Discovery of an unknown diversity of Leucinodes species damaging Solanaceae fruits in sub-Saharan Africa and moving in trade (Insecta, Lepidoptera, Pyraloidea)

Abstract The larvae of the Old World genera Leucinodes Guenée, 1854 and Sceliodes Guenée, 1854 are internal feeders in the fruits of Solanaceae, causing economic damage to cultivated plants like Solanum melongena and Solanum aethiopicum. In sub-Saharan Africa five nominal species of Leucinodes and one of Sceliodes occur. One of these species, the eggplant fruit and shoot borer Leucinodes orbonalis Guenée, 1854, is regarded as regularly intercepted from Africa and Asia in Europe, North and South America and is therefore a quarantine pest on these continents. We investigate the taxonomy of African Leucinodes and Sceliodes based on morphological characters in wing pattern, genitalia and larvae, as well as mitochondrial DNA, providing these data for identification of all life stages. The results suggest that both genera are congeneric, with Sceliodes syn. n. established as junior subjective synonym of Leucinodes. Leucinodes orbonalis is described from Asia and none of the samples investigated from Africa belong to this species. Instead, sub-Saharan Africa harbours a complex of eight endemic Leucinodes species. Among the former nominal species of Leucinodes (and Sceliodes) from Africa, only Leucinodes laisalis (Walker, 1859), comb. n. (Sceliodes) is confirmed, with Leucinodes translucidalis Gaede, 1917, syn. n. as a junior subjective synonym. The other African Leucinodes species were unknown to science and are described as new: Leucinodes africensis sp. n., Leucinodes ethiopica sp. n., Leucinodes kenyensis sp. n., Leucinodes malawiensis sp. n., Leucinodes pseudorbonalis sp. n., Leucinodes rimavallis sp. n. and Leucinodes ugandensis sp. n. An identification key based on male genitalia is provided for the African Leucinodes species. Most imports of Leucinodes specimens from Africa into Europe refer to Leucinodes africensis, which has been frequently imported with fruits during the last 50 years. In contrast, Leucinodes laisalis has been much less frequently recorded, and Leucinodes pseudorbonalis as well as Leucinodes rimavallis only very recently in fruit imports from Uganda. Accordingly, interceptions of Leucinodes from Africa into other continents will need to be re-investigated for their species identity and will likely require, at least in parts, revisions of the quarantine regulations. The following African taxa are excluded from Leucinodes: Hyperanalyta Strand, 1918, syn. rev. as revised synonym of Analyta Lederer, 1863; Analyta apicalis (Hampson, 1896), comb. n. (Leucinodes); Lygropia aureomarginalis (Gaede, 1916), comb. n. (Leucinodes); Syllepte hemichionalis Mabille, 1900, comb. rev., Syllepte hemichionalis idalis Viette, 1958, comb. rev. and Syllepte vagans (Tutt, 1890), comb. n. (Aphytoceros). Deanolis iriocapna (Meyrick, 1938), comb. n. from Indonesia is originally described and misplaced in Sceliodes, and Leucinodes cordalis (Doubleday, 1843), comb. n. (Margaritia) from New Zealand, Leucinodes raondry (Viette, 1981), comb. n. (Daraba) from Madagascar as well as Leucinodes grisealis (Kenrick, 1912), comb. n. (Sceliodes) from New Guinea are transferred from Sceliodes to Leucinodes. While Leucinodes is now revised from Africa, it still needs further revision in Asia.


Introduction
Leucinodes orbonalis Guenée, 1854, the eggplant fruit and shoot borer, is a species of moth that was first described from specimens from India and Java (Guenée 1854). According to current knowledge, it is widely distributed in tropical and subtropical Asia (CABI 2012a) and sub-Saharan Africa (Walker 1859, Frempong 1979, CABI 2012a. The larvae are pests of Solanaceae, especially Solanum melongena L. (aubergine, eggplant or brinjal) fruits and stems where they feed internally. Their infestation can substantially reduce yields from aubergine cultivation, and yield losses of more than 65% have been recorded from Asia (EPPO 2008).
The larvae are commonly moved in international trade with plants and fruits, as their internal feeding and the resulting damage may not be visible during pre-export inspections. Thus, Leucinodes orbonalis is a quarantine pest of concern to a number of countries outside its native range. This includes the member countries of the European and Mediterranean Plant Protection Organisation (EPPO), where it was recommended as an addition to the alert list of pests in 2008 (EPPO 2008), and in 2012 transferred to the A1 list of pests recommended for statutory regulation (EPPO 2012(EPPO , 2013. It is an A1 pest for several South American countries including Uruguay and Argentina (COSAVE 2006) and has repeatedly been intercepted in the USA (Whittle and Ferguson 1987, Solis 1999, 2006. In England and Wales, the Plant Health and Seeds Inspectorate (PHSI) regularly intercept Leucinodes Guenée, 1854 larvae inside aubergines from South Asia and West Africa (The Food and Environment Research Agency (Fera), unpublished data).
Due to the economic impact of Leucinodes orbonalis, the development of a genetically modified eggplant was initiated in 2005 in India by introducing a crystal protein gene (Cry1Ac) from the bacterium Bacillus thuringiensis Berliner, 1915 (Bt) into the plant (Sood 2012). The insecticidal effect of the crystal proteins makes eggplant less susceptible to infestations by larvae of Leucinodes orbonalis. After field trials and approval for commercial cultivation from government scientists in 2009, a moratorium on the commercialisation of Bt brinjal was imposed due to public concerns on food safety (Pandey 2010).
Until recently, all pyraloid larvae damaging Solanaceae fruits in Asia and Africa and intercepted from imports to Europe have been regarded as two species, Leucinodes orbonalis and Sceliodes laisalis (Walker, 1859). Hayden et al. (2013) and Gilligan and Passoa (2014) pointed out that Leucinodes orbonalis is restricted to Asia and that there are "three species in the L. orbonalis complex in Africa that are not conspecific with the Asian species. Our investigations reveal that an even largernumber of Leucinodes species are intercepted from Africa. This points to the question of the identity of the intercepted species as well as on further four species of Leucinodes known from Africa, L. aureomarginalis Gaede, 1916, L. hemichionalis (Mabille, 1900), L. translucidalis Gaede, 1917 and L. vagans (Tutt, 1890) (Nuss et al. 2003(Nuss et al. -2014. Besides Leucinodes, there is the similar genus Sceliodes Guenée, 1854, of which Sceliodes cordalis (Doubleday, 1843) and S. laisalis (Walker, 1859) are also pests on solanaceous crops in Australia (Davis 1964), New Zealand (Martin 2010) and Africa (CABI 2012b).
Here we taxonomically revise Leucinodes and Sceliodes and their species from continental sub-Saharan Africa, in order to delimit species and to allow their proper identification. Larvae were sourced through quarantine interceptions of eggplant fruit from Africa and Asia at several ports of entry in England. Most larvae were studied alive and subsequently reared to adults in order to confirm the species identity. In large containments a few larvae were preserved by boiling them in water for 30-90 seconds, then transferred to 70% ethanol. After 3-5 days, the ethanol was replaced to limit dilution from body contents. Reared adults were killed soon after emergence, with cyanide, ammonia or by freezing at -20 °C for a minimum of 2 hours. Field-collected adults were attracted by artificial light and killed with cyanide. All adult specimens were subsequently pinned. Genital dissections of thoroughly dried specimens were performed according to Robinson (1976). Setal nomenclature follows Hinton (1946). The chaetotaxic descriptions and the setal map focus on those microscopic setae visible at 60 × magnification.

Specimens
DNA was extracted using either the NucleoSpin tissue kit (Macherey-Nagel) according to the manufacturer's instructions or using the Chelex-100 resin based method (Boonham et al. 2002). DNA extraction with the NucleoSpin tissue kit was performed following the procedure of Knölke et al. (2005), extracting DNA from the abdomen of adult specimens and subsequent dissection of the genitalia from the macerated abdomen. Extracted abdomina were stored in 70% ethanol until genitalia were dissected. For the Chelex-100 resin based method single legs or wings were removed from dried, pinned specimens using fine forceps and placed in individual 0.6 ml Eppendorf tubes. Briefly, 100 µl moleculargrade water was added to the tissue sample and ground using a micropestle. 100 µl of a 50% w/v chelex resin:water slurry was added, the sample heated to 95 °C for 5 minutes, centrifuged for 5 minutes and the supernatant transferred and stored at -20 °C prior to use.
We apply the morphospecies concept to our study. The DNA Barcode is used as additional source of information and as an identification tool for all developmental stages of African Leucinodes species. The Solanaceae species names mentioned in this study refer to their former context and do not necessarily correspond to the revised Solanum taxonomy by Knapp et al. (2013).
Label data of studied specimens were compiled in order to generate a distribution map. Geographical coordinates, if not given on the label, were obtained via Google Earth, Version 5.2.1.1588 and subsequently plotted on a map using DIVA-GIS, Version 7.2.3 (Hijmans et al. 2004).

Data resources
The data underpinning the analyses reported in this paper are deposited in the Dryad Data Repository at doi: 10.5061/dryad.kk0n9.

Leucinodes Guenée, 1854
Leucinodes Guenée, 1854. Type species: Leucinodes orbonalis Guenée, 1854 = Sceliodes Guenée, 1854, syn. n. Type species: Sceliodes mucidalis Guenée, 1854= Daraba Walker, 1859(synonymised by Hampson, 1899. Type species: Daraba idmonealis Walker, 1859 = Eretria Snellen, 1880 (synonymised by Hampson, 1899;Shaffer et al. 1996: junior homonym of Eretria Robineau-Desvoidy, 1863). Type species: Eretria obsistalis Snellen, 1880= Leuctinodes South, 1897 Diagnosis. Leucinodes is characterized by a forewing pattern which includes a brown base, a white antemedian line which is distally brown edged; a median area that is ochreous or brown from the costa to the middle of wing, and red-brown from the middle of wing towards the dorsum; below the apex is a black-brown half moon-shaped patch (missing in L. malawiensis sp. n.), edged by a thin white postmedian line and a white line at the margin of wing. The hindwings are white with inconspicuous pattern elements. Leucinodes females with only one frenular bristle in the hindwing, female labial palps with elongated 3 rd meron, male genitalia with identical location of the fibula-sacculus process-complex (process lacking in L. cordalis (Doubleday, 1843), L. laisalis and L. malawiensis), female genitalia with fine granular sclerotization of ductus bursae (in most species), antrum with thickened mesocuticula, presence of lateral antrum pockets. Larvae are internal feeders in Solanaceae.
Redescription of adults. Head. Frons conically bulged (Figs 11-12) to flat; labial palps porrect, brownish, 1 st meron on ventral side with forward-directed tuft, 3rd meron in males half as long as 2 nd meron, longer in females (Figs 11-12); maxillary palps minute or missing; haustellum well developed; eyes large, hemispherical; ocelli present; antennae ciliate, pedicel white to brown, flagellum light brown, cilia in males longer than basal antennal radius (except in L. malawiensis), in females shorter than antennal radius; vertex with whitish to brown scales at the collar and brown scales directed forward; chaetosemata absent.
Wings. Forewing white translucent, light brown or orange-to grey-brown, basal area light to dark brown, delimited by white and dark brown double line or in species with brown forewing ground colour by dark brown antemedian line; median area with pale to dark brown, sometimes very faint proximal discoidal stigma (absent in L. malawiensis); distal discoidal stigma pale to dark brown, reaching from costa to forewing centre; central dorsum with prominent orange to dark brown, broadly L-shaped or triangular spot connected or disconnected with distal discoidal stigma; postmedian line sinuate, faint and grey to grey-brown, white edged, with prominent subcostal bulge; apex brown to grey-brown coloured (absent in L. malawiensis), with slim strip of white at outer margin; margin dotted at veins, with large dots at apex and M 3 ; fringe white to pale brown with dark interruption at apex and at M 3 (absent in L. malawiensis). Hindwing in both sexes with one frenular bristle, ground colour whitish, middle of wing with one or two spots, often faint; postmedian line inconspicuous, bent towards spot at middle of wing; area below apex suffused by pale brown to grey; margin dotted at end of veins, with large dot at end of M 3 .
Abdomen. First segment whitish, remainder light-, orange-or dark brown to grey. Male genitalia. Uncus neck constricted, head circular, with dorsal agglomeration of thick setae; narrow transtilla arms with central notch, in L. ethiopica with dorsad spike on each arm; vinculum saccus round to V-shaped, short to more or less elongated, with or without keeled tip; juxta oval, subulate, short rhombical or tongueshaped, with semicircular base; valvae elongate triangular, tapering posteriorly, costa and posterioventral margin loosely covered with long setae; fibula (fi in Fig. 13) arising at central part of mesal wall of valva or near costa; sacculus (sa in Fig. 13) large, elongate oval, with distal sclerotized process (sp in Fig. 13), often in close association with fibula, process absent in L. laisalis, L. malawiensis; ventral margin of distal valvae with or without granulated area (ga in Fig. 13); phallus simple, with variously shaped sclerites at posterior apodeme, vesica with or without cornuti.
Female genitalia. Corpus bursae ovoid, membranous, without signa; ductus bursae membranous with delicate granulation, partly reaching into posterior corpus bursae; antrum short to long, slim to broader than ductus bursae, anterior part sometimes coiled, mesocuticula thickened (strongly stained with Chlorazol Black) and exocuticula (inner layer) partly sclerotized; ostium bursae with lateral membranous pockets, with or without oval sclerites; both apophyses pairs simple, apophyses anteriores normally stronger developed than posterior apophyses, with or without broadened central portion.
Immature stages. Larva. Last instar larvae with pink dorsal integument, intersegmental areas cream or light pink, the ventral integument cream; strength of the colouration very variable, pink colour on majority of abdominal segments often interrupted laterally by a transverse cream line; head, prothoracic and anal shields mid brown with variable black markings; early instar larvae white or cream with brown pinacula and black head, prothoracic and anal shields. In older larvae the dorsal integument turns beige, then increasingly deeper pink as the moults progress, head and prothoracic shield brown; pinacula pale brown and prominent against the integument in all instars. The chaetotaxy of the thorax and first nine abdominal segments of the last instar is illustrated in Fig. 35. The relative size of pinacula and positions of setae are very variable intraspecifically. The head is mid to light brown, with variable black markings around ocelli and at genal angle; relative positions of cranial setae very unstable in the specimens examined. The prothoracic shield is light to dark brown with pale median sulcus and two variable dark markings: one along part of the posterior margin, strongest medially, and the other mediolaterally; usually additional darker spots bordering the median sulcus and extending laterally, spots very variable in extent and position; prothoracic L pinaculum crescent shaped with variable posterior extension, L setae anterior to the spiracle, usually vertically aligned; microscopic seta MV3 clearly visible in most specimens, can be almost as prominent as the V seta; MV3 setae share a mid-ventral pinaculum or are on separate pinacula; meso-and metathorax with clearly visible dorsal and subdorsal microscopic pinacula at 60 × magnification; three ventral microscopic setae less prominent, with MV3 usually being the least evident, these with or without small pinacula. Many larvae are asymmetrical in this feature, with a pinaculum on one side, and seta only on the other. On the abdomen, there is one SV seta on segment 1, three SV setae on a single pinaculum on segment 2; microscopic seta MV3 visible on both segments 1 and 2, not prominent; microscopic setae on segments 3-8 mostly not visible at 60 × magnification; prolegs with crochets in a mesopenellipse; anal shield often lighter brown than pinacula, usually darker pigmented in anterior half.
Pupa.  Yellow to pale brown, lightly sclerotized, developing adult clearly visible as development proceeds; two distinct, raised hood-like structures dorsal to spiracles on abdominal segments 2 and 3 (Figs 42, 44); four pairs of long hooked setae ventral to cremaster; cocoon stout leathery, made of silk, firmly attached to the substrate.
Remarks. Sceliodes and Leucinodes have traditionally been distinguished by their forewing ground colour, which is predominantly orange-brown to greyish-brown in Sceliodes and white translucent in Leucinodes. The newly discovered Leucinodes ethi-opica sp. n. is intermediate in this character whereas all other wing pattern elements are homologous among Leucinodes and Sceliodes species. Study of the genitalia showed that cornuti are present in Sceliodes species, but are absent in Leucinodes, including L. ethiopica. The female genitalia contain oval to semicircular sclerites in the lateral antrum pockets of L. ethiopica, African Sceliodes and S. cordalis, the type-species of Sceliodes, which is distributed in Australia and New Zealand. Thus, there is a continuous variation between Leucinodes and Sceliodes and we here synonymise Sceliodes syn. n. with Leucinodes. As Leucinodes and Sceliodes have been published on the same date and in the same work, we here give precedence to Leucinodes as it is the better known of the two names, acting as first reviser according to ICZN 24.2.2. Guenée, 1854 Figs 1, 11-13, 23, 31, 35-37 Leucinodes orbonalis Guenée, 1854: 223. Diagnosis. Wing pattern indistinguishable from those of L. africensis sp. n., L. rimavallis sp. n., L. pseudorbonalis sp. n., L. kenyensis sp. n. and "Leucinodes spp.", but distinguished from L. malawiensis by the absence of the forewing basal transversal streak and the presence of the apical half moon-shaped patch, and from L. laisalis, L. ethiopica and L. ugandensis sp. n. by the predominantly white forewing ground colour. Frons usually more strongly bulged than in other Leucinodes species, but L. pseudorbonalis can be very similar in this feature. In male genitalia distinguishable by: dorsal margin of valval sacculus concave; apical sclerotized sacculus process elongated cone-shaped and crossing with the similar-sized fibula (as in L. pseudorbonalis); juxta slender, tapering (similar in L. africensis and L. rimavallis); saccus of vinculum short, less prominent. Female genitalia: antrum only slightly bulged, exocuticula without sclerotized strip.

Redescription of adults. Head.
As for the genus, with frons strongly bulged .
Thorax. As for the genus, with dorsal side brown. Wings. Forewing length ♂ 8.5-10.5 m, ♀ 9.5-12.0 m; forewing ground colour white, basal area light-to dark brown, delimited by dark brown to grey antemedial line; median area with pale brown, faint proximal discoidal stigma; distal discoidal stigma pale brown, reaching from costa to forewing centre; central dorsum with prominent orange to dark brown L-shaped or triangular spot leading to forewing centre and often meeting with distal discoidal stigma; antemedial line sinuate, more or less distinct, but with prominent subcostal bulge; subapical half of termen with half moon-shaped brown to grey-grown spot; marginal line dotted; fringe and marginal line darkened at the tips of the half moon-shaped spot; hindwing ground colour white, internal area white, with discoidal spot, basicostally often with auxiliary spot; medial line sinuate, distal half approaching the discoidal spot, then turning towards dorsum; external area pale brown to gray; marginal line dotted.
Female genitalia. As for the genus, apart from: anterior antrum with spoonshaped posteriad indentation, flanked on either side by a sclerotized portion (as in Fig.  31); sternite 8 anterior edge arched (ae in Fig. 31), with a sclerite process leading in each of the lateral pockets; both apophysis pairs simple, slightly curved. Immature stages. Larva. MSD1 and MSD2 of meso-and metathorax usually on a shared pinaculum, earlier instars frequently have the MSD setae on separate pinacula on one or both segments; dorsal abdominal pinacula show apparent differentiation between West-and East-Asian populations: in live western specimens (e.g., from Pakistan), the abdominal D1 pinacula usually have an unpigmented cream coloured area near to their anteriomedian margin. This unpigmented area may be contiguous with the unmelanized cuticle surrounding the pinaculum or be surrounded by the melanized cuticle of the pinaculum (illustrated on A3-6 in Fig.  35); this cream white, unpigmented area may darken in preserved specimens, and in pre-pupae may be ringed in black. Eastern specimens (e.g., from Thailand) often have dark spots on at least some of the D1 pinacula (illustrated on A2 in Fig. 35); geographically intermediate populations (e.g., Sri Lanka) often show an intermediate form with black spots on occasional pinacula, and any unpigmented area is usually ringed with black pigmentation (illustrated on A7 in Fig. 35). East-Asian populations usually have mesally triordinal crochets, whereas the crochets of West-Asian populations are mesally biordinal. Pupa. Cremaster forms a variable shelf-like, sub-rectangular structure, much wider than long, usually with distinct distal corners and median notch; dorsal surface spinulose, with additional small but distinct spines which are variable in extent, form and number; cocoon of dark brown silk, may be white or beige when newly spun.
Distribution. India, Indonesia: Java (Guenée 1854), Sri Lanka (Walker 1859, Moore 1885), Myanmar (Burma), Andaman Islands (Pagenstecher 1900), Bangladesh, Brunei, Cambodia, China, Japan, Laos, Malaysia, Nepal, Pakistan, Philippines, Singapore, Taiwan, Thailand, Vietnam (CABI 2012a), Australia (Shaffer et al. 1996); imported to Great Britain, the Netherlands (pers. comm. M. van der Straten), Denmark (pers. comm. O. Karsholt) and the U.S.A. (Boateng et al. 2005, Solis 2006). Remarks. Leucinodes orbonalis has previously been reported in Europe from the Netherlands (van der Gaag et al. 2005) and Great Britain (Agassiz 1983;Higgott 2009) as well as from the following African countries due to misidentification: South Africa (Walker 1859; Pagenstecher 1900; Kemal and Koçak 2007); Kenya (Poulton 1916); Ghana (for example, Frempong and Buahin 1977, Frempong 1979, Duodu 1986, Horna et al. 2007); Lesotho, Zambia and Zimbabwe (Kopij 2005); Burundi, Cameroon, Congo, Democratic Republic of the Congo, Ethiopia, Malawi, Mozam-bique, Nigeria, Rwanda, Sao Tome and Principe, Sierra Leone, Somalia, Tanzania and Uganda (CABI 2012a). We have not found a single specimen from Africa belonging to this species and therefore postulate that L. orbonalis does not occur in Africa. Diagnosis. The frons is less strongly bulged than in L. orbonalis. In wing pattern this species is indistinguishable from those of L. orbonalis, L. rimavallis, L. pseudorbonalis, L. kenyensis and "Leucinodes spp.", but distinguished from L. malawiensis by the absence of the forewing basal transversal streak and the presence of the apical half moon-shaped patch, and from L. laisalis, L. ethiopica and L. ugandensis by the predominantly white forewing ground colour. In male genitalia it is distinguished by: the long ventrad fibula (as in L. rimavallis, short and triangular in L. malawiensis, broad and stout in L. laisalis); the elongate, straight or hook-shaped, sometimes branching distal sacculus process projecting towards the valva apex (similar in L. rimavallis); the apically thin, subulate juxta (similar in L. rimavallis); the prominent oval saw blade-shaped sclerotization of the posterior phallus apodeme (as in L. rimavallis); it is distinguished from L. rimavallis by the longer, more curved fibula with a slender base, the elongate distal sacculus process, which spans more than half the distance fibula base-valva apex, is straight or hook-shaped and sometimes exhibits a side branch, and the pointed valva apex (rounded in L. rimavallis). Female genitalia resemble those of L. pseudorbonalis in having a swollen antrum, but they lack the posterior constriction of the ostium bursae.

Leucinodes africensis
Description of adults. Head. As for the genus, with frons moderately bulged, base of each meron of labial palps with white scales.
Thorax. As for the genus, with dorsal side brown.
Immature stages. Larva. Final-instar larvae of L. africensis and L. orbonalis cannot be definitively separated. In final instars of live specimens of L. africensis, the majority of the abdominal D1 pinacula have a dark pigmented spot on the anteriomedian margin (illustrated on A2 in Fig. 35), although in the occasional pinaculum this is replaced by an unpigmented area, which can be contiguous with the unmelanized integument surrounding the pinaculum or separated from it by the melanized cuticle of the pinaculum; crochets are mesally triordinal, as in the East-Asian populations of L. orbonalis. Pupa. length ca. 8.5 m; no consistent features separate the pupae of L. orbonalis and L. africensis.
Etymology. Latinized africensis, derived from the continent of Africa from where the type material originates and referring to the widespread distribution of this species on the African continent. Remarks. This species is very similar to L. rimavallis, but both COI Barcoding data and constant morphological differences in genitalia separate the two species. Diagnosis. Frons is moderately bulged; in wing pattern this species is indistinguishable from L. orbonalis, L. africensis, L. pseudorbonalis, L. kenyensis and "Leucinodes spp.", but distinguished from L. malawiensis by the absence of the forewing basal transversal streak and the presence of the apical half moon-shaped patch, and from L. laisalis, L. ethiopica and L. ugandensis by the predominantly white forewing ground colour. In male genitalia it is distinguished by: the long ventrad fibula (as in L. africensis, short and triangular in L. malawiensis, broad and stout in L. laisalis); the granulate, hook-shaped distal sacculus process (similar in L. africensis, process smooth in L. kenyensis); the apically thin, subulate juxta (as in L. africensis, broad subulate in L. orbonalis); the prominent oval saw blade-shaped sclerotization of the posterior phallus apodeme (as in L. africensis); distinguished from L. africensis by the shorter, straight to slightly curved fibula with a broader base, the shorter, always hook-shaped distal sacculus process, and the rounded valva apex (pointed in L. africensis). Female genitalia have slender apophyses anteriores (as in L. orbonalis), the central antrum tube with a short, strongly sclerotized section (long strip in L. africensis and L. pseudorbonalis), and the anteriolateral edges of sternite 8 with triangular processes extending into the lateral antrum pockets (as in L. kenyensis).
Abdomen. First segment whitish, remainder light to dark brown. Male genitalia. As in L. africensis, but with the fibula short, more triangular and robust, straight or slightly curved; distal sacculus process short and always bent apically; valva apex rounded or stout.
Female genitalia. As for the genus, apart from: anterior antrum with short sclerotized section, central posterior antrum with diffuse weak sclerotization; sternite 8 on each side with anteriad triangular process extending into the lateral antrum pockets.
Etymology. From latin rima for 'rift' and vallis for 'valley', referring to the African Rift Valley, the main distributional area of this species (as far as known).
Distribution. Burundi, Eastern and Southern Democratic Republic of the Congo, Kenya, Rwanda, South Africa, Uganda (import).  Diagnosis. Frons is moderately to strongly bulged; Wing pattern indistinguishable from those of L. orbonalis, L. africensis, L. rimavallis, L. kenyensis and "Leucinodes spp.", but distinguished from L. malawiensis by the absence of the forewing basal transversal streak and the presence of the apical half moon-shaped patch, and from L. laisalis, L. ethiopica and L. ugandensis by the predominantly white forewing ground colour. In male genitalia the prominent dorsad fibula and the fibula-like sacculus process are roughly of same size and run parallel or cross each other (as in L. orbonalis, fibula and fibula-like sacculus process very small in L. ugandensis). Very similar to male genitalia of L. orbonalis, but valva tips slimmer and more acute, juxta with larger hemicircular base, elongated saccus tip and more prominent oval sclerite at posterior phallus apodeme. In female genitalia discriminated by the globular, posteriorly somewhat constricted antrum with a longitudinal, sclerotized exocuticular strip bearing transversal ridges (as in L. africensis).
Description of adults. Head. As for the genus, with frons moderately to strongly bulged.
Female genitalia. As for the genus, apart from: anterior antrum shortly coiled in coronal plane, with the exoculticle exhibiting a longitudinal sclerotized strip bearing transverse ridges (Fig. 33); sternite 8 intruding into the posteriorly somewhat constricted antrum, giving it a globular appearence.
Immature stages. Larva. Only one specimen available, examined live. Black spots present on dorsal pinacula; in the final instar, MSD1 and MSD2 on a shared pinaculum on both meso-and metathorax; crochets mesally triordinal.
Etymology. Composition of greek pseud(o) 'false' and orbonalis, meaning 'false orbonalis', referring to the similarities in external and male genital characters with L. orbonalis.
Remarks. We found this species among material from Senegal, Uganda and Angola, leaving a considerable distribution gap in Central Africa.
Recently, several interceptions of larvae in solanaceous fruits imported from Uganda have been recorded in England (own observation) and the Netherlands (Marja van der Straten, pers. comm.). L. pseudorbonalis is one of the three African Leucinodes species intercepted at European ports of entry. Diagnosis. Frons less strongly bulged than in L. orbonalis. Wing pattern indistinguishable from those of L. orbonalis, L. africensis, L. rimavallis, L. pseudorbonalis and Leucinodes spp., but distinguished from L. malawiensis by the absence of the forewing basal transversal streak and the presence of the apical half moon-shaped patch, and from L. laisalis, L. ethiopica and L. ugandensis by the predominantly white forewing ground colour. Distinguished in male genitalia from all other Leucinodes species except "Leucinodes spp." (see below) by the prominent distal sacculus process arching anteriodorsally above the shorter, conical fibula. Distinguished from Leucinodes spp. by the more or less bulged subapical portion of costa (straight in Leucinodes spp.) and the weakly sclerotized basal section of the sacculus process (between sacculus and distal hook-shaped process) which is as wide as base of hook-shaped process (strongly sclerotized and narrower than base of hook-shaped process in Leucinodes spp.). Female genitalia resemble those of L. rimavallis in having an anteriad triangular process on each side of sternite 8 extending into the lateral antrum pockets, but the exocuticle of the anterior antrum forms a sclerotized tube (short sclerotized section in L. rimavallis).

Leucinodes kenyensis
Description of adults. Head. As for the genus, with frons slightly bulged. Thorax. As for the genus, with dorsal side brown. Wings. Forewing length ♂ 9.0 mm, ♀ 9.0 mm; wing pattern as in L. orbonalis. Abdomen. First segment whitish, remainder light to dark brown.

Male genitalia.
As for the genus, apart from: saccus bent posteriodorsad (not an artifact of embedding); juxta oval to rectangular; valvae roughly triangular; costa subapically more or less bulged; ventral valva apex flipped over, covered with small tubercles; ventral valva edge smoothly rounded at sacculus; base of sacculus process broad, weakly sclerotized, leading over to a large, strongly sclerotized hook which encompasses the fibula dorsally; fibula conical, slightly curved, its base somewhat constricted, projecting dorsad; phallus similar to L. orbonalis, posteriodorsal apodeme with short dentate sclerite (sometimes indistinct), vesica with area of minute teeth.
Female genitalia. As for the genus, apart from: anterior antrum with tubular sclerotized exocuticle; sclerotized wall at antero-ventral edge of the ostium bursae which at rest closes the ostium bursae against abdominal segment 8. This wall is not melanized and can be stained with chlorazol black. It is delimited dorso-laterally by sclerotized and melanized lobes arising from the anterior edge of segment 8 just ventral of the apophyses anteriores. Anterior to the sclerotized wall there is a small, melanized colliculum.
Etymology. The species is named after Kenya, the only country from where it is confidently recorded so far.
Distribution. So far only known from Kenya. The record from Zimbabwe needs confirmation by investigation of female specimens and molecular analysis.

Remarks.
There are further male specimens with indistinctive wing pattern and very similar genitalia, but DNA Barcode data suggest that among them are at least two further species. For more information, see under Leucinodes spp. Diagnosis. Distinguished from the other Leucinodes species by the dark, straightframed forewing base and the absence of the subapical mark of the forewing termen. The male genitalia are similar to those of Leucinodes ethiopica and L. ugandensis, but are distinct in the long spinoid process of the posterior phallus apodeme.

Leucinodes malawiensis
Description of adults. Head. As for the genus, with frons flat. Thorax. As for the genus, with dorsal side brown. Wings. Forewing length ♂ 8.5 mm; forewing base dark brown, its outer margin a straight diagonal line from the costa to the maculation of the central hind margin, a triangular patch leading lateroposteriad from the costa with the costal half reddishbrown and the central tip white; outer median area with a faint brownish transverse Abdomen. Pale ochreous, first abdominal segment white, terminal segments brown. Male genitalia. As for the genus, apart from: saccus elongated, U-shaped; juxta short, oval, twice as broad as long; valvae slender, tapering towards the dorsad bent apex; fibula small, triangular, oriented ventrad; sacculus process absent; phallus with a spinoid posteriad sclerotization emerging from the ventroposterior apodeme.
Female genitalia. Unknown. Etymology. Latinized malawiensis from the country Malawi where the holotype originates.

Leucinodes laisalis
Redescription of adults. Head. Frons slightly bulged; labial palps upturned, greyish to brown, first meron on ventral side with forward-directed tuft, third meron in males half as long as second meron, considerably longer in females; maxillary palps minute; haustellum well developed; eyes large, hemispherical; ocelli present; antennae ciliate, cilia considerably longer in males; vertex with creamy white scales; chaetosemata absent.
Thorax. Dorsal side brown with greyish and dark brown scales mixed in; ventral side grey to whitish; legs predominantly whitish or grey, epiphysis present; tibial spurs 0, 2, 4 with outer spur 2/3 the length of inner spur.
Wings. Forewing length 7.0-11.5 mm, the females being somewhat larger; both sexes with one frenular bristle; forewing ground colour orange-to grey-brown, with the general Sceliodes wing pattern.
Abdomen. First segment whitish, remainder light-brown; older specimens often with darkened abdomen due to degeneration of abdominal fat body.
Male genitalia. As for the genus, apart from: vinculum saccus conspicuously elongated anteriad; juxta usually with small notch at median base; valvae emerging in narrow angle from vinculum; phallus with keeled coecum, posteriodorsal apodeme with slim, fingerlike, slightly curved and well sclerotized process, vesica with a short line of tiny cornuti.
Female genitalia. As for the genus, apart from: antrum long, tubular, anterior end coiled; apophysis pairs straight, apophyses anteriores with somewhat broader muscle attachment area at posterior quarter.
Distribution. In Africa known from Côte d'Ivoire, Ghana, Kenya, Morocco, Nigeria, Senegal, South Africa, Tanzania (own observations). Externally, L. laisalis is similar to L. ugandensis (see below), therefore literature records from other African countries than those listed here need verification. In Europe recorded from Spain, Portugal (Speidel 1996, Huertas Dionisio 2000own observations) and Great Britain (own observations). The records from Great Britain certainly refer to interceptions and it is assumed that those from the Iberian Penninsula also do not refer to a native occurrence. Huertas Dionisio (2000) recorded the species from Solanum linnaeanum, a species native to southern Africa. Immature stages. Larva. Generally very similar to L. orbonalis and L. africensis. On the metathorax the MSD setae are usually on separate pinacula, while the mesothoracic MSD setae are usually on the same pinaculum. The abdominal D1 pinacula are often smaller than those of L. orbonalis and L. africensis, and lack dark spots or unpigmented areas (see Ogunwolu (1978) for a detailed larval description and chaetotaxy). Pupa. length ca. 8.5 mm; distal margins of cremaster usually evenly rounded, without distinct corners; spinulation of cremaster's dorsal surface a little coarser than dorsal spinulation on abdominal segment 9 dorsal surface, no distinct small spines as in L. orbonalis or L. africensis; cocoon of beige coloured silk that does not darken significantly over time.
Remarks. We found a significant DNA Barcode difference of 2.4-2.8% uncorrected p-distance between the single South African specimen and the Kenyan and Ghanan/Nigerian Barcode clusters ( Fig. 47; see also section 'DNA Barcoding' below). These differences in the DNA Barcode are not reflected in a divergent morphology of the clusters.
The record of L. laisalis from Belgium by Nyst (2004) is most probably a misidentification, since the illustrated imago resembles much more the whitish Leucinodes species. Apart from that, there is a European record of L. laisalis from Spain. Additionally, it is frequently intercepted with the import of solanaceous fruits in Great Britain.
Despite repeated search in the collection of the ZMHB, original material of Leucinodes translucidalis Gaede, 1917 from Tanzania, Tendaguru, could not be traced. According to the original description, this taxon can be regarded as conspecific with L. laisalis due to all details given in the original description. Especially the white triangle at the anterior line, another white triangle, though often somewhat inconspicuous, at the middle of costa, and the dark brown area below apex support the conspecifity with L. laisalis. Diagnosis. This species' forewing colour has more ochreous than the whitish species of Leucinodes but less orange-brown to greyish than in L. laisalis and L. ugandensis. From L. ugandensis and L. laisalis it can be distinguished by the genitalia: in the male genitalia the transtilla arms each bear a dorsad spine; in the female genitalia the ductus bursae lacks the fine granular sclerotization, the antrum is strongly sclerotized, tubular and widest at its anterior end, and the oval ostial sclerites in the lateral antrum pockets are larger.
Wings. Forewing length 6.0-8.0 mm. Forewing mixed ochreous and white, an oblique dark ochreous fascia from above dorsum reaching halfway across wing, a blackish crescent before ochreous subterminal line, black dots along termen. Hindwing white, a small black discal spot, a faint irregular dark subterminal line, ochreous suffusion in outer part of wing in middle and towards apex.
Abdomen. Pale ochreous, first abdominal segment white. Male genitalia. As for the genus, apart from: transtilla with short central notch, each transtilla arm with a dorsad spine; vinculum saccus with rounded tip; juxta oval to rectangular, apex with a short central notch; apex of valvae dorsally curved; small, spine-like dorsad fibula emerging from the central inner side of valva; sacculus with a fibula-like spiny dorsad process emerging ventrodistally of the fibula; posteriodorsal phallus apodeme with semicircular dentate sclerite.
Female genitalia. As for the genus, apart from: ductus bursae with fine longitudinal ripples; antrum tubular, with broader anterior end; apophyses anteriores at posterior half laterally broadened for muscle attachment.
Etymology. Latinized ethiopica from the country Ethiopia where the holotype originates.

Leucinodes ugandensis
Diagnosis. Distinguished from the whitish species of Leucinodes and L. ethiopica by the predominantly brown forewing ground colour with minor white patches. Distinguished from L. laisalis in the male genitalia: less strongly sclerotized, valvae triangular, fibula small, tooth-like, a similarly shaped distal sacculus process present, saccus process shorter, phallus much shorter, dorsoposterior apodeme without slim, finger-like process.
Thorax. Pale fuscous, metathorax blackish fuscous. Wings. Forewing length 6.5-11.5 mm. Forewing pale fuscous, an oblique brown partial fascia at halfway with white markings on either side, an orange triangle on dorsum beyond halfway, apex deep brown, separated by a whitish line. Hindwing whitish, fuscous suffused near margin in middle and towards apex, a faint subterminal line in costal part of wing.
Abdomen. Abdomen first segment whitish, remainder orange-brown. Male genitalia. As for the genus, apart from: small, hooked dorsad fibula emerging from the ventrocentral inner side of valva; distal sacculus with dorsad ridge forming a bulge, followed by a spiny, curved terminal process overlapping with the fibula; phallus vesica with several small spiny cornuti.
Female genitalia. As for the genus, apart from: diffusely sclerotized exocuticula reaching into posterior ductus bursae; lateral antrum pockets rather small.
Etymology. Latinized ugandensis from the country Uganda where the type specimens originate.

Remarks.
No COI Barcode sequences were obtained for this species.
Note. The following material contains male specimens from southern Africa with indistinctive wing pattern and very similar male genitalia to Leucinodes kenyensis. According to morphology, we could not separate these specimens from L. kenyensis. In contrast to the morphological data, two of these specimens, the single records from Namibia and Swaziland, have distinctive DNA Barcodes from L. kenyensis as well as from each other (Fig. 47). For the remaining specimens listed below, we did not obtain DNA Barcodes. In spite of the absence of further specimens for comparison, especially females, and the lack of convincing morphological differences, we are not going to describe these possibly distinct species here. This complex needs further study.
Remarks. This species was described in Daraba Walker, 1859, a genus that has previously been synonymised with Sceliodes Guenée, 1854 by Hampson (1899). The type specimen of Leucinodes raondry (Viette, 1981), comb. n. from Madagascar could not be traced at MNHN. According to the original description and the illustration of the species given therein (Viette 1981), this species agrees with the diagnostic wing pattern elements of Leucinodes, and we therefore consider it as correctly placed in this genus. It differs from L. laisalis in the larger size, the more ochreous grey tone and the reduced half moon-shaped subapical patch of the forewings (Viette 1981). None of the species described here as new have the prominent dark subapical patch in the hindwings of L. raondry, so that conspecifity with any of them can be ruled out. (Kenrick, 1912) | 1928-34." (BMNH). Female unknown.

Leucinodes grisealis
Remarks. Due to the synonymisation of Sceliodes, this species is provisionally transferred to Leucinodes, as no proper generic placement has been found. Compared to Leucinodes, several differences can be found in wing pattern of grisealis Kenrick, 1912: In the fore wing, the postmedian line is originating in the apex, and its median protrusion is closely approaching the termen; the half moon-shaped patch below apex is protruding beyond M 3 ; in the hind wing, the postmedian line is originating closer to the apex and is running closer to the termen.

Syllepte Hübner, 1823
Syllepte Hübner, 1823. Type species: Syllepte incomptalis Hübner, 1823. Syllepte hemichionalis (Mabille, 1900), comb. rev. (Sylepta [sic]). Type locality: Madagascar, baie d'Antongil = Syllepte hemichionalis idalis Viette, 1958, comb. rev. (Syllepta [sic]). Type locality: Comoros, Mohéli. Syllepte vagans (Tutt, 1890), comb. n. (Aphytoceros). Type locality: Great Britain, Chepstow. = Aphytoceros longipalpis Warren, 1892(synonymised by South, 1897 Remarks. Syllepte hemichionalis and S. vagans are similar to each other in wing pattern elements, the presence of two frenular bristles in the female hindwings and features of male genitalia (see Suppl. material 2I), but not so to Leucinodes. Their wing pattern differs from Leucinodes in the following: forewing without the L-shaped or triangular spot at central dorsum of Leucinodes and without the half moon-shaped subapical patch; postmedian line broad, straight, but with long loop before dorsum, reaching to central part of forewing; dorsum with broad brown line leading from brown antemedian area to distal end of postmedian line; in the hindwing they have a comma-or V-shaped distal discoidal sigma, light brown with dark brown border. This homologous wing pattern is also found in Syllepte chalybifascia Hampson, 1896 from India and S. dottoalis Schaus, 1927 from the Philippines, and we consider them congeneric with each other as well as with S. hemichionalis and S. vagans. The genus Syllepte comprises 193 species (Nuss et al. 2003(Nuss et al. -2014, is probably polyphyletic and in need for taxonomic revision.

Deanolis Snellen, 1899
Type species: Deanolis sublimbalis Snellen, 1899 Deanolis iriocapna (Meyrick, 1938) Remarks. The wing pattern of iriocapna Meyrick, 1938 exhibits none of the features found in Leucinodes. Instead, the fore wings are pale yellow, with a yellowish costa, a dark spot in both outer edges of the cell, and a reddish undulating margin along the termen. The hind wings are of the same pale yellow ground colour, and the anterior half of the termen exhibits a similar margin as found in the fore wings. This wing pattern is common to the genus Deanolis Snellen, 1899, where this species is correctly placed (pers. comm. James E. Hayden).
Identification key for African Leucinodes species based on male genitalia:

DNA Barcoding
The Neighbor Joining (NJ) analysis based on uncorrected p-distances ( Fig. 47) groups taxa which are excluded from Leucinodes in this study outside of the Leucinodes + Neoleucinodes group. Deanolis iriocapna, Leucinodes ethiopica and L. ugandensis have not been barcoded and are therefore not included in the NJ tree. The Leucinodes + Neoleucinodes group forms a polytomy comprising Neoleucinodes, L. malawiensis, L. laisalis and a group containing the remaining Leucinodes species. Within this last group, 11 barcode clusters are revealed. Leucinodes laisalis clusters in two Barcode groups: One group containing all specimens imported with fruits from Kenya, Ghana and Nigeria, and a second group comprising a single South African specimen. This single specimen shows high uncorr-p distances of 2.4-2.8% to the other L. laisalis specimens.
L. orbonalis clusters in two groups, separated by 1.1-1.8% uncorr-p distance. Within-subcluster distances are 0-0.5% for the smaller and 0-0.9% for the larger of the two subclusters.

Discussion
The investigation of the Leucinodes species from sub-Saharan Africa reveals a new picture on the taxonomy of the Solanaceae fruit feeders. Instead of one widely distributed species in Asia and Africa, there is an endemic species complex in Africa and L. orbonalis is restricted to Asia, corroborating the statement by Hayden et al. (2013). Among the African species, L. africensis and L. laisalis are widely distributed on the continent, whereas L. ethiopica, L. ugandensis, L. rimavallis and L. kenyensis seem to have a rather restricted range. L. malawiensis is only known from one specimen from Malawi. These data may indicate different, though partly overlapping patterns of distribution as seen in Fig. 48, but this is still an incomplete dataset, resulting in fragmentary distribution patterns for L. pseudorbonalis and L. rimavallis. The possibility of transport by man also has to be taken into consideration. Food plant data are only available for L. africensis, L. laisalis, L. pseudorbonalis, L. rimavallis and L. kenyensis (Table 2). While L. laisalis is known to feed on a variety of cultivated and non-cultivated Solanaceae, the other four species are known from cultivated Solanaceae only. This may rather refer to a bias of investigation, which so far has concentrated on fruits of economic importance. In nature, the Leucinodes species may have different host plants to which they were originally adapted. Those wild hosts may also have certain areas of origin. Such a scenario has to be expected, considering the fact that several species of Leucinodes have evolved in Africa. Some of these species might be restricted to their wild host plants and unable to switch to other, cultivated Solanaceae species. In contrast, species able to adapt to cultivated Solanaceae might be transported with fruits in trade, resulting in a larger range within Africa.
After movement of larvae with fruits in African trade, they may be more frequently intercepted by trade with those fruits into other continents. Most imports of Leucinodes specimens from Africa into Europe refer to Leucinodes africensis, which has been frequently imported with fruits during the last 50 years. In contrast, L. laisalis has been much less frequently recorded, and L. pseudorbonalis as well as L. rimavallis only very recently in fruit imports from Uganda. Since our investigations show that L. orbonalis does not occur in Africa, interceptions of Leucinodes from Africa into other continents will need to be re-investigated for their species identity and will likely require, at least in parts, revisions of the quarantine regulations. Furthermore, if Leucinodes species are transported in trade, it has to be considered that species of Leucinodes and related South American genera, e.g. Euleucinodes Capps, 1948, Neoleucinodes and Proleucinodes, might also become introduced from one to another of the southern continents. Analysis of the COI gene of the Leucinodes species demonstrated that interspecific differences allow the use of the marker as a DNA barcode for species identification. However, for L. kenyensis and "Leucinodes spp.", we found little morphological differences but two distinct barcode species within Leucinodes spp. Moreover, we observed high intraspecific divergence in Leucinodes laisalis with one specimen from South Africa exhibiting a COI distance of 2.8%.
Outside Africa, the taxonomy and phylogeny of Leucinodes requires further research. The characteristic forewing pattern elements of the Old World Leucinodes is also found in the New World genera Euleucinodes, Neoleucinodes and Proleucinodes, all described by Capps (1948). In male genitalia, Leucinodes is distinguished from these three genera by the absence of cornuti in the phallus, from Euleucinodes and Proleucinodes also by the presence of a fibula, and further from Euleucinodes by the dorsal location of the uncus spines. In female genitalia, Leucinodes is distinguished from Proleucinodes by the presence of lateral antrum pockets (condition unknown in Euleucinodes), but cannot be clearly distinguished from Neoleucinodes, in which antrum pockets can be absent (e.g., N. elegantalis (Guenée, 1854)) or present (Neoleucinodes prophetica (Dyar, 1914), N. torvis Capps, 1948), and pocket sclerites can be absent (e.g., N. elegantalis) or present (N. imperialis (Guenée, 1854), N. torvis). Whether these differences refer to a typological classification or justify the maintenance of the current generic classification needs to be investigated by phylogenetic analysis.
For Austral-Asia, there remain twelve nominal Leucinodes species (Nuss et al. 2003(Nuss et al. -2014. Other than L. orbonalis, at least some of these species are certainly misplaced in Leucinodes, e.g. Leucinodes labefactalis Swinhoe, 1904 from Borneo and Leucinodes perlucidalis Caradja, 1933 from China. Therefore, the Asian Leucinodes are in need of taxonomic revision. This also points to the question whether all Leucinodes samples intercepted from Asian exports refer to L. orbonalis, or whether there are several Leucinodes species of economic importance in Asia as well (Hayden et al. 2013, Chang et al. 2014, Gilligan and Passoa 2014. Our work contributes to the identification of African Leucinodes species, based on adult characters and on DNA Barcodes, by which also the immature stages can be efficiently distinguished. This may help to systematically survey the continent for distribution of species, in order to discover their wild host plants and their movements in trade. At this stage of knowledge it wouldn't be a surprise to discover additional, still unknown species. The results of our completely revised taxonomy of African Leucinodes suggests that a revision of the EPPO A1 List of pests recommended for regulation as quarantine pests (EPPO 2013) will be necessary. During the period from 2004 to 2007, 121 interceptions of Solanum fruits infested by Leucinodes were recorded by several EPPO member countries. The majority (94 consignments) originated from Thailand, Ghana accounted for 18 infested consignments (EPPO 2008). L. orbonalis is also ranked as quarantine species important for the USA, but specimens intercepted from Africa cannot be L. orbonalis, as stated by Hayden et al. (2013) and Gilligan and Passoa (2014) and as shown by our results. The USA has recorded 1745 interceptions from Ghana to the USA between 1985 and 2004 (Boateng et al. 2005).

Conclusion
A careful revision of Leucinodes in sub-Saharan Africa resulted in the new synonymy of the genus Sceliodes syn. n., the revised synonymy of Hyperanalyta with Analyta, the transfer of four species to Leucinodes, the description of seven new Leucinodes species, the new synonymy of one species, the omission of the type species L. orbonalis from the African list and the generic transfer of five species found to be misplaced in Leucinodes. Of the eight species recognized from Africa now, at least four are frequently intercepted among imports of solanaceous fruits at European ports of entry. We provide the DNA Barcode for these four and two additional African Leucinodes species, allowing the identification of all life stages of these species.
Our work shows that typological concepts of taxonomy based on superficial similarity were still the state of the art in the genus Leucinodes. The discovery of a complex of highly similar species demonstrates that traditional morphological methods and DNA Barcoding are helpful tools to detect species diversity and to improve their classification based on sound arguments.
Similar revisionary work remains to be done for Austral-Asian Leucinodes. A phylogenetic study including Leucinodes and the New World Euleucinodes, Neoleucinodes and Proleucinodes is needed in order to test the monophyly of these genera (Hayden and Mally, in prep.). This study was supported by a SYNTHESYS grant for the visit of RM at the BMNH London. Fera acknowledges the support of Defra Plant Health Policy. Material reared at Fera was under the authority of Plant Health Licence number 33173/220099-2 and its predecessors.