Review of the genus Togoda Melichar, 1906 (Hemiptera, Auchenorrhyncha, Fulgoroidea, Tropiduchidae), with descriptions of three new species from tropical Africa and Madagascar

Directeur De la publication / Publication director : Gilles Bloch Président du Muséum national d’Histoire naturelle réDactrice en chef / editor-in-chief : Laure Desutter-Grandcolas assistante De réDaction / assistant editor : Anne Mabille ([email protected]) Mise en page / Page layout : Anne Mabille coMité scientifique / scientific board : Nesrine Akkari (Naturhistorisches Museum, Vienne, Autriche) Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine) Serge Gofas (Universidad de Málaga, Málaga, Espagne) Sylvain Hugel (CNRS, Université de Strasbourg, France) Marco Isaia (Università degli Studi di Torino, Turin, Italie) Rafael Marquez (CSIC, Madrid, Espagne) Jose Christopher E. Mendoza (Lee Kong Chian Natural History Museum, Singapour) Annemarie Ohler (MNHN, Paris, France) Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France) Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne) couverture / cover : Togoda sp. (Nigeria: Ekiti, Ikere). Credits: Babajide Agboola. 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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / https://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387 473 ZOOSYSTEMA • 2025 • 47 (23) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:44B9F8A0-391B-4C00-AE1B-23D3E5494409 Gnezdilov V. M. 2025. — Review of the genus Togoda Melichar, 1906 (Hemiptera, Auchenorrhyncha, Fulgoroidea, Tropiduchidae), with descriptions of three new species from tropical Africa and Madagascar. Zoosystema 47 (23): 473-488. https://doi.org/10.5252/zoosystema2025v47a23. http://zoosystema.com/47/23 ABSTRACT The tropiduchid genus Togoda Melichar, 1906 is reviewed, with a type species, Togoda africana Melichar, 1906, redescribed and three new species from the Central African Republic, Mozambique, and Madagascar described and illustrated. A key to species is given. Wing morphology of Trienopini Fennah, 1954 is discussed. RÉSUMÉ Révision du genre Togoda Melichar, 1906 (Hemiptera, Auchenorrhyncha, Fulgoroidea, Tropiduchidae), avec la description de trois nouvelles espèces d’Afrique tropicale et de Madagascar. Le genre de tropiduchidés Togoda Melichar, 1906 est révisé, l’espèce type, Togoda africana Melichar, 1906, est redécrite et trois nouvelles espèces de la République centrafricaine, du Mozambique et de Madagascar sont décrites et illustrées. Une clé des espèces est donnée. La morphologie des ailes des Trienopini Fennah, 1954 est discutée. Vladimir M. GNEZDILOV Zoological Institute, Russian Academy of Sciences, Universitetskaya Emb. 1, 199034 Saint Petersburg (Russia) [email protected] [email protected] Submitted on 12 February 2025 | Accepted on 21 March 2025 | Published on 30 September 2025 Review of the genus Togoda Melichar, 1906 (Hemiptera, Auchenorrhyncha, Fulgoroidea, Tropiduchidae), with descriptions of three new species from tropical Africa and Madagascar MOTS CLÉS Trienopini, clé d’identification, morphologie, nouvelle espèce, région afrotropicale. KEY WORDS Trienopini, identification key, morphology, new species, Afrotropical Region. 474 ZOOSYSTEMA • 2025 • 47 (23) Gnezdilov V. M. INTRODUCTION The genus Togoda Melichar, 1906 belongs to the tribe Trienopini Fennah, 1954 of the subfamily Tropiduchinae Stål, 1866 represented by five Afrotropical genera with 21species (Metcalf 1958; Gnezdilov 2007, 2013; Bourgoin 2024). Togoda clearly differs from other genera of the tribe by large metopial proboscis. The genus was established by L. Melichar for a single species, Togoda africana Melichar, 1906, described after long series of specimens collected by L. Conradt in Bismarckburg in Togo (Melichar 1906). Soon after Schmidt (1910) described another species, T.transvaalensis Schmidt, 1910, from Zoutpansberg (Transvaal) in South Africa. Since that time the genus was never in a focus of the taxonomic study. Togoda africana was erroneously recorded by Metcalf (1958) from Angola referring to Melichar (1906) who mentioned “Westafrica” in fact referring to Togo. Schmidt (1910) mentioned this species also from Cameroon, but noted that these specimens are twice larger than those from Togo. Finally Gnezdilov (2022) recorded T.africana from the canopies of Ghana, but the reexamination of these specimens (females) did not confirm our previous identification and currently this material is identified as Togoda sp. until the males will be available for study. Thus currently T.africana is authentically known only from the type locality in Togo. T.africana is here redescribed based on the specimens from Bismarckburg (topotypes) and the male genitalia structures are illustrated for the first time. Three new species of the genus are described from the Central African Republic, Mozambique, and Madagascar. I had no material available for study on T.transvaalensis which is included in the key based on Schmidt’s original description. Thus up to date the genus Togoda Melichar comprising five species known from tropical Africa and here is recorded for the first time from Madagascar. MATERIAL AND METHODS Morphological terminology follows Anufriev& Emeljanov (1988) for structures of the head, Gnezdilov& Bartlett (2018) – for wing venation, and Gnezdilov (2013) – for male and female genitalia. Photographs were taken using a Canon 5D Mark IV digital camera equipped with a lens Canon MP-E 65mm f/2.8 1-5x macro and Canon Macro Twin Lite MT–26EX-RT flash and SEM Hitachi TM3000. Images are produced using Helicon Focus v. 7.6.4 and Adobe Photoshop software. The genital segments of male specimens examined were macerated in 10% KOH and figured in glycerin jelly (Brunel Micro Ltd, UK) using Leica MZ9.5 light microscope with camera lucida attached. The map of Africa is designed by freepik.com (https:// ru.freepik.com/free-photos-vectors/map-of-africa). Label information is quoted, with “/” indicating new line and “//” indicating next label. Right/left sides of genital structures are designated based on the position of genital structures inside the insect in the direction from its head to apex of abdomen (see the legends to Figs8-10). The material examined, including the type specimens of the species described below, are deposited in the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg, Russia (ZIN) and in the Muséum national d’Histoire naturelle, Paris, France (MNHN). SYSTEMATICS Family Tropiduchidae Stål, 1866 Subfamily Tropiduchinae Stål, 1866 Tribe Trienopini Fennah, 1954 Genus Togoda Melichar, 1906 Togoda Melichar, 1906: 233. Type species.— Togoda africana Melichar, 1906, by monotypy. disTribuTion.— Tropical Africa (Nigeria, Ghana, Togo, Central African Republic, South African Republic, Mozambique, Tanzania: Zanzibar Island) and Madagascar (Fig.11). composiTion.— Five species. d iagnosis .— Coryphe transverse. Metope long and wide, nearly parallel-sided (in dorsal and frontal views), protruding in shape of wide proboscis (in lateral view). Coryphe and metope joint at obtuse angle in lateral view. Postclypeus large, with median carina. Metopoclypeal suture straight. Scapus short, cylindrical. Pedicel 4-5times longer than scapus, elongate, twice as long as wide, slightly convex dorsally, covered with setae and sensory organs (Fig.1D). Ocelli absent. Rostrum long, narrow, reaching hind coxae or slightly surpassing them; third segment slightly longer than second one, narrowing and bent apically (Fig.7B). Pronotum large, with distinct lateral carinae diverging posteriorly, wide paradiscal fields behind the eyes and wide paranotal lobes; posterior margin strongly concave. Median part of pronotum hanging over coryphe. Mesonotum 1.5times as long as pronotum, without carinae. Forewing with loop-shaped process of precostal margin basally and with semicircular lobe on wing margin behind claw-shaped apex of clavus, both with marginal setae (Figs1; 2). Marginal cell below the semicircular lobe distinctly concave. Tegulae large. Fore wings without hypocostal plate, but with narrow precostal field, with transverse veins and with a wide subcostal field in its basal fifth; clavus open. Hind wings well developed, slightly shorter than fore wings (Fig.7A). Hind tibia with two large lateral spines on the apical half and one small lateral spine on the proximal half. First and second metatarsomeres nearly equal in length, with dense long hair-shaped setae ventrally. Second metatarsomere with only two latero-apical spines. Pygofer with widely and slightly concave lower margin (Fig.9J). Penis strongly curved in lateral view. Apical part of aedeagus weakly sclerotized and apparently sometimes swollen. Styles massive, fused at its basal halves (Fig.9I), with short capitulum, without neck and with a strong comb on the margin below capitulum. Finger-shaped process of the style large, curved and turned posteriorly and ventrally. Abdominal sternite III large, with triangularly protruding hind margin overhanging above other sternites (Figs4B, D; 5B, D). Female sternite VII with weakly convex medially hind margin. Gonoplacs large, triangular-shaped, flat, with large marginal teeth (Fig.3C, D). 475 Review of the genus Togoda Melichar, 1906 ZOOSYSTEMA • 2025 • 47 (23) Togoda africana Melichar, 1906 (Figs2A, B; 4A, B; 6A, B; 7; 8) Togoda africana Melichar, 1906: 233, fig.55. m aTerial examined .— Togo • 1 ♂ ; “Togo / Bismarckburg / 22.IX-1.X.92 / L. Conradt S.”; ZIN; 1 ♀ ; “Togo / Bismarckburg / 23.XI-2.XII.92 / L. Conradt S.”; ZIN; 1 ♂ ; “Togo / Bismarckburg/ 9-11.V.93 / L. Conradt S.”; ZIN; 1 ♀ ; “Togo / Bismarckburg / 1214.IV.93 / L. Conradt S.”; ZIN. diagnosis.— Coryphe-metope margin keel-shaped. Metopial proboscis weakly concave frontally. Fore wings dark brown to black, with rich elongate venation. Phallobase short, aedeagus wide in lateral view, with four narrow subapical processes – three on left side and one on right side. Male anal tube narrowing apically in dorsal view. disTribuTion.— Central Togo. descripTion Structure (Figs2A, B; 4A, B; 6A, B; 7) Coryphe transverse, 1.5 times as wide as long medially, smooth, without carinae, with slightly convex or nearly straight anterior margin (Fig.4A). Coryphe-metope margin keel-shaped (Fig.6A). Eyes large, diameter of each eye nearly as large as width of metope between the eyes. Metope, proand mesonotum densely covered with traces of larval sensory pits (pustules). Metopial proboscis weakly concave ventrally in frontal view (Fig.6B). Postand anteclypeus with distinct median carina. Pedicel slightly flattened laterally, with area of dense setae dorsally (Fig.6A). Fore wings large, nearly oval, narrowing apically, with 1/3 of its length behind apex of clavus in lateral view (Fig.4B). Basal cell small, loop-shaped. Radius and median running from the basal cell separately, but nearly from one point. Fore wings with small number of transverse veins. Forewing vein branching sequence: R 5-8, firstly furcating closely to basal cell; M 10-14, firstly furcating closely to basal cell; M1 and M2 furcating again in basal wing third – M2 before M1; CuA 2-3, furcating at the same point as M1; CuA1 and CuA2 terminating at concave cell behind apex of clavus, CuA 1 sometimes furcating also apically (in one specimen right wing with CuA 2 and left wing with CuA 3). Pcu joint A1 in apical fourth of clavus. Clavus long, 2/3 of wing length. Hind wings bi-lobed, without cubital cleft and with weak vannal cleft; anterior margin with long and narrow coupling lobe (Fig.7A). Precostal margin with dense macrosetae at its basal third and with sparse marginal microsetae. Hind wing vein branching sequence (Fig.7A): R2, furcating before wing middle; r-m 1; M 1; m-cua 1; CuA3, firstly furcating at wing middle and CuA2 furcating apically; cua-cup1; CuP 1; cup-pcu 1; Pcu 1; A1 1; A2 1. Legs not flattened neither foliated. Hind tibia with 9-12 spines apically. First metatarsomere with two latero-apical and 4-6intermediate spines arranged in an arc apically. Arolium of pretarsus not protruding beyond claw apices, sometimes with notched hind margin (in dorsal view). Dorso-lateral plates of arolium long and narrow. Coloration (Fig.4A, B) Metope, pronotum, mesonotum, and fore wings dark brown to black (on fore wings). Head laterally light yellow to light brown and black above brown scapi. Anteclypeus and rostrum yellowish light brown. Pedicels dark brown to black, with green area of setae dorsally. Third segment of rostrum Key To Togoda melichar, 1906 1. Fore wings dull green, with transverse double broken brown stripe in its basal third. Metope distinctly narrower between eyes. Metopial proboscis and postclypeus joint at obtuse angle in lateral view. Distribution: South African Republic ............................................................................................... T.transvaalensis Schmidt, 1910 — Fore wings yellowish light brown to black. Metope with nearly parallel lateral margins. Metopial proboscis and postclypeus joint at nearly right angle in lateral view (Fig.6) ....................................................................... 2 2. Metope with two weak carinae medially and groove between them (Fig.6G). Postclypeus with strong triangularshaped median carina (Figs5D, 6H). Fore and middle femora flattened or nearly foliate. Distribution: Madagascar ................................................................................................................................... T.carinata n.sp. — Metope without carinae. Postclypeus with tiny median carina. Legs not flattened neither foliated ............... 3 3. Fore wings large, with 1/3 of its length behind apex of clavus in lateral view (Figs4). Metopial proboscis distinctly notched frontally (Fig.6B, D). Phallobase short, 1/3 of aedeagus length, symmetric (Figs8D, E; 9D, F .......................................................................................................................................................... 4 — Fore wings small, with 1/5 of its length behind apex of clavus in lateral view (Fig.5A, B). Metopial proboscis without notch frontally (Fig.6F). Phallobase long, nearly as long as aedeagus, asymmetric (Fig.10D, E). Distribution: Mozambique ........................................................................................................ T.minima n.sp. 4. Coryphe-metope margin keel-shaped. Metopial proboscis weakly concave frontally (Fig.6B). Male anal tube narrowing apically in dorsal view (Fig.8B). Aedeagus wide, with four subapical processes (Fig.8D, E, F). Distribution: Togo ...................................................................................................... T.africana Melichar, 1906 — Coryphe-metope margin smooth, without keel. Metopial proboscis deeply notched frontally (Fig.6D). Male anal tube deeply concave apically in dorsal view (Fig.9C). Aedeagus narrow, with two subapical processes (Fig.9D, E, F). Distribution: Central African Republic ........................................................................... T.excisa n.sp. 476 ZOOSYSTEMA • 2025 • 47 (23) Gnezdilov V. M. A CD EF B Fig. 1. — Forewing and pedicel structure of Trienopini: A, Trienopa flavida Signoret, 1860 (Madagascar), male, basal part of forewing, with ear-shaped process of precostal margin; B, Togoda sp. (Ghana), female, pronotum and basal part of forewing, with ear-shaped process of precostal margin; C, Togoda sp. (Ghana), female, ear-shaped process of precostal margin dorsally; D, Togoda sp. (Ghana), female, pedicel; E, Togoda sp. (Ghana), female, precostal margin of forewing at its middle; F, Togoda sp. (Ghana), female, ear-shaped process margin. Scale bars: A, 400 μm; B, 500 μm; C, 200 μm; D, E, 100 μm; F, 50 μm. 477 Review of the genus Togoda Melichar, 1906 ZOOSYSTEMA • 2025 • 47 (23) and flagellum dark brown to black. Postclypeus dark brown, sometimes pale laterally. Precostal margins of fore wings and semicircular lobes behind apex of clavus reddish. Hind wings brown to black. Legs greenish brown to dark brown or black, with yellowish light brown trochanters. Apices of leg spines black. Claws dark brown to black. Dorso-lateral plates of arolium dark brown to black. Abdomen light yellow to dark brown. Male terminalia (Fig.8) Pygofer vertically elongate, with right upper angle (below anal tube), hind margins straight, lower margin weakly concave (Fig.8A). Anal tube short, wide, narrowing apically in a convex margin (in dorsal view) (Fig.8B, C). Anal column short, 0.3 as long as anal tube. Phallobase short, nearly 0.3 as long as aedeagus, without processes. Aedeagus curved, horseshoe-shaped, asymmetrical, with straight apical part, cylindrical, with small denticles ventrally and laterally. Aedeagus with four subapical horn-shaped narrow processes directed downwards – three on left side (longest one near to ventral surface (Fig.8E, c), twice shorter one near to dorsal surface (Fig.8E, b), and smallest one above them at the middle (Fig.8E, d) and one long (as long as longest left one) on right side (Fig.8D, a). Style with short apical tooth, large lateral tooth, and distinctly convex hind margin (Fig.8G, hm, at, lt). Connective with widely opened short cup (Fig.8D, cp). Measurement Total length. Males: 5.0-5.5mm. Females: 5.5-6.0mm. AB CD Fig. 2. — Forewing structure of Togoda spp.: A, B, Togoda africana Melichar, 1906, Togo (Bismarckburg), male: A, apex of forewing clavus, dorsal view; B, semicircular process of forewing margin behind claval apex; C, D, Togoda excisa n. sp., female paratype: C, semicircular process of forewing margin behind claval apex; D, margin of semicircular process. Scale bars: A, 500 μm; B, 100 μm; C, 400 μm; D, 50 μm. 478 ZOOSYSTEMA • 2025 • 47 (23) Gnezdilov V. M. Togoda excisa n.sp. (Figs2C, D; 4C, D; 6C, D; 9) urn:lsid:zoobank.org:act:583816E8-B828-4066-AE0C-B8B92486619C Type maTerial.— Holotype. Central African Republic • ♂; “Boukoko / Rep. Centrafric. [print] / 20 IV.1967 [hand written in ink] / Michael Boulard [print]“ [blue] // “MoLoFo” [hand written in ball pen] // “Museum Paris / MNHN(EH) / 16569” [print, white]; MNHN. Paratype. Central African Republic • ♀; “Boukoko–RCA [print] / 30-1-1969 [hand written in ink] / Michael Boulard [print]” [blue] // “Museum Paris / MNHN(EH) / 16568 [print, white]”; MNHN. d iagnosis .— Coryphe-metope margin smooth, without keel. Metopial proboscis deeply notched frontally. Male anal tube deeply concave apically in dorsal view. Aedeagus narrow, with two long subapical processes. eTymology.— Species name referring to deeply concave apically male anal tube. disTribuTion.— Southwestern Central African Republic (Lobaye Prefecture). descripTion Structure (Figs2C, D; 4C, D; 6C, D) Coryphe twice as wide as long medially, with weak median carina and nearly straight anterior margin (Fig.4C). Metope densely covered with pustules (Fig.6D). Coryphe-metope margin smooth, without keel (Fig.6C). Metopial proboscis deeply notched frontally (Fig.6D). Diameter of each eye nearly as wide as metope. Postand anteclypeus with tiny median carina. Fore wings large and wide, with 1/3 of its length behind apex of clavus (Fig.4D). Pcu joint A1 in AB DC Fig. 3. — Habitus of Trienopini: A, Trienopa typica (Distant, 1909) (Seychelles); B, Togoda sp. (Nigeria: Ekiti, Ikere); C, Togoda sp. (Ghana), ovipositor, lateral view; D, Togoda sp. (Ghana), female, abdomen, ventral view. Scale bars: 1 mm. Credits: A, Gernot Kunz; B, Babajide Agboola. 479 Review of the genus Togoda Melichar, 1906 ZOOSYSTEMA • 2025 • 47 (23) apical fourth of clavus. Clavus long, 2/3 of wing length. Basal cell narrow. Radius and median running from the same point on basal cell, both firstly furcating closely to basal cell in basal wing third. Forewing vein branching sequence: R 5-7; M13-16, M1 and M2 furcating before wing middle and again apically; CuA 2, furcating after M in basal wing third. Legs not flattened neither foliated. Hind tibia with ten spines apically. First metatarsomere with two latero-apical and five intermediate spines arranged in an arc. Coloration (Fig.4C, D) Coryphe yellowish light brown. Metope brown to dark brown. Head laterally dark brown to black above antenna and light brown below it. Scapi and pedicels brown. Postclypeus light brown to brown frontally. Anteclypeus yellow. Rostrum yellow, with black apex. Proand mesonotum brown to dark brown. Fore wings dark brown to black. Basal process of forewing precostal margin yellowish light brown to dark brown. Fore and middle coxae and hind trochanters brown. Fore femora and tibiae yellowish brown. Fore and middle tarsi brown. Middle femora with brown stripes in female. Third tarsomeres and claws brown. Apices of leg spines black. Female abdominal sternites light yellow except black sternite VII. Hind margins of gonoplacs and anterior connective laminae of gonapophyses VIII dark brown to black. Male terminalia (Fig.9) Pygofer elongated dorso-ventrally, with hind margins deeply concave in their basal thirds (in lateral view) (Fig.9A). Anal tube elongated, nearly twice as long as wide medially, convex laterally and deeply concave apically (in dorsal view) (Fig.9B, C). Anal column small and wide. Penis narrow (Fig.9D, E, F). Phallobase short, 1/3 of aedeagus length, symmetric. Aedeagus with denticles in its proximal half and with two long narrow subapical processes directed downwards – one on left side (Fig.9D, E, c) and one – on right side, almost dorsally (Fig.9D, E, F, b). Style with weakly convex hind margin and large apical tooth (Fig.9H, G, at). Connective with small elongate cup (Fig.9D). Measurement Total length. Male: 5.5mm. Female: 6.5mm. AB CD Fig. 4. — Habitus of Togoda spp.: A, B, Togoda africana Melichar, 1906, Togo (Bismarckburg), male, dorsal view (A) and lateral view (B); C, D, Togoda excisa n. sp., female paratype, dorsal view (C) and lateral view (D). Scale bar: 1 mm. 486 ZOOSYSTEMA • 2025 • 47 (23) Gnezdilov V. M. descripTion Structure (Figs5C, D; 6H, G) Coryphe three times as wide as long medially, smooth, without carinae; anterior margin convex; posterior margin concave (Fig.5C). Metope with two weak carinae medially and groove between them, with sparse pustules (Fig.6G). Coryphe-metope margin keel-shaped. Postclypeus with strong triangularshaped median carina (Fig.5D; 6H). Anteclypeus with fine median carina. Eyes large, diameter of each eye 1.5-2 times as width of coryphe. Fore wings nearly oval, protruding beyond ovipositor apex (Fig.5D). Basal cell small and oval. Radius and median running from the same point on the basal cell. Radius and median firstly furcating closely to basal cell in basal wing fifth. Many transverse veins between branches of median vein. Forewing vein branching sequence: R 5; r-m 6; M 5; CuA 2, furcating near to wing middle. Fore and middle femora flattened or nearly foliate. Hind tibia with ten spines apically. First metatarsomere with two latero-apical and five intermediate spines arranged in arc apically. Coloration (Fig.5C, D) Generally brown to dark brown. Coryphe reddish brown. Metope dark brown, with yellowish pustules. Head laterally with wide black stripe above antennae. Scapi, pedicels, and postclypeus dark brown to black. Anteclypeus light yellow. Rostrum light yellow, with black apex. Proand mesonotum reddish brown to dark brown. Fore wings reddish brown, with large oval black spot on each clavus, between Pcu and first anal vein, and with several large black spots inside of cells on corium including oval one between posterior branch of median and CuA in front of claval spot. Hind wings dark brown. Fore and middle coxae, femora, and tibiae dark brown. Trochanters light yellow. Hind femora dark brown. Hind tibiae dark brown proximally and light yellow distally. First and second tarsomeres of fore and middle tarsi light yellow; third tarsomeres dark brown. Hind tarsi light yellow. Claws dark brown to black. Apices of leg spines black. Abdominal sternites dark brown. Ovipositor yellowish brown. MADAGASCAR SOUTH AFRICA MOZAMBIQUE ZANZIBAR GHANA TOGO NIGERIA Indian ocean Atlantic ocean CENTRAL AFRICAN REPUBLIC Fig. 11. — Distribution of the genus Togoda Melichar, 1906. Abbreviations: ��, Togoda sp.; ��, T. africana Melichar, 1906; ��, T. carinata n. sp.; ��, T. excisa n. sp.; ��, T. minima n. sp.; ��, T. transvaalensis Schmidt, 1910. Not to scale. 487 Review of the genus Togoda Melichar, 1906 ZOOSYSTEMA • 2025 • 47 (23) Measurement Total length. Female: 5.5mm. Togoda sp. (Figs1B-F; 3C, D) maTerial examined.— Ghana • 2 ♀ ; Angu village; 98m; 5°02.007’N, 1°45.602’W; sweeping trees and shrubs in forest and forest edge; 1.V.2022; Z. Assan& V.M. Gnezdilov leg.; ZIN • 2 ♀ ; Kakum National Park; 5°21.198’N, 1°23.034’W; 202m; sweeping in the forest canopy at 11-40m; 4-6.V.2022; V.M. Gnezdilov leg.; ZIN. noTes. —The specimens from Ghana are close to Togoda africana Melichar and T. excisa n.sp. based on metopial proboscis with frontal concavity. Final identification needs examination of male. Togoda sp. maTerial examined.— Tanzania • 2 ♀ ; “Zanzibar / Africa or. 1892 / N 143-1924”; ZIN. noTes.— Final identification needs examination of male. DISCUSSION The tribe Trienopini is characterized by unique synapomorphy – the presence of ear-shaped (or loop-shaped) process (or projection) of forewing precostal margin basally bearing dense marginal setae (Fig.1 A, B, C, F) same as on forewing precostal margin along its whole length (Fig.1E). Another character of the tribe is the presence of semicircular lobe of forewing margin behind claw-shaped apex of clavus also bearing marginal setae (Fig.2). Both ear-shaped processes and semicircular processes of forewings apparently may stick out perpendicularly to forewing plane in alive specimens (Fig.3A, B), but became pressed to fore wings in dry specimens. There is no data on behavior neither ecology of Trienopini taxa and currently it is impossible to explain the role of these forewing processes and lobes. Apparently the center of origin and speciation of the tribe Trienopini is located in Western Equatorial Africa as main part of genera (Fritzruehlia Schmidt, 1924, Ingoma Fennah, 1954, Togoda Melichar, 1906, and Trienopa Signoret, 1860) are found in this region with subsequent distribution to Eastern and Southern Africa and Madagascar, with surrounding islands like Zanzibar and the Seychelles (Schmidt 1924; Melichar 1906; Metcalf 1958; Gnezdilov in press). Currently Madagascar is one of the hot spots for Trienopa biodiversity with an outshift to the Seychelles from where only one species, Trienopa typica (Distant, 1909), apparently closely related to Madagascan T.flavida Signoret, 1860, is recorded. As Trienopini species have rather well developed fore and hind wings the ancestors of modern island taxa, e.g. Togoda and Trienopa species, could be carried from continental Africa by wind dispersal across water barriers to the islands of Indian Ocean. Such a possibility was hypothesized by Ross (1968) to explain the origins of New World species of the genus Exitianus Ball, 1929 (Cicadellidae) by long-distance trans-Atlantic dispersal from Africa received the most support from the analyses according to Zahniser& Dietrich (2015). Distinct metopial proboscis of Togoda species makes them superficially similar to some members of the tribe Parahiraciini Cheng & Yang, 1991 of the family Issidae, e.g. the genus Bardunia Stål, 1863 (Gnezdilov 2011, figs1-18) which is apparently convergent in these planthopper families. While cuspidal apex of forewing clavus also known for Parahiraciini, e.g. the genera Thabena Stål, 1866 and Pseudochoutagus Che, Zhang & Wang, 2011 (Gnezdilov 2015, figs7, 8), may be connected with coleopterization of fore wings. The genus Togoda Melichar is currently distributed across continental tropical Africa from its west to south-east and to Zanzibar and Madagascar, with five species described so far. Males are needed to find out which species are presented in Nigeria, Ghana, and Zanzibar as well as to confirm species status of T.transvaalensis Schmidt, 1910. Acknowledgements I am glad to thank Prof. DrThierry Bourgoin and DrAdeline Soulier-Perkins (Paris, France) for their kind permissions to study the specimens, DrVladimir V. Neimorovets (Saint Petersburg, Russia) and Mr. Konstantin A. Benken (Equipment Sharing Centre “Taxon” (https://www.ckp-rf.ru/ckp/3038/), Saint Petersburg, Russia) for taking photos, DrBabajide Agboola (University of Ibadan, Nigeria, Ekiti Biodiversity Survey Project, https://www.inaturalist.org/projects/ekiti-biodiversitysurvey-project and Ekiti State Forestry Commission) and Mr. Gernot Kunz (Graz, Austria) for their permissions to use the photos of Togoda sp. from Nigeria and Trienopa typica (Distant, 1909) from the Seychelles, and Prof. DrHannelore Hoch (Berlin, Germany) and anonymous reviewer for their valuable comments on the manuscript. The study was performed in the framework of the Russian State Research project No. 125012901042-9. 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Zoological Journal of the Linnean Society 175(3): 473-495. https://doi. org/10.1111/zoj.12292 Submitted on12February 2025; accepted on 21March 2025; published on 30 September 2025.