Phyllonorycter Hübner, 1822

Prins, Jurate De & Kawahara, Akito Y., 2012, Systematics, revisionary taxonomy, and biodiversity of Afrotropical Lithocolletinae (Lepidoptera: Gracillariidae), Zootaxa 3594 (1), pp. 1-283 : 56-60

Phyllonorycter Hübner, 1822

Phyllonorycter Hübner, 1822: 66–74 , 76–80.

Type species: Phalaena rajella Linnaeus, 1758 by subsequent designation by Walsingham 1908: 976. The type species was cited by Walsingham as rayella, an incorrect subsequent spelling ( Nye & Fletcher 1991: 240–241) (see also Bradley 1966: 218).

Phyllonorycter Hübner, 1806 : [2]. A work rejected for nomenclatural purposes by the International Commission on Zoological Nomenclature, 1926 (Opinion 97): 19–30. A nomenclaturally unavailable name.

Lithocolletis Hübner, 1825: 423 . Type species: Phalaena rajella Linnaeus, 1758 by subsequent designation by Walsingham 1908: 976. A junior objective synonym of Phyllonorycter Hübner, 1822 .

Eucestis Hübner, 1825: 423 . Type species: Tinea ulmifoliella Hübner, 1817 by subsequent designation by Hampson 1918: 387. A junior subjective synonym of Phyllonorycter Hübner, 1822 .

Eucesta Hübner, 1826: 67 . An incorrect subsequent spelling of Eucestis Hübner, 1825 . A nomenclaturally unavailable name.

Hirsuta Bruand, 1851: 50 . A nomenclaturally unavailable name as the genus was not described and the only included species was denoted by the undescribed manuscript name fritilella Tischer.

Lithocolletes Hübner, Dyar 1903: 549 . An incorrect subsequent spelling of Lithocolletis Hübner, 1825 . A nomenclaturally unavailable name.

Phyllorycter Walsingham (de Grey), 1914: 336 . An unjustified emendation of Phyllonorycter Hübner, 1822 .

Hirsuta Fletcher 1929: 110 . Type species: Elachista populifoliella Treitschke, 1833 , by original designation. By citing " Hirsuta, Bruand 1847 (non-descr.)" together with a type species, Fletcher unintentionally established Bruand's nomenclaturally unavailable name. A junior subjective synonym of Phyllonorycter Hübner, 1822 ( Nye & Fletcher 1991: 150).

Подрод [Subgenus] Asymmetrivalva Kuznetzov et Baryshnikova, 2004: 630–633 . Type species: Lithocolletis acerifoliella Zeller, 1839 , by original designation. A junior subjective synonym of Phyllonorycter Hübner, 1822 , synonimized by De Prins & De Prins 2005: 264. Asymmetrivalva was established to denote a subgenus of Phyllonorycter Hübner, 1822 .

The genus Phyllonorycter , known as Lithocolletis for many years, is one of the most successful lineages within Gracillariidae and accommodates 401 species world wide ( De Prins & De Prins 2012). The concept of the genus based on adult and larval morphology, as well as biology is well established and defined by many authors ( Braun 1908; Meyrick 1912b, 1927; Ely 1918; Kumata 1959, 1963, 1967, 1973, 1982, 1993; Patočka 1980; Kuznetzov 1978, 1979a –c, 1981; Emmet et al. 1985; Davis & Robinson 1998; Davis & Deschka 2001; Laštůvka & Laštůvka 2006; Liblikas et al. 2009; Bengtsson & Johansson, 2011).

Diagnosis. We define the genus Phyllonorycter as the assemblage of species-group taxa which fall into the clade Phyllonorycter ( Fig. 4 ). Morphologically Phyllonorycter is very similar to Cameraria in adult external features (except for the basal-edged white markings of the forewing), and wing venation (it is identical with Cameraria with the exception of C. fasciata ). Sternum VIII in males forms a characteristic large flap laying under valva like in Cameraria , Cremastobombycia , Hyloconis , Neolithocolletis , and Porphyrosela , but differently from Chrysaster , Leucanthiza , Macrosaccus , and Protolithocolletis where sternum VIII in males is not protruded. Phyllonorycter can be distinguished from other genera of Lithocolletinae including Cameraria and Macrosaccus by the simple conical tegumen without apical setae, which is an apomorphy for this genus. However, the preimaginal stages might propose more generic differential diagnostic characters in Lithocolletinae ( Kumata 1993; Davis & Deschka 2001). Phyllonorycter differs from other genera of Lithocolletinae by the single lateral seta (L1) on the mesothorax, metathorac and all abdominal segments in the last instar larva ( Kumata 1993; Davis & Deschka 2001). The loss of setae L2 and L3 on the body segments of the last instar larva is possibly an apomorphy that would suggest that the genus is an advanced group in the subfamily ( Kumata 1993; Davis & Deschka 2001). However, the larvae of the Afrotropical Phyllonorycter have not been examined yet.

Phyllonorycter also differs from Cameraria in COI sequence data. These two groups are separated by at least 10% sequence divergence for the taxa that were sampled in this study ( Fig. 3 ).

Diagnosis of Afrotropical Phyllonorycter . Afrotropical Phyllonorycter are generally small moths, having a forewing length of 2.7±0.5 mm with a few species falling outside this range. Forewing markings are very attractive, often showing white fasciate or strigulate patterns with silvery shine contrasting sharply with golden yellowish, ochreous or metallic brown ground colour coloration. The melanosparta group is the only one among Afrotropical Phyllonororycter without white markings on the forewing. The following distinctive features characterize Afrotropical Phyllonorycter :

Head: Vertex tufted with erected long piliform scales, being bicoloured at their tips and frequently varying in shading and length from frons to occiput [the rough appearance of vertex is due to the neck plumes which are long and project forwards above the vertex between antennae ( Vári 1961)]; frons covered with appressed smooth scales, frequently white, sometimes with golden sheen or strong metallic gloss, or mottled with brownish scales ( Figs 132, 133 ); labial palpus moderate, porrect, filiform, drooping, straight, terminal palpomere about 1.5× longer than second palpomere, sharply pointed; maxillary palpus minute, porrect, proboscis well developed, about twice as long as labial palpus; antenna simple, about as long as forewing, not clearly ringed; scape short, thickened ( Figs 7, 8 ), bearing pecten of different length.

Thorax: Forewing with black or blackish brown scales arranged in a row(s) edge fasciate or strigulate whitish markings basally or mixed (except melanosparta group, where white markings absent); fringe long, particularly near tornus and dorsum, reaching width of wing in forewing and ca. 3× width of wing in hindwing. Descaled forewing lanceolate, slender, and pointed. Venation with 8 veins ( Figs 16, 17 ), discal cell closed, ca. 0.75 of wing length, apical part with 5 separate veins: R3, R4, R5, M1 and Cu1; CuP indistinct for entire length, A1 strong, to margin. Hindwing greyish, dirty white, lustrous, silvery, lanceolate, maximum width 0.11 that of length, venation reduced, Sc very short, Rs very long running almost to apex, basal 2/3 of M1 indistinct, parallel to Rs, distal 1/3 of M1 ends at 3/4 of dorsum, Cu1 strong, ends about 2/5 of dorsum ( Ely, 1918; Vári, 1961; Kumata, 1993). Legs with darker rings, spots or markings aiding identification ( Figs 126–131 ); fore- and mid-legs slender, midtibia bears scales which project alongside a pair of short tibial spurs; hindtibia thickened, with long fine loose hairs and medial and apical spurs, hindtarsus smooth, slender and ca.1.5× as long as tibia.

Male genitalia. Sternum VIII developed, flap-like, extended. Tegumen simple, without apical setae, in grewiaecola group with lateral appendages, valvae symmetrical, whereas valvae of many Phyllonorycter species of northern hemisphere asymmetrical, much elongated, mostly bar shaped with filament-like setae, in P. loxozona valvae sinuate, sometimes with shorter or longer spines ( P. achilleus , P. jabalshamsi ), in the rhynchosiae group ventral surfaces of valva bears flap-like projection(s). Transtilla in most of the species complete, quadrate, or with thicked lateral lobes or protruded projections on cephalic margin. In one species of the Afrotropical Phyllonorycter — P. achilleus —transtilla incomplete with long dorsal basal valval projections present which abute each other but not joining. In P. grewiaecola incomplete transtilla connected with sclerotized fultura superior. Vinculum mostly crescent or U-shaped, thickly sclerotized. Saccus varies from tiny pointed tip on vinculum in P. chionopa ( Fig. 182 ) to extremely long slender projection, almost as long as abdomen in P. obandai and P. farensis ( Figs 254 , 257 ). In most Afrotropical Phyllonorycter , saccus slender, straight, of medium length, projecting anteriad. In leucaspis group, saccus characteristically sinuoid, turned at basal part ( Figs 224 , 227 , 230 , 231, 234 ). Anellus weakly sclerotized, except P. grewiaecola where anellus with sclerotized fultura superior bearing two sinuate horns posteriorly. Aedoeagus usually straight or slightly curved, but mostly slender, cylindrical, slightly enlarged at coecum. In chionopa species group, girth of coecum and vesica strikingly differing ( Fig. 182 ). Length of aedoeagus varying from short in P. brachylaenae and P. melhaniae ( Figs 207, 210 , 253 ) to extremely long, almost as long as abdomen in P. farensis ( Fig. 258 ). Apical part of aedoeagus usually with slender rod-like cornuti, sometimes tiny barbs on vesica.

Female genitalia. Papillae anales slightly compressed laterally, attached to each other laterally, usually as long as wide, covered with long sparsely set setae. Stronger or weaker sclerotized basal bar may be present or absent or partly present. In most of the species groups, a needle-like projection streches from basal part of papillae anales to posterior ca. 1/3 of segment VIII in the encaeria , gato , hibiscina , hibiscola , lemarchandi , melhaniae , mida , rhynchosiae , silvicola and umukarus groups. Segment VIII is well connected with segment VII. Posterior apophyses usually slender in most of the informal species groups, and with slightly enlarged basal halves in the leucaspis and jabalshamsi groups. In most cases anterior apophyses initiate at basal plate in segment VIII, except in the melanosparta and achilleus groups. In the melanosparta group anterior apophyses are replaced by caudal sterigmatic appendages and in the achilleus group the anterior apophyses are short, straight and initiate at the joint of segment VIII and VII. In the Afrotropical Phyllonorycter ostium bursae opens either at the joint of segments VII and VIII like in the chionopa , encaeria , grewiella , jabalshamsi , lemarchandi , leucaspis , melhaniae , ruwenzori and silvicola groups or in segment VII like in the remaining informal species groups. Sterigmatic sclerotizations on segment VII in Afrotropical Phyllonorycter are either not developed like in the achilleus , chionopa , grewiella , jabalshamsi , leucaspis , and ruwenzori informal species groups or are well developed as volcano, bumerang, or triangular cuticle fold or arc-shaped suture like in the encaeria , gato , hibiscina , hibiscola , mida , rhynchosiae , silvicola and umukarus species groups. In the mida group lamella post-vaginalis possesses very complicatedshaped sclerotizations covered with sharp large spines ( Fig. 338 ); in the melanosparta group sterigma formed as a plate-shaped appendage with the extended caudal processes serving as anterior apophyses ( Figs 334, 335 ); the similar sterigmatic appendages present in the grewiaecola group, in the obandai group sterigma is shaped as moderate M-shaped suture with opening of ostium bursae at its depression ( Fig. 340 ); in the melhaniae group sterigma is partly attached tube-shaped sclerotization ( Fig. 336 ). The anterior margin of segment VII can be encircled by a heavily sclerotized ring-like structure in P. hibiscola . Ductus bursae varies from very short in the leucaspis group to very long in the obandai group. Corpus bursae is variable in size and shape, in most species with the corpus bursae and ductus bursae clearly differentiated, although in some Afrotropical Phyllonorycter species (the encaeria , obandai , didymopa , melanosparta groups) the junction of the ductus and the corpus bursae is gradual and indistinct. The corpus bursae often bears signa areas and a sclerotized signa, but often lacks a distinct signum. There can be one or two signa areas on corpus bursae; sclerotized signum can have the shape of a star with extended rays (in the mida and rhynchosiae groups), a band consisting of sharp barbs arranged in 3–4 rows (in the hibiscina group), a long sharp needle (in the umukarus , grewiaecola groups) or other shapes.

Biology. The life history of Phyllonorycter is described by Braun 1908; Needham et al. 1928; Vári 1961; Kumata 1963, 1993; Davis 1987; Landry & Wagner 1995; Davis & Robinson 1998; Davis & Deschka 2001). Phyllonorycter species are believed to possess three early sap-feeding and two subsequent tissue feeding instars ( Davis & Deschka 2001), while Kumata (1993) proposes that the last instar might be also the sixth. The range of host plant choice in Phyllonorycter is very broad; no less than 32 families of plants serve as foodplants (for a full account of Phyllonorycter host plants, see De Prins & De Prins 2012). As with most of the Afrotropical microlepidoptera, the pattern on biology, host plant preferences and host plant specialization obtained from limited rearing records of Phyllonorycter is still rather obscure. Four families of plants: Asteraceae , Fabaceae , Malvaceae , and Verbenaceae were reported as hosts of Afrotropical Phyllonorycter ( Viette 1951; Vári 1961; Kroon 1999; Dall’Asta et al. 2001; De Prins & De Prins 2005, 2012). The rearing record of P. lemarchandi on Solanaceae ( Paulian & Viette 1955) needs confirmation. In this study we add Convolvulaceae , Lamiaceae , and Rosaceae to the list of host-plant families for Afrotropical Phyllonorycter . All informal species groups belonging to Afrotropical Phyllonorycter exhibit high host plant specialization at least at the generic level. In the hibiscina , lemarchandi , leucaspis , melanosparta , rhynchosiae groups restricted oligophagy is documented. Species-related host-plant specifity needs a special study for the grewiella and hibiscina species groups, since an unusually high number of hosts for one species has been recorded, 7 for P. grewiella , and 5 for P. hibiscina . Furthermore, even the entire informal hibiscina species group is characterized by a great span of host plants,—eight, belonging to three plant families: Asteraceae , Convolvulaceae , and Malvaceae , an unusual biological character for other Phyllonorycter species groups. Twenty seven Afrotropical Phyllonorycter species (41.5%) have rearing records reporting 38 plant species as hosts. Twenty-three species of Phyllonorycter in the Afrotropical region feed on Malvaceae .

The great majority of reared Afrotropical Phyllonorycter make underside mines, however one species— P. grewiella —mines the upper side of Grewia spp. leaves. The prognathous sap-feeding instar initiates a slender, serpentine subepidermal gallery which is enlarged by the third instar to form a flat blotch. The hypognathous tissue feeding larva (fourth and fifth instars) do not enlarge the area of the mine but contracts the edges of the mine closer together by silk, thus finally constructing a tentiform blotch mine ( Hering 1951; Kumata 1963, Emmet et al. 1985; Davis & Deschka 2001). The parenchymal tissues within the mine are consumed almost wholly, or in spots, larval frass of tissue feeding instars becomes granullar and is typically collected at one end of the mine. Pupation occurs within the mine cavity either without a cocoon or it is enclosed by a slender, ellipsoid cocoon of which colour, size, shape and position within the mine is usually constant and species-linked ( Emmet 1985 et al.).

Distribution. Afrotropical Phyllonorycter species are largely distributed in southern Africa (Botswana, Namibia, South Africa, Zimbabwe) and in eastern Africa (Kenya, Tanzania, Uganda) and the Arabian Peninsula (southern Oman, southern Yemen. Several species are recorded from Madagascar and the Mascarene Islands, and four species from the regions bordering Central Africa (northern Cameroon, Nigeria and Rwanda). Until the present study, no Phyllonorycter species has been recorded from coastal western Africa and the high inlands of Central Africa. The greatest concentration of Phyllonorycter diversity is documented from eastern Africa (23 species). The distributional pattern we report might be biased by extensive collecting efforts in eastern Africa by many microlepidopterists (e.g., Vári 1961; Triberti 2004), but we believe that the distributional pattern concentrated in the eastcoast of Africa is the general pattern. Only hibiscina and leucaspis groups showed pan- (trans-) Afrotropical distribution. The remaining groups of Afrotropical Phyllonorycter were recorded from restricted woodland and savannah biomes. Fabaceae and especially Malvaceae , the most common host plant families for Phyllonorycter , play a critical role in woodland communities. Savannah communities are dominated by grasses ( Poaceae ) as they are everywhere, but the woody component is strongly influenced by the presence of Fabaceae and Malvaceae species (J. De Prins, pers. observation).

Afrotropical Phyllonorycter species are not restricted to particular elevations; they vary from almost sea level ( P. mida ) up to almost 3000 m ( P. acutulus ) in Kenya. However, the greatest species richness can be found between 0–500 m and 1500–2000 m elevations. Nevertheless, the mida and grewiella species groups showed a large range in elevation preference: P. mida was recorded from sea level in Kenya to 1300 m in Yemen, P. grewiella ranges from 80 and 500 m in eastern Kenya up to 1300 m in Namibia and 1800 m in Yemen. Most species are represented by too few individuals to draw any meaningful conclusions.

In general, Phyllonorycter is particularly species-rich in the northern hemisphere, and poorly represented in the southern hemisphere. Thirteen species assigned to Phyllonorycter are known from the Neotropics and only four are known from the Australian region. Here we present 48 species of Phyllonorycter from the Afrotropical region grouped into 22 informal species groups. Some of these groups can be distinguished superficially by forewing pattern and forewing length: i) the melanosparta group shows only black markings on forewing without white markings; ii) first and second fasciae on forewing in the grewiaecola group sinuate and are linearly bordered by black scales apically and basally; iii) the ground colour in chionopa group is bright yellow with big, broad, semiround or rectangular shiny white patches; iv) wings in the lemarchandi group have metallic lustre and small white dots similar to those in Porphyrosela , however, the specimens of the lemarchandi group are slightly bigger than those of Porphyrosela ; v) the grewiella species group possesses large strigulae on dorsal margin of forewing whereas distinctive strigulae on the costal margin are absent; vi) the obandai , agassizi and achilleus groups show a unique and recognizable pattern combined from strigulae of different length and orientation to apex with the length of basal streak (see appropriate descriptions). However, wing pattern is not a reliable character for species group differentiation in Afrotropical Phyllonorycter , since the wing pattern of the informal species groups can be confused with that of other Lithocolletinae genera. For instance, the wing pattern of P. lemanchandi is almost indistinguishable from many Porphyrosela species , the melanosparta group can be confused with Cameraria hexalobina and Cremastobombycia kipepeo due to the absence of white markings on forewings, the grewiaecola group can be confused with the landryi and sokoke groups of Cameraria since these groups possess sinuate fasciae etc. Only male and female genitalia provide the most reliable features for discriminating the species groups and the species. Valva provides some of the best characters to discriminate the informal groups of Afrotropical Phyllonorycter , like proportial valval length and width in comparison with other male genital structures, valval size, shape of cucullus, location of ventral appendices, presence of distal spines and setation. In female genitalia, the shape of sterigma, location and position of ostium bursae, form of corpus bursae, the configuration and shape of signa are diagnostic for species or species groups. Based on genitalia, nine distinct multi-species groups plus 15 single-species groups are recognized. Morphologically different genitalia demand that each of the 15 single-species groups is assigned to its own group.

Relationships to other genera. Phylogenetically, Phyllonorycter is most closely related to Cremastobombycia ( Fig. 4 ). These two genera together form a well-supported clade (BP = 100%, PP = 1.0) with the nuclear genes sampled for this study. Phyllonorycter and Cremastobombycia share a similar preimaginal morphology, host plant, and male genital morphology. Specifically, larvae of both genera: i) are cylindrical, whereas larvae of Cameraria are flat; ii) construct strongly contracted mainly abaxial tentiform mines, whereas the larva of Cameraria builds mainly an adaxial blotch-shaped mine; iii) feed on Asteraceae and Verbenaceae , whereas Asteraceae and Verbenaceae plants are not utilized by Cameraria ( Table 3 ). The transtilla in male genitalia is complete in both genera Phyllonorycter and Cremastobombycia whereas it is incomplete in Cameraria .

Figures

FIGURE 3. Maximum-likelihood tree of Cameraria + Phyllonorycter based on 40 taxa, for the COI data set alone. African species are genetically distinct, as shown by the long internal branch lengths among taxa boxed in grey. ML bootstrap values show above branches. Scale bar = 0.08 substitutions/site.

FIGURE 4. Molecular phylogeny of Gracillariidae, showing relationships among the Lithocolletinae species sampled for eight nuclear genes in this study. ML bootstrap values are shown above branches, Bayesian posterior probabilities shown below. African taxa are indicated with an asterisk. Scale bar = 0.09 substitutions/site.

FIGURE 230. Male genitalia. Scale bar 100 µm. 230, Phyllonorycter ololua, holotype, genitalia prep. MRAC/KMMA 00369, in RMCA.

FIGURES 132–133. Diagnostic characters in frons between two related Afrotropical Lithocolletinae species. 132, Phyllonorycter melhaniae, frons shiny white with irrorated dispersed dark brown scales. 133, Phyllonorycter pavoniae, frons whitish with a very faint pale ochreous suffusion.

FIGURES 5–10. Adult head morphology of the Afrotropical Lithocolletinae genera. 5, Neolithocolletis nsengai, descaled head prep. MRAC/KMMA 00537. 6, Cameraria landryi, descaled head prep. MRAC/KMMA 00529. 7, Phyllonorycter grewiaecola, descaled head prep. MRAC/KMMA 00532. 8, Phyllonorycter hibiscina, descaled head prep. MRAC/KMMA 00531. 9, Cremastobombycia solidaginis, descaled head prep. De Prins 3791. 10, Porphyrosela teramni, descaled head prep. MRAC/KMMA 00433.

FIGURES 11–20. Wing morphology of the Afrotropical Lithocolletinae genera. 11, Hyloconis luki, wing venation prep. MRAC/KMMA 00454. 12, Neolithocolletis mayumbe, wing venation prep. MRAC/KMMA 00535. 13, Neolithocolletis nsengai, wing venation prep. MRAC/KMMA 00427. 14, Cameraria perodeaui, wing venation prep. MRAC/KMMA 00453. 15, Cameraria torridella, wing venation prep. MRAC/KMMA 00490. 16, Phyllonorycter grewiaecola, wing venation prep. MRAC/KMMA 00528. 17, Phyllonorycter melanosparta, wing venation prep. MRAC/KMMA 00497. 18, Cremastobombycia kipepeo, wing venation prep. MRAC/KMMA 00494. 19, Cremastobombycia morogorene, wing venation prep. MRAC/KMMA 00667. 20, Porphyrosela teramni, wing venation prep. MRAC/KMMA 00434.

FIGURES 123–131. Diagnostic characters in hind legs between congeneric Afrotropical Lithocolletinae species. 123, Cameraria fara, hind tarsus white with a brownish spot subbasally and two blackish rings: narrow medially and broad subapically, tarsomere V white. 124, Cameraria landryi, hind tarsus white with two blackish small spots basally and two blackish rings: narrow medially and broad subapically, tarsomere V white with dark fuscous tip. 125, Cameraria varii, hind tarsus white with a blackish small spot basally and three blackish rings of median width, tarsomere V white. 126, Phyllonorycter encaeria, hind tarsomeres I–III with faint fuscous subapical patches. 127, Phyllonorycter lantanae, hind tarsomere I with subbasal and subapical dark fuscous patches, tarsomere II with dark fuscous apical half, tarsomere III with a dark fuscous basal half. 128, Phyllonorycter anchistea, hind tarsomere I dotted. 129, Phyllonorycter melanosparta, hind tarsomere I not dotted. 130, Phyllonorycter hibiscina, hind tarsomere I white with a dark fuscous apex. 131, Phyllonorycter pavoniae, hind tarsomere I with a faint subapical ochreous patch.

FIGURES 182–184. Male genitalia. Scale bar 100 µm. 182–184, Phyllonorycter chionopa, the drawings are made by Willy De Prins after the additional description and the drawing of Triberti (2004: 81; fig. 6: A–C). 182, ventral view. 183, tip of valva. 184, aedoeagus.

FIGURES 254–256. Male genitalia. Scale bar 100 µm. Phyllonorycter obandai, holotype, genitalia prep. MRAC/KMMA 00251, in RMCA. 254, ventral view. 255, sternum VIII. 256, aedoeagus.

FIGURES 257–258. Male genitalia. Scale bar 100 µm. Phyllonorycter farensis, holotype, genitalia prep. MRAC/KMMA 00279, in RMCA. 257, ventral view. 258, aedoeagus.

FIGURES 224–226. Male genitalia. Scale bar 100 µm. 224–226, Phyllonorycter albertinus, holotype, genitalia prep. De Prins 3504, in BMNH. 224, ventral view. 225, sternum VIII. 226, aedoeagus.

FIGURES 227–229. Male genitalia. Scale bar 100 µm. Phyllonorycter leucaspis, paratype, genitalia prep. MRAC/KMMA 00424, in RMCA. 227, ventral view. 228, sternum VIII. 229, aedoeagus.

FIGURES 231–236. Male genitalia. Scale bar 100 µm. 231–233, Phyllonorycter ruizivorus, holotype, genitalia prep. De Prins 3723, in BMNH. 231, ventral view. 232, sternum VIII. 233, aedoeagus. 234–236, Phyllonorycter trochetellus, holotype, genitalia prep. De Prins 3720, in BMNH. 234, ventral view. 235, sternum VIII. 236, aedoeagus.

FIGURES 206–210. Male genitalia. Scale bar 100 µm. 206–210, Phyllonorycter brachylaenae. 206–207, the drawing is made by Willy De Prins after the genitalia prep. Vári 7523, in TMSA. 206, lateral view. 207, aedoeagus. 208–210, genitalia prep. De Prins 3709, in ZMHB. 208, ventral view. 209, sternum VIII. 210, aedoeagus.

FIGURES 252–253. Male genitalia. Scale bar 100 µm. 252–253, Phyllonorycter melhaniae, holotype, the drawing is made by Willy De Prins after the genitalia prep. Vári 7495, in TMSA. 252, lateral view. 253, aedoeagus.

FIGURES 338–339. Female genitalia. Scale bar 200 µm. 338, Phyllonorycter mida, paratype, genitalia prep. 3715, in ZMUC. 339, Phyllonorycter tsavensis, paratype, genitalia prep. Vári 7501, in TMSA.

FIGURES 334–335. Female genitalia. Scale bar 200 µm. 334, Phyllonorycter anchistea, paratype, the drawing is made by Willy De Prins after the genitalia prep. Vári 7139, in TMSA. 335, Phyllonorycter melanosparta, genitalia prep. MRAC/ KMMA 00374, in RMCA.

FIGURES 340–342. Female genitalia. Scale bar 200 µm. 340–341, Phyllonorycter obandai, genitalia prep. De Prins 3503, in BMNH. 340, segments VII–X, ventral view. 341, corpus bursae. 342, Phyllonorycter farensis, paratype, genitalia prep. MRAC/ KMMA 00282, in RMCA.

FIGURES 336–337. Female genitalia. Scale bar 200 µm. 336, Phyllonorycter melhaniae, paratype, the drawing is made by Willy De Prins after the genitalia prep. Vári 7496, in TMSA. 337, Phyllonorycter rongensis, holotype, genitalia prep. MRAC/ KMMA 00383, in RMCA.