutescens for the sample from 52 Proceedings of the 6th International Fruit Fly Symposium

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