Afrachilini trib. nov. of Achilidae from Southern Africa (Hemiptera: Fulgoromorpha: Fulgoroidea)

275 European Journal of Taxonomy 1026: 275–303 https://doi.org/10.5852/ejt.2025.1026.3123 europeanjournaloftaxonomy.eu ISSN 2118-9773 2025 · Stroiński A. et al. This work is licensed under a Creative Commons Attribution License (CC BY 4.0) Received: 28 March 2025 • Accepted: 28 August 2025 • Published: 28 November 2025 Topic editor: Tony Robillard • Section editor: Christopher Dietrich • Desk editor: Pepe Fernández Research article urn:lsid:zoobank.org:pub:41780D0A-0651-4A03-BC44-23276972C747 Afrachilini trib. nov. of Achilidae from Southern Africa (Hemiptera: Fulgoromorpha: Fulgoroidea) Adam STROIŃSKI 1 , Alicja M. BRYSZ 2 & Jacek SZWEDO 3,* 1 Museum and Institute of Zoology, Polish Academy of Sciences, 51/55, Twarda St, 00-818 Warsaw, Poland. 2,3 Department of Invertebrate Zoology and Parasitology, University of Gdańsk, PL80-308 Gdańsk, Poland. * Corresponding author: [email protected] 1 Email: [email protected].pl 2 Email: [email protected] Abstract. This paper describes a new species of Afrachilus Fennah, 1965 (Hemiptera: Fulgoromorpha) – Afrachilus montanifynbosensis sp. nov., from the tribe Afrachilini trib. nov., subfamily Myconinae, found in the West Cape of South Africa. This region is one of the Earth’s biologically most diverse areas, also characterized by the phylogenetic antiquity of its invertebrates. A brief discussion is provided on the host plant records of the Afrachilini trib. nov. Morphological peculiarities and diagnostic features of the new tribe are discussed, especially a modification of the head capsule with a subdivision of the frontal area, which is unique in Achilidae, the presence of platellae on the first two metatarsomeres, open cell C1 on tegmen, and the presence of two terminals of CuA on the hind wing, as well as the presence of a subvaginal plate in females, which is another peculiar feature not commonly present among Achilidae. Further distributional data on Achiplecton stilleri (Achiplectini) are provided herein. Keywords. Afrachilini trib. nov., Afrachilus montanifynbosensis sp. nov., planthopper, new species, new tribe, distribution, South Africa. Stroiński A., Brysz A.M. & Szwedo J. 2025. Afrachilini trib. nov. of Achilidae from Southern Africa (Hemiptera: Fulgoromorpha: Fulgoroidea). European Journal of Taxonomy 1026: 275–303. https://doi.org/10.5852/ejt.2025.1026.3123 Introduction The family Achilidae Stål, 1866 is one of the most widespread families of planthoppers, currently present on all continents, with the exception of Arctica and Antarctica. It is distinguished by its high level of diversity, particularly in the northern regions of the subtropics and across moderate climates. The family encompasses a total of 164 genera and 525 species, classified into 12 extant and 3 extinct tribes. The extinct representatives of this family are found in fossil resins (amber) deposits and as adpression fossils in sedimentary rocks (Bourgoin 2024; Brysz et al. 2024; Deng et al. 2024). The classification, content and concepts of the groups within the family are subjects of vivid and ongoing discussion European Journal of Taxonomy 1026: 275–303 (2025) 276 (Brysz et al. 2024; Deng et al. 2024). The family Achilidae in South Africa has received only limited attention from researchers, with eight genera and nine species documented to date, the majority of which are endemic. The following taxa are representative of the Achilidae family found in South Africa: Apatesoninae Metcalf, 1938: Ilvini Emeljanov, 1991 – Ilva nigrosignata Stål, 1866; Mycarini Emeljanov, 1991 – Katbergella griseobrunnea Fennah, 1950, K. bastet Emeljanov, 2005; Myconinae Fennah, 1950: Plectoderini Fennah, 1950 – Caffropyrrhyllis bicuspidata bicuspidata Fennah, 1950 and Caffropyrrhyllis bicuspidata epona Fennah, 1958, Cnidus variegatus (Stål, 1855), Phypia albipennis (Stål, 1855); tribal placement of Brachypyrrhyllis Fennah, 1967a, placed once in Plectoderini is questionable. The Myconinae tribe Achiplectini Brysz, Stroiński & Szwedo, 2024 has been added recently, along with Achiplecton stilleri Brysz, Stroiński & Szwedo, 2024. The Afrachilini trib. nov. is herein described, with Afrachilus mirabilis Fennah, 1965 and Afrachilus montanifynbosensis sp. nov., completing the list of taxa already reported. The present contribution serves to broaden our understanding of the taxonomic diversity and morphological disparity of the modern Achilidae, thereby augmenting our knowledge of the endemicity of insects in the Cape Floristic Region (CFR), which is biologically the most diverse area on Earth (Cowling et al. 1992; Allsop et al. 2014). Material and methods Observations and documentation were conducted at the Laboratory of Evolutionary Entomology and Museum of Amber Inclusions at the University of Gdańsk in Gdańsk, and at the Museum and Institute of Zoology at the Polish Academy of Sciences in Warsaw. The observations and documentation were conducted utilising stereoscopic microscopes, namely the Leica M205A equipped with the Leica DM6000 camera, the Olympus SZX10 with the EP50 camera attached, and the Olympus BX51 with the Canon EOS 90D camera. The photographs of the habitus and internal structures were captured using a stereo microscope Leica MZ 16 with an IC3 D camera. The final images were subjected to adjustments using Helicon. The software in question is ver. 5.0, and the Adobe Photoshop software is ver. 7.0. The compositions were rendered using CorelDRAWX7 software. The SEM photographs of uncoated specimens were taken in the Laboratory of Scanning Microscopy, MIZ PAS (Warsaw), using a scanning electron microscope HITACHI S-3400N under low vacuum conditions. In order to reveal the genital structures, it was necessary to excise the whole abdomen of the specimen examined and then clear it for 30 minutes in a warm (50°C) 10% potassium hydroxide (KOH) solution, with the addition of a few drops of black chlorazol (CAS No. 1937-37-7) for the staining of the ectodermic genital structures. This method was based on the technique introduced by Carayon (1969). The dissection and cleansing of the genital structures was conducted in distilled water. The utilisation of morphological terminology in this context is consistent with the proposals of Anufriev & Emeljanov (1988), Emeljanov (1995), Bartlett et al. (2014), and Asche (2015). Differences in terminology used in the Emeljanov’s descriptions and interpretations of head topological structures (Emeljanov 1991, 1992; Dmitriev 2009) are as follows: coryphe = vertex auct.; metope = frons auct.; upper portion of ‘frons’ separated by carinations – acrometope; lower section of ‘frons’ – eumetope. The nomenclature of the fore wing (tegmen) follows the interpretation proposed by Bourgoin et al. (2015) and Stroiński (2020), for hind wings after Anufriev & Emeljanov (1988) and Emeljanov (1991, 1992). The nomenclature of the antennal structures is in accordance with Stroiński et al. (2011). The terminology of the genitalia follows proposals of Bourgoin (1993) and Asche (2015) for the female. The quotations from the examined material labels are verbatim and are enclosed in double quotation marks. Repositories Material is deposited in: MAIG = collection of the Laboratory of Evolutionary Entomology and Museum of Amber Inclusions, University of Gdańsk STROIŃSKI A. et al., New achilid planthoppers tribe Afrachilini from Southern Africa (Hemiptera) 277 SANC = Southern African Collection of Insects, Pretoria TMP = Ditsong National Museum of Natural History (formerly Transvaal Museum), Pretoria, Southern Africa Results Systematics Class Insecta Linnaeus, 1758 Order Hemiptera Linnaeus, 1758 Suborder Fulgoromorpha Evans, 1946 Superfamily Fulgoroidea Latreille, 1807 Family Achilidae Stål, 1866 Subfamily Myconinae Fennah, 1950 Diagnostic characters (modified after Emeljanov 1991) Coryphe visibly exceeding anterior margin of the eyes. Lora visible in ventral view. Compound eye with an indentation. Metatibial subgenual lateral spine present. Tegmina at rest folded flat with membranes overlapping, postcostal area narrow, vein MP3+4 usually not branched. Hind wing vein ScP+R and MP veins never exit basal cell far apart, vein A2 usually widened apically, never reaches wing margin, median fold usually reaching wing margin. Composition Achiplectini Brysz, Stroiński & Szwedo, 2024: [Achiplecton Brysz, Stroiński & Szwedo, 2024]; Afrachilini trib. nov. [Afrachilus Fennah, 1965]; Amphignomini Emeljanov, 1991: [Amphignoma Emeljanov, 1991; †Amphignokachinia Brysz & Szwedo, 2023]; Mycarini Emeljanov, 1991: [Acocarinus Emeljanov, 1991; Emeljanocarinus Bourgoin & Soulier-Perkins, 2006; Katbergella Fennah, 1950; Mycarinus Emeljanov, 1991; Mycarus Emeljanov, 1991]; Myconini Fennah, 1950: [Ganachilla Wang & Huang, 1989; Haicixidia Wang, 1989; Myconellus Fennah, 1950; Myconus Stål, 1862]; †Niryasaburniini Wang & Bourgoin, 2024 in Deng et al. 2024: [†Niryasaburnia Szwedo, 2004, †Sinuovenaxius Wang & Bourgoin, 2024 in Deng et al. 2024]; Plectoderini Fennah, 1950: [Abas Fennah, 1950; Agandecca White, 1879; Akotropis Matsumura, 1914; Amblycratus Uhler, 1895; Aphypia Melichar, 1908; Argeleusa Kirkaldy, 1906; Aristyllis Kirkaldy, 1906; Ballomarius Jacobi, 1941; Bathycephala Fennah, 1950; Benella Kirkaldy, 1906; Betatropis Matsumura, 1914; Caffropyrrhyllis Fennah, 1950; Calerda Signoret, 1863; Callichlamys Kirkaldy, 1907; Callinesia Kirkaldy, 1907; Caristianus Distant, 1916; Catonia Uhler, 1895 (subgen. Catonia Uhler, 1895; subgen. Pyren Fennah, 1950); Catonoides Metcalf, 1938; Cenophron Fennah, 1969; Cernea Williams, 1977; Chroneba Stål, 1859; Cionoderella Fennah, 1950; Clidonisma Fennah, 1969; Clusivius Distant, 1917; Cnidus Stål, 1866; Cocottea Williams, 1977; Cythna Kirkaldy, 1906; Deferunda Distant, 1912 (subgen. Deferunda Distant, 1912; subgen. Tugaia Dlabola, 1961); Epirama Melichar, 1903; Epiusana Fennah, 1950; Epiusanella Synave, 1959; Eudeferunda Chen, Yang & Wilson, 1989; Eurynomella Fennah, 1967b; Eurynomeus Kirkaldy, 1906; Francesca Kirkaldy, 1906; Gongistes Fennah, 1969; Gordiacea Metcalf, 1948; Haitiana Dozier, 1936; Hamba Distant, 1907; Hemiplectoderes Fennah, 1950; Horcomotes Fennah, 1969; Indorupex Fennah, 1965; Isodaemon Fennah, 1969; Juniperthia O’Brien, 1985; Kardopocephalus Metcalf, 1938; Kawanda Fennah, 1950; Kawandella Synave, 1959; Kempiana Muir, 1922; Koloptera Metcalf, 1938; Kosalya Distant, 1906; Kurandella Fennah, 1950; Lanuvia Stål, 1866 (subgen. Lanuvia Stål, 1866; subgen. Nulavia Emeljanov & Gnezdilov, 2023); Magadha Distant, 1906; Magadhaideus Long & Chen, 2017; Mahuna Distant, 1907; Martorella Caldwell & Martorell, 1951; Metalticeps Dmitriev, 2020; Mlanjella Fennah, 1950; Momar Fennah, 1950; Moraballia Fennah, 1950; Neoacus Dmitriev, 2020; Nephelesia Fennah, 1965; Nephelia Kirkaldy, 1907; Nyonga Synave, 1959; Opsiplanon Fennah, 1945; Paracatonia Fennah, 1950; Paraclusivius Fennah, European Journal of Taxonomy 1026: 275–303 (2025) 278 1950; Paragandecca Fennah, 1950; Parakosalya Distant, 1917; Paraphypia Synave, 1960; Parargeleusa Fennah, 1950; Paratangia Melichar, 1903; Phenelia Kirkaldy, 1906; Phypia Stål, 1862; Plectoderes Spinola, 1839 (subgen. Plectoderes Spinola, 1839; subgen. Plectoderella Fennah, 1950); Plectoderoides Matsumura, 1914; Plectoringa Fennah, 1950; Prosagandecca Fennah, 1950; Pseudhelicoptera Fowler, 1904; Pyrrhyllis Kirkaldy, 1906; Quadrana Caldwell & Martorell, 1951; Remosachilus Fennah, 1950; Rhinocolura Fennah, 1950; Rhotaloides Fennah, 1965; Rupex Fennah, 1950; Salemina Kirkaldy, 1906; Semibetatropis Chen, Yang & Wilson, 1989; Spino Fennah, 1950; Symplegadella Fennah, 1950; Synecdoche O’Brien, 1971; Taloka Distant, 1907; Tangina Melichar, 1903; Thectoceps Williams, 1977; Usana Distant, 1906; Williamsus Özdikmen & Demir, 2007; Xerbus O’Brien, 1971; Zathauma Fennah, 1949; Rhotalini Fennah, 1950: [Errada Walker, 1870; Errotasa Emeljanov, 2005; Hebrotasa Melichar, 1914: Rhotala Walker, 1857; Taractellus Metcalf, 1948]; †Waghildini Szwedo, 2006: [†Waghilde Szwedo, 2006]. Key to the tribes of Myconinae Fennah, 1950 1. Subantennal carina present on genae; mesonotum mediolateral carinae absent; tegmen RP vein with 1 terminal, CuA with 3 terminals; hind wing median carina not exceeding mp-cua crossvein ......... ....................................................................................................... Amphignomini Emeljanov, 1991 – Subantennal carina absent; mesonotum mediolateral carinae present; tegmen RP vein with more than 1 terminal, CuA never with 3 terminalia; hind wing median carina exceeding mp-cua crossvein .. 2 2. Body flattened laterally; coryphe extremely elongated, triangular; tegmen held curved over body, longitudinal veins carinated or frosted depending on wing region ..................................................... ................................................................................. Achiplectini Brysz, Stroiński & Szwedo, 2024 – Body flattened dorso-ventrally; coryphe not extremely elongated, never triangular; tegmen held flat over body, longitudinal veins smooth ............................................................................................... 3 3. Acrometope with lateral compartments; eumetope with reduced median carina; tibia without lateral spines (including subgenual one); tegmen’s cell C1 cell open ......................... Afrachilini trib. nov. – Acrometope without lateral compartments; eumetope with full median carina; tibia with at least 1 lateral spine; tegmen’s cell C1 cell closed ........................................................................................ 4 4. Tegmen with vein RP with 3 terminals; hind wing veins ScP+R and MP leaving basal cell from the same point ......................................................................................................................................... 5 – Tegmen with vein RP with 2 terminals; hind wing veins ScP+R and MP leaving basal cell close to each other but not from the same point ............................................................................................ 6 5. Pronotum narrowed; tibia with 2–3 lateral spines and a subgenual one; tegmen with vein MP with 6+ terminals, vein CuA with 4 to 5 terminals; hind wing median fold singular ................................. ...................................................................................................................... Myconini Fennah, 1950 – Pronotum streamer-like; tibia with 1 lateral spine, subgenual one absent; tegmen with MP vein with 3 terminals, vein CuA with 2 terminals; hind wing median fold branched ........................................ ..................................................................................................................Plectoderini Fennah, 1950 6. Coryphe not strongly elongated, anterior margin never rounded; pronotum narrowed; tibia with 1–3 lateral spines (not including the subgenual one); tegmen with vein CuA with 2 terminalia ............ 7 – Coryphe strongly elongated with a rounded anterior margin; pronotum robust, trapezoidal; tibia with 5–7 lateral spines and a subgenual one; tegmen with vein CuA vein with 5+ terminalia ................ 8 7. Tibia with 2–3 lateral spines; tegmen with vein MP with 4 terminalia; hind wing median fold reaching wing margin ............................................................................................ Mycarini Emeljanov, 1991 STROIŃSKI A. et al., New achilid planthoppers tribe Afrachilini from Southern Africa (Hemiptera) 279 – Tibia with 1 lateral spine; tegmen with vein MP vein with 3 terminalia; hind wing median fold not reaching wing margin ......................Niryasaburniini Wang & Bourgoin, 2024 in Deng et al. 2024 8. Body flattened laterally; acrometope with lateral compartments; pronotum not strongly elongated into coryphe, postocular carinae absent; tegmen RP vein with 3+ terminalia, CuA with 5 terminalia; subapical platellae present on apical teeth of metatarsomeres I and II ............................................... ...................................................................................................................Waghildini Szwedo, 2006 – Body flattened dorso-ventrally; acrometope without lateral compartments; pronotum strongly elongated into coryphe, postocular carinae present; tegmen RP vein with 2 terminalia, CuA with 2–3 terminalia; subapical platellae present only on apical teeth of metatarsomere II ............................... ...................................................................................................................... Rhotalini Fennah, 1950 Tribe Afrachilini trib. nov. urn:lsid:zoobank.org:act:1D612C99-2D0B-47BB-9888-746E967990EE Type genus Afrachilus Fennah, 1965, here designated. Diagnosis The body distinctly dorso-ventrally flattened, fore wings (tegmina) membranes overlapping. The head capsule with separated acrometope; lateral carinae of frons (eumetope) incomplete, obsolete in lower portion; frons (eumetope) divided by complete transverse carination into upper portion (aneumetope) and lower portion (kateumetope); lateral carinae of postclypeus absent (as in Amphignomini), postclypeus, lora and maxillary plates fused, loral suture partly preserved, anteclypeus wedged in between lora; postclypeus and anteclypeus without median and lateral carinae; ocellus in touch with lower margin of compound eye. The tegmen with open cell C1 (exclusive character among Achilidae); stem MP very long, forked distinctly apicad of claval apex (similar pattern is to be observed in some Plectoderini, stem MP forked beyond claval apex in Achiplectini, Amphignomini and Niryasaburniini); claval fold directed to tornus, not reaching tornal veinlet; clavus open. The hind wing with basal cell (present in Plectoderini, basal cell absent in Achiplectini), median fold simple, not intersecting cubital area (median fold forked and intersecting cubital area in Plectoderini), CuA 1 single (forked in other tribes). The metatibia without subgenual and lateral spines, basimetatarsomere and midmetatarsomere with subapical platellae. The female terminalia with subvaginal plate; bursa copulatrix single pouched, ornamented, without sclerite. Distribution Afrotropical region, Southern Africa, the Cape. Composition Only type genus so far. Genus Afrachilus Fennah, 1965 Afrachilus Fennah, 1965: 81, figs 11–16 [described, female]. Afrachilus – Fennah 1967a: 659, fig. 2a–e [described, male]; 1969: 87, 115. — O’Brien 1971: 4. — Koçak & Kemal 2010: 11. — Dmitriev 2022: 12. Type species Afrachilus mirabilis Fennah, 1965; by original designation and monotypy. European Journal of Taxonomy 1026: 275–303 (2025) 280 Revised diagnosis Superficially resembles Haitiana Dozier, 1936 (Plectoderini) by virtue of its elongated antennae and deep incision on lower margin of its compound eye. The head with acrometope, and eumetope distinctly separated by transverse carination; eumetope subdivided into upper portion (aneumetope) and lower portion (kateumetope), differing in sculpture and coloration. The lora lying almost in same plane as disc of postclypeal part of the face. The rostrum surpassing the mesocoxae but not reaching the metacoxae. Compound eyes deeply incised ventrally. Pronotum with at least one carina placed laterally on each side between eye and tegula; pustulae present between lateral margin of disc and carina collateralis. Metatibiae laterally unarmed, apically with seven or eight teeth. Male terminalia (after type species Afrachilus mirabilis in Fennah 1967a: 659): anal segment of male in dorsal view more than twice as long as broad at apex, lateral margins straight, diverging distally, apical margin truncate or very shallowly convex, anal foramen at apex, anal style long, longer than pygofer, in dorsal view about twice as long as broad at widest part, in lateral view depressed. Pygofer relatively short, dorsolateral angles produced caudad; no medioventral process developed. Aedeagus long, subequal in width throughout, shallowly curved upward distad, tapering to a point at apex; a pair of slender spinose processes emerging ventrolaterally, directed dorsocaudad. Genital styles rather more than twice as long as broad, ventral margin convex, apical margin truncate, dorsal margin thickly callused, produced laterad at one quarter from apex in a short spinose process, and mesad in a broad thumb-shaped lobe. Diversity and distribution The genus Afrachilus and its two known so far species: A. mirabilis Fennah, 1965 and A. montanifynbosensis sp. nov. are known from localities in Western and Eastern Cape Province, Republic of Southern Africa. Afrachilus montanifynbosensis sp. nov. urn:lsid:zoobank.org:act:EDA72A64-A1E4-44D3-8C7B-9D27F50BDCA2 Figs 1–11 Diagnosis Smaller than Afrachilus mirabilis; anterior margin of head arcuate (anterior margin of the head more acute in A. mirabilis); mesonotum with median carina present in anterior and median section (anterior carina present only in median section of mesonotum in A. mirabilis), metatibio-tarsal formula is 8 : 8 : 8 (metatibio-tarsal formula 9 : 9 : 9 in A. mirabilis); subvaginal plate subquadrate (subvaginal plate subrectangular with distinctly rounded apex in A. mirabilis). Etymology Specific epithet derived from the habitat wherefrom the examined material originates. Type material Holotype SOUTH AFRICA • ♀; “Western Cape, Gifberg Pass summit, Vanrhynsdorp; 31°45ʹ S, 18°42ʹ E; 360 m a.s.l.; 9 Oct. 2002; M. Stiller leg.; sweeping, low grass, forbs & restios, at FM tower; National Coll., of Insects, Pretoria, S. Afr.”; SANC. Paratypes (9 ♀♀) SOUTH AFRICA – Northern Cape • 1 ♀; “Northern Cape, Vanrhyns Pass, summit view, Nieuwoudtville; 31°22′ S, 19°01′ E; 7.x.2002; 832 m a.s.l.; leg. M. Stiller leg.; Sweeping grass, forbs, shrubs, around parking, area at view point; National Coll. of Insects, Pretoria, S. Afr.”; SANC • 1 ♀; “Van STROIŃSKI A. et al., New achilid planthoppers tribe Afrachilini from Southern Africa (Hemiptera) 281 RHYN’S-, PASS; 4-5.11’33; leg. G. van Son”; TMP. – Western Cape • 1 ♀; same data as for holotype; SANC • 1 ♀; “Western Cape, Prov. Wiedouw, Farm foot, Gifberg Pass, SE Vanrhynsdorp; 31°44′ S, 18°46′ E; 3–10.x.Oct. 2002; 120 m; leg. M. Stiller; swept off Dicerothamnus rhinocerotis, Asteraceae; National Coll. of Insects, Pretoria, S. Afr.”; SANC • 1 ♀; same data as for preceding; MAIG • 1 ♀; same data as for preceding; “Swept off Nylandtia scoparia, Polygalaceae; National Coll. of Insects, Pretoria, S. Afr.”; SANC • 1 ♀; same data as for preceding; “Swept off Diosma hirsute, Rutaceae; National Coll. of Insects, Pretoria, S. Afr.”; SANC • 1 ♀; same data as for preceding; “Swept off, Galenia africana, AIZOACEAE; National Coll. of Insects, Pretoria, S. Afr.; SANC • 1 ♀; “Western Cape, Prov. Driehoek, Farm Cedarberg, SE of Clanwilliam; 32°26′ S, 18°13′ E; 1200 m; 13.x.2002; leg. M. Stiller leg.; Swept off Leucadendron pubescens, Proteaceae; National Coll. of Insects, Pretoria, S. Afr.”; SANC. Note The coordinates on the label of the specimen collected 13 Oct. 2002 are wrong. The correct coordinates are 32°26ʹ S, 19°11ʹ E (M. Stiller pers. com.) Description Coloration (Fig. 1A–F). General coloration brownish-ochraceous, with darker anterior part of body: head, pronotum and mesonotum. Vertex with anterior section dirty apricot hue to light penny brown, median and posterior section dark blackish-brown; lateral margins in posterior section ivory, two ivory stains at anterolateral angles, posterior margin ivory medially to brownish posteriad; acrometope caramel, aneumetope light caramel, kateumetope ivory, with two light tawny fuzzy stains; upper portion of postclypeus ivory, lower portion tawny, anteclypeus tawny mediad, lighter laterad; lora ivory in upper section, more tawny in lower section, margins light caramel; lateral portion of head capsule and antennal fovea ivory, scapus ivory at base with caramel ring in upper section, pedicel dark caramel. Pronotum with disc blackish in anterior section, penny brown near posterior margin; median and lateral carinae of pronotum ivory, lateral portions of pronotum blackish-penny brown posteriad, pustulae ivory; pectoral lobes light caramel dorsally to ivory ventrad. Mesonotum, tar-black, with five elongated stains of a burnt orange hue located along the posterior margins and at the tip of the scutellum. Tegmina semitransparent, veins with alternation of walnut brownish-ivory and whitish hues, costal area and anterior apical cells with oblique, alternate tawny – whitish transparent stains, membrane light tawny, fourth apical cell with irregular chocolate colored elongated spot. Legs dark straw to tawny, apical teeth of metatibia and basiand midmetatarsomere dark chocolate-blackish. Abdominal sternites ranging from dark-straw to light tawny laterad, median portions chockolate brown, posterior margins ivory to light straw; median section of pregenital sternite with subtriangular caramel stain and caramel lateral portions; subvaginal plate dirty ivory; gonoplacs light tawny, anal tube light tawny-caramel. MeasureMents. Body with wings 4.2–5.4 mm in length. appearanCe. Body distinctly dorso-ventrally flattened, tegminal membranes overlapping. Head with compound eyes, slightly narrower than pronotum. Coryphe (vertex) with same wide at posterior and at anterior lateral angles and 1.46–1.63 times as wide as long in mid line slated. Head (Figs 1A–D, 2A–F, 3A–F, 4A–F). Anterior margin of coryphe angulately convex, lateral margins sinusoidal, converging anteriad; posterior margin slightly concave, elevated, reaching ⅔ of the compound eye length; all margins carinate; disc of vertex concave, divergently strigate, with median carina almost reaching anterior margin. The acrometope with two lateral compartments, elongately triangular, lower margin of acrometope distinctly arcuate, almost fused with upper carina between acrometope and coryphe (vertex). The frons (eumetope) divided by transverse carination; upper portion, aneumetope with subvertically costulate sculpture; lower portion, kateumetope with strigate sculpture; kateumetope with arcuate incision above the frontoclypeal suture, and median eminence, prolonged to postclypeus. European Journal of Taxonomy 1026: 275–303 (2025) 282 Fig. 1. Afrachilus montanifynbosensis sp. nov., holotype, ♀ (SANC). A. Habitus, lateral view. B. Habitus, dorsal view. C. Anterior part of body, frontal view. D. Anterior part of body, dorsal view. E. Hind legs and abdomen, ventral view. F. Hind wing. G. Distribution map, physical. H. Distribution map, ecoregions. STROIŃSKI A. et al., New achilid planthoppers tribe Afrachilini from Southern Africa (Hemiptera) 283 The genal suture indistinct, posterior margins of lora marked by coloration. The frontoclypeal suture angulately-arcuate. The postclypeus subhexagonal, convex, weakly and sparsely striolate; the anteclypeus about twice as long as wide at base, more convex in upper section, the clypellus elongately triangular. The rostrum reaching to the anterior margin of metacoxae, apical segment shorter than subapical, apical segment more than 3 times as long as wide. The compound eye kidney-like, with median indentation to half of its width, without subocular callosity; lateral ocellus adjacent to compound eye, behind its anterior Fig. 2. Afrachilus montanifynbosensis sp. nov., holotype, ♀ (SANC), SEM photographs. A. Habitus, lateral view. B. Habitus, dorsal view. C. Head and pronotum, dorsal view. D. Mesonotum, dorsal view. E. Anterior part of body, latero-dorsal view. F. Anterior part of body, dorsal view. European Journal of Taxonomy 1026: 275–303 (2025) 290 rounded, apical margin widely rounded, posteroapical angle obtusely rounded, tornus straight, claval margin straight, the angle between A 2 and tornus ca 165°; apex of clavus not reaching ⅔ of tegmen length; the basal cell narrow, about 8 times as long as wide; the stem ScP+R+MP leaving basal cell at same point, with a short stalk, less than half of the basal cell length; the stem ScP+R subparallel to costal margin, merely diverging mediad, first branching of ScP+RA distinctly apicad of apex of clavus, slightly apicad of mp-cua veinlet; branch RP with arcuate base, reaching margin with 3 terminals, basad of anteroapical angle of tegmen; the stem MP slightly sigmoid, forked on membrane, distinctly apicad of apex of clavus, apicad of ScP+R forking, branch MP 1+2 arcuate, with two terminals reaching margin at anteroapical angle, branch MP3+4 not forked, reaching apical margin; the stem CuA straight, parallel to claval margin, forked slightly apicad of claval veins junction; branch CuA 1 directed mediad, then, after mp-cua veinlet, posteriad with shift on membrane reaching margin basad of posteroapical angle; branch CuA2, sigmoid, then shifted on membrane, reaching margin at posteroapical angle; claval vein CuP distinct, claval fold prolonged on membrane, not reaching veinlet icu; claval veins Pcu and A 1 fused apicad of half of clavus length, apicad of ⅔ of tegmen length; apex of clavus obtuse; appendix with transverse wrinkles, wider in anterior and apical portion of membrane; postcostal and radial cell of similar width; cell C1 open, call C3 short, closed with apical imp veinlet on membrane; cell C5 long, nearly 4 times as long as cell C3, closed by apical veinlet icu on membrane; first veinlet rp-mp on membrane, apicad of claval apex, second veinlet rp-mp1+2 short, apicad of stem MP forking; first veinlet mp-cua1, apicad of apex of clavus, slightly basad of stem ScP+R forking; sclerotised macula on fourth apical cell. Fig. 9. Afrachilus montanifynbosensis sp. nov., holotype, ♀ (SANC), SEM photographs. A. Abdomen and terminalia, latero-ventral view. B. Terminalia, latero-ventral view. C. Terminalia, ventral view. D. Median part of gonoplac, ventral view. STROIŃSKI A. et al., New achilid planthoppers tribe Afrachilini from Southern Africa (Hemiptera) 291 Fig. 10. Afrachilus montanifynbosensis sp. nov., holotype, ♀ (SANC). A. Pregenital sternit (flattened) ventral view. B. Anal tube, lateral view. C. Anal tube, dorsal view. D. Anal tube, ventral view. E. Gonoplac, lateral view. F. Gonoplac, dorsal view. G. Subvaginal plate and gonapophysis VIII with endogonocoxal process, ventral view. H. Gonapophysis VIII and endogonocoxal process, lateral external view. I. Gonapophysis VIII and endogonocoxal process, lateral internal view. European Journal of Taxonomy 1026: 275–303 (2025) 292 H ing wing (Figs 1F, 6D). Membranous, shorter than fore wing, with distinct anal lobe, separated by incision; costal margin arcuate at base then almost straight, with the small connecting apparatus lobe at level of ScP+R forking; basal cell present; the stems ScP+R and MP leaving basal cell at point, the stem ScP+R parallel to costal margin, forked in apicad ⅓ of hind wing length; branch ScP+RA short reaching margin well basad of apex; branch RP shifted mediad, then curved, reaching margin basad of hind wing apex; the stem MP straight, diverging mediad, then shifted, reaching margin with two terminals, forked well apicad of veinlets rp-mp and mp-cua1; stem CuA forked slightly apicad of half of hind wing length, with the two terminals CuA1 and CuA2 reaching margin; stem CuP slightly sinuate, diverging in apical section from postcubital fold; Pcu sinuate, A1 forked at ⅓ of its length, A2 arcuate, slightly thickened, not reaching margin of anal lobe, median field present, intersecting mp-cua1 veinlet; cell C1 open, longer than cell C3; cell C3 subtriangular, slightly longer than wide; cell C5 the largest and the longest, open, with margins distinctly diverging towards margin. Fig. 11. Afrachilus montanifynbosensis sp. nov., holotype, ♀ (SANC). A. Gonapophyses IX and gonospiculum bridge, dorsal view. B. Gonapophyses IX and gonospiculum bridge, ventral view. C. Gonapophyses IX and gonospiculum bridge, fronto-latero-dorsal view. D. Bursa copulatrix and spermatheca, lateral view. STROIŃSKI A. et al., New achilid planthoppers tribe Afrachilini from Southern Africa (Hemiptera) 293 aBdoMen. Flattened, with sternites not divided medially. Male terMinalia. Male unknown. FeMale terMinalia (Figs 8–11). The pregenital sternite (Figs 8E–F, 9A–B, 10A), almost rectangular in ventral view, with anterior margin almost straight with small wide incision medially; posterior margin almost straight. The anal tube (Figs 8A–D, 10B–D), in lateral view triangular and bit extending posterior margin of gonoplac; in dorsal view cup-like, upper posterior margin weakly concave, medially shallowly convex; posterior-ventral margin deeply concave; the epiproct distinctly shorter than the paraproct, in dorsal view triangular with median split, not extending ventro-posterior margin of anal tube; the paraproct fusiform, distinctly longer than epiproct and distinctly extending posterior-ventral margin of anal tube; anus placed distinctly after the middle. Whole anal tube and with numerous long setae. Gonoplac (Figs 8A–B, 9B–D, 10E–F) unilobate, well sclerotized, fully covering gonapophysis VIII; posterior margin with numerous setae (bristles) and membranous part forming close line/margin. Gonapophysis VIII (Fig. 10 G–I) wide, distinctly tapering distally; anterior connective lamina (ACL) with two strongly sclerotized arms, apically with 5 strong teeth of similar sizes, apical part strongly curved ventrad. Endogonocoxal process as long as gonapophysis, unilobate; endogonocoxal lobe narrowly sclerotized apically with short bluntly process oriented basally. Gonospiculum as in Fig. 11A–C. Subvaginal plate (Figs 8E–F, 9A–C, 10G) well sclerotised, in ventral view, subquadrate with posterior margin widely arcuate. Bursa copulatrix (Fig. 11D) with membranous single elongate pouch, without visible cell but with sclerotized plates on whole wall; sclerite of bursa absent. Spermatheca well developed; ductus receptaculi narrow and smooth, distinctly shorter than diverticulum ductus; diverticulum ductus with very long smooth and narrow (but wider than ductus receptaculi) ductus and with ovoid and smooth bulla apically (Fig. 11D). Additional localities for Achiplecton stilleri Brysz, Stroiński & Szwedo, 2024 Note: utilising the data collated through citizen science initiatives, notably iNaturalist, we have successfully identified supplementary locations for the species that were documented in the preceding year. Photographs of live specimens of Achiplecton stilleri taken in the wild indicate a wider occurrence of this species in the Western Cape Province. Concurrently, the colouration of the examined dead specimens (Fig. 12A–B) indicates a stable colouration of this species when compared to living specimens. All photographs were taken by Cecile Roux and published with her permission. The original description of Achiplecton stilleri (Brysz et al. 2023) demonstrated that this species occurs in the Montane Fynbos and Renosterveld ecoregion (AT1203). Newly obtained data indicate the occurrence of this taxon in the Lowland Fynbos and Renosterveld ecoregion (AT1202). However, its distribution remains confined to the Western Cape Province and the Cape Floristic Region. SOUTH AFRICA – Western Cape Province, West Coast district • Swartland Municipality; 33°28′12.0″ S, 18°44′14.5″ E; 23 Sep. 2024; Klipkoppie Nature Reserve [https://www.inaturalist.org/observations/244140803] • Bergriver Municipality; 32°36′57.8″ S, 18°18′04.4″ E; 21 Sep. 2024; Rocherpan Nature Reserve [https://www.inaturalist.org/observations/243927735] • Bergriver Municipality; 32°41′18.4″ S, 18°14′14.9″ E; 8 Oct. 2023; Berg River [https://www.inaturalist.org/observations/187225824] • Saldanha Bay Municipality; 33°00'38.9″ S, 18°21′24.7 E; 24 Sep. 2024; Hopefield [https://www.inaturalist.org/observations/243745437]. Discussion The above newly described tribe Afrachilini trib. nov. represents the third endemic tribe of Achilidae for the Cape Province (following Ilvini Emeljanov, 1991 and Achiplectini Brysz, Stroiński & Szwedo, 2024) and exhibits a distinctive set of characters clearly separating it from other Myconinae tribes. A new tribe, Afrachilini trib. nov., with its sole genus Afrachilus Fennah, 1965 somewhat resembles European Journal of Taxonomy 1026: 275–303 (2025) 294 Fig. 12. Achiplecton stilleri Brysz, Stroiński & Szwedo, 2024. A–D. Photos of living specimens. A. Specimen from Klipkoppie Nature Reserve. B. Specimen from Rocherpan Nature Reserve. C. Specimen from Berg River. D. Specimen from Hopefield. E. Distribution map, physical. F. Distribution map, ecoregions. Colors in physical map: red dots = original records; purple dots = new records. Colors in ecoregions map: black dots = original records; purple dots = new records; red color of ecoregions = Montane Fynbos and Renosterveld ecoregion; pink color of ecoregion = Lowland Fynbos and Renosterveld. STROIŃSKI A. et al., New achilid planthoppers tribe Afrachilini from Southern Africa (Hemiptera) 295 some representatives placed in the tribe Plectoderini (e.g., genus Haitiana Dozier, 1936) by the elongated antennae and the lower margin of compound eye deeply incised. The head capsule with separated acrometope and eumetope constitutes a unique attribute, not common among other Achilidae (except Haitiana). The postclypeus without lateral carinae (another exceptional feature) is also to be found in the Amphignomini Emeljanov, 1991 and the extinct Waghildini Szwedo, 2006. The fusion of postclypeus with lora is a rather exceptional feature, present in Amphignomini (also in extinct Ptychoptilini Emeljanov, 1990 of incertae sedis position in Achilidae); however, it is also observed in a few other planthoppers, e.g., in some Flatidae: Antillormenis Fennah, 1942 and Flatoidinus Melichar, 1923. Tegmina with an open cell C1 is a very exceptional character, not known among the Achilidae. The hind wing with only two terminals of CuA is a feature of the Afrachilini shared with the extinct Niryasaburniini Wang & Bourgoin, 2024 in Deng et al. (2024). The structure of hind tibiae with subapical platellae is the same as in Achiplectini. The subapical setae, which vary in their modification among the Achilidae, present on basiand midmetatarsomeres are observed, e.g., in Achilini: Cixidiina Emeljanov, 1992 – in form of platellae – Cixidia Fieber, 1866, extinct Gedanochila Brysz & Szwedo, 2022, extinct Waghilde Szwedo, 2006 (Waghildini Szwedo, 2006) (Asche 2005; Szwedo 2006; Brysz et al. 2022), as platellae in Tropiphlepsini Emeljanov, 1991, as setae in Seviini Emeljanov, 1991 (Emeljanov 1992) and extinct genera of Achilini: Protomenocria Emeljanov & Shcherbakov, 2009, Psycheona Emeljanov & Shcherbakov, 2009, Protepiptera Usinger, 1939 (Emeljanov & Shcherbakov 2009) from the Eocene amber of the Gulf of Gdańsk. As recently the structure of metatibiae and tarsal structures have been redefined among Cixiidae (Brożek et al. 2024), the same comparative structures are necessary to clear the patterns and models among the Achilidae. Fennah (1950) provided a definition of the subvaginal plate as a sclerotized structure, typically manifesting as a moderately broad, transverse plate of the membrane situated between the pregenital sternite and the external orifice of the vagina. This structure is not frequently observed among the known Achilidae and achilid-related taxa (see Table 1). It is present in the genus Afrachilus Fennah, 1965 as a distinct, large structure, much more developed than in other genera (see Fennah 1950, 1967a). The subvaginal plate is a distinctive feature present in a number of taxa, but is predominantly in these distributed in the New World (Table 1). The vast majority of the specimens described above were collected in 2002 by Michael Stiller in the western part of the Republic of South Africa, by sweeping the vegetation. The single specimen of Afrachilus montanifynbosensis sp. nov. was collected in 1933. The area of collecting has been classified as the Montane Fynbos and Renosterveld (AT1203) ecoregion (World Wildlife Fund 2021a, 2021b), which is part of the Cape Floristic Region (CFR). That region is widely acknowledged to be one of the biologically most diverse areas on Earth (Cowling et al. 1992; Allsop et al. 2014). The specimens were collected on five plants of the families Fabaceae Lindl., Polygalaceae Hoffmanns. & Link (Fabales), Asteraceae Bercht. & J.Presl (Asterales), Proteaceae Juss. (Proteales) and Rutaceae Juss. (Sapindales). According to the synopsis of the host plants of Achilidae (Bourgoin 2024), these are the first records for the order Proteales, family Proteaceae and family Polygalaceae of Fabales (Table 2). The distribution of Afrachilini trib. nov. coincides with that of Achiplectini (Brysz et al. 2024) and with the Cape Faunal Center (CFC), a discrete zoogeographic zone characterized by the phylogenetic antiquity of much of its invertebrate fauna (Struckenberg 1962). The level of knowledge concerning Achilidae in the African continent, the Afrotropical region, and the adjacent Madagascan region remains inadequate. The subfamily Achilinae Stål, 1866 is represented by Achilini Stål, 1866 incertae sedis genus Mabira Fennah, 1950 with 4 species, Achillini Emeljanov, 1991 genera Achilla Haglund, 1899 with 3 species and monospecific genus Maurisca Emeljanov, 2005. The subfamily Apatesoninae Metcalf, 1938 is represented by the tribe Ilvini Emeljanov, 1991 with monospecific genus Ilva Stål, 1866. The largest subfamily Myconinae Fennah, 1950 may be not European Journal of Taxonomy 1026: 275–303 (2025) 296 monophyletic (Brysz et al. in prep.), as well as some lower rank units delimited within this subfamily. The distribution of the two tribes, Achiplectini Brysz, Stroiński & Szwedo, 2024 and Afrachilini trib. nov., is confined to the Southern Africa. The tribe Mycarini Emeljanov, 1991 is represented by Katbergella Fennah, 1950 with 2 species from South Africa and Madagascan monospecific genera Acocarinus Emeljanov, 1991, Emeljanocarinus Bourgoin & Soulier-Perkins, 2006, Mycarinus Emeljanov, 1991 and Mycarus Emeljanov, 1991. The Plectoderini Fennah, 1950 may not be monophyletic, its African representatives covers single species of the genus Akotropis Matsumura, 1914, Aphypia Melichar, 1908 with 31 species, Ballomarius Jacobi, 1941 with 21 species, monospecific Bathycephala Fennah, 1950, monospecific Caffropyrrhyllis Fennah, 1950, Cnidus Stål, 1866 with 12 species, Epiusana Fennah, 1950 with 2 species, monospecific genera Epiusanella Synave, 1959, Kawanda Fennah, 1950 and Kawandella Synave, 1959, Lanuvia Stål, 1866 with 5 species, Mlanjella Fennah, 1950 with 7 species, monospecific genera Moraballia Fennah, 1950, Nyonga Synave, 1959 and Paraclusivius Fennah, 1950, Paraphypia Synave, 1960 with 4 species, Phypia Stål, 1862 with 2 species, and monospecific Prosagandecca Fennah, 1950. The placement of the monospecific genus Brachypyrrhyllis Fennah, 1967a, from South Africa, in the subfamily Plectoderini is, at this time, uncertain. It can, therefore, be provisionally placed in the subfamily Myconinae incertae sedis. Some other genera, Leptarciella Fennah, 1958 with 9 species and monospecific genera Parasabecoides Synave, 1965 and Sabecoides Fennah, 1958, are to be placed as Achilidae incertae sedis. In addition several monospecific genera of Plectoderini reported from the Madagascan region must be noted: Cernea Williams, 1977, Metalticeps Dmitriev, 2020 and Thectoceps Table 1. Distribution of the subvaginal plate in Achilidae Stål, 1866. Genus Distribution Derbidae: Breddiniolinae Breddiniola Muir, 1934 Cameroon Achilidae: Achilinae: Achilini Paraphradmon Fennah, 1950 Brazil Parelidiptera Fennah, 1950 Brazil Achilidae: Myconinae: Plectoderini Amblycratus Uhler, 1895 Mesoamerica and the Caribbean Bathycephala Fennah, 1950 Guyana Catonia Uhler, 1895 New World Hemiplectoderes Fennah, 1950 Trinidad and Tobago Kurandella Fennah, 1950 Australia (Queensland) Mlanjella Fennah, 1950 Africa Democratic Republic of the Congo, Malawi Paracatonia Fennah, 1950 Dominica, Grenada Paraclusivius Fennah, 1950 Ghana, Sierra Leone Remosachilus Fennah, 1950 Indonesia (Western New Guinea) (Irian Jaya; Papua New Guinea) Symplegadella Fennah, 1950 Mesoamerica Achilidae: Myconinae: Afrachilini trib. nov. Afrachilus Fennah, 1965 South Africa Achilidae: Myconinae incertae sedis Brachypyrrhyllis Fennah, 1967a South Africa STROIŃSKI A. et al., New achilid planthoppers tribe Afrachilini from Southern Africa (Hemiptera) 297 Williams, 1977 from Mauritius, Clusivius Distant, 1917 reported from Mauritius and the Seychelles, Cocottea Williams, 1977 with 3 species from Mauritius, and Williamsus Özdikmen & Demir, 2007 reported from Mauritius and Reunion. Therefore, South African records of the Achilidae embrace the tribes Achiplectini (1 genus with 1 species; Western Cape Province), Afrachilini trib. nov. (1 genus with 2 species; Western Cape Province, Eastern Cape Province), Ilvini (1 genus with 1 species; Western Cape Province), Plectoderini (4 genera with 5 species; Western Cape Province, Eastern Cape Province, KwaZulu-Natal) and Myconinae incertae sedis genus Brachypyrrhyllis Fennah, 1967a (1 species; Western Cape Province). The description of a new tribe of Achilidae, the third endemic to Southern Africa, showing specific, uncommon morphological features, enriches our knowledge of the classification and morphology of this insect family. Simultaneously, this study gives rise to additional inquiries concerning the diversification of the Achilidae fauna and ecology on the African continent, the morphological adaptations exhibited by these planthoppers, and the similarities and phylogenetic affinities observed among them. Acknowledgments We would like to thank Mrs Cecile Roux for providing the Achiplecton stilleri photos and for allowing us to use them in this publication. We would like to thank Dr Michael Stiller for lending us the specimens for research. We would also like to thank two anonymous reviewers for the comments and suggestions improving this work. Table 2. List of the host plants of Afrachilus montanifynbosensis sp. nov. Order Family Species Common names Fabales Bromhead Fabaceae Lindl. Aspalathus costulata Benth. Ribbed Rooibos Strandtee, Bokkeveld Tee, Burger Tee, Oranjerivier Tee Fabales Bromhead Polygalaceae Hoffmanns. & Link Nylandtia scoparia (Eckl. & Zeyh.) Goldblatt & J.C. Manning Tortoise Berry, Dune-berry, Skilpadbessie, Duinebessie Asterales Link Asteraceae Bercht. & J. Presl Dicerothamnus rhinocerotis (L.f.) Koek. rhinoceros bush, rhenoster bush, Renosterbos, rhenosterbos Proteales Juss. ex Bercht. & J. Presl Proteaceae Juss. Leucadendron pubescens R. Br. Silky Conebush, knokkerbos, pitjiebos, knopbos, Syhaartolbos, pitjieknopbos, syhaar-tolbos Sapindales Juss. ex Bercht. & J. Presl Rutaceae Juss. Diosma hirsuta L. wild buchu, rooiboegoe, rooi Buchu wildeboegoe, Hottentotsboegoe Caryophyllales Juss. ex Bercht. & J. Presl Aizoaceae Martinov Galenia africana L. 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