Description of new Ceratitis MacLeay (Diptera, Tephritidae) species from Africa

As an addition to earlier published systematic revisions of the genus Ceratitis MacLeay, five new species are hereby described: Ceratitis (Pardalaspis) millicentae sp. n., C. (Ceratalaspis) oraria sp. n., C. (C.) perisae sp. n., and C. (C.) perseus, sp. n. all from Kenya, and C. (C.) ealensis sp. n. from the Democratic Republic of Congo. Their relationship with closely allied species within the respective subgenera is discussed and differentiating characters given.


Introduction
Tephritidae are picture-winged flies of variable size and worldwide distribution. Although commonly named ''fruit flies'', larval development can also take place in other parts of the host plants, including flowers, seeds and stems. Fruit flies of economic significance were treated by White and Elson-Harris (1994), and the currently acknowledged classification is presented by Norrbom et al. (1999). The genus Ceratitis MacLeay belongs to the tribe Ceratitidini (subfamily Dacinae) which is predominantly an Afrotropical group. For a discussion of classification and relationship with other genera, see White and Elson-Harris (1994), De Meyer (1999b) and Norrbom et al. (1999). The larvae of most Ceratitidini species develop in fruit, and several species of agricultural importance are known, especially within the genera Ceratitis, Capparimyia Bezzi, Neoceratitis Hendel, and Trirhithrum Bezzi.
Since 1999 recent surveys in Kenya have been undertaken by the second author, as part of USAID and USDA projects, funded through Texas A&M University. In total 3257 collections (834,040 fruits) were collected from ca 800 plant species, representing 109 families (Copeland et al. 2002;R. Copeland, unpublished data). Twenty-five per cent of the collections produced tephritids, including at least 20 species new to science. New species belonging to the subgenus Pterandrus are included in above-mentioned paper. New species of the subgenera Ceratalaspis and Pardalaspis are described below. Also included is another new Ceratalaspis species from the Democratic Republic of Congo, that was found in the recently revised collection of the Royal Institute for Natural Sciences at Brussels, Belgium (KBIN).

Material and methods
Terminology follows McAlpine (1981) and White et al. (1999). A specific term used for describing the leg ornamentations is ''feathering''. This refers to a regular row of long, stout and flattened setae along margins of a femur or tibia, giving that part the appearance of a bird's feather (see De Meyer and Freidberg forthcoming). Drawings were made with a camera lucida attached to either a dissecting or compound microscope. Measurements of wing length and body length (average and range) are based on 10 specimens of each sex (whenever available) and are given in mm. Body length measurements do not include oviscape. Plant names follow Mziray (1992), Brummitt (1992), Beentje (1994), and Mabberley (1998).

Subgenus Ceratitits (Ceratalaspis) Hancock
A diagnostic description of the subgenus Ceratalaspis is given in De Meyer (1996).

Description
Female. Head: antenna yellow to yellow-orange. First flagellomere three times as long as pedicel. Arista with short to medium long rays at most twice as long as basal width of arista; ventral rays shorter than dorsal rays basally. Frons yellow to yellow-orange; with short scattered black setulae. Face white, lower half and below lower eye margin more yellow. Genal seta and setulae orange to red, well developed. Thorax: postpronotal lobe pale yellowish white; with black spot. Scutum (Figure 1), ground colour yellow-orange, with distinct black markings including median presutural and sutural spots; prescutellar markings yellowish white, merged; with silver microtrichosity and yellow pilosity. Scapular setae yellow. Scutellum white to yellow, basally with small black spots, apically with three separate black spots, extending to basal third. Subscutellum yellow with three separate black spots, sometimes black spots touching along dorsal margin. Anepisternum yellow, upper margin white with black patch; one black anepisternal seta, setulae pale.
Legs: yellow; setation yellow to orange except last tarsal segments black. Fore femur with ventral setae orange.
Wing: pattern as in Figure 7; bands mainly yellowish with some brown patches. Anterior apical and discal bands not separated; subapical band free; posterior apical band absent; crossvein R-M at, or just proximal to, middle of discal medial cell.
Abdomen: yellow to orange; pilosity mixed yellow and black. Tergites 2 and 4 with pale grey band occupying posterior third of tergite. Tergite 5 with longer black setulae along posterior margin. Oviscape about as long as preabdomen. Aculeus 10 times longer than wide, tip ( Figure 12) slightly bilobed and lateral margin slightly sinuate.
Male. As female except for the following characters. Arista with medium long rays, longest dorsal rays about three times as long as basal width of arista. Face completely white. Fore femur posteriorly with longer and somewhat denser pilosity.

Etymology
After the Latin ''orarius'', meaning coastal dweller, and referring to the type locality Arabuko Sokoke Forest which is the largest remnant of coastal forest in Kenya.

Remarks
This species is very similar to C. stictica Bezzi and keys out as such in De Meyer (1998). The following characteristics can be used to differentiate both species. Arista with medium long rays in C. oraria, at most three times basal width; distinctly plumose in C. stictica with longest rays at least five times as long as width. Presutural median spot less developed in C. oraria. Basal scutellar spots small, reduced in C. oraria. Also abdominal pilosity largely yellow in stictica while mixed in oraria, and grey band on tergite 4 more distinct in oraria. C. stictica often with variable wing pattern, with subapical band joined with discal band. This is not the case in C. oraria which has the typical Ceratitis pattern with subapical band separate from discal band. The species can be identified with the key provided by De Meyer (1998) given the following adjustment. Confirmed records for C. stictica so far are all from Central Africa, as well as Ukerewe Island in Lake Victoria and Kakamega forest (the easternmost relict of Guineo-Congolian rain forest, situated in the extreme west of Kenya, close to the border with Uganda). C. oraria is a coastal species, known from a plant host whose distribution is also restricted to the coastal region in Kenya. The only specimens of C. stictica from Kenya, on the other hand (Kakamega forest, vide supra), were reared from fruits of Craterispermum schweinfurthii Hiern. (Rubiaceae), a plant that is restricted in Kenya to Kakamega forest.
Ceratitis (Ceratalaspis) perisae sp. nov. (Figures 2,8,13) Diagnosis Arista with short rays; frons pale yellow, without silvery microtrichosity; two frontal setae; postpronotal lobe without a spot; scutal pattern with distinct black spots but latter strongly reduced; anepisternum completely yellow, with one anepisternal seta; wing bands with subapical band joined to discal band, and posterior apical band joined to anterior apical band; legs without feathering; abdomen without distinct spots, with grey band on tergites 2 and 4; apical part of aculeus with three large rounded indentations.

Description
Female. Head: antenna pale yellow. First flagellomere darker yellow; three times as long as pedicel. Arista with short rays, longest dorsal rays about as long as basal width of arista. Frons pale yellow; with short scattered yellow setulae; two frontal setae. Face white. Genal setulae darkish, poorly developed; genal seta reddish.
Thorax: postpronotal lobe yellowish white; without spot. Scutum (Figure 2), ground colour yellow-orange, with distinct brown to black markings but these restricted to intra-alar spot and spot linking dorsocentral and acrostichal prescutellar setae, sutural spot brownish, other dark markings pale brownish; prescutellar markings white, separate; with yellow microtrichosity and pilosity. Scapular setae black. Scutellum white, basally without black spots but with yellow-brown marking, apically with three separate black spots, extending to basal third. Subscutellum yellow with three separate black spots, touching along dorsal margin. Anepisternum yellow; one black anepisternal seta, setulae pale.
Legs: yellow; setation yellow to orange except last tarsal segments black. Fore femur with ventral setae yellow, posteriorly with longer pilosity.
Wing: pattern as in Figure 8; bands mainly yellowish with some brown patches. Anterior apical and discal bands not separate; subapical band joined to discal band; posterior apical band joined to anterior apical band; crossvein R-M at proximal two-fifths of discal medial cell.
Abdomen: yellow to orange; pilosity mainly black. Tergites 2 and 4 with pale grey band occupying posterior third of tergite. Tergite 5 with longer black setulae along posterior margin. Oviscape shorter than preabdomen. Aculeus six to seven times longer than wide, apical part ( Figure 13) with three large rounded indentations, tip with small depression.
Male. As female except for the following characters. First flagellomere pale yellow. Mesonotal pattern with sutural spot also pale brownish, male paratype of Ngaia Forest with very weak mediolongitudinal line postsuturally. Scutellum basally with yellow marking. Subscutellum with black spots separate. Wing, crossvein R-M at proximal third of discal medial cell.

Etymology
Named for Ms Peris Machera, an extraordinarily dedicated technician whose work was largely responsible for the success of the USAID and USDA projects that resulted in this new material.

Remarks
This species keys out to the connexa-argenteobrunnea couplet in the key of De Meyer (1998). It can be differentiated from the former by differences in number of frontal seta (always two pairs in perisae, one or three pairs in C. connexa (Bezzi)) and from the latter by the different mesonotal pattern. The aculeus shape indicates a relationship with C. argenteobrunnea Munro and with the C. andranotobaka Hancock species group as defined in De Meyer (1998) within the subgenus Ceratalaspis as well as with C. (Hoplolophomyia) cristata (Bezzi). The host plant of perisae is also known to host C. cristata in the same region. It can be identified with the key provided by De Meyer (1998)  Diagnosis Arista with medium long rays; frons yellow, without silvery microtrichosity; postpronotal lobe with distinct black spot; scutal pattern with distinct black markings; anepisternum yellow with upper half white, with one anepisternal seta; wing bands with subapical band separate, posterior apical band absent; male mid tibia with black feathering; abdomen without distinct spots, with grey band on tergites 2 and 4; aculeus tip with median step.

Description
Male. Head: antenna yellow to yellow-orange. First flagellomere three times as long as pedicel. Arista with medium long rays, longest dorsal rays about three times as long as basal width of arista; ventral rays shorter than dorsal rays basally. Frons yellow to pale yellow; with short scattered black and yellow setulae. Face white. Genal seta and setulae black, well developed. Thorax: postpronotal lobe pale yellowish white; with black spot. Scutal pattern (Figure 3), ground colour yellow to dark orange, with distinct black markings, including median sutural spot, prescutellar markings white, separated by pale area; with golden microtrichosity and yellow pilosity. Scapular setae black. Scutellum white to yellow, basally with black spots, apically with three separate black spots, extending to basal third. Subscutellum black. Anepisternum yellow, upper half white, one anepisternal seta, setulae white to yellow.
Legs: yellow; setation yellow to orange. Fore femur with ventral setae mainly black; posteriorly with dense yellow to silvery pilosity. Mid leg (Figure 6), femur ventrally with few conspicuously longer yellow setulae along basal third; tibia with black feathering at apical outer half and inner two-fifths, area of tibia with feathering coloured black.
Wing: pattern as in Figure 9; bands mainly yellowish with some brown patches. Anterior apical and discal bands largely but incompletely separated, narrowly joined; subapical band free; posterior apical band absent; crossvein R-M at, or just distal to, middle of discal medial cell.
Abdomen: yellow to orange; pilosity mixed yellow and black. Tergites 2 and 4 with pale grey band occupying posterior third of tergite. Tergite 5 with longer black setulae along posterior margin.
Female. As male except for the following characters. Frons only with short black setulae. Fore femur with less dense pilosity posteriorly; mid tibia without black feathering, completely yellow. Oviscape shorter than preabdomen. Aculeus eight times longer than wide, tip ( Figure 14) with median step and lateral margin slightly sinuate.

Etymology
After the Greek mythological figure Perseus, who borrowed the God Hermes' winged shoes when killing the Medusa.

Remarks
This is an enigmatic species, showing characteristics of the subgenus Ceratalaspis, and apparently related to the stictica group as defined in De Meyer (1998). However, the male has secondary sexual characters typical for the subgenus Pterandrus in that the mid tibia has distinct black feathering (the only other Ceratalaspis species with feathering, C. neostictica De Meyer, has orange feathering on the mid tibia). C. perseus lacks, however, the black band on abdominal tergite 3 which is found in most Pterandrus species. C. perseus shows some affinities with C. brucei Munro. The latter is only known from the female holotype of which the aculeus tip is damaged. However, the general shape of the aculeus in the latter is different from that of C. perseus. In brucei it is slender, and gradually narrowing towards the apex, while in C. perseus it is much broader (cf. Figures 14 and 15). Currently, we place this species in the subgenus Ceratalaspis. The species can be identified with the key provided by De Meyer (1998)  Diagnosis Arista with short rays; frons yellow, without silvery microtrichosity; postpronotal lobe without a spot; scutal pattern with distinct black markings but spots restricted; anepisternum yellow, with two anepisternal setae; wing bands with subapical band joined to discal band, posterior apical band absent; legs without feathering; abdomen without distinct spots, with grey band on tergites 2 and 4; apical part of aculeus with two large rounded indentations.

Description
Female. Head: antenna yellow. First flagellomere two times as long as pedicel. Arista with short rays, longest dorsal rays usually about as long as basal width of arista; if somewhat longer never twice as long as basal width. Frons yellow; with short scattered yellow setulae. Face yellowish white, below lower eye margin darker yellow. Genal setulae and genal seta yellow to orange, well developed. Thorax: postpronotal lobe yellow; without spot. Scutal pattern (Figure 4), ground colour yellow, with distinct black markings but the latter strongly reduced; prescutellar markings white, separate or merged; with yellow microtrichosity and pilosity. Scapular setae yelloworange. Scutellum yellow-white, basally without black spots, apically with three separate black spots, extending to basal half. Subscutellum yellow with three separate black spots. Anepisternum yellow; setulae yellow; two black anepisternal setae, occasionally with a third, less developed and dark orange, seta below two black ones.
Legs: yellow; setation yellow to orange. Fore femur with ventral setae orange.
Wing: pattern as in Figure 10; bands mainly yellowish with some brown patches. Anterior apical and discal bands not separate; subapical band joined to discal band; posterior apical band absent; crossvein R-M at basal third of discal medial cell.
Abdomen: yellow; pilosity mainly yellow. Tergites 2 and 4 with pale grey band occupying posterior third to half of tergite. Tergite 5 with longer black setulae along posterior margin.
Oviscape about as long as preabdomen. Aculeus eight times longer than wide, apical part ( Figure 16) with two large rounded indentations, tip with small depression.
Male. As female.

Etymology
Named after the type locality, Eala.

Remarks
C. ealensis keys out to C. hancocki De Meyer in De Meyer (1998). It shows some affinities with this species, but lacks a black spot on the postpronotal lobe, has lateral presutural black spots that do not reach the postpronotal lobe, and possesses an aculeus that lacks the large blade-like serrations. It seems to be more closely related to C. paradumeti De Meyer. The latter has a similar strongly reduced mesonotal pattern with restricted black spots, and an aculeus with rounded indentations. C. ealensis can be differentiated by the depression at the tip of the aculeus (rounded and without depression in C. paradumeti), and by the subapical band joining the discal band (subapical band isolated in C. paradumeti). More material of both species is, however, needed to unambiguously confirm the status of both taxa. This species can be identified with the key provided by De Meyer (1998)

Subgenus Ceratitis (Pardalaspis) Bezzi
A diagnostic description for the subgenus can be found in De Meyer (1996).

Description
Male: Head: antenna orange. First flagellomere two times as long as pedicel. Arista with short rays, longest dorsal rays usually about twice as long as basal width of arista; ventral rays shorter. Frons completey silvery, less dense in front of ocellar triangle, with short scattered darker setulae. Face orange. Genal setulae and genal seta black, well developed. Thorax: postpronotal lobe pale greyish white; without spot. Scutum ( Figure 5), ground colour greyish, with darker stripes and streaks but no distinct black spots except near intra-alar and prescutellar acrostichal setae; prescutellar markings white, separated by pale area or merged; with silver-grey microtrichosity and pilosity. Scapular setae black. Scutellum pale yellow, basally with two distinct black spots, apically with three separate black spots, extending to basal third. Subscutellum completely black. Anepisternum lower half yellow-brown, upper half white; two black anepisternal setae, setulae white, upper margin with black setulae.
Legs: pale yellowish brown; setation mixed black and white. Fore femur with ventral setae black.
Wing: pattern as in Figure 11; bands mainly brown. Anterior apical and discal bands not separate; subapical band separate; posterior apical band absent; crossvein R-M at, or just distal to, middle of discal medial cell.
Abdomen: greyish, lateral margin and tergite 5 largely with orange tinge; pilosity mixed black and white. Tergites with darker spots.
Female. As male except for the following characters. Frons yellow-brown, weakly silvery. Face white, lower margin yellow. Oviscape about as long as preabdomen. Aculeus at least 10 times longer than wide, tip ( Figure 17) bilobed and lateral margin slightly sinuate.

Etymology
Named after the late Ms Millicent Okumu, the chief laboratory technician for the USAID and USDA projects at ICIPE, who supervised the rearing of this and other new species described in this paper.

Remarks
This species is closely related to C. punctata (Wiedemann). The females can be easily differentiated by the shape of the aculeus tip which is simply pointed in C. punctata while bilobed in C. millicentae. The males are very similar, the only difference noted is in the extent of the frontal microtrichosity. In C. punctata the silvery microtrichosity is restricted to the lower third of the frons (not along entire length as first stated in De Meyer 1996, see correction in De Meyer 1999a, not extending beyond the frontal setae. In C. millicentae the microtrichosity extends all the way up to the medial vertical seta, but is less conspicuous in the upper median part in front of the ocellar triangle. Confusion in earlier identified material of C. punctata cannot be excluded and records, especially from eastern and southern Africa, should be reconfirmed. The species can be identified with the key provided by De Meyer (1996)