BIOTAXONOMY OF TEPHRITOIDEA Isr. J. Entomol. Vol. 35-36, 2005/6, pp. 367-422

A Revision of the Gymnosagena group of genera (Diptera: : )

Amnon Freidberg1 and Bernhard Merz2

ABSTRACT The three Afrotropical genera of Tephritidae comprising the Gymnosagena group are revised: Elgonina Munro (including E. fuscana Munro, E. refulgens Munro, and seven new species: E. dimorphica, E. flavicornis, E. inexpectata, E. infusctata, E. pollinosa, E. splendida and E. yaromi); Gymnosagena Munro (including G. unicornuta Munro and four new species: G. campiglossina, G. kakamega, G. longicauda and G. nyikaensis), and Marriottella Munro (including M. exquisita Munro and M. sepsoides n. sp.). A lectotype is designated for Gymnosagena unicornuta. A cladistic analysis of the group supports the current classification.

1Department of Zoology, The George S. Wise Faculty of Life Sciences, Tel Aviv University, Tel Aviv 69978, Israel. E-mail: [email protected]; 2Departement d’Entomologie, Museum d’Histoire Naturelle, C.P. 6434, 1211 Geneve, Switzerland. E-mail: [email protected] BIOTAXONOMY OF TEPHRITOIDEA

INTRODUCTION The three small, -feeding, Afrotropical genera of , Gymnosagena, Marriottella and Elgonina, revised here, were all originally described by Munro (in 1935, 1939 and 1957, respectively) and later included in the Spathulina genus group of Tephritini (Tephritinae) (Norrbom et al., 1999). To date Gymnosagena includes one species, Marriottella – one species, and Elgonina – two species, and nothing on the of these genera has been published subsequent to their original description. All three genera are characterized by a shiny black abdomen, a character shared with only a few other genera of Tephritini (see key below). Outside Tephritini, a shiny black abdomen is known in most Tephrellini (Tephritinae)(including the Platensinini sensu Norrbom et al. (1999)), a non-Asteraceae- feeding group restricted to the Old World. Outside Tephritinae there are many species with a shiny black abdomen, especially within the frugivorous and leaf-mining Trypetinae and the biologically diverse Acanthonevrini (Phytalmiinae). Species of Gymnosagena, Marriottella and Elgonina are also characterized by a short posterodistal lobe of cell bcu, or no such lobe at all; often only one orbital seta; and often a reduced wing pattern or no pattern at all. Together with nine other, mostly Afrotropical, genera they comprise the Spathulina group (Norrbom et al, 1999). The Spathulina group, however, has never been studied comprehensively, and no clear synapomorphies have been identified. Rather, relationships have been established in a so-called “serial” fashion, with one genus (e.g., Gymnosagena) considered similar and closely related to Spathulina Rondani, and another genus (e.g., Elgonina) added because it was considered related to Gymnosagena, and so on. Our working hypothesis was that these three genera together form a monophyletic subgroup within the Spathulina group that is tentatively called here the Gymnosagena subgroup. Nevertheless, the phylogenetic relationships within this subgroup and between it and other Tephritini are not well understood, and the exact composition of the Spathulina group will probably be redetermined only after a revision of Spathulina and other related genera. As Spathulina is species rich (Norrbom et al, 1999; Freidberg, unpublished data), it is not included in the present study. However, it is highly likely that after a comprehensive revision of the Spathulina group, the Gymnosagena subgroup will no longer prevail as an independent group. In the course of our on-going studies on Afrotropical Tephritidae, it has become evident that all three genera contain additional undescribed species. The purpose of the present publication is to revise the Gymnosagena subgroup on both species and generic levels, including a cladistic analysis. We also provide a key to Tephritini genera containing species with a shiny black abdomen. Labels are cited verbatim, except when annotated differently, e.g., collector names are spelled with only first letter capitalized. Authors of host are cited in Table 1. The following acronyms appearing on labels are cited verbatim, as are entire holotype labels: N = North, NE = North East, etc. Rt. = Route, nr = near, Km = Kilometer, m = meter; ft = feet, Mt(s) = Mountain(s). Terminology essentially follows McAlpine et al. (1981) and White et al. (1999). The tergal- oviscapal measure [“oviscapal measure” in this publication] of females is the number of terga immediately preceding the oviscape with combined length equal to length of the oviscape (Freidberg and Mathis, 1986). Two ratios are used for the pterostigma: 1. pterostigma ratio = pterostigma length divided by pterostigma width, and 2. pterostigma-costal-cell ratio = pterostigma length divided by costal cell length. The term ‘caudal flange’ refers to the flange borne on the posteroventral aspect of

368 Freidberg and Merz / Revision of the Gymnosagena group the epandrium and seen best when the epandrium is viewed in lateral view. The term ‘pre-glans part’ of the distiphallus refers to the distalmost part of the distiphallus immediately proximal to the glans. The word “dark”, unless accompanied by a color, usually means (e.g., for setae, setulae and wing pattern) brown to black. Likewise the word “pale” usually means white to yellow or gray. Body size is represented by wing length which is given at the end of each species description. Scutum width is measured at the level of postalar setae. Descriptions are composite; characters common to all species of a genus are mentioned in the generic description and not repeated in the species description. The cladistic analysis, including explanation of the methodology, comprises the last part of this chapter. The collections that loaned material for this study and their acronyms (curator names in parenthesis) are as follows: BMNH — Natural History Museum, London, UK (Mr. N. Wyatt) IST — Institute Scientifique, Antananarivo, Madagascar (Curator unknown) MNNG — Naturhistorisches Museum, Geneva, Switzerland NMB — Naturhistorisches Museum, Basel, Switzerland (Dr. M. Brancucci) NMP — Natal Museum, Pietermaritzburg, (Dr. B. Stuckenberg) SANC — National Collection of , Pretoria, South Africa (Dr. M. W. Mansell and Ms. Ros Urban) TAUI — Zoological Museum, Tel Aviv University, Tel Aviv, Israel ZIL — Museum of Zoology, Lund University, Lund, Sweden (Dr. R. Danielsson)

Paratypes of species represented by long series will be distributed among the following three collections: National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA; Natural History Museum, London, England; and National Collection of Insects, Protection Research Institute, Pretoria, South Africa.

TAXONOMY The genera, species groups and species in this section are treated alphabetically. For the construction of the following key, specimens of all the genera except Paraspathulina were studied by the authors. The large included genera, and , contain only very few species with shiny abdomen.

Key to genera of Tephritini with partly or completely shiny black abdomen 1. Basal section of costa strongly arched; frons, face, parafacial and gena unusually wide; thorax mostly shiny, with indistinct bands of microtomentum medially; male with lateral membrane of abdomen seen in dorsal view and tergites unusually narrow (as in many Myopitini) (Palaearctic)...... Placaciura Hendel -. Different combination of characters; costa and male abdomen different...... 2 2. Head about 1.5 times as long as high; proboscis long geniculate and nearly as long as thorax and abdomen combined (Namibia) ...... Euryphalara Munro -. Head and proboscis not so elongate; proboscis at most about half as long as body ...... 3 3. Thorax and abdomen entirely and strongly shiny black; wing pattern extensively black, comprised of partly connected transverse and oblique bands alternating with hyaline transverse and oblique bands (Lesotho) ...... Peratomixis Munro

369 BIOTAXONOMY OF TEPHRITOIDEA

-. Thorax usually distinctly microtrichose, not strongly shiny; in doubtful cases, wing not extensively black ...... 4 4. Palpus black; only basal scutellar seta present; only one orbital seta present; halter black; posterior notopleural seta dark brown to black; wing pattern reduced: at most cells bc, c,

pterostigma, spot at tip of vein R2+3 and border around crossvein DM-Cu infuscated (South Africa)...... Marriottella Munro -. Palpus mostly or entirely yellow; halter yellow (black only in some Tephritis, but then with 2 orbital and 2 scutellar setae); other characters variable...... 5 5. 3 concolorous frontal setae present; only basal scutellar seta present; 2 orbital setae present; posterior notopleural seta pale (Cosmopolitan) ...... Trupanea Schrank (shiny abdomen only in species of the former Urelliosoma Hendel; mostly south and east Mediterranean) -. 2 concolorous frontal setae present; other characters variable ...... 6 6. Anterior and posterior notopleural setae concolorous (black) or nearly concolorous (anterior seta black, posterior seta brown), both acuminate ...... 7 -. Anterior and posterior notopleural setae contrasted in color (anterior seta black, posterior seta whitish); anterior seta acuminate, posterior seta lanceolate ...... 9 7. Wing with Spathulina-type pattern: predominantly black or blackish, banded or reticulate banded, with only 0-2 isolated round hyaline spots in addition to marginal hyaline spots; 2

such marginal spots in cell r1 and wide hyaline spot at apex of cells r2+3 and r4+5 with small

dark spot at apex of vein R4+5; 2 scutellar setae present, apical seta at least 0.5 times as long as basal seta (Australian) ...... Paraspathulina Hardy and Drew -. Wing pattern different, either entirely or almost entirely lacking or, if conspicuous, then reticulate, with numerous isolated round hyaline spots and with different pattern at apex; 1 or 2 scutellar setae present, if 2, then apical seta at most 0.33 times as long as basal seta .... 8 8. Wing more or less infuscated, but not with reticulate pattern, usually with milky-white apical spot (absent in E. fuscana); femora black at least on basal half; lateral vertical seta dark; all postocular setae uniformly brown to black (Afrotropical) ...... Elgonina Munro -. Wing with reticulate pattern; color of femora variable; lateral vertical seta white; postocular setae mixed white and black (Afrotropical) ...... Gymnosagena Munro 9. Only basal scutellar seta present; proboscis spatulate or capitate ...... 10 -. 2 scutellar setae present, apical seta at least 0.33 times as long as basal seta; proboscis capitate ...... 12

10. Proboscis spatulate; mesofrons bare; vein R2+3 straight (Palaeotropic and Palaearctic) ...... Spathulina Rondani -. Proboscis capitate; other characters variable ...... 11

11. Vein R2+3 sinuous (Palaearctic)...... Hendrella Munro

-. Vein R2+3 straight (Neotropical) ...... Lamproxynella Hering and Pseudoedaspis Hendel

12. Wing with large, bulla-like strong infuscation in anterior half of cell r4+5 distal to

continuation of crossvein DM-Cu; vein R4+5 densely setulose ventrally at least 0.66 of its length; thorax with thick gray microtrichia, strongly contrasted with black abdomen (West Palaearctic) ...... Heringina Aczél

-. Wing more or less evenly dark, without bulla in cell r4+5; vein R4+5 usually setulose ventrally only to crossvein R-M (in T. dilacerata Loew also distally); thorax and abdomen with similar microtrichia, not strongly contrasted (Cosmopolitan) ...... Tephritis Latreille (shiny abdomen only in group of T. corolla Richter, T. sauterina Merz; central European)

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The Gymnosagena group Diagnosis Small (wing length 1.6-3.5 mm), predominantly black, with parts of head and legs yellow; 2 frontal setae; 1-2 orbital setae, anterior orbital seta dark, posterior orbital seta, if present, varies between yellowish and lanceolate to blackish and acuminate; lateral vertical seta pale or dark; postocular setae dark, pale, or mixed dark and pale; proboscis usually capitate or spatulate, rarely geniculate; scutum with pale or dark setulae; scapular setae differentiated or undifferentiated; dorsocentral seta near suture; 1-2 scutellar setae, apical scutellar seta, when present, no more than 0.3 times as long as basal scutellar seta; wing pattern reticulate, or mostly hyaline and usually with small dark spots and a white apical spot; tergites predominantly shiny or subshiny black, usually with only brownish setae and setulae, sometimes also with whitish or yellowish setulae; preglans part of distiphallus either setulose, or spinose, or with two large lateral setulose lobes, or with none; oviscape shiny black; aculeus: distal part often shaped as an arrowhead, tip with or without preapical step; two, more or less tuberculate, spermathecae.

Elgonina Munro Elgonina Munro, 1957: 890. Type species: Elgonina refulgens Munro, 1957, by original designation; Cogan and Munro, 1980: 542 (Afrotropical Catalog); Freidberg and Hancock, 1989: 49 (Comment on biology); Norrbom et al., 1999: 143 (World Catalog). Diagnosis Small flies (wing length 1.9-3.5 mm), predominantly black, with parts of head and legs and halter yellow; 1-2 orbital setae, all postocular setae dark, except in one species mixed whitish and black; 1-2 scutellar setae; wing hyaline, usually slightly infuscate, with 0-1 white apical spots and 0-2 white spots in cell r1; tergites predominantly shiny black, usually with only brownish setae and setulae; male: preglans part of distiphallus either with two large lateral setulose lobes or setulose without lobes; female: aculeus distally arrowhead in shape; tip with or without preapical step; sternite 8 preapically with lateral projection slightly extending beyond margin of tergite 8. Redescription Head (Figs. 1-3, 8-9). Structure: Slightly higher than long, with prominent occiput. Face strongly concave, without facial ridge or groove ; ventral facial margin and frontofacial angle about equally protuberant in lateral view. Frons bare, margins subparallel, usually about 1.25 times as wide at level of anterior ocellus as long (length measured between lunule and posterior ocelli). Eye suboval, about 1.3 times as high as long, with short, sparse setulae. Gena narrow, about 0.12-0.20 times as high as eye. Parafacial narrow, at narrowest point about as wide as thickest part of arista. Antenna: 1st flagellomere sometimes slightly pointed dorsoapically, about 1.5-2.0 times as long as pedicel, about 1.7 times as long as wide; rays of arista shorter than diameter of arista at base. Proboscis spatulate, labellum about 0.3-0.5 times as long as vertical diameter of eye. Palpus broad, usually about as broad as gena at narrowest point. Coloration and vestiture: Predominantly yellow, covered by dense microtrichia, only vibrissal angle shiny; occiput, postgena, orbital plate and ocellar triangle black, microtrichose. Antenna black or yellow, or bicolorous, similar in sexes or sexually dimorphic, in the latter case with 1st flagellomere yellow in male, black in female; scape and pedicel dorsally with short setulae;

371 BIOTAXONOMY OF TEPHRITOIDEA second aristomere yellow, third aristomere yellowish at base, darker towards apex. Palpus yellow, with some spiny, dark setulae on apical half. Labellum yellow. Chaetotaxy: 1-2 orbital setae, anterior orbital seta black, posterior orbital seta, if present, about 0.66 times as long as anterior seta, black or whitish; 2 black frontal setae; 1 black ocellar seta; 1 black medial vertical seta; 1 whitish to black lateral vertical seta; 1 whitish to brown postocellar seta; paravertical seta setula-like or about as long as postocellar seta, acuminate and brown or lanceolate and yellowish; postocular setae all dark and acuminate or mixed with some yellow lanceolate setae; genal seta and all postgenal and occipital setulae yellowish. Thorax. Structure: Scutum moderately convex, about 1.13-1.18 times as long as wide. Scutellum about 0.30-0.33 times as long as scutum, 0.42-0.50 times as long as wide. Coloration and vestiture: Black, shiny or dull; microtrichia sparse, not obscuring entirely underlying black cuticule, or dense. Mesonotum with brown or yellow setulae arranged somewhat irregularly. Propleuron with fan of 4-6 whitish setulae. Prosternum with some long, white setulae. Chaetotaxy: Scapular setae undifferentiated; 1 black postpronotal seta; 2 usually black notopleural setae, posterior seta sometimes slightly paler (dark brown), but not yellow, about 0.50-0.66 times as long as anterior seta; 1 black presutural seta; 1 black supra-alar seta; 1 postalar seta, either much less than half as long as intra-alar seta and pale (yellow or brown) or well developed (about half as long as intra-alar seta) and black; 1 long, black intra-alar seta; 1 dark dorsocentral seta, almost on suture; 1 dark prescutellar acrostichal seta, aligned slightly anterior to intra-alar seta; 1 black basal scutellar seta; apical scutellar seta lacking or present, if present then varies between 1.5-5.0 times as long as scutellar setulae, brown; 1 dark anepisternal seta; 1 paler (brown to yellow) anepimeral seta; 1 yellow to black katepisternal seta. Halter yellow. Calypteres yellow, with yellow fringe, ventral calypter narrow, stripe-like. Legs. Without overt features; coxae usually blackish (only coxae of E. fuscana predominantly yellow), with dense, silvery microtrichia; trochanters yellow; femora black, microtrichose, but ventrally often shiny, apical 0.2 yellow; tibiae yellow except hindtibia sometimes black at about basal half; tarsi yellow with darkened distal tarsomeres.

Wing (Figs. 12-19). Anal lobe well developed. Venation: Vein R1 setulose dorsally, row of setulae with gap at level of end of vein sc, other veins bare; crossvein R-M situated at about distal 0.66 of cell dm; distance between crossveins R-M and DM-Cu about equal to length of DM-Cu; cell bcu without posterodistal lobe; anal lobe well developed; veins usually pale at base, darker distally, vein C yellow to costagial or humeral break, then dark. Costal spine short, scarcely longer than surrounding setulae. Pattern: Wing usually without distinct pattern, although sometimes partly infuscated; cell r4+5 with milky-hyaline apical spot in most species; cell r1 with or without 1-2 milky-hyaline spots; pterostigma yellow to black. Abdomen. Predominantly shiny black, partly or entirely microtrichose on syntergite 1+2 and often at anterior margin of tergites 3 and 4; setulae mostly fine and brownish; marginal setae on tergites usually barely longer than setulae, sometimes distinctly longer; E. pollinosa has some whitish lanceolate setae laterally on most tergites. Male terminalia: Sternite 5 apically excised; syntergosternite 6-8 (Figs. 40-47) uniform in all species; epandrium oval in posterior view (Figs. 26-33), narrow, parallel-sided in lateral view, ventrally more or less pointed; inconspicuous to distinct caudal flanges present; lateral surstylus rather broad, distinctly curved; medial surstylus with 2 conspicuous setae medially and two dark brown prensisetae, lateral prensiseta smaller than

372 Freidberg and Merz / Revision of the Gymnosagena group medial, usually directed ventrally; hypandrium U-shaped, with or without lappet-like sclerotizations at articulation with epandrium; phallapodeme triangular in lateral view, asymmetrical as in other Tephritini, with additional branch articulating with hypandrium on one side; ejaculatory apodeme almost as long as epandrium, distally expanded, fan-like; basiphallus short, shovel-like. Three types of distiphallus, corresponding to three species groups, may be distinguished: (1) pollinosa type (Fig. 54): Distiphallus at preglans part with short appressed setulae; glans elongate, with distinct vesica; elongate internal sclerotized tube-like structure protruding from sclerotized sheath; acrophallus simple, in form of open tube, about half as long as vesica. (2) refulgens type (Figs. 55-57): Distiphallus at preglans part with few appressed setulae; glans rather short, without distinct vesica at tip; sheath-like sclerotization encircling almost entire acrophallus distally, with sclerotized, apically enlarged tube protruding distally; acrophallus without distinct sclerotized plate in middle of basal half. (3) splendida type (Figs. 58-61): Distiphallus at preglans part with two more or less symmetrical, recurrent, cushion-like lobes, densely covered by setulae; glans elongated, with distinct apical vesica; incomplete pipe-like sclerotization forming sheath with straight sclerotized tube inside; acrophallus tube-like, with sclerotized plate internally separting between two ducts. Female terminalia: Oviscape shiny black, covered by fine brown setulae; oviscapal measure about 2-3. Aculeus (Figs. 68-76) gradually tapered towards tip, tip either with preapical step or evenly acute and without such step; sternite 8 preapically with lateral projection slightly extending beyond margin of tergite 8, giving aculeus arrow-head shape. Spermathecae two, long oval, bearing variable number of tubercles on surface. Discussion This is a distinctive and homogenous genus, characterized, among other things, by a considerable or complete reduction of the wing pattern, an uncommon trait in Tephritidae. Within Tephritini it might only be confused with Marriottella. However, Marriottella can readily be separated from Elgonina by the black halter, the bicolored calypteres and the more discrete wing pattern. Other Tephritini with black abdomen differ from Elgonina by having a reticulate wing pattern. Based on external characters and the structure of the phallus (glans and pre-glans part), Elgonina may be divided into 3 well-defined species-groups: (1) pollinosa group - comprises only one species which is unique in having the lateral vertical seta, posterior orbital seta and postocellar seta whitish (blackish in other species), the mesonotum and abdominal tergites with white coarse setulae, and in the structure of the male terminalia (see above). (2) refulgens group - 2 black orbital setae and 2 scutellar setae present; glans with rather complicated structure of acrophallus and without distinct vesica. (3) splendida group - 1 orbital and only basal scutellar seta present; preglans part with 2 distinct setulose lobes; glans more parallel-sided and elongate, with distinct vesica and rather simple acrophallus.

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Figs. 1-7. Head, lateral view. 1. Elgonina pollinosa. 2. E. refulgens. 3. E. splendida. 4. Gymnosagena longicauda. 5 G. unicornuta. 6. Marriottella exqusita. 7. M. sepsoides.

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Figs. 8-11. Head, frontal view. 8. Elgonina refulgens. 9. E. splendida. 10. Marriottella exqusita. 11. M. sepsoides.

Biology The larvae develop in flowerheads of various Conyza spp. and Microglossa pyrifolia (Table 1, p. 415). They either pupate inside the flowerhead or leave it before pupation. Distribution The genus is distributed mainly in the highlands of eastern Africa from to southern Africa, and in Cameroon (only on Mt. Cameroon).

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Key to the species of Elgonina 1. Mesonotum and lateral margin of tergites with whitish, coarse setulae (although most of abdomen covered by fine and dark setulae); 2 orbital setae present, posterior seta whitish; postocellar seta whitish; lateral vertical seta whitish (Fig. 1); only basal scutellar seta present; wing uniformly hyaline, without milky-white spot at apex (Fig. 12) (Host: Conyza hypoleuca; Ethiopia; pollinosa group) ...... E. pollinosa, n. sp. -. Mesonotum and abdomen covered uniformly by fine, acuminate dark setulae, although setulae on mesonotum sometimes with yellow tinge; both orbital setae (if posterior orbital seta present) acuminate and usually dark, although posterior orbital seta sometimes yellow; postocellar seta brown to black; lateral vertical seta black (Figs. 2-3); other characters variable, but wing usually with apical milky-white spot (absent in E. fuscana) (Figs. 13-19) ...... 2 2. Coxae almost entirely yellow; wing uniformly hyaline, without milky-white apical spot (as in Fig. 12); 2 orbital and 2 scutellar setae present; 1st flagellomere black (only known) (Uganda; refulgens group) ...... E. fuscana Munro () -. Coxae almost entirely black; tip of wing with milky-white spot (Figs. 13-19); other characters variable ...... 3 3. Pterostigma more or less uniformly black; wing distinctly darker towards tip, where strongly contrasted with milky-white apical spot (Fig. 17); one orbital seta and only basal scutellar seta present; male with yellow and female with black 1st flagellomere (Host: Microglossa pyrifolia; Kenya; splendida group) ...... E. infuscata, n. sp. -. Pterostigma either uniformly yellow or darkened only in apical half; wing evenly infuscated, milky-white apical spot less conspicuous (Figs. 13-16, 18-19); other characters variable ...... 4 4. 2 orbital and 2 scutellar setae present; apical scutellar seta seta-like or setula-like, usually 3- 5 times as long as adjacent setulae; postalar seta dark brown to black, at least twice as long as posterior notopleural seta; preglans part of distiphallus with short appressed setulae, without lobes (refulgens group) (Figs. 55-57) ...... 5 -. One orbital seta and only basal scutellar seta present; apical scutellar seta not seta-like, if present, similar to, and usually 1.5-2.0 times as long as, adjacent setulae; postalar seta short, yellow to brown, about as long as posterior notopleural seta; preglans part of distiphallus with setulose lobes (splendida group) (Figs. 58-61) ...... 10 5. Male; 1st flagellomere yellow or black ...... 6 -. Female; 1st flagellomere black ...... 8 6. 1st flagellomere yellow (Ethiopia) ...... E. dimorphica, n. sp. () -. 1st flagellomere black...... 7 7. Glans with apical pore surrounded by wide sclerotization (Fig. 56) ...... E. inexpectata, n. sp. () -. Glans with apical pore surrounded by narrow sclerotization (Fig. 57) ...... E. refulgens Munro () 8. Aculeus without preapical step (Fig. 72) (Host: possibly Conyza ruwenzoriensis; Kenya) ...... E. refulgens Munro () -. Aculeus with preapical step (Figs. 69, 71) ...... 9 9. Known from Ethiopia ...... E. dimorphica, n. sp. ()

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-. Known from Kenya, Uganda and Tanzania ...... E. inexpectata, n. sp. ()

10. Pterostigma bicolored, yellow in basal half and brown in apical half; cell r1 with 1-2 distinct

hyaline spots touching vein R1 (Fig. 19); syntergite 1+2 entirely microtrichose; 1st flagellomere black in both sexes (Host: Conyza newii; Kenya, Tanzania) ... E. yaromi, n. sp.

-. Pterostigma yellow, at most indistinctly darker in apical quarter; cell r1 either uniformly

hyaline (Figs. 16, 18) or with small milky-white area at base which does not touch vein R1; syntergite 1+2 microtrichose at most on basal 0.75; color of 1st flagellomere variable ... 11 11. Male; 1st flagellomere yellow ...... 12 -. Female; 1st flagellomere variable ...... 13 12. Known from Tanzania, and South Africa ...... E. flavicornis, n. sp. () -. Known from Ethiopia, Kenya and Tanzania ...... E. splendida, n. sp. () 13. 1st flagellomere yellow (Tanzania, Malawi and South Africa) .... E. flavicornis, n. sp. () -. 1st flagellomere black (Ethiopia, Kenya and Tanzania) ...... E. splendida, n. sp. ()

Elgonina pollinosa group This group is characterized by the presence of 2 orbital setae, with the posterior orbital seta whitish and lanceolate; postocular setae mixed whitish and black; mesonotum rather densely microtrichose, with coarse whitish setulae; postalar seta long, only basal scutellar seta present; tergites laterally with some long, coarse, whitish setulae; distiphallus pollinosa type: preglans part with short appressed setulae, without lobes; glans elongate, with distinct vesica; elongate internal sclerotized tube-like structure protruding from sclerotized sheath. This group contains only Elgonina pollinosa n. sp.

Elgonina pollinosa Freidberg and Merz, n. sp. (Figs. 1, 12, 26, 40, 54, 68, 83)

Diagnosis Differs from all congeners by the diagnostic characters of the group, in particular the whitish, lanceolate setae on the head, the coarse whitish setulae on the thorax and the elongate glans. Description Male: Head (Fig. 1). Mostly yellow; ocellar triangle, orbital plate, occiput and postgena black; microtrichose; antenna yellow; anterior orbital seta, frontal setae, ocellar seta and medial vertical seta black; posterior orbital seta, postocellar seta, genal seta and postgenal seta whitish, lanceolate; postocular setae mixed whitish and black; proboscis mostly yellow, prementum brownish. Thorax. Black, with whitish-golden microtrichia; scutum and scutellum with coarse white setulae; posterior notopleural seta black; postalar seta black, about half as long as intra-alar seta; apical scutellar seta lacking. Legs. Coxae black; femora, except distal part, black; hindtibia with black ring occupying 0.33- 0.50 of length basally, except narrow basal yellow ring; distal 1-2 tarsomeres of all legs brownish; all other parts yellow. Wing (Fig. 12). Entirely hyaline, without dark pattern and milky-white spots; pterostigma pale

377 BIOTAXONOMY OF TEPHRITOIDEA yellow, somewhat darker toward tip. Abdomen. Syntergite 1+2 at about basal 0.75, and anterior margin of tergites 3 and 4, microtrichose; tergites 2-5 laterally with rather long white coarse setulae, setulae otherwise fine and dark; tergite 5 with dark brown setae along posterior margin. Terminalia: Epandrium in posterior view (Fig. 26) rounded, broader than high, with dense group of pale, elongate, club- shaped, dorsomedially curved setulae situated near apex of lateral surstylus, in lateral view (Fig. 40) arched posteriorly, strongly tapered ventrally, with tip of lateral surstylus at different level than remaining part, with shallow, toothed caudal flange; medial prensiseta carried on elongate projection; distiphallus pollinosa type (Fig. 54), at preglans part with short appressed setulae; glans elongate, with distinct vesica; elongate internal sclerotized tube-like structure protruding from sclerotized sheath. Female. Similar to male, except 1st flagellomere black; tergites 2-6 laterally with rather long white coarse setulae, tergites 5 and 6 with dark brown setae along posterior margins. Terminalia: Oviscapal measure about 2; tip of aculeus pointed rather abruptly, without preapical step (Fig. 68); spermatheca about 4 times as long as wide (Fig. 83). Wing length. Male: 2.4- 2.8 mm; female: 2.6-2.9 mm. Material Examined Holotype , ETHIOPIA: BALE, Bale Mountains, 10 kmS Goba, 3200m, 1.ii.2000, A. Freidberg & I. Yarom. The holotype is double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in TAUI. Paratypes: same locality data as holotype (8, 7, of which 1 collected on Hagenia abyssinica; TAUI and MHNG); same locality data but also: ex flowerhead Conyza hypoleuca (5, 6; TAUI); same locality data but also: Micha River, 3100m (1; TAUI). Etymology Named after the densely microtrichose (=pollinose) thorax. Biology Reared from flowerheads of Conyza hypoleuca. All specimens were collected at an altitude of about 3200 m. Distribution Ethiopia (only on Bale mountains). Discussion Although this species undoubtedly belongs to Elgonina, it differs from all other congeners by the denser microtrichia and coarse, whitish setae and setulae, and thus shows greater affinities with other, more typical, tephritine genera, such as .

Elgonina refulgens group This group is characterized by the presence of 2 orbital setae, the anterior seta black, the posterior seta at most 0.66 times as long as the anterior seta, black to brown; black antenna (except for males of E. dimorphica); large postalar seta; conspicuous apical scutellar seta; distiphallus refulgens type: preglans part with short appressed setulae; glans short and wide, without distinct vesica; internal sclerotized tube protruding from sclerotized sheath,

378 Freidberg and Merz / Revision of the Gymnosagena group considerably widened distally. This group contains Elgonina fuscana Munro, E. refulgens Munro, E. dimorphica n. sp. and E. inexpectata n. sp.

Elgonina dimorphica Freidberg & Merz, n. sp. (Figs. 13, 27, 41, 55, 69, 84) Diagnosis This is the only species of the E. refulgens group in which the male has yellow 1st flagellomere. The male of E. fuscana, which is unknown, might also have yellow 1st flagellomere; however, we would expect it to resemble the conspecific female by not having a white apical spot on the wing, which would distinguish it from the male of E. dimorphica. Description Male: Head. Antenna yellow; 2 orbital setae present. Thorax. Scutum subshiny, with sparse gray microtrichia; postalar seta long, black; apical scutellar seta thin, about 0.2 times as long as basal scutellar seta, and about as long as posterior notopleural seta. Legs. Tibiae predominantly yellow, hindtibia with black ring on basal half.

Wing (Fig. 13). Slightly infuscated, pterostigma yellow, cell r1 without white spots, apex of wing with white spot. Abdomen. Syntergite 1+2 sparsely microtrichose over about 0.66 of area. Terminalia: Epandrium (Figs. 27, 41) with shallow but distinct caudal flange; lateral surstylus broad, more or less evenly rounded towards tip in lateral view (Fig. 41); distiphallus with apical duct of glans conspicuously enlarged apically, not surrounded by second sclerotized sheath (Fig. 55). Female. Differs from male in having 1st flagellomere black; abdomen with basal 0.66 of syntergite 1+2 and narrow stripes at bases of tergites 3-5 microtrichose. Terminalia: Oviscapal measure 2-3; Aculeus with distinct preapical step (Fig. 69); spermatheca with numerous small tubercles (Fig. 84). Wing length. Male: 2.9-3.0 mm; female: 2.9-3.0 mm. Material Examined Holotype , ETHIOPIA: Bale Mts., Goba, 2800m, 15.xii.1989, A. Freidberg & F. Kaplan (TAUI). The holotype is double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in TAUI. Paratypes: Ethiopia, Bale Mts., 3000-3300m, 15.xii.1989, A. Freidberg & F. Kaplan (2; TAUI). Bale Mountains, 10KmS Goba, 31.i.2000, A. Freidberg & I. Yarom (3, 2; TAUI), 1.ii.2000 (2; TAUI). Gamo Gofa, Chencha, 40KmNW Arba Minch, 2800m, 6.ii.2000, I Yarom & A. Freidberg (1; TAUI). Kefa: Jimma, 35KmS, 2700m, 12.ii.2000, A. Freidberg & I. Yarom (5, 2; TAUI). Etymology Named after the sexually dimorphic color of the 1st flagellomere. Biology Host plants unknown. A high-altitude species, collected between 2700-3300 m.

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Distribution Ethiopia (Bale mountains and Jimma area). Discussion Elgonina dimorphica resembles closely the other three members of its species group, although the male differs readily from the males of E. inexpectata and E. refulgens in antenna color and glans structure. The male of the fourth member of the group, E. fuscana, is unknown. The discovery in Ethiopia of two endemic species of Elgonina, E. dimorphica and E. pollinosa, in a cursory survey may suggest that additional endemic species occur there.

Elgonina fuscana Munro (Fig. 70, 85) Elgonina fuscana Munro, 1957: 891; Cogan & Munro, 1980: 542 (Afrotropical Catalog); Norrbom et al., 1999: 143 (World Catalog). Diagnosis Elgonina fuscana and E. pollinosa are the only species of Elgonina that lack the apical white spot on the wing. However, they belong to two different species groups, differing in the diagnostic characters of their respective groups, including the coarse whitish setulae on the thorax of E. pollinosa and the relatively long apical scutellar seta in E. fuscana. Within Elgonina, E. fuscana is unique in having yellow coxae. Redescription Female: Head. Gena about 0.2 times as high as eye; labellum about half as long as eye height; antenna with 1st flagellomere black, pedicel brownish-yellow; paravertical seta slightly longer than postocellar seta; all postocular setae black. Thorax. Scutum dull, with dense gray microtrichia; posterior notopleural seta about 0.5 times as long as anterior notopleural seta; postalar seta long, brown; apical scutellar seta about 0.33 times as long as basal scutellar seta. Legs. Coxae and tibiae predominantly yellow, hindtibia with narrow black ring on basal half. Wing. Slightly infuscated, without white spots, pterostigma yellow. Abdomen. Tergites 5 and 6 with few long brown setae; oviscapal measure about 3. Terminalia: Aculeus with distinct preapical step (Fig. 70). Spermatheca with numerous rather large tubercles (Fig. 85). Wing length. 3.5 mm. Male. unknown. Material Examined Holotype , UGANDA: Kigezi district, Mt. Muhavura, 10,000-12,000 ft., xi.1934, F.W. Edwards (BMNH). The holotype is double mounted on a minuten pin and is in good condition. The abdomen has been dissected and is stored in a vial on another pin. Biology Host plants unknown. The holotype was collected at an altitude between 3000-3700 m.

380 Freidberg and Merz / Revision of the Gymnosagena group

Distribution Uganda. Discussion This species is still known from the female holotype only, and the male terminalia remain unknown. Nevertheless it appears to be distinct and is probably a member of the E. refulgens group, based on the presence of 2 orbital setae and relatively long apical scutellar setae. We assume that the yellow color of the coxae, diagnostic of the female, would also characterize the males. The five-line original description (Munro, 1957) merely contains a comparison with E. refulgens, noting differences in wing pattern, leg coloration, oviscape and size.

Elgonina inexpectata Freidberg & Merz, n. sp. (Figs. 14, 28, 42, 56, 71, 86) Diagnosis Within the refulgens group, this species is similar to E. refulgens by the 1st flagellomere being black in both sexes, and apical white spot present on wing. It differs from E. refulgens by the glans having large sclerotization of the apical tube and the aculeus having a preapical step. Description Male: Head. Antenna with scape and pedicel yellow and 1st flagellomere black. Thorax. Scutum subshiny, covered by sparse, brownish-gray microtrichia; postalar seta long, black; apical scutellar seta about 0.25 times as long as basal scutellar seta. Legs. Tibiae mostly yellow, but hindtibia in most specimens with more or less distinct black ring on basal half. Wing (Fig. 14). Slightly infuscated, with white apical spot; pterostigma yellow, slightly darkened towards tip. Abdomen. Basal 0.66 of syntergite 1+2 and sometimes also small medial stripe on tergites 3 and 4 microtrichose. Terminalia: differ from E. refulgens primarily in tip of glans (Fig. 56): apical sclerotization around apical pore broader, consisting of two sclerotized layers. Epandrium: Posterior view as in Fig. 28; quadrangular at dorsal 0.66 in lateral view (Fig. 42). Female. Similar to male, but abdominal tergites 3 and 4 basally often with narrow microtrichose stripe, without medial stripe. Terminalia: Oviscapal measure about 3; aculeus with distinct preapical step (Fig. 71); spermatheca with rather large and close-set tubercles (Fig. 86). Wing length. Male: 2.5-3.0 mm; female: 2.6-3.2 mm. Material examined Holotype , KENYA: Equator, Rt. A104, 2500m, 20.ix.1992, A. Freidberg (TAUI). The holotype is double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in TAUI. Paratypes: Same data as holotype (25, 21; TAUI); KENYA: Rt. A104, nr. Uplands, 18.xi.1989, A. Freidberg & F. Kaplan (7, 7; TAUI); Rt. A104, Uplands, 2500m, 18.ix.1992, A. Freidberg (1, 1; TAUI); Aberdare, 3000-4000m, 1.xii.1986, A. Freidberg (11, 3;

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TAUI), Aberdare, 2600-3000m, 8.xii.1989, A. Freidberg & Fini Kaplan (3, 1; TAUI); Uplands, 15.xi.1986, A. Freidberg and I. Susman (8, 3; TAUI); KENYA (West): Timboroa, 26.xi.1989, A. Freidberg & Fini Kaplan (7, 5; TAUI); Equator, 30.x.[19]83, A. Freidberg and I. Yarom (11, 8; TAUI); Molo, 19, 27.xi.1989, A. Freidberg & Fini Kaplan (1; TAUI); 10 Km North Kabarnet, 29.xi.1989, A. Freidberg & Fini Kaplan (2; TAUI); Mau, East Mau, 28.viii.[19]83, A. Freidberg (1; TAUI). Nabkoi, 50km SE Eldoret, 15.iii.1993, Merz (3, 4; MNHG); Tarakwa, 35km SE Eldoret, 15.iii.1993, Merz (1; MNHG). TANZANIA: 1800m, Usmabara Mts., Rt. B124, Gologolo, 12-13.ix.1992, A. Freidberg (4, 3; TAUI); Usambara Mts., Gologolo, 1900m, 23.viii.1996, A. Freidberg (11, 9; TAUI); UGANDA: S.W., Ichuya Forest, Kanaba Gap, 2500m, 25.xii.1995, I. Yarom & A. Freidberg (1; TAUI). Etymology Named after its unexpected discovery which was made possible only after close examination of both male and female terminalia. Biology Host plants unknown, although consistently swept from Conyza steudelii and C. newii. All specimens were collected between 1800 m and approximately 3000 m. Distribution Kenya, Uganda and Tanzania. Discussion This species can be distinguished from E. refulgens only by characters of the male and female terminalia. Examples of most series recorded under these species therefore required dissection before their identity could be established.

Elgonina refulgens Munro (Figs. 2, 8, 15, 29, 43, 57, 72, 87) Elgonina refulgens Munro, 1957: 890; Cogan & Munro, 1980: 542 (Afrotropical Catalog); Norrbom et al., 1999: 143 (World Catalog). Diagnosis Within the refulgens group, this species is similar to E. inexpectata in the 1st flagellomere being black in both sexes, and the presence of a white apical spot on the wing. It differs from E. inexpectata by the narrow sclerotization of the apical tube of the glans, and by the aculeus being evenly pointed, without preapical step. Redescription Male: Head (Figs. 2, 8). Antenna with scape and pedicel brownish-yellow and 1st flagellomere black. Thorax. Scutum subshiny, covered by dense, brownish-gray microtrichia; postalar seta long, black; apical scutellar seta about 0.2-0.3 times as long as basal scutellar seta. Legs. Tibiae mostly yellow, but hindtibia in most specimens with more or less distinct black ring on basal half.

382 Freidberg and Merz / Revision of the Gymnosagena group

Wing (Fig. 15). Slightly infuscated, with distinct white apical spot and sometimes less distinct, small white spots at base of cell r1 and apex of cells r1 and r2+3. Pterostigma yellow, but slightly darkened towards tip. Abdomen. Basal 0.66 of syntergite 1+2 and sometimes also small median stripe on tergites 3 and 4 microtrichose. Terminalia: Epandrium with distinct serrate caudal flange; lateral surstylus triangular and pointed distally in lateral view (Fig. 43) and indistinctly split in posterior view (Fig. 29); glans (Fig. 57) consisting of sheath-like external tube with strongly sclerotized internal duct; internal duct narrow, extending beyond external tube distally. In contrast to E. inexpectata and E. dimorphica, this internal duct is not surrounded at its tip by second, large sclerotization. Female. Similar to male, but abdominal tergites 3 and 4 basally often with narrow microtrichose stripe, without median stripe. Terminalia: Oviscapal measure about 3; aculeus without preapical step (Fig. 72); spermatheca with rather large and close-set tubercles (Fig. 87). Wing length. Male: 2.5-3.2 mm; female: 2.6-3.2 mm. Material Examined Holotype , KENYA: Mt. Elgon, 10,500-12,500 ft., ii.1935, on flowers of Conyza ruwenzoriensis, F.W. Edwards. The holotype is double mounted (minuten in a celluloid strip), is in excellent condition, and is deposited in BMNH. Paratypes: same data as holotype, and also “B.M. E. Afr. Exp., B.M. 1935-203” (5, 5 - BMNH; 2, 3 - SANC); same data as holotype, but on Erlangia [sic.] sp. (1; BMNH). Additional specimens: KENYA: Mt. Elgon, 3800m, 25.xi.1986, A. Freidberg (7, 4; TAUI); same collecting data, ex flowerhead Conyza sp., 15.xii.1986 (1, associated puparium, 1 intact puparium; TAUI); Mt. Elgon, 3500m, 31.x.[19]83, A. Freidberg (3; TAUI), I. Yarom (1, 1; TAUI); Aberdares N.P., 2800-3000m, 6.iii.1993, B. Merz (3ff, MNHG); Aberdares N[ational]. P[ark]., 2800-3000m, 7.iii.1993, B. Merz (6, 2ff; MHNG). Biology The type series was collected on flowers of Conyza ruwenzoriensis (Munro, 1957), except for one specimen, which was found on Erlangea sp. One female emerged from a puparium that was found inside a rearing bag with flowerheads of Conyza sp. (Mt. Elgon). A second larva also left the flowerheads, but did not yield an adult. We assume that this Conyza sp. may actually be C. ruwenzoriensis. Distribution Kenya. Only known from Mt. Elgon and the Aberdare mountains above the timberline (2800-3800 m). Discussion An excellent description of the external morphology is provided by Munro (1957), which is primarily supplemented here by characters of the terminalia. However, his Fig. 19 exaggerates the contrast between the apical band and the remaining parts of the wing. The wing is strongly grayish, but the apical area is at most only slightly darker. The terminalia (of a topotypical male and a female paratype) are described and illustrated here for the first time. This species can be distinguished from E. inexpectata only by characters of the male and female terminalia. Examples of most series recorded under these species therefore required dissection before their

383 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 12-19. Wing. 12. Elgonina pollinosa. 13. E. dimorphica. 14. E. inexpectata. 15. E. refulgens. 16. E. flavicornis. 17. E. infuscata. 18. E. splendida. 19. E. yaromi. identity could be established. We have studied almost the entire type series that was divided between the BMNH and SANC.

Elgonina splendida group This group is characterized by the presence of only one orbital setae; short and relatively pale (yellow to brown) postalar seta; absence of distinct apical scutellar setae, at most represented by a pair of tiny setulae that are scarely longer than surrounding setulae; distiphallus splendida type: preglans part of distiphallus with two prolonged cushion-like, strongly setulose lobes; glans rather elongated, with distinct vesica; sclerotization simple, with straight tube centrally not protruding distally beyond engulfing, sclerotized sheath. This group contains the following four species, all new: Elgonina flavicornis n. sp., E. infuscata n. sp., E. splendida n. sp., and E. yaromi n. sp.

384 Freidberg and Merz / Revision of the Gymnosagena group

Figs. 20-25. Wing. 20. Gymnosagena campiglossina. 21. G. kakamega. 22. G. longicauda. 23. G. unicornuta. 24. Marriottella exquisita. 25. M. sepsoides.

Elgonina flavicornis Freidberg & Merz, n. sp. (Figs.16, 30, 44, 58, 73, 88) Diagnosis The only species of Elgonina that has the antenna entirely yellow in both sexes; wing, including pterostigma, rather pale, without dark markings. Description Male: Head. Antenna yellow, sometimes slightly infuscated only around base of arista. Thorax. Scutum subshiny black, with sparse gray microtrichia; postalar seta short and yellowish; apical scutellar seta tiny, scarcely longer than scutal setulae. Legs. Hindtibia either entirely yellow or with indistinct black ring on basal half.

Wing (Fig. 16). With faint infuscation; pterostigma yellow; cell r1 without white spots; apical white spot present but indistinct. Abdomen. Basal 0.75 of syntergite 1+2 and narrow stripe at base of tergites 3 and 4 microtrichose. Terminalia: Epandrium with prominent, serrate caudal flange (Fig. 44); lateral surstylus slightly split apically in posterior view (Fig. 30); glans with distinct apical vesica and membranous lateral process (Fig. 58).

385 BIOTAXONOMY OF TEPHRITOIDEA

Female. Terminalia: Oviscapal measure about 3; aculeus with small preapical step (Fig. 73); spermatheca with rather few tubercles (Fig. 88). Wing length. Male: 1.9-2.1 mm; female: 2.0-2.2 mm. Material Examined Holotype , TANZANIA: Njombe, 20KmSE, Rt. B4, 2000m, 27-28.viii.1996, A. Freidberg. The holotype is double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in TAUI. Paratypes: Same collecting data (6, 3; TAUI, MHNG). TANZANIA: Rt. A345, near Ngozi Crater, 1900m, 1.ix.1996, A. Freidberg (1; TAUI). MALAWI: Zomba Plateau, Chiradzulu Forest, 22-23.x.1983, A. Freidberg (1, 1; TAUI). Zomba Plateau, Emperor’s View, 22-23.x.1983, A. Freidberg (1; TAUI). SOUTH AFRICA: Robbers’ Pass, 1500m, 29.xii.1994, A. Freidberg (1; TAUI); Natal, World View, Pietermaritzburg, 25.ix-2.x.1983, A. Freidberg (1, 1, TAUI). Etymology Named after the yellow antenna in both sexes. In Latin ‘flavus’ = yellow; ‘cornu’ = horn. Biology Host plants are unknown. Found in relatively low altitudes and high latitudes. Distribution Tanzania, Malawi and South Africa (Natal). Discussion This is the only Elgonina species known from the southern part of the Afrotropical region (southern Tanzania to South Africa). The females can usually be separated easily from other species of the splendida group by the yellow antenna, although females of E. splendida from Njombe, Tanzania, are intermediate between typical E. flavicornis and typical E. splendida (see also Discussion under E. splendida). The males of the two species are similar and difficult to separate. Further studies, including data on the host plants, are therefore needed in order to clarify the relationships between these species.

Elgonina infuscata Freidberg & Merz, n. sp. (Figs. 17, 31, 45, 59, 74, 89)

Diagnosis This species is easily distinguished from all congeners by the strongly infuscated wings with entirely black pterostigma and contrasting milky-white apical spot. Description Male: Head. Antenna entirely yellow. Thorax. Scutum subshiny black, with sparse microtrichia, not obscuring underlying cuticule; postalar seta short and yellow; apical scutellar seta tiny, scarcely longer than scutal setulae. Legs. Tibiae entirely yellow.

386 Freidberg and Merz / Revision of the Gymnosagena group

Wing (Fig. 17). Generally dark, more so towards apex, with dark band-like spot wider and more distinct along costal margin of cell r2+3, extending into apex of cell r1 and across cell r4+5 into cell m, strongly contrasted with large white apical spot; no other white spots present; pterostigma blackish to black, slightly paler at base. Abdomen. Basal 0.75 of syntergite 1+2 and base of tergite 3 microtrichose. Terminalia: Epandrium with shallow caudal flange (Fig. 45); lateral surstylus slightly split apically in posterior view (Fig. 31); distiphallus splendida-type, with distinct apical vesica (Fig. 59). Female. similar to male, except antenna darker, in particular 1st flagellomere entirely black. Terminalia: Oviscapal measure about 3; tip of aculeus with small preapical step (Fig. 74); spermatheca with numerous, medium-sized tubercles (Fig. 89). Wing length. Male and female: 2.0-2.2 mm. Material Examined Holotype , KENYA: Molo, 19, 27.xi.1989, A. Freidberg & F. Kaplan (TAUI). The holotype is double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in TAUI. Paratypes: same collecting data as holotype (4; TAUI); same collecting data as holotype, but also: ex flowerhead Microglossa pyrifolia, 21.xi.1989 (5; TAUI); Miti ya Hunter, 30kmS. Eldoret, 11.v.1991, A. Freidberg & F. Kaplan (3, 2; TAUI); 8km NE Kericho, 18.xi.1986, A. Freidberg (2; TAUI), 26.viii.2003, A. Freidberg (1, 2; TAUI); 15km SW Kericho, 16.xi.1986, A. Freidberg (1; TAUI); 30km NE Kericho, 10.xi.[19]83, A. Freidberg (1; TAUI); Aberdare, 3000-4000m, 1.xii.1986, A. Freidberg (1; TAUI); Kenya West, Kapenguria, 24; 25.xi.1989, A. Freidberg & F. Kaplan (1; TAUI); 10km NE Kericho, 2200m, 9.iii.1993, B. Merz (1; MHNG). Etymology Named after the strongly infuscated wing. Biology Reared from Microglossa pyrifolia. All specimens were collected between 2000 m and about 3500 m. Distribution Kenya (Only from central Kenyan highlands). Discussion Although this species is a typical member of Elgonina, the strongly infuscated wing differs from that of all other congeners and resembles the wings of Marriottella spp.

Elgonina splendida Freidberg & Merz, n. sp. (Figs. 3, 9, 18, 32, 46, 60, 75, 90) Diagnosis

Rather pale-winged species with pale pterostigma and only indistinct white spots in cell r1; antenna sexually dimorphic: 1st flagellomere yellow in male, brown to black in female.

387 BIOTAXONOMY OF TEPHRITOIDEA

Description Male: Head (Figs. 3, 9). Antenna entirely yellow; 1 orbital seta; postocellar seta fine, paler than medial vertical seta. Thorax. Scutum with rather dense microtrichia, covering underlying cuticule almost entirely; postalar seta short and brown; apical scutellar seta tiny, scarcely longer than scutal setulae. Legs. Hindtibia predominantly yellow, often with dark ring on basal half, sometimes rather indistinct. Wing (Fig. 18). With rather faint infuscation; apical white spot present but indistinct; pterostigma yellow with tip sometimes slightly darkened. Abdomen. Basal 0.75 of syntergite 1+2 and base of tergite 3 sparsely microtrichose. Terminalia: Epandrium with large serrate caudal flange (Fig. 46); lateral surstylus indistinctly split apically in posterior view (Fig. 32); distiphallus splendida type (Fig. 60); glans parallel- sided, with large vesica and some small lateral, membranous processes. Female. Similar to male, except 1st flagellomere browinish to black. Terminalia: Oviscapal measure about 3; aculeus tapered evenly at tip, without preapical step (Fig. 75); spermatheca with only few, large tubercles (Fig. 90). Wing length. Male: 1.9-2.3 mm; female: 2.1-2.5 mm. Material Examined Holotype , KENYA: (West) Kapenguria-tartar road, 24.xi.1989, A. Freidberg & Fini Kaplan. The holotype is double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in TAUI. Paratypes: Same collecting data as holotype (33, 23; TAUI); KENYA: Kapenguria, 5.xi.[19]83, A. Friedberg (1; TAUI), 25-25.xi.1989, A. Freidberg & Fini Kaplan (2, 1; TAUI); Mt. Kenya, Timau, 22.viii.[19]83, A. Freidberg (9, 6; TAUI); Menengai Crater, 27.viii.[19]83, A. Friedberg (3; TAUI); West Pokot, Chepareria, 4-5.xi.[19]83, I. Yarom (1, 1; TAUI); Tartar, 30 Km N[orth] Kitale, 4-5.xi.[19]83, I. Yarom (1; TAUI); Chewele, 15 Km N Bungoma, 7.xi.[19]83, A. Freidberg (2; TAUI); Cherangani Hills, Labot, 3000m, 2.xi.[19]83. A. Freidberg (2; TAUI); Cherangani Hills, Kapcherop, 2.xi.[19]83, A. Freidberg and I. Yarom (3, 7; TAUI); Eldoret, 30.x.[19]83, A. Freidberg (1; TAUI); Aberdare, 2000-2600m, 8.xii.1989, A. Freidberg & Fini Kaplan (1, 2; TAUI), 3000-4000m, 1.xii.1986, A. Freidberg (1; TAUI); 10 Km N Maralal, 28.xi.1986, A. Freidberg (1; TAUI); Rt. A104, 15 Km SE Nairobi, 29.iv-15.v.1991, A. Freidberg & Fini Kaplan (1; TAUI); Athi River, 18.viii.[19]83, A. Friedberg (2; TAUI); W-Nairobi, ILRAD, 1900m, 16.iii.1993, B. Merz (3, 2; MHNG); W-Nairobi, ILRAD, 1900m, 8.iii.1993, B. Merz (3, 2; MHNG); Tarakwa, 35km SE Eldoret, 2350m, 15.iii.1993, B. Merz (2, 2; MHNG); Nabkoi, 50km SE Eldoret, 2550m, 15.iii.1993, B. Merz (1; MHNG); 75km SE Nairobi, A109, 1700m, 18.iii.1993, B. Merz (1, 2; MHNG); Magadi Road, 1000-1600m, 3.iii.1993, B. Merz (1, 1; MHNG). Additional specimens (not included as paratypes): ETHIOPIA: Debra Libanos, 2.i.[19]72, A. Freidberg (1, 1 ; TAUI), 12.xii.1989, A. Freidberg & Fini Kaplan (1,1; TAUI); Mt. Managesha, 3000m, 10.xii.1989, A. Freidberg & Fini Kaplan (2; TAUI). KENYA: Equator,

388 Freidberg and Merz / Revision of the Gymnosagena group

Rt. A104, 2500m, 20.ix.1992, A. Freidberg (1; TAUI). TANZANIA: Mt. Rungwe, Rt. A345, 2200m, 30.viii.1996, A. Freidberg (1, 1; TAUI); Njombe, 10kmSE, Rt. B4, 2000m, 27- 28.viii.1996, A. Freidberg (3; TAUI). Etymology Named after its splendid appearance. Biology Host plants unknown, although collected on Conyza pyrrhopappa, and there is also a doubtful rearing record from this potential host. The specimens at hand were collected between 1000 and 4000 m. This is the only species of Elgonina which can be found regularly below 2000 m. This wide amplitude in altitude may indicate that this species probably has several host plants. Distribution Ethiopia, Kenya and Tanzania. Discussion This species is similar to E. flavicornis, differing only in the color of the 1st flagellomere in the female. No consistent difference could be found in the terminalia. Moreover, females from the series collected in Njombe (Tanzania) have the antenna paler than the usual (brown or brownish), and thus resemble closely specimens of E. flavicornis collected at the same time and place. Additional data, especially on the host plants, are required to resolve the relationships between these species.

Elgonina yaromi Freidberg & Merz, n. sp. (Figs. 19, 33, 47, 61, 76, 91)

Diagnosis The only species of Elgonina that has the entire syntergite 1+2 covered by microtrichia; wing with 1-2 rather conspicuous white spots in cell r1; pterostigma bicolored. Description Male: Head. Antenna rather darkened, in particular 1st flagellomere black, scape and pedicel often infuscated, sometimes only pedicel dorsoapically pale; postocellar seta paler than medial vertical seta. Thorax. Scutum with rather dense microtrichia, underlying cuticule hardly visible; postalar seta short and yellowish; apical scutellar seta tiny, scarcely longer than scutal setulae. Legs. Hindtibia with black ring on basal half. Wing (Fig. 19). Generally evenly infuscated; pterostigma bicolored, with base yellowish and distal half brownish; cell r1 with 1-2 white spots; white apical spot present at wing tip. Abdomen. Syntergite 1+2, basal half of tergites 3 and basal 0.25 of tergites 4 and 5 with sparse microtrichia. Terminalia: Epandrium (Figs. 33, 47) with large serrate caudal flange; glans with conspicuous vesica (Fig. 61). Female. Similar to male, including darkened 1st flagellomere; anterior stripes of sparse

389 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 26-33. Epandrium, posterior view. 26. Elgonina pollinosa. 27. E. dimorphica. 28. E. inexpectata. 29. E. refulgens. 30. E. flavicornis. 31. E. infuscata. 32. E. splendida. 33. E. yaromi.

390 Freidberg and Merz / Revision of the Gymnosagena group

Figs. 34-39. Epandrium, posterior view. 34. Gymnosagena campiglossina. 35. G. kakamega. 36. G. longicauda. 37. G. unicornuta. 38. Marriottella exquisita. 39. M. sepsoides. microtrichia on tergites 3-5 narrower. Terminalia: Oviscapal measure about 3; aculeus tip with preapical step (Fig. 76); spermatheca with rather large tubercles (Fig. 91). Wing length. Male: 2.2-2.5 mm; female: 2.3-2.6 mm. Material examined Holotype , TANZANIA: 1800m, Usambara Mts., Rt. B124, Gologolo, 12-13.ix.1992, A. Freidberg. The holotype is double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in TAUI.

391 BIOTAXONOMY OF TEPHRITOIDEA

Paratypes: Same collecting data as holotype (24, 26; TAUI); same collecting data as holotype, but also: ex flowerhead Conyza newii, 15.ix.1992 (1; TAUI); Usambara Mts., Gologolo, 1900m, 23.viii.1996, A. Freidberg (12, 6; TAUI); same collecting data, but also: ex flowerhead Conyza newii, 27-28.viii.1996 (2; TAUI). KENYA: 10 km North Kabarnet, 29.xi.1989, A. Freidberg & F. Kaplan (1; TAUI); Equator, 30.x.[19]83, I. Yarom (16, 19 ; TAUI); KENYA (West): Timboroa, 26.xi.1989, A. Freidberg & Fini Kaplan (2, 1; TAUI); 10km North Kabarnet, 29.xi.1989, A. Freidberg & Fini Kaplan (1; TAUI); Nabkoi, 50km SE Eldoret, 2550m, 15.iii.1993, B. Merz (4, 2; MHNG); Tarakwa, 35km SE Eldoret, 2350m, 15.iii.1993, B. Merz (1; MHNG). UGANDA: S.W. Ichuya Forest, Kanaba Gap, 2500m, 25.xii.1995, I. Yarom & A. Freidberg (11, 7; TAUI); same collecting data, but also: ex flowerhead Conyza newii?, 1-5.i.1995 (3, 2; TAUI). Etymology Named after our colleague, Dr. Ilan Yarom, who collected part of the type series. Biology Reared from flowerheads of Conyza newii. All specimens were collected between 1800- 2800 m. Distribution Kenya, Uganda and Tanzania. Discussion This species is readily distinguished from its congeners in both sexes by the characters given in the key, of which the entirely microtrichose syntergite 1+2 and the 2 conspicuous white spots in cell r1 are the best diagnostic characters. The male terminalia are similar to those of other species of the splendida group.

Elgonina sp. Two females labelled: CAMEROON: Mt. Cameroon, 2100m, 11-13.xi.1987, A. Freidberg (TAUI), extend the known distribution of the genus considerably toward the west. They belong to the splendida group by lacking the posterior orbital seta and apical scutellar seta and by the postalar seta being small and brownish. In the key to species, they run to E. yaromi, from which they differ by not having any white spots in cell r1. In addition, the aculeus is tapered more strongly preapically, at level of the lateral projection of sternite 8, and widened again more distally before finally narrowed toward the very tip. A preapical step is visible with difficulty in these intact specimens. Males and knowledge about the host plants are needed in order to clarify the taxonomic status of these specimens.

Gymnosagena Munro Gymnosagena Munro, 1935: 5. Type species: Gymnosagena unicornuta Munro, 1935, by original designation and monotypy); Munro, 1947: 244 (Generic relationships); Munro, 1957: 890 (comparison with Elgonina); Cogan & Munro, 1980: 541 (Afrotropical Catalog); Norrbom et al., 1999: 156 (World Catalog). Diagnosis Small flies (wing length 1.8-3.0 mm), predominantly black, with most of head, part of legs and

392 Freidberg and Merz / Revision of the Gymnosagena group halter yellow; 1-2 orbital setae; postocular setae mixed pale and dark; 1-2 scutellar setae; wing pattern reticulate, Campiglossa-type, usually dark, sometimes pale or even reduced; tergites mostly shiny black, with brownish setulae, and in two species also with some coarse, whitish setulae; preglans part of distiphallus not setulose and without lobes; glans rather strongly sclerotized, with a basal cylindrical acrophallus and a narrow tube extending into vesica; aculeus evenly pointed; tip with preapical step; sternite 8 without lateral projection extending beyond margin of tergite 8. Redescription Head (Figs. 4-5). Structure: Slightly higher than long. Occiput moderately prominent. Frons flat, usually about as wide as long, margins subparallel. Fronto-facial angle about 100º-135º, distinctly projecting anterior to eye. Frontal stripe bare. Face slightly concave, without facial ridge or groove, ventral facial margin barely to moderately protuberant. Parafacial narrow, at narrowest point about 1-2 times as wide as thickest part of arista. Gena about 0.50-0.75 times as high as antenna, about 0.15 times as high as eye. Eye suboval, about 1.3 times as high as long, without setulae. Antenna slightly shorter than face; 1st flagellomere usually rounded at apex, sometimes slightly pointed dorsoapically, about twice as long as pedicel, and about twice as long as high; rays of arista shorter than diameter of arista at base. Proboscis capitate or spatulate. Palpus normal, about as broad as gena at lowest point. Coloration and vestiture: Head predominantly yellow, covered by dense microtrichia; occiput, postgena and ocellar triangle black, microtrichose. Antenna yellow, scape and pedicel dorsally with short setulae; second aristomere yellow; third aristomere yellowish at base, darker towards apex. Palpus yellow, with some dark, spiny setulae on apical half; labellum yellow. Chaetotaxy: 1-2 orbital setae, usually black, posterior seta, if present, about 0.5-0.7 times as long as anterior seta, sometimes yellow; 2 black frontal setae, 1 black ocellar seta, 1 black medial vertical seta, 1 yellow lateral vertical seta, 1 yellow postocellar seta, paravertical seta setula-like, yellowish or blackish; postocular setae mixed dark and acuminate, with 2-3 longer and whitish setae near middle of row; genal seta and all postgenal and occipital setulae yellowish. Thorax. Structure, coloration and vestiture: Scutum rather flat, about 1.11-1.12 times as long as wide. Scutellum about 0.3 times as long as scutum, 0.43 times as long as wide. Mesonotum dull black, microtrichia gray to brown, dense, slightly more conspicuous on scutum than on scutellum. Mesonotum with whitish and coarse to brown and fine setulae arranged somewhat irregularly. Propleuron with fan of 2-5 whitish setulae. Prosternum with some long, white setulae. Chaetotaxy (major setae mostly black): Scapular setae more or less well differentiated; 1 postpronotal seta, 2 notopleural setae, posterior seta sometimes dark brown, about 0.50-0.66 times as long as anterior seta, 1 presutural seta, 1 supra-alar seta, 1 postalar seta, short and whitish to brown or long and black, 1 long intra-alar seta, 1 dorsocentral seta, aligned slightly but distinctly posterior to suture, 1 prescutellar acrostichal seta, aligned somewhat anterior to intra-alar seta, 1 basal scutellar seta, apical scutellar seta someties lacking, when present no more than about 0.25 times as long as basal seta, brown, 1 black anepisternal seta, 1 black katepisternal seta, 1 yellow anepimeral seta. Halter yellow. Calypteres yellow with yellow fringe, ventral calypter narrow, stripe-like. Legs. Without overt features, entirely yellow or partly blackish, in latter case coxae with dense, gray microtrichia; trochanters yellow or brown; femora usually predominantly black, with yellow apex, microtrichose, but somewhat shiny; tibiae yellow; tarsi yellow, sometimes with darkened distal tarsomere.

393 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 40-47. Epandrium, lateral view (most species also with syntergosternum 6-8). 40. Elgonina pollinosa. 41. E. dimorphica. 42. E. inexpectata. 43. E. refulgens. 44. E. flavicornis. 45. E. infuscata. 46. E. splendida. 47. E. yaromi.

394 Freidberg and Merz / Revision of the Gymnosagena group

Figs. 48-53. Epandrium, lateral view. 48. Gymnosagena campiglossina. 49. G. kakamega. 50. G. longicauda. 51. G. unicornuta. 52. Marriottella exquisita. 53. M. sepsoides.

Wing (Figs. 20-23). Anal lobe well developed. Venation: Veins darkened within dark spots of pattern. Pterostigma ratio varies from 2.2-2.7, and pterostigma-costal-cell ratio varies from

0.52-0.72. Longitudinal veins mostly straight, although vein R2+3 sometimes slightly sinuous; distal sections of veins R4+5 and M parallel or slightly divergent; only vein R1 setulose dorsally, with gap at level of end of subcostal vein, other veins bare; crossvein R-M situated approximately opposite distal 0.66 of cell dm; distance between crossveins about 1.5 times as long as crossvein DM-Cu; cell bcu with short, sometimes indistinct posterodistal lobe. Costal spine short, but slightly longer than surrounding setulae. Pattern: reticulate, Campiglossa-type: usually with several conspicuous dark areas, in particular area over pterostigma extending to, or

395 BIOTAXONOMY OF TEPHRITOIDEA

nearly to, crossvein R-M, or subbasally across cell r2+3 to preapical part of cell br; area over crossvein DM-Cu; and area occupying apex of cells r1, r2+3, r4+5 and m, containing also some large and small hyaline spots; pattern occasionally more or less reduced. Abdomen. Syntergite 1+2 partly microtrichose; anterior margin of tergites 3-5 often microtrichose; setulae mostly brownish; tergites 3-6 sometimes with long whitish marginal setae; setae usually about as long as setulae, sometimes distinctly longer. Male terminalia: Sternite 5 apically excised. Syntergosternite 6-8 uniform in all species. Epandrium oval in posterior view (Figs. 34-37), narrow and parallel-sided in lateral view (Figs. 48-51), more or less pointed ventrally; caudal flange large; lateral surstylus rather broad, convex posteriorly; medial surstylus with 2 conspicuous setae on medial surface and two brownish prensisetae, lateral prensiseta smaller than medial, usually directed ventrally. Hypandrium semicircular, with or without lappet-like sclerotizations at articulation with epandrium. Phallapodeme triangular in lateral view, asymmetrical as in other Tephritini, with additional branch articulating with hypandrium on one side. Ejaculatory apodeme almost as long as epandrium, distally expanded. Basiphallus short, shovel-like. Distiphallus at preglans part without setulae (Figs. 62-65). Glans short or elongate, with distinct vesica; elongate internal sclerotized tube- like structure protruding from sclerotized sheath. Female terminalia: Oviscape shiny black, covered by fine brown setulae; oviscapal measure 2-4. Aculeus gradually tapered towards tip, not arrow-like, tip with preapical step (Figs. 77-80). Spermatheca pyriform to nearly cylindrical, bearing variable number of tubercles on surface (Figs. 92-95). Biology The known host genera are: Matricaria L. (Anthemideae), Nidorella Cass. and Psiadia Jacq. (Astereae), and Thouars () – a plethora of taxa unusually diverse for such a small and homogenous group (Table 1, p. 415). Distribution The genus is distributed mainly from the highlands of to southern Africa, and on Madagascar. Discussion Gymnosagena is similar to Elgonina, especially structurally (such as in the head shape); but differs in the more complex wing pattern. Gymnosagena is also similar to Spathulina based on the shiny black abdomen and the reticulate wing pattern; but differs from this genus in the black posterior notopleural seta, details of the wing pattern (in Gymnosagena there are three hyaline costal spots in cell r1, and the dark apical fork is less discrete, whereas in Spathulina there are usually 0-2 such costal spots, and the apical fork is usually discrete), and in the generally shorter labella. Gymnosagena is also similar to Campiglossa, based on plesiomorphic characters, such as the wing pattern; but differs from this genus by the shiny black abdomen and lack of setulae on the preglans part of the distiphallus.

Key to the species of Gymnosagena 1. Wing pattern reduced and faint, only pterostigma distinctly dark (Fig. 23); 1 orbital seta and 1 scutellar seta present; legs entirely yellow, or femora mostly black ...... 2 -. Wing pattern dark and extensive (Figs. 20-22); 2 orbital and 2 scutellar setae usually present; legs partly black, femora mostly black ...... 3

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2. Legs usually entirely yellow, femora at most with small brown spots; abdomen with whitish, lanceolate setae (South Africa, ; hosts: Nidorella resedefolia and Matricaria nigellifolia) ...... G. unicornuta Munro -. Femora predominantly black; abdomen with pale brown setae and setulae, without whitish, lanceolate setae (Malawi; hosts unknown) ...... G. nyikaensis, n. sp. 3. Wing pattern uneven, with two large and more or less isolated dark areas (one around pterostigma and one preapically), mostly hyaline base (extending to oblique line from base

of pterostigma, through basal third of cell dm, to near apex of cell cu1) and large, predominantly hyaline area between these two dark areas; black area from pterostigma to

vein R4+5 widened posteriorly (Kenya, Zimbabwe, Madagascar) ...... G. kakamega, n. sp. -. Wing pattern more or less even, without such distinct and isolated dark and hyaline areas...... 4 4. Pterostigma and pterostigmal spot about as long as crossvein DM-Cu; abdomen with some whitish, lanceolate setae or setulae; posterior orbital seta pale brown; abdominal tergites 1- 3 entirely and tergite 4 medially subshiny, microtrichose; oviscapal measure about 2 (Madagascar; host: Psiadia salviiflora)...... G. campiglossina, n. sp. -. Pterostigma and pterostigmal spot about 1.5 times as long as crossvein DM-Cu; abdomen without any whitish, lanceolate setae; posterior orbital seta usually dark brown, rarely pale, minute or absent; only anterior 0.7 of syntergite 1+2 sparsely microtrichose, posterior tergites shiny black, sometimes except extreme base; oviscape about as long as preabdomen (Uganda; host: Aspilia mossambicensis) ...... G. longicauda, n. sp.

Gymnosagena campiglossina Freidberg & Merz, n. sp. (Figs. 20, 34, 48, 62, 77, 92) Diagnosis This species is distinguished from all other congeners by the following combination of characters: An evenly and strongly reticulate wing pattern; 2 orbital setae of which the posterior seta is pale brown; 2 scutellar setae; abdominal tergites with some white lanceolate setulae along posterior margin; and oviscape much shorter than preabdomen. Description Similar to G. unicornuta, except as noted below. Head. Ventral facial margin slightly protuberant; 2 orbital setae present, anterior seta blackish, posterior seta pale brown, lanceolate; 1st flagellomere yellow to brownish; proboscis capitate or short spatulate. Thorax. Mesonotum with predominantly brown microtrichia, darker dorsocentral and median vittae, gray microtrichia only at anterior margin, and bases of major setae in dark spots; setulae yellowish and coarse; major setae on mesonotum black; postalar seta short and brown; apical scutellar seta brown, about 0.2 times as long as basal seta. Legs. Predominantly yellow, but all femora black on about basal 0.6-0.8. Wing (Fig. 20). Pattern reticulate, rather uniform, similar to that of G. unicornuta, but blackish, not predominantly grayish, with distinct markings on, and basal to level of, basal crossveins; pterostigmal spot about 1.5 times as long and as wide as hyaline spot in cell r1 distal to it; hyaline

397 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 54-61. Distiphallus. 54. Elgonina pollinosa. 55. E. dimorphica. 56. E. inexpectata. 57. E. refulgens. 58. E. flavicornis. 59. E. infuscata. 60. E. splendida. 61. E. yaromi.

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Figs. 62-67. Distiphallus. 62. Gymnosagena campiglossina. 63. G. kakamega. 64. G. longicauda. 65. G. unicornuta. 66. Marriottella exquisita (two enlargements). 67. M. sepsoides.

spot at end of vein R2+3 extends to vein R4+5; hyaline spot at apex of cell r4+5 relatively small, about half as wide as cell. Pterostigma ratio about 2.5; pterostigma-costal-cell ratio about 0.57. Abdomen. Tergites 1-4 in both sexes entirely or at least anteriorly and laterally fine grayly microtrichose; tergites 3-4 posterolaterally with few conspicuous, white, lanceolate setulae. Male terminalia: Epandrium (Figs. 34, 48) similar to that of G. unicornuta, although somewhat different in proportions; lateral surstylus widened apically in posterior view (Fig. 34); medial surstylus rather broad, with apical part apparently not strongly fused to basal part, prensisetae subequal; acrophallus of glans opens distally in wide tube (Fig. 62), medial duct shorter than in

399 BIOTAXONOMY OF TEPHRITOIDEA other studied species. Female terminalia: Aculeus similar to that of G. unicornuta, but preapical step less conspicuous (Fig. 77). Spermatheca (Fig. 92) pyriform, with pointed tubercles. Wing Length. Male: 1.9-2.2 mm; female: 1.8-2.4 mm. Material Examined Holotype , MADAGASCAR: C[enter], Ivato, Rt. 7, 1600m, 15kmS Ambositra, 25.iv.1991, A. Freidberg & F. Kaplan (TAUI). The holotype is double mounted (minuten in a block of plastic), is in excellent condition, and is deposited in TAUI. Paratypes: same collecting data as holotype (44, 26; TAUI); same data, but also ex flowerhead of Psiadia salviiflora (Asteraceae), v.1991 (1; TAUI). C[enter], Rt. 7, 35KmS Ambositra, 27.iv.1991, A. Freidberg & F. Kaplan (5, 12; TAUI); C[enter], Rt. 45, 30kmNE Fianarantsoa, 26.iv.1991, A. Freidberg & F. Kaplan (11, 4; TAUI); Fia., Ambalamanakana, 18.i.[19]58, F. Keiser (2, 5; NHMB); Ivato, 17.i.[19]58, F. Keiser (2; NHMB); Tan., Ambatolampy, 1.i.[19]58, F. Keiser (1; NHMB); Manjakatompo, 6.i.[19]58, F. Keiser (1; NHMB); Andriampilany, 31.xii.[19]57, F. Keiser (1; NHMB); Nord, Prairie de lisières, 840m, Joffreville Diego Suarez, 4.xii.[19]57, B. Stuckenberg (1; NMP); Centre, Rivière Zomandoa, 1650m, Andringitra Ambalavao, 10.i.[19]57, B. Stuckenberg (1; NMP); dct Ambatolampy, Manjakatompo, 1700m, 11-15.xii.[19]57, B. Stuckenberg (1; NMP); Col Mahafompeno, 2200-2400m, dct Ambatolampy, 11-15.xii.[19]57, B. Stuckenberg (1, 2, NMP); Plateau Soaindrana, 2060m, Andringitra-Ambalavao, 14-17.i.[19]58, B. Stuckenberg (5, 2; NMP); Ankaratra Massif, Manjakatompo, forest station, i.1956, B. Stuckenberg (1, 3; NMP); Ankaratra Massif, Tsiafajavona Peak, i.1956, B. Stuckenberg (2, 5; NMP); Ankasoka, 1130m, Route Lakato, 2.xii.[19]56, P. Griv. (1; IST); La Mandraka, 1250m, Manjakandriana, 30.x.[19]56, E.R. (1, 2; IST); Pèrinet (1; IST). Etymology Named after the similarity to species of Campiglossa. Biology Most of the specimens of the type series were collected on Psiadia salviiflora, and one paratype was reared from flowerheads of this plant. Distribution Madagascar. Discussion This species and specimens temporarily assigned to the next species are the only known representatives of the group in Madagascar.

Gymnosagena kakamega Freidberg & Merz, n. sp. (Figs. 21, 35, 49, 63, 78, 93) Diagnosis Differs from other congeners by the 3 discrete dark areas in the wing pattern (Fig. 21), 2 concolorous orbital setae and 2 scutellar setae of which the apical seta is much shorter. Description Similar to G. campiglossina, except as noted below.

400 Freidberg and Merz / Revision of the Gymnosagena group

Head. Ventral facial margin barely or not protuberant; 2 acuminate orbital setae present, anterior seta blackish, posterior seta brownish; 1st flagellomere yellow; proboscis capitate or short spatulate. Thorax. Microtrichia gray, with slight tinge of brown on dorsum, although in the specimen from Manjakatompo more pronounced and similar to G. campiglossina; setulae brownish and fine; major setae on mesonotum black; postalar seta long and black; apical scutellar seta acuminate but yellowish, 0.15 times as long as basal seta. Legs. Predominantly yellow, but femora black on basal 0.5-0.8, least blackened on midfemur and usually on anterior aspect of forefemur and hindfemur. Wing (Fig. 21). Pattern blackish, reticulate, but less uniform than in other congeners, with 3 large dark areas more obvious than in other congeners: one at pterostigma, extending to vein M, one around crossvein DM-Cu, and one occupying apical 0.25 of wing; markings on basal 0.33 of wing, including most of cell cu1, grayish, rather indistinct; dark pterostigmal spot widened posteriorly and contains small hyaline spots only in cells br and r4+5; large apical area contains mostly smaller hyaline spots, and often larger spot at end of vein R2+3 ; hyaline spot at apex of cell r4+5 even larger, almost as wide as cell. Pterostigma ratio about 2.7; pterostigma-costal-cell ratio about 0.72 (the highest ratios for the genus). Abdomen. Syntergite 1+2 mostly (except posterolaterally) and tergites 3 and 4 basally grayly microtrichose; setae and setulae brown, fine; without whitish, lanceolate setae. Male terminalia: Epandrium (Figs. 35, 49) similar to G. campiglossina, although lateral surstylus less distinctly separated from epandrium, and medial surstylus more evenly concave (Fig. 35); glans (Fig. 63) with long sclerotized tube extending deeply into vesica. Female terminalia (Figs. 78, 93): Almost identical to G. campiglossina; oviscapal measure about 2.5. Wing length. Male: 2.1-2.4 mm; female: 2.3-2.6 mm. Material Examined Holotype , KENYA: Kakamega Forest, 20-21.xi.1986, A. Freidberg (TAUI). The holotype is double mounted (minuten in a plastic block), is in excellent condition, and is deposited in TAUI. Paratypes: KENYA: 25km NE Kericho, 17.xi.1986, A. Freidberg (1; TAUI); ZIMBABWE [S. RHODESIA]: N[orth] Vumba, 21, 24.ix.1964, D. Cookson (2; NMP). Additional material, not included as paratypes: MADAGASCAR: Tan., Manjakatompo, 4.i.[19]58, F. Keiser (1; NHMB); La Mandraka, 1250m, Manjakandriana, 30.x.[19]56, E.R. (1, 1; IST). Etymology Named after the type locality, Kakamega forest, a noun in apposition. Biology Unknown. Distribution Kenya, Zimbabwe; possibly Madagascar. Discussion The specimens from Madagascar are similar to those from mainland Africa but differ

401 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 68-76. Aculeus (whole – left; enlarged tip – right). 68. Elgonina pollinosa. 69. E. dimorphica. 70. E. fuscana. 71. E. inexpectata. 72. E. refulgens. 73. E. flavicornis. 74. E. infuscata. 75. E. splendida (inversion membrane and a third, lateral view also included). 76. E. yaromi.

402 Freidberg and Merz / Revision of the Gymnosagena group

Figs. 77-82. Aculeus (whole – left; enlarged tip – right). 77. Gymnosagena campiglossina. 78. G. kakamega. 79. G. longicauda. 80. G. unicornuta. 81. Marriottella exquisita. 82. M. sepsoides. primarily in the dark pterostigmal spot, which is parallel-sided and not widened posteriorly. These specimens therefore are not included in the type series.

Gymnosagena longicauda Freidberg & Merz, n. sp. (Figs. 4, 22, 36, 50, 64, 79, 94)

Diagnosis This species is distinguished from all other congeners by the following combination of characters: Wing pattern evenly dark on entire surface; 2 orbital setae present, the posterior seta short and of variable color; 2 scutellar setae, the apical seta fine and short; abdominal tergites shiny black, only basal 0.7 of syntergite 1+2 grayly microtrichose; lateral margins of tergites 3

403 BIOTAXONOMY OF TEPHRITOIDEA and 4 without white, lanceolate setae. The female of this species has the oviscape as long as the preabdomen, the longest among all Gymnosagena species. Description Similar to G. campiglossina except as noted below. Head (Fig. 4). Ventral facial margin barely or not protuberant; 1st flagellomere yellow; proboscis capitate; 2 orbital setae, anterior seta acuminate, blackish, posterior seta yellowish, brownish or black, often varying in the same specimen. Thorax. Grayish microtrichose, with slight tinge of brown on dorsum and with 3 indistinctly browner stripes in middle and dorsocentrally when viewed from in front; setulae brownish and fine; postalar seta long and black; apical scutellar seta acuminate, brown to black, 0.15-0.20 times as long as basal seta. Legs. Predominantly yellow, but coxae black, grayish microtrichose, and femora black on basal 0.65-0.90, more on forefemur than on other femora and slightly more on posterior aspect than on anterior aspect. Wing (Fig. 22). Pattern blackish, reticulate, less uniform than in C. campiglossina and more uniform than in C. kakamega, with 3 large and more or less discrete dark areas: one at pterostigma, extending to vein m, one on crossvein DM-Cu and nearby, and one occupying apical 0.25 of wing; markings on basal 0.33 of wing, including most of cell cu1, almost as dark as remaining pattern; dark pterostigmal spot widened posteriorly and contains small hyaline spots only in cells br and r4+5; large apical area mostly contains medium-sized hyaline spots, including spot at end of vein R2+3, but hyaline spot at apex of cell r4+5 larger, almost as wide as cell. Pterostigma ratio about 2.6; pterostigma-costal-cell ratio about 0.68. Abdomen. Basal 0.7 of syntergite 1+2 grayly microtrichose; anterior margins of tergites 3 and 4 sometimes narrowly microtrichose; setae and setulae brown, fine; without whitish, lanceolate setae at posterolateral margins of tergites 3 and 4. Male terminalia: Epandrium and surstyli (Figs. 36, 50) almost identical to G. campiglossina; glans similar to G. kakamega (Fig. 64), with acrophallus forming a basal sclerotized sheath and distally a narrow duct which is penetrating into the long and narrow vesica; this tube shorter than in G. kakamega, but vesica narrower and longer. Female terminalia: Oviscapal measure 5. Aculeus similar to but more elongate than that of G. campiglossina (Fig. 79). Spermatheca (Fig. 94) cylindrical, with pointed tubercles. Wing length. Male: 2.3-2.6 mm; female: 2.6-3.0 mm. Material Examined Holotype , UGANDA: S.W. Matiri, 70kmE Fort Portal, 17.i.1996, I. Yarom & A. Freidberg. The holotype is double mounted (minuten in a plastic block), is in excellent condition, and is deposited in TAUI. Paratypes: Same collecting data (20, 6; TAUI, MHNG); same collecting data, but also ex flowerhead Aspilia mossambicensis (1, 1; TAUI). UGANDA: S.W. Kabale-Katuna Rd., 1900m, 23.xii.1995, I. Yarom & A. Freidberg (1; TAUI). Etymology Named after the long ovipositor.

404 Freidberg and Merz / Revision of the Gymnosagena group

Biology Reared from flowerheads of Aspilia mossambicensis. Distribution Uganda. Discussion This species is characterized by the long ovipositor, the longest in the entire group, and by its host plant, Aspilia mossambicensis, which is the only representative of the tribe Heliantheae among hosts of this group. Otherwise, it is a typical Gymnosagena.

Gymnosagena nyikaensis Freidberg & Merz, n. sp.

Diagnosis This species is similar to G. unicornuta, differing from it in the predominantly black femora, lack of whitish, lanceolate setae on the abdomen and more reduced wing pattern. Description Similar to G. unicornuta except as noted below. Head. Frontofacial angle 135º; gena about 0.75 times as high as antenna; ventral facial margin not protuberant; proboscis capitate or short spatulate; only one, right, orbital seta present, dark brown; left orbital seta missing. Thorax. Microtrichia less conspicuous, and thorax appearing distinctly darker; postalar seta short and yellow. Legs. Coxae blackish; femora predominantly black, with only apical 0.2 of forefemur, 0.4 of midfemur and 0.25 of hindfemur yellow; remaining parts mostly yellow, but distalmost tarsomeres brownish. Wing. Almost entirely grayish hyaline, with reduced dark pattern comprising following spots: almost entirely black pterostigma (with basal 0.25 hyaline), extending as blackish crescent across vein R1 to or almost to vein R2+3; three gray spots along cell r1, appearing darker on costa, narrower than hyaline spots between them, distalmost gray spot extending (vaguely) into cell r2+3; similar spot over apex of vein R4+5, mostly in cell r2+3, bordered by large hyaline area extending across cell r2+3 preapically and forming round hyaline spot at apex of cell r4+5; most of wing otherwise lightly grayish, but no dark spots present around crossveins R-M and DM-Cu. Other details of pattern difficult to discern partly due to its reduction and partly due to dirt cover. Pterostigma ratio about

2.5; pterostigma-costal-cell ratio about 0.58. Veins R4+5 and M slightly but distinctly divergent. Abdomen. Syntergite 1+2 grayly microtrichose, except on posterior margin; anterior margins of tergites 3-5 medially slightly microtrichose; Setae and setulae brown, fine; without whitish, lanceolate setae. Terminalia not studied. Wing length. 2.33 mm. Material Examined Holotype , MALAWI: North, Nyika National Park, 2000m, 10kmS Chelinda, 24.ix.1998, F. Kaplan & A. Freidberg. The holotype is double mounted (minuten in a plastic block), is in good condition, and is deposited in TAUI.

405 BIOTAXONOMY OF TEPHRITOIDEA

Etymology Named after the type locality, the Nyika Plateau in Malawi. Biology Unknown. Distribution Malawi. Discussion Although this species is a typical member of Gymnosagena, the reduced wing pattern makes it superficially similar to species of Elgonina. Wing and terminalia preparations have not been made in order to preserve the single available specimen as intact as possible

Gymnosagena unicornuta Munro (Figs. 5, 23, 37, 51, 65, 80, 95) Gymnosagena unicornuta Munro, 1935: 5; Cogan & Munro, 1980: 541 (Afrotropical Catalog); Hancock, 1986: 29 (Zimbabwe record); Norrbom et al, 1999: 156 (World Catalog). Diagnosis This species is distinguished from all other congeners by the following combination of characters: Only 1, black, orbital seta; apical scutellar seta absent; legs yellow; and white, lanceolate setulae at posterior margin of tergites 3-4 present. Redescription Similar to G. campiglossina except as noted below. Head (Fig. 5). Fronto-facial angle about 120º-135º; gena about 0.6-0.7 times as high as antenna; ventral facial margin moderately protuberant; 1st flagellomere yellow; proboscis spatulate; only anterior orbital seta present. Thorax. Mesonotum silvery gray, setulae whitish and coarse; postalar seta short and whitish; only basal scutellar seta present. Legs. Entirely yellow including coxae, or femora irregularly brownish at about basal 0.5. Wing (Fig. 23). Pattern reduced and generally pale, with following darker and more conspicuous areas: blackish pterostigmal area, extending into cell r1, and slightly crossing vein

R2+3; three lighter, more or less equally spaced spots along cell r1; distalmost spot wider, filling apex of cell, comprising part of large preapical area that extends clearly at least to middle of cell r4+5; apical spot comprising wide apical fork over tips of veins R4+5 and M; remaining pattern gray, with relatively non-contrasting hyaline spots, not extending more proximally than level of basal crossveins; crossveins R-M and DM-Cu clearly within gray areas. Pterostigma ratio about

2.2; pterostigma-costal-cell ratio about 0.52 (the lowest ratios for the genus). Veins R4+5 and M parallel or slightly divergent. Abdomen. Microtrichia present on entire syntergite 1+2 and more or less narrowly on anterior margins of tergites 3-5, except medially, where microtrichia sometimes occupy entire width of tergite; coarse whitish setae and setulae conspicuous along posterior margins of tergites 3-4 in

406 Freidberg and Merz / Revision of the Gymnosagena group male, 3-6 in female, occasionally few on tergite 2, especially in female. Male terminalia: Epandrium and surstyli (Figs. 37, 51) as in G. campiglossina although lateral surstylus tapered, not widened, in posterior view (Fig. 37) and lateral prensiseta relatively smaller; glans (Fig. 65) sclerotized almost to apex, duct emerging from acrophallus short; vesica relatively shorter than in other studied congeners. Female terminalia: Oviscapal measure 2.0-2.5; aculeus (Fig. 80) gradually narrowed at distal half, with distinct preapical step; spermatheca (Fig. 95) with moderately large number of short and long tubercles. Wing length. Male: 2.0-2.2 mm; female: 2.3-2.5 mm. Material Examined Lectotype (here designated) labeled: “Mazoe, S.R. [= SOUTH RHODESIA = Zimbabwe], Aug. 1932, W.K. Ford/ Type. Gymnosagena unicornuta Mro., det. HKMunro 1934 (on red label)/ M. 372". This lectotype is designated in order to fix the status of the species. The lectotype is double mounted (minuten and a polyporus block), is in good condition, and is deposited in SANC. Paralectotypes: same labels as lectotype, except female labelled as paratype (on yellow) (1, 1; SANC). Additional specimens: [Zimbabwe:] Salisbury, S.R., Dunstan Estate, Mar. 1951., H. K. Munro. M. 1071. (3, 1; SANC). SOUTH AFRICA: Natal, 10kmNE Drumond, 18.ix.[19]83, A. Freidberg (20, 9; TAUI); same data, but also: ex flowerhead Matricaria nigellifolia, ix.1983 (9; TAUI); Cape Province, 8Km N Fort Beaufort, Blinkwater, 32º43’S 26º35’E, 750m, 21.x.1994, R. Danielsson (1; NRS). Biology Reared from flowerheads of Nidorella resedifolia (Munro, 1935) and Matricaria nigellifolia. Distribution Zimbabwe, South Africa. Discussion Munro’s (1935) original description is generally adequate. However, he erred in noting that the frons is 2.5 times as wide as the eye; it is actually only 1.5 times as wide. Furthermore, in addition to the extreme base of the abdomen that he recorded as being microtrichose, sparse microtrichia are also present medially and on the anterior margins of most terga. The terminalia are described and illustrated here for the first time. In addition to the eleven paratypes, Munro (1935) recorded in his original description: “Type, and ,”, although no holotype was labeled as such. Of this type series we have studied a male and female labeled as “Type” and another female labeled as “paratype”; we selected the former female as lectotype.

Marriottella Munro Marriottella Munro, 1939: 147. Type species: Marriottella exquisita Munro, 1939, by original designation; Steyskal, 1979: 6 (Review of ); Norrbom et al., 1999: 165 (World catalog); Freidberg and Norrbom, 1999: 582 (Revision of Myopitini). Marriotella (misspelling). Cogan and Munro, 1980: 523 (Afrotropical catalog); Freidberg, 1984: 133 (Gall Tephritidae).

407 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 83-91. Spermatheca. 83. Elgonina pollinosa. 84. E. dimorphica. 85. E. fuscana. 86. E. inexpectata. 87. E. refulgens. 88. E. flavicornis. 89. E. infuscata. 90. E. splendida. 91. E. yaromi.

Diagnosis Small flies (wing length 1.6-3.4 mm), predominantly black, including palpus and halter, with parts of head and legs yellow to brown; 1 orbital seta; postocular setae dark or pale; 1 scutellar seta; wing predominantly hyaline, with blackish costal cells, pterostigma and subapical spot,

408 Freidberg and Merz / Revision of the Gymnosagena group

Figs. 92-97. Spermatheca. 92. Gymnosagena campiglossina. 93. G. kakamega. 94. G. longicauda. 95. G. unicornuta. 96. Marriottella exquisita. 97. M. sepsoides. and with white apical spot; tergites shiny or subshiny black, with setulae coarse and whitish, or fine and brown; distiphallus with two large spines at preglans part or with few small spines at preglans part and two large spines near middle; aculeus more or less tapered evenly, pointed, tip attenuated, without preapical step; sternite 8 without lateral projection. Marriottella can easily be separated from related Tephritini genera by using the key presented in this publication. Superficially it also resembles some Myopitini genera (such as Asimoneura Czerny and Freidberg and Norrbom) by the shiny black body, single orbital seta, elongate proboscis, cell bcu without posterodistal lobe, and reduced wing pattern. However, it differs from all Myopitini genera by having only the basal scutellar seta. Redescription Head (Figs. 6-7, 10-11). Structure: Higher than long, with prominent occiput. Frontofacial angle 90º-110º. Frons bare, about as wide as long or wider than long, margins subparallel. Face strongly protuberant, especially ventrally, slightly concave, without facial ridge or groove. Eye suboval, about 1.5-1.7 times as high as long, with short, sparse setulae. Gena about 0.20-0.25 times as high as eye. Parafacial about half as wide as 1st flagellomere. Antenna elongated; 1st flagellomere about 2.5-3 times as long as high, attenuated, tip pointed or nearly so; scape and

409 BIOTAXONOMY OF TEPHRITOIDEA pedicel dorsally with short setulae; arista with rays shorter than diameter of arista at base. Proboscis spatulate or geniculate, labellum about 0.4-0.5 times as long as vertical diameter of eye; palpus broad. Coloration and vestiture: Predominantly brown to black, shiny or covered by more or less dense microtrichia (and then dull, except face). Arista more or less unicolored. Proboscis blackish; palpus black or blackish, with some spiny, dark setulae on apical half. Chaetotaxy: 1 black orbital seta, 2 black frontal setae, 1 black ocellar seta, 1 black medial vertical seta, 1 black lateral vertical seta, 1 postocellar seta, brown to black or white, paravertical seta setula-like and black or about as long as postocellar seta, lanceolate and yellowish; postocular setae dark and acuminate or yellow; genal and postgenal setae and setulae yellow to black. Thorax. Structure, coloration and vestiture: Scutum strongly convex, short, 1.02-1.03 times as long as wide. Scutellum short, triangular, dorsally convex, about 0.25 times as long as scutum and 0.36 times as long as wide. Thorax black, mostly shiny, or dull; microtrichia sparse, not obscuring entirely underlying black cuticule, or dense. Mesonotum with sparse brown or whitish setulae arranged irregularly. Propleuron with fan of 3-5 whitish setulae. Prosternum with some long, white setulae. Chaetotaxy (all major setae black): Scapular setae barely or well differentiated; 1 postpronotal seta, 2 notopleural setae, posterior seta about 0.5-0.7 times as long as anterior seta, 1 presutural seta, 1 supra-alar seta, 1 long and black or blackish postalar seta, 1 long intra-alar seta, 1 dorsocentral seta, almost on suture, 1 prescutellar acrostichal seta, aligned about halfway between dorsocentral seta and scutellum, 1 (basal) scutellar seta, apical seta lacking, 1 anepisternal seta, 1 anepimeral seta, 1 katepisternal seta. Halter blackish. Calypteres narrow, dorsal calypter mostly hyaline, but black basally, with yellow fringe, ventral calypter stripe-like, blackish on margin. Legs. Without overt features; coxae brownish or blackish, with dense, silvery microtrichia; trochanters black; femora, except tips, black, microtrichose, but ventrally often shiny; tibiae partly yellow, progressively darker toward hindtibia; hindtibia sometimes entirely black; tarsi more or less yellow with darkened distal tarsomeres. Wing (Figs. 24-25). Elongate (length to width ratio 2.8-3.0); anal lobe somewhat reduced.

Venation: Vein R1 setulose dorsally, with gap at level of end of subcostal vein; other veins bare; vein R1 runs close to vein C, especially at pterostigma; pterostigma unusually narrow, pointed apically; crossvein R-M situated opposite distal 0.66-0.75 of cell dm; distance between crossveins about equal to length of crossvein DM-Cu; cell bcu without posterodistal lobe; terminal sections of veins R4+5 and M divergent; longitudinal veins pale at base, otherwise dark, vein C black or blackish throughout. Costal spine short and undifferentiated, or well developed.

Pattern: Comprising only costal cells and pterostigma black; cell r4+5 apically with milky- hyaline spot delimited by blackish subapical spot. Abdomen. Black, shiny or dull, microtrichia uniform, setulae fine and brown or coarse and whitish, semi-erect; marginal setae on tergites barely longer than setulae. Male terminalia: Epandrium oval in posterior view; narrow, more or less parallel-sided in lateral view, with ventral end tapered; caudal flange well developed; lateral surstylus rather broad, distinctly curved; medial surstylus with 2 conspicuous setae on medial surface and 2 dark brown prensisetae, lateral prensiseta smaller than medial, directed ventrally; distiphallus with 2 large

410 Freidberg and Merz / Revision of the Gymnosagena group spines either near middle or at preglans part; in former case with few smaller spines at preglans part. Female terminalia: Oviscape shiny black, covered by fine blackish setulae, oviscapal measure 2. Aculeus more or less evenly pointed, sternite 8 without lateral projection, tip of aculeus attenuated, without preapical step. Two oval to clavate spermathecae present bearing variable number of long or short tubercles on surface. Biology One of the species infests flowerheads of Helichrysum Mill. (Inuleae) without inducing galls; the other induces terminal twig galls on another species of the same genus (Table 1, p. 415). Distribution South Africa. Discussion Both species of Marriottella have unusual habitus and characters for Tephritidae. Based on the cladistic analysis (Fig. 99), Marriottella is the sister group of Elgonina, and indeed the two genera share several important characters. Marriottella is restricted to South Africa, and it shares with Spathulina infestation of host plants that belong to the tribe Inuleae. Spathulina is especially species-rich in South Africa, and the relationships between the two genera should be investigated further.

Key to the species of Marriottella Munro 1. Larger species (wing length over 3.0 mm); proboscis spatulate (Fig. 6); labellum distinctly shorter than ventral margin of head; wing length to width ratio about 3.0 (Fig. 24); ultimate section of vein M 3 times as long as penultimate section; white apical spot occupies apex of

cells r2+3 and r4+5, bordered proximally by dark, narrow band; postocular setae white; ocellar triangle matt due to microtrichia, reaching at most halfway to lunule (South Africa; bud galls on Helichrysum splendidum) ...... M. exquisita Munro -. Smaller species (wing length less than 2.0 mm); proboscis geniculate (Fig. 7); labellum as long as ventral margin of head; wing length to width ratio about 2.8 (Fig. 25); ultimate section of vein M about 2.5 times as long as penultimate section; white apical spot occupies

apex of cell r4+5 only, bodered anteriorly by dark wide triangle that fills apex of cell r2+3; postocular setae dark; ocellar triangle large, shiny, reaching lunule (South Africa; flowerheads of Helichrysum probably dregeanum) ...... M. sepsoides, n. sp.

Marriottella exquisita Munro (Figs. 6, 10, 24, 38, 52, 66, 81, 96) Marriottella exquisita Munro, 1939: 148; Norrbom et al., 1999: 165 (World catalog). Marriotella exquisita (misspelling). Cogan & Munro, 1980: 523 (Afrotropical catalog); Freidberg, 1984: 133 (Biology). Diagnosis This species is unmistakable based on the characters given in the key. The elongated wing with the unusually large white spot at the wing tip, which is proximally bordered by a narrow black crossband, in combination with the chaetotaxy, are especially diagnostic within the Tephritidae.

411 BIOTAXONOMY OF TEPHRITOIDEA

Redescription Head (Figs. 6, 10). Coloration and vestiture: mostly brownish-black; frons, parafacial, and pedicel brown, face subshiny; ocellar triangle blackish, but borders not always sharp, 0.4 times as long as frons; most head parts with fine white microtrichia; microtrichia less conspicuous on mesofrons and face; setulae mostly yellowish. Structure: frons width to eye width ratio 2.5-3.0; frontofacial angle about 100o, angulate and moderately strongly protuberant; parafacial 0.33, gena 1.5 times as wide (or as high) as 1st flagellomere; face in profile concave, with ventral margin strongly projected, but ventral half of face medially convex, and ventral margin deeply and widely evaginate; occiput rather strongly and evenly convex; antenna elongate, nearly as long as face; pedicel about 1.5 times as long as wide; 1st flagellomere about 2-2.5 times as long as wide, tapered distally, pointed or narrowly rounded apically; arista short-pubescent; proboscis short, capitate or spatulate; haustellum slightly longer and labellum slightly shorter than half length of ventral margin of head; palpus large. Chaetotaxy: major setae blackish, except postocellar, postvertical and postoculars whitish to yellowish. Thorax. black, dorsum matt, entirely covered by dense, fine, whitish microtrichia; pleura, especially anepisternum, subshiny, with sparser microtrichia; setulae sparse, whitish; major setae black except scapular setae whitish, thick, about twice as long as adjacent setulae; dorsocentral setae aligned slightly posterior to transverse suture. Legs. predominantly black and brown; forecoxa yellow, with dense grayish microtrichia; tibiae increasingly yellow anteriorly; tarsi sometimes entirely yellow, but distal tarsomeres sometimes darkened. Wing (Fig. 24). elongate (length to width ratio about 3). Venation: Longitudinal veins generally straight; vein R1 dorsally with wide gap in row of setulae opposite end of subcosta, ventrally with 0-2 setulae distally; vein R4+5 bare; veins R4+5 and M distinctly divergent apically; crossvein R-M at about distal 0.20-0.25 of cell dm; cell bcu closed distally by straight vein, without posterodistal lobe. Costal spine well developed. Pattern: costal cells dark brown or blackish; some brown slightly extending posterior to root of vein R; pterostigma entirely blackish; narrow subapical band from end of cell r2+3 usually extending to vein M, sometimes to, or nearly to, hind margin; distal border of band generally straighter and smoother than proximal border; apex of wing white beyond dark band, including apex of cells r2+3 and r4+5, and tip of cell m; posterior half of crossvein DM-Cu bordered by brown patch; small gray spot over crossvein r-m (as in Fig. 4 in Munro, 1939) present only in one specimen; remaining wing surface grayish; veins mostly brownish-yellow. Abdomen. blackish-brown, subshiny, with indistinct microtrichia and long, coarse, whitish, semi-erect, rather dense setulae; setae mostly indistinguishable from setulae, except marginally on posteriormost tergite where sometimes longer, finer and darker; tergite 6 in female slightly longer than tergite 5. Male terminalia: Epandrium oval in posteior view, widest below mid- height (Fig. 38), wider dorsally and with small but protuberant caudal flange in lateral view (Fig. 52); distiphallus (Fig. 66) with 2 large spines at about middle and series of 3 smaller spines at preglans part. Female terminalia: Oviscape conical, but often flattened in preserved specimens, blackish, with fine brown setulae; oviscapal measure 2; aculeus tapered from base to apex (Fig. 81), indistinctly arrow-like, distal 0.15 strongly attenuated; spermatheca (Fig. 96) pyriform, sparsely covered by small tubercles.

412 Freidberg and Merz / Revision of the Gymnosagena group

Wing length: Male: 3.0-3.3 mm; female: 3.2-3.4 mm. Material Examined Holotype , [SOUTH AFRICA:] Drakensberg, Natal, Rockeries Section, Ntonyelana River (Puma Kwelanga), July 1938, W. E. Marriott. M. 645. The holotype is double mounted on minute pin and polyporus block, is in excellent condition and is deposited in SANC. Allotype female and paratypes (9, 5): same collecting data as holotype. In addition, two puparia, one with same collecting data as holotype and labelled as paratype, and one labelled: National Park, Natal, July 1939, W.E. Marriott. M. 699. New records: Transvaal : Inn on Robbers’ Pass, 1400m, 27.xii.1994, A. Freidberg (10 galls, each with puparium; TAUI) ; E[astern] Cape Prov[ince], Katberg, 4000ft., xii.1932, R.E. Turner (1; BMNH). Paratypes are deposited in SANC, MHNG and TAUI. Biology This species forms terminal twig galls on Helichrysum splendidum. The gall and puparium were described by Munro (1939), and a photograph of the gall (Fig. 5) was also included. Distribution South Africa. Discussion This species and its newly described congener show a striking habitus for tephritids. Munro’s (1939) description of this species is generally adequate, but the gena is about 0.33-0.4, not 0.2, as high as the eye, as is obvious from his Fig. 3. The terminalia are described and illustrated here for the first time. The two new locality records (see Material Examined) are significant, despite the fact that one of them is based solely on galls and puparia, because this species has hitherto been known from one or two localities only. We have studied 15 specimens, including the holotype, out of the 39 specimens originally included in the type series. An unusual aberration observed in three of the specimens (2 and 1) is a complete supernumerary crossvein, between veins R2+3 and R4+5, that is an extension of crossvein R-M. It was observed on both wings of two specimens and one wing of a third specimen.

Marriottella sepsoides Freidberg & Merz, n. sp. (Figs. 7, 11, 25, 39, 53, 67, 82, 97) Diagnosis A predominantly blackish, sepsid-like , similar to M. exquisita, but differing readily by its smaller size (wing length less than 2 mm), large and shiny ocellar triangle, geniculate proboscis, brownish postocular setae, and the slightly but distinctly different wing pattern. Description Head (Figs. 7, 11). Coloration: Dull to subshiny brown to black; frons, parafacial, and antenna brown, face shiny black; discrete shiny ocellar triangle approximately as long as frons, extending to lunule; dorsal occiput with more or less distinct, white, fine microtrichia; major setae blackish, postocellar and postgenal setae whitish or yellowish; setulae mostly yellowish. Structure: Frons width to eye width ratio about 2; frontofacial angle 90o-110o, angulate but only slightly protuberant; parafacial 0.15, gena 1.5 times as wide (as high) as 1st flagellomere; face in profile slightly concave, with ventral margin strongly protubebrant, but ventral half of face

413 BIOTAXONOMY OF TEPHRITOIDEA medially convex, and ventral margin deeply and widely evaginate; occiput rather strongly and evenly convex; antenna elongate, as long as face; pedicel about 1.5 times as long as wide; 1st flagellomere about 3 times as long as wide, tapered, rounded or pointed apically; arista with short rays; proboscis geniculate; haustellum and labellum, each, about as long as ventral border of head; palpus large. Thorax. Scutum squarish; scutellum small, triangular, dorsally convex, about 0.35 times as long as wide. Thorax brownish-black, mesonotum dull, entirely covered by sparse, fine, grayish microtrichia; dorsal pleura, especially anepisternum and anepimeron, shiny, with sparser microtrichia; setulae sparse, whitish; major setae blackish, except scapular setae undifferentiated, setula-like; dorsocentral setae aligned slightly posterior to transverse suture. Legs. predominantly blackish or brown; forecoxa brownish, with dense grayish microtrichia; femora black except apex; tibiae usually black, except apex, foretibia sometimes predominantly yellow; tarsi vary from predominantly yellow, with distal tarsomeres dark, to almost entirely dark. Wing (Fig. 25). elongate (length-width ratio about 2.8). Venation: Longitudinal veins generally straight; vein R1 dorsally with wide gap in row of setulae opposite hyaline gap in dark

“marginal” band, ventrally with 0-2 setulae distally; vein R4+5 without setulae; veins R4+5 and M distinctly divergent apically; crossvein r-m at about distal 0.33-0.40 of cell dm; cell bcu closed distally by straight vein, without posterodistal lobe. Costal spine short, undifferentiated. Pattern: Costal cells, except hyaline distal 0.25-0.30 of (distal) costal cell, dark brown; some brown slightly extending posterior to root of vein R; pterostigma entirely black; apex of cell r2+3 from end of vein R2+3 to end of vein R4+5 entirely blackish, spot penetrating slightly and subapically into cell r4+5; apex of cell r4+5, beyond dark spot with milky-white microtrichia, forming rather indistinct white apical spot; remaining wing surface grayish; veins mostly brownish yellow. Calypteres narrow, whitish, ventral calypter with dark margin. Halter blackish. Abdomen. blackish-brown, shiny, with fine, whitish, barely distinct microtrichia and long and fine, brown but tinged with yellow, semi-erect, sparse setulae; setae laterally and marginally on posteriormost tergite slightly longer but otherwise indistinguishable from setulae; tergites 5 and 6 subequal in female. Male terminalia: Epandrium oval in posterior view (Fig. 39), more or less uniformly narrow and with large caudal flage in lateral view (Fig. 53); distiphallus (Fig. 67) with two spines approximated to each other and to glans. Female terminalia: Oviscape conical, but often flattened in preserved specimens, brownish, with setulae as on preabdomen; oviscapal measure 2; aculeus tapered from base to apex (Fig. 82), indistinctly arrowlike; margin of basal 0.66 with rasperlike structure; spermatheca (Fig. 97) elongate, more or less densely covered by elongate tubercles. Wing length. Male and female: 1.60-1.85 mm (width 0.57-0.68 mm). Material Examined Holotype , SOUTH AFRICA, Glen, OFS [Orange Free State], Dec. 1951, E.K. Hartwig, M. 1109. The holotype is double mounted (minuten and polyporus block), is in excellent condition, and is deposited in SANC.

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Table 1 Host list Host plants Tephritid Infested Association New/old (by tribe) species part status record

Anthemideae Matricaria nigellifolia DC. Gymnosagena unicornuta FH + N Astereae Conyza hypoleuca A. Rich. Elgonina pollinosa FH + N Conyza newii Oliv. et Hierm. Elgonina inexpectata FH ? N Elgonina yaromi FH + N Conyza pyrrhopappa Sch. Bip. Elgonina splendida FH + N ex A. Rich. ssp. pyrrhopappa Conyza ruwenzoriensis Elgonina refulgens FH ? O (S. Moore) R.E.Fr. Conyza steudelii Sch. Bip. Elgonina inexpectata FH ? N ex A. Rich. Conyza sp. Elgonina refulgens FH + N Microglossa pyrifolia (Lam.) Elgonina infuscata FH + N O. Kuntze Nidorella resedefolia DC. Gymnosagena unicornuta FH + O Psiadia salviiflora Backer Gymnosagena campiglossina FH + N Heliantheae Aspilia mossambicensis (Oliver) Gymnosagena longicauda FH + N Wild Inuleae Helichrysum ?dreganeanum Marriottella sepsoides FH + N Sonder et Harvey Helichrysum splendidum Marriottella exquisita Gall + O (Thunberg) Lessing Phagnalon rupestre L. Spathulina sicula Gall + O FH = flower head; N = new; O = old; + = reared; ? = not actually reared.

Paratypes: same data as holotype (15, 19; SANC, MHNG, TAUI); SOUTH AFRICA : Middelburg, C.P. [Cape Province], 15.v.[19]24, H.K. Munro (1, 1; SANC). Many of the paratypes were partly damaged by pests. Etymology Named after its habitus, which is similar to some common sepsids. Biology and host plants This species was reared from flowerheads of a Helichrysum species. One of the flowerheads that were preserved with the flies was sent for identification to Prof. O.M. Hilliard (Royal Botanic Gardens Edinburgh), who noted (in litt.) that it is probably Helichrysum dregeanum. The black puparium is glued to the remains of the achenes on which it fed; one puparium per flowerhead. No gall is formed. Distribution South Africa.

415 BIOTAXONOMY OF TEPHRITOIDEA

Discussion This species exhibits a number of unusual characters for Tephritids, in particular the presence of a shiny, large ocellar triangle reaching the lunule, and the general sepsid-like appearance. Its placement in Marriottella together with M. exquisita is corroborated in the phylogenetic analysis by five synapomorphies.

CLADISTIC ANALYSIS Methods Phylogenetic relationships between the species of the Gymnosagena subgroup were analyzed using the ie option of Hennig86 software (Farris, 1988; Fitzhugh, 1989). We chose Spathulina sicula Rondani (the type species of Spathulina) as an outgroup because we consider Spathulina to be the sister group of the Gymnosagena subgroup. Characters used in the analysis are elaborated in Appendix 1. Most of these characters are straightforward and do not require additional explanation. Qualifying comments are added as deemed necessary. A ‘0’ indicates the more plesiomorphic state, with ‘1’ and ‘2’ indicating, respectively, more derived states. The coding for nonadditive characters (in which there is no obvious indication of the polarity of the derived states) is reviewed on a character by character basis as indicated in the character list (Appendix 1). The numbers used in the list are the same as those on the illustrated cladograms (Fig. 98, 99), and the sequence of taxa is the same as noted in the data matrix (Table 2). Results, discussion and conclusions The ie option of Hennig86 yielded four similar trees, two of which are fully resolved (e.g., Fig. 98) and two of which are almost fully resolved. The Nelsen consensus tree (length 62; consistency index 51; retention index 74), produced from these trees (Fig. 99), is also largely resolved, except for two clades within Elgonina. These two clades correspond nicely to two of the species groups proposed in Elgonina (refulgens and splendida, respectively), whereas the third group, pollinosa, containing only one species (pollinosa), was represented in all five trees as a separate basal clade to all other Elgonina. The clade representing the splendida group appeared as fully resolved in all four initial trees, although there were two options: flavicornis (yaromi (infusctata, splendida)) (i.e., Fig. 98) and splendida (yaromi (flavicornis, infusctata)). Hence, the three species groups within Elgonina appear as monophyletic clades in all five trees, as do the three genera analyzed, Elgonina, Gymnosagena and Marriottella. In all five trees Elgonina and Marriottella are sister groups, and Gymnosagena is a sister group to both of them together. Within Gymnosagena, G. longicauda + G. kakamega form a sister group to all the other species. Almost all major clades of the Nelsen tree are supported by at least one synapomorphy. The monophyly of the Gymnosagena subgroup is supported by three synapomorphies (characters #11; 15; 28), of which the posterior notopleural seta (brown to black and acuminate) appears to be the most significant. The monophyly of the genus Gymnosagena is supported by four synapomorphies, of which the weakly protuberant ventral facial margin, the capitate proboscis, and the non-spinulose, non-setulose, preglans part of the distiphallus (#1, 7, 21) are especially significant. None of the two main clades within Gymnosagena is supported by synapomorphies. Sister-group relationships between Marriottella and Elgonina is supported by a synapomorphy

416 Freidberg and Merz / Revision of the Gymnosagena group

Table 2 Character matrix for the Gymnosagena subgroup Characters 1 1111111112 222222222 Taxon 1234567890 1234567890 123456789 Spathulina sicula 0000000000 0000000000 000000000 Elgonina pollinosa 0000000010 0100010040 000001011 E. dimorphica 0001110010 1100110030 1001111?1 E. fuscana 0?011100?0 1?01110040 ?????11?1 E. inexpectata 0001110000 1100110030 100111111 E. refulgens 0001100000 1100110030 100111011 E. flavicornis 0011100000 1101010030 1010011?1 E. infuscata 0011110010 1101010020 101001111 E. splendida 0011110010 1100010030 101001011 E. yaromi 0011110000 1100010030 101001111 Marriottella exquisita 0011001001 0100011111 000000000 M. sepsoides 0011111201 1110011111 100000001 Gymnosagena campiglossina 1100000100 0111110000 010000111 G. kakamega 1100000100 1100110000 1100001?1 G. longicuda 1100000100 1100110000 110000131 G. nyikaensis 1110000100 0112110000 010?00??1 G. unicornuta 1110000100 0112000000 0101001?1

of the wing pattern. The monophyly of Marriottella is supported by five synapomorphies, whereas that of Elgonina by two synapomorphies (#18 and 25: reduced wing patern, and sternite 8 in the aculeus extending laterally beyond margin of tergite 8, respectively). Finally, the monophyly of the refulgens and splendida groups (of Elgonina) are each supported by a single synapomorphy of the male phallus (#24 and 22, respectively). The conclusions of the current analysis strongly support the accepted classification of the Gymnosagena subgroup that was used in this publication. However, this analysis is preliminary in the sense that only one external member of the Spathulina group was included, and only as outgroup; no weighting procedures have been carried out; and only the relatively weak ie option was used. A more rigorous analysis is required, especially with other members of the group included, notably the numerous species of Spathulina.

ACKNOWLEDGMENTS We are grateful to the curators listed in the introduction who arranged the loans that made this study possible. We are grateful to the following persons for critically reviewing various versions of the manuscript: Netta Dorchin, A.L. Leonid Friedman, Wayne N. Mathis, Allen Norrbom, Ilan Yarom, and Irina Zonstein. We thank Henk J. Beentje for identification of several of the host plants recorded in this paper, and Olive M. Hilliard for providing an identification for the flowerhead of Helichrysum dreganeanum. Thanks are due to Walter Ferguson who made some of the drawings, Amikam Shoob who prepared the photographs, and Naomi Paz who corrected the style of a final version.

417 BIOTAXONOMY OF TEPHRITOIDEA

Fig. 98. Reconstructed phylogeny for the Gymnosagena group: one of four trees obtained by using the ie option. Character numbers refer to Appendix 1. ● = forward changes without homoplasy; ⅙ = changes with homoplasy.

418 Freidberg and Merz / Revision of the Gymnosagena group

Fig. 99. Reconstructed phylogeny for the Gymnosagena group: Nelsen tree. Character numbers refer to Appendix 1. ● = forward changes without homoplasy; ⅙ = changes with homoplasy.

419 BIOTAXONOMY OF TEPHRITOIDEA

REFERENCES Cogan, B.H. & Munro, H.K. 1980. 40. Family Tephritidae, pp. 518-554. In: Crosskey, R.W. et al. (eds.). Catalog of the Diptera of the Afrotropical region. British Museum (Natural History), London. 1437 pp. Farris, J.S. 1988. Hennig86 reference. Documentation for version 1.5. Fitzhugh, K. 1989. Cladistics in the fast lane. Journal of the New York Entomological Society 97: 234-241. Freidberg, A. 1984. Gall Tephritidae, pp. 129-167. In: Ananthakrishnan, T.N. (ed.). Biology of Gall Insects. Oxford and IBH Publishing Co., New Dehli. 362 pp. Freidberg, A. & Hancock, D.L. 1989. Cryptophorellia, a remarkable new genus of Afrotropical Tephritidae (Diptera: Tephritidae). Annals of the Natal Museum 30: 15-52. Freidberg, A. and Mathis, W.N. 1986. Studies of Terelliinae (Diptera: Tephritidae): a revision of the genus Neaspilota Osten Sacken. Smithsonian Contributions to Zoology 439: 1-75. Freidberg A. & Norrbom, A.L. 1999. 23. A generic reclassification and phylogeny of the tribe Myopitini (Tephritinae), pp. 581-627. In: Aluja, M. and Norrbom, A.L. (eds.). Fruit flies (Tephritidae): Phylogeny and evolution of behavior. CRC Press, Boca Raton. 944 pp. Hancock, D.L. 1986. New genera and species of African Tephritinae (Diptera: Tephritidae). with com- ments on some currently unplaced or misplaced taxa and on classification. Transactions of the Zimbabwe Scientific Association 63 (3): 16-34. McAlpine, J.F. (1981) Morphology and terminology - adults. In: J.F. McAlpine et al. (ed.). Manual of Nearctic Diptera, Vol. 1, Agriculture Canada, Research Branch, Hull (Quebec). (Monograph No. 27). Pp. 9-63. Munro, H.K. 1935. Some interesting new African Trypetidae (Dipt.). Stylops 4 (1): 4-7. Munro, H.K. 1939. Some new species of South African Trypetidae (Diptera), including one from Madagascar. Journal of the Entomological Society of Southern Africa 2: 139-153. Munro, H.K. 1947. African Trypetidae (Diptera). Memoirs of the Entomological Society of Southern Africa 1: 1-300. Munro, H.K. 1957. Trypetidae. Ruwenzori Expedition, 1934-35, Vol. II, No. 9: 853-1054. Norrbom, A.L., Carroll, L.E., Thompson, F.C., White, I.M. and Freidberg, A. 1999. Systematic database of names, pp. 65-299. In: Thompson, F.C. (ed.). Fruit fly expert identification system and systematic information database. Myia 9: 1-524 . Backhuys Publishers, Leiden. Steyskal, G.C. 1979. Taxonomic studies on fruit flies of the genus Urophora (Diptera: Tephritidae). Entomological Society of Washington, Washington, D.C. 61 pp. White, I.M., Headrick, D.H., Norrbom, A.L. and Carroll, L.E. 1999. Glossary. pp. 881-924. In: Aluja, M. and Norrbom, A.L. (eds.). Fruit flies (Tephritidae): Phylogeny and evolution of behavior. CRC Press. Boca Raton. [16] + 944 pp.

420 Freidberg and Merz / Revision of the Gymnosagena group

APPENDIX 1 Characters used in the phylogenetic analysis Head 1. Head height to length ratio: 0) 1.00-1.15; 1) 1.16-1.30. 2. Ventral facial margin: 0) Strongly protuberant; 1) Weakly or not protuberant. No objective measure is provided. However, all Elgonina and Marriottella species have the ventral facial margin strongly protuberant, more or less as in S. sicula, whereas in Gymnosagena it varies from barely to moderately protuberant (most strongly in G. uniconuta) and coded 1 in all these species. 3. Orbital setae: 0) 2 pairs; 1) 1 pair. The number of orbital setae is rarely variable intraspecifically, and in those cases only the predominant situation was scored. 4. Lateral vertical seta: 0) Whitish; 1) black. 5. Postocular setae: 0) Whitish, or mixed whitish and black; 1) Black. 6. Postocellar seta: 0) Whitish or yellow; 1) Black. 7. Palpus: 0) Entirely yellow or partly brownish-yellow; 1) Black or blackish. 8. Proboscis: 0) Spatulate; 1) Capitate; 2) Geniculate. No objective measure has been provided, and the separation between state 0 and 1 is difficult. State 2 is clearly represented only in M. sepsoides. This character is treated as non-additive because a logical transformation series would require re-shuffling of states 0 and 1. 9. Antenna coloration: 0) Not sexually dimorphic; 1) Sexually dimorphic. Sexually dimorphic antenna, in which the 1st flagellomere is black in the female and yellow in the male, occurs sporadically in all three groups of Elgonina. Non sexually dimorphic antennae may either be yellow (E. flavicornis) or black (E. yaromi). Thorax 10. Scutellum: 0) Long (0.3 times or more as long as scutum); 1) Short (0.25 times as long as scutum). 11. Scutal setulae: 0) Whitish, coarse, rarely fine; 1) Brownish to black, fine. No objective measure has been provided, and some degree of overlap between the two states has been observed in both species of Marriottella. 12. Posterior notopleural seta: 0) Whitish and lanceolate; 1) Brown to black and acuminate. 13. Postalar seta: 0) Long, about half as long as posterior supra-alar seta or more; 1) Short, less than half as long as posterior supra-alar seta. 14. Postalar seta: 0) Blackish; 1) Brownish; 2) Whitish. This character is treated as additive. 15. Apical scutellar seta: 0) Indistinguishable from setulae or lacking, at most 0.15 times as long as basal scutellar seta; 1) Seta-like, conspicuous, about 0.25 times as long as basal scutellar seta. Legs 16. Femora: 0) Pale yellow to brownish-yellow or slightly spotted by dark spots; 1) Dark brown to black on approximately basal 0.66-0.90, sometimes irregularly. Wing 17. Wing shape: 0) Normal (length to width ratio below 2.8); 1) Elongate (length to width ratio above 2.8).

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18. Pterostigma at base (between, but not including, veins Sc and R1): 0) at least three times as wide as either vein; 1) 1-2 times as wide as either vein. The unusually narrow pterostigma of Marriottella is exceptional in Tephritinae. 19. Wing pattern: 0) Reticulate (whether grayish or blackish); 1) Comprising marginal band at base, preapical dark spot and milky-white apical spot; 2) Comprising a few dark spots, including pterostigma and subapical spot; 3) Comprising a milky-white apical spot and sometimes other whitish spots (on a generally pale gray background); 4) Lacking (no dark or whitish spots). The above-mentioned five states seem to form a transformation series reflecting an evolutionary pathway of wing pattern reduction, and the character is treated as additive. 20. Halter: 0) Yellow; 1) Blackish. Abdomen 21. Setulae: 0) Partly or entirely coarse and whitish; 1) Entirely fine and brown to black. 22. Preglans part of distiphallus: 0) Spinulose or setulose; 1) Non-spinulose and non- setulose. ‘Spinulose’ and ‘setulose’ should perhaps be treated as different characters. 23. Preglans part of distiphallus: 0) Without setulae-bearing lobes 1) With setulae-bearing lobes. 24. Acrophallus: 0) Cylindrical; 1) Greatly enlarged apically. 25. Vesica: 0) Large and conspicuous; 1) Almost entirely reduced, indistinct. 26. Aculeus: 0) Sternite 8 not extending laterally beyond margin of tergite 8; 1) Sternite 8 extending laterally beyond margin of tergite 8. 27. Aculeus: 0) Without preapical step; 1) With preapical step. Ecology 28. Host tribe: 0) Inuleae; 1) Astereae; 2) Anthemideae; 3) Heliantheae. This character is treated as non-additive. 29. Gall inducer: 0) Yes; 1) No. The species were coded based partly on rearing records, partly on speculation.

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