Biodiversity Data Journal :
Taxonomy & Inventories
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Corresponding author: Arianna Thomas-Cabianca (athomasbio@gmail.com)
Academic editor: Pierfilippo Cerretti
Received: 09 Aug 2021 | Accepted: 11 Nov 2022 | Published: 13 Jan 2023
© 2023 Arianna Thomas-Cabianca, Martin Villet, Anabel Martínez-Sánchez, Santos Rojo
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Thomas-Cabianca A, Villet MH, Martínez-Sánchez A, Rojo S (2023) South African nose flies (Diptera, Calliphoridae, Rhiniinae): taxonomy, diversity, distribution and biology. Biodiversity Data Journal 11: e72764. https://doi.org/10.3897/BDJ.11.e72764
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Rhiniinae (Diptera, Calliphoridae) is a taxon of nearly 400 known species, many of them termitophilous. Approximatelly 160 valid species in 16 genera are Afrotropical, with over 60 of them occurring in South Africa. The taxonomy of this group is outdated, as most studies of the South African taxa were conducted 40 to 70 years ago (mostly by Salvador Peris and Fritz Zumpt). Published information on their biology and ecology is also scarce.
An annotated checklist of 73 species of Rhiniinae for South Africa was developed, based on the holdings of sixteen entomological collections in Africa, Europe and North America. Over 3,700 specimens were examined, revealing nine new species records for South Africa (Cosmina undulata Malloch, 1926, Isomyia cuthbertsoni (Curran, 1938), Rhyncomya botswana Zumpt, 1974, R. tristis Séguy, 1933, Stomorhina apta Curran, 1931, S. malobana (Lehrer, 2007), Thoracites kirkspriggsi Kurahashi, 2001, Th. sarcophagoides Kurahashi, 2001 and Trichoberia lanata (Villeneuve, 1920)). We propose one new combination Eurhyncomyia metzi (Zumpt, 1981) comb. nov. (= Rhyncomya metzi Zumpt, 1981)). Additionally, evidence is presented to remove Rhyncomya viduella Villeneuve, 1927 stat. rev. from synonymy with Rhyncomya cassotis (Walker, 1849). Relevant novel biological and seasonality information, historical occurrence maps and high-definition photographs for each species are compiled.
biology, checklist, distribution maps, new combinations, new synonym, Oestroidea, Rhiniinae
Rhiniines, recently re-established as a subfamily of Calliphoridae, comprise around 400 recognised species within 30-39 genera (
Rhiniinae are represented in South Africa by 12 genera belonging to two tribes, Rhiniini and Cosminini. Rhiniini include the genera Rhinia Robineau-Desvoidy, 1830, Rhyncomya Robineau-Desvoidy, 1830, Isomyia Walker, 1859, Stomorhina Rondani, 1861, and Fainia Zumpt, 1958, while Cosminini include Cosmina Robineau-Desvoidy, 1830, Stegosoma Loew, 1863, Thoracites Brauer & Bergenstamm, 1891, Eurhyncomyia Malloch, 1926, Trichoberia Townsend, 1933, Zumba Peris, 1951 and Pseudorhyncomyia Peris, 1952 (
The latest reviews of the taxonomy and diversity of South African Rhiniinae were carried out between the 1950s and 1980s by Dr. S.V. Peris and Dr. F.K.E. Zumpt, who made excellent contributions for the entire African continent. They generated keys for most Afrotropical genera and species, discussed known species and described 13 new species from South Africa (
The biology (e.g. immature stages and habits) of most species of Rhiniinae remain poorly known (
In order to update and augment the knowledge of the Rhiniinae of South Africa, we assembled the first checklist of species for the country. We also provide information on the current taxonomic and nomenclatural status of the species, occurrence maps, a thorough compilation of the known and novel biological information and high definition habitus photographs for each species.
This study is based on the examination of about 3,000 specimens from South Africa and 700 specimens from other countries in the Afrotropical Region housed in sixteen entomological collections in ten countries. Acronyms used in the text for the museums and institutions are as follows: AMGS (Department of Entomology, Albany Museum, Grahamstown, South Africa); BMSA (Department of Entomology, National Museum, Bloemfontein, South Africa); CEUA (Entomological Collection, University of Alicante, Alicante, Spain); DMSA (Durban Natural Science Museum, Durban, South Africa); MNHN (Muséum national d’Histoire Naturelle, Paris, France); MZSUR (Zoology Museum, La Sapienza University of Rome, Rome, Italy); NHMUK (Natural History Museum, London, United Kingdom); NMSA (KwaZulu-Natal Museum, Pietermaritzburg, South Africa); RMCA (Musée Royal de l’Afrique Centrale, Tervuren, Belgium); SAMC (Iziko South African Museum, Cape Town, South Africa); SANC (South African National Collection of Insects, Agricultural Research Council, Pretoria, South Africa); SMNHTAU (Steinhardt Museum of Natural History, Tel Aviv University, Tel Aviv, Israel); UCME (Entomological Museum, Biology School, Complutense University of Madrid, Madrid, Spain); USNM-SM (United States National Museum, Smithsonian Institution, Washington D.C., United States of America); ZMHB (Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Berlin, Germany); ZMUC (Natural History Museum of Denmark, Copenhagen, Denmark).
Pinned specimens or specimens preserved in ethanol were examined using stereoscopic microscopes with ocular micrometres (Leica M80 or Leica MZ95). Identifications were based on the works of
For each genus and species studied, the following sections are provided: generic: synonyms and type species; species: specific synonyms (considered from
The material examined for South Africa is included in Suppl. material
Distribution data were obtained from
A distribution gazetteer (Suppl. material
= Fainia Zumpt, 1958: 83. Type species: Idia albitarsis Macquart, 1846, by original designation.
= Idia albitarsis Macquart, 1846: 321. Type locality: South Africa, Cafreria.
= Idia eupoda Loew, 1852: 660 [1862: 24]. Type locality: Mozambique, Inhambane.
= Idia extensa Walker, 1858: 211. Type locality: South Africa, Port Natal [= Durban].
= Fainia sambura Lehrer, 2008: 16. Type locality: Kenya, Taita Hills, 1000–2000 m, Wyundani Rd., 3º24'S 38º23'E.
Afrotropical: Central African Republic, Democratic Republic of Congo, ?Ghana, Kenya, Malawi, Mozambique, Namibia, Sierra Leone, South Africa (Fig.
Preferred environment: Afromontane forests, sand forests, dry scrub forest, open savannah/grassland, coastal bush and Ficus L. forest. In Namibia, a single specimen was reported from the Arid Savannah Biome (
Material examined: Suppl. materials
= Stomatorrhina elongata Bezzi, 1908: 38. Type locality: Bas-Congo [= Democratic Republic of Congo].
= Idiella major Malloch, 1926: 510. Type locality: Sierra Leone, Masimera to Yonnibanna.
Afrotropical: Cameroon, Central African Republic, Democratic Republic of Congo, Equatorial Guinea, Côte d'Ivoire, Kenya, Madagascar, Malawi, Mozambique, Namibia, ?Nigeria, Rwanda, Sierra Leone, South Africa (Fig.
Preferred environment: in Namibia, apparently restricted to the Arid and Mesic Savannah Biomes (
Material examined: Suppl. materials
= Rhinia Robineau-Desvoidy 1830: 422. Type species: Rhinia testacea Robineau-Desvoidy, 1830 (= Rhinia apicalis (Wiedemann, 1830)), by monotypy.
= Beccarimyia Rondani 1873: 287. Type species: Beccarimyia glossina Rondani, 1873 (= Rhinia apicalis (Wiedemann, 1830)), by monotypy.
= Idia apicalis Wiedemann, 1830: 354. Type locality: Canary Island, Tenerife.
= Rhinia testacea Robineau-Desvoidy, 1830: 423. Type locality: France, I'lle de France [= Mauritius].
= Idia flavipennis Macquart, 1844: 125. Type locality: Indonesia, Java.
= Idia simulatrix Loew, 1852: 660. Type locality: Olifant-River, South Africa.
= Idia punctata Bigot, 1858: 369. Type locality: Gabon.
= Idia bigoti Coquere, 1862: 96. Type locality: Senegal.
= Idia pleuralis Thomson, 1869: 542. Type locality: Australia, Keeling [= Cocos (Keeling)] Islands.
= Beccarimyia glossina Rondani, 1873: 287. Type locality: Abyssinia [= Ethiopia].
= Rhinia fulvipes Bigot, 1874: 239. Type locality: Ceylon [= Sri Lanka].
= Idiella trineuriformis Speiser, 1910: 153. Type locality: Tanzania, Kilimandjaro.
Afrotropical: Aldabra Island (Seychelles), Amirante Island (Seychelles), Angola, Benin*, Botswana, Burundi, Cameroon, Cosmoledo Island (Seychelles), Democratic Republic of Congo, Ethiopia, Gabon, Gambia, Ghana, Réunion Island (France)*, Madagascar, Malawi, Mauritus Island (Mauritius), Mozambique, Namibia, Nigeria, Oman, Rodriguez Island (Mauritius), Rwanda, Senegal, Sierra Leone, Socotra Island (Yemen), South Africa (Fig.
Preferred environment: associated with a variety of anthropogenic and natural environments including poultry farms and gardens, dune and sand forests, dry scrub forests, Ficus forest, grassy floodplain, woodland savannah, broad-leaved deciduous woodland and Succulent Karoo. In Namibia, it occurs in all biomes, being especially abundant in the Mesic Savannah Biome (
Material examined: Suppl. materials
= Idia coxendix Villeneuve, 1915: 204. Type locality: South Africa, Cap [= Cape Province].
= Rhinia pallidula Curran, 1927: 1. Type locality: Belgian Congo, Haut-Congo [Democratic Republic of Congo], Stanleyville [Kisangani].
Afrotropical: Angola*, Burundi*, Cameroon, ?Cosmoledo Island (Seychelles), Democratic Republic of Congo, Kenya, South Africa (Fig.
Preferred environment: associated with dune forest, coast scarp forest and coastal bush. In Democratic Republic of Congo, it was reported in lowland evergreen swamp forest. Recorded elevations: 50–1400 m a.s.l. Seasonality: uncommon species with highest numbers observed between January and February. No records in June, August and November; the rest of the year only one or two specimens per month. Behaviour and ecology: one specimen was collected inside the nest a Sphex tormentosus Fabricius (Hymenoptera). Collection methods: Malaise traps. In Democratic Republic of Congo, collected with Malaise traps and in Tanzania, with hand net. Life cycle and developmental stages: oviparous; immature stages and life history unknown. Illustrations and photographs: male habitus as in Fig.
Material examined: Suppl. materials
= Idia nigricornis Macquart, 1843: 281. Type locality: Senegal.
= Rhinia winthemi Villeneuve, 1915: 203. Type locality: Guinea.
Afrotropical: Botswana*, Cameroon, Democratic Republic of Congo, Equatorial Guinea*, Gambia, Ghana, Côte d'Ivoire, Lesotho*, Liberia, Madagascar, Malawi, Mozambique, Namibia, Nigeria, Senegal, Sierra Leone, South Africa (Fig.
Preferred environment: dry scrub forest and on the edge of a coastal forest in KwaZulu-Natal. In Namibia, apparently restricted to the Arid and Mesic Savannah Biomes (
Material examined: Suppl. materials
= Idia Wiedemann, 1820: 21. Type species: Musca lunata Fabricius, 1805 by subsequent designation of Townsend (1916: 7).
= Stomorhina Rondani, 1861: 9 (replacement name for Idia Meigen by Wiedemann, 1820 (nec Hübner 1813)).
= Stomorhyna (misspelling of Stomorhina in Stomorhyna maculata Rondani 1865: 228)
= Stomathorrhina Bezzi, 1906: 53 (replacement name for Idia Wiedemann).
= Stomatorrhina Bezzi, 1906: 144 (unjustified emendation of Stomathorrhina Bezzi).
= Stomatorhina Speiser, 1910: 153 (misspelling of Stomorhina).
= Lomwerhina Lehrer, 2007: 12. Type species: Lomwerhina malobana Lehrer, 2007, by original designation.
= Stomorhina apta Curran, 1931: 17. Type locality: Kenya Colony [Kenya], S. Masai Reserve [Maasai Mara National Reserve].
Preferred environment: associated with different environments, including: indigenous afromontane forest, indigenous mixed afromontane forest, indigenous forest, forest at the stream edge and Fynbos. In Burundi, it was associated with indigenous Afromontane forest and cloud forest. Recorded elevations: 30–1186 m a.s.l. Seasonality: uncommon species, most abundant in January with four specimens and present in September, November, December and April. The rest of the year it was absent. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: unknown. Illustrations and photographs: female habitus as in Fig.
Material examined: Suppl. materials
= Stomatorrhina armatipes Malloch, 1926: 500. Type locality: South Africa, Natal [KwaZulu-Natal].
= Stomorhina fasciculata Curran, 1927: 528. Type locality: South Africa, Natal [KwaZulu-Natal], Willow Grange.
Preferred environment: no data. Recorded elevations: 1440–1750 m a.s.l. Seasonality: most abundant in September; absent or scarce for the rest of the year. Behaviour and ecology: adults were observed visiting flowers and collected on Cassine sp., Protea caffra Meisn. and Cussonia Thunb. sp. Additionally, two males were found as prey of Oxybelus lingula Gerstaecker (Hymenoptera) in Grahamstown (Eastern Cape). During field work in September 2016 at the Kogelberg Nature Reserve, females were observed ovipositing on soil surrounding a termite nest. Gravid females and termites were collected and oviposition was achieved using a mixture of soil and live termites. Larvae were reared to adulthood and were observed preying on termites. Life cycle and developmental stages: oviparous. Immature stages (egg, larva and pupa) and adult females will be described (Thomas-Cabianca et al., unpublished). Collection methods: Malaise traps and sweeping. Illustrations and photographs: male habitus as in Fig.
Material examined: Suppl. material
= Stomorhina chapini Curran, 1931: 16. Type locality: Zaire [Democratic Republic of Congo], Lukulela.
= Rhinia patrizii Peris, 1952: 29. Type locality: Kenya, Ngong. Remarks: Zumpt (1962) indicated that S. patrizzi is identical to S. chapini after he examined the type series at NHMUK, but did not suggest it as Syn. nov. Subsequently,
Afrotropical: Cameroon, Democratic Republic of Congo, Kenya, Liberia, Namibia, South Africa (Fig.
Preferred environment: associated with Ficus forest, Acacia thornveld, indigenous forest and margin, sand forest and broad-leafed deciduous forest and long grass and woodland areas. In the Democratic Republic of Congo, the species was associated with lowland evergreen primary forest (disturbed). In Namibia, a single record was reported from the Mesic Savannah Biome (
Material examined: Suppl. materials
= Rhinia cribrata Bigot, 1874: 239. Type locality: Sierra Leone, Yiraia; Sierra Leone, Dilijuli.
= Rhinia vertebrata Bigot, 1891: 378. Type locality: Ivory Coast [Côte d'Ivoire], Assinie.
= Rhinia tricincta Bigot, 1891: 379. Type locality: Ivory Coast [Côte d'Ivoire], Assinie.
= Rhinia striata Becker, 1912: 626. Type locality: Iran, Pers-Beludshistan [Sistan and Baluchestan Province].
Afrotropical: Botswana, ?Burundi (plain Ruzizi), Cameroon, Democratic Republic of Congo, Gambia*, Ghana, Côte d'Ivoire, Kenya, Madagascar, Malawi, Mali, Namibia, Nigeria, Oman, Rwanda, Sierra Leone, South Africa (Fig.
Preferred environment: Indigenous forests (mixed woodland, margin of a dune forest), in grassveld near a stream and thornveld camp grounds. In Namibia, it is apparently restricted to the Arid and Mesic Savannah Biome (
Material examined: Suppl. materials
= Rhinia guttata Villeneuve, 1914: 384. Type locality: South Africa, Natal [KwaZulu-Natal], Willow Grange, Mooi River.
Preferred environments: Acacia savannah, Succulent Karoo Scrub, white dune and coastal sand dunes biomes and to river banks, plains and slopes. Recorded in all Namibian biomes (
Material examined: Suppl. materials
= Musca lunata Fabricius, 1805: 292. Type locality: Portugal, Madeira Island. Remarks: type series specimen in ZMUC.
= Idia rostrata Wiedemann, 1820: 22. Type locality: South Africa, Cape of Good Hope - Promontorio bonae spei [Western Cape]. Remarks: type series specimen in ZMUC.
= Idia fasciata Meigen, 1826: 9. Type locality: France, Marseilles.
= Idia syrphoidea Robineau-Desvoidy, 1830: 421. Type locality: Mauritius.
= Idia cinerea Robineau-Desvoidy, 1830: 422. Type locality: Isles de la mer d'Africa [Indian Ocean d'Africa].
= Stomorhyna maculata Rondani, 1865: 228. Type locality: Italy, Parma.
= Stomorhina melanorhina Bigot, 1888: 592. Type locality: South Africa, Cape of Good Hope [Western Cape].
= Stomorhina muscoidea Brauer, 1899: 516. Type locality: Madagascar.
= Stomorhina selgae Lehrer, 1979: 89. Type locality: Bermuda.
Afrotropical: Angola, Burundi, Democratic Republic of Congo, Eritrea, Ethiopia, Kenya, Lesotho, Madagascar, Malawi, Mauricio Island (Mauritius), Oman, Namibia, Réunion Island (France), Rodriguez Island (Mauritius), South Africa (Fig.
Preferred environment: montane grass and woodlands, montane meadows, grasslands, rocky hillside, indigenous montane forest and forest margins, slopes, ravines, streams and cascade areas. Different kind of biomes such as: Macchia vegetation and old lands, mesic mountain Fynbos, false Macchia slopes and coastal Macchia. Associated with human environments such as houses, a university campus, caravan parks and main tracks through forest. Recorded in low numbers in Namibia, where it is virtually restricted to the Brandberg Massif and occurs at high elevations on the edge of Nama-Karoo (
Type material examined: I. rostrata: 1 ? / Mus. / Westerm. // Type // I. rostrata / Weid. / Cape of Good Hope / Jan: 1817 // [ZMUC 00025098]. M. lunata: 1 ? // [ZMUC 00027332].
Material examined: Suppl. materials
= Lomwerhina malobana Lehrer, 2007: 12. Type locality: Malawi, Mulanje Mt. near Likabula. Remarks: HT in SMNHTAU (TAUI) .
Preferred environment: no data. Recorded elevations: 1050–1880 m a.s.l. Seasonality: the single specimen was collected in November. Behaviour and ecology: collected on avocado flowers (Gauteng). Life cycle and developmental stages: females collected in Tanzania contained a completely developed 3rd instar larva that occupied all of the abdomen (four females were dissected). This suggests an unilarviparous biology, an exclusive and new trait for the Afrotropical Stomorhina (Thomas-Cabianca et al., unpublished). Collection methods: in Tanzania, it was collected with Malaise and pitfall traps and a Malaise trap in Malawi. Illustrations and photographs: male habitus as in Fig.
Type material examined: L. malobana: 1 ? Malawi, 1500 m a.s.l., Mulanje Mt. nr. Likabula, 26-27.x.83, A. Freidberg // Holotype // Lomwerhina malobana Det. Dr. A. Z. Lehrer, 2006 // SMNHTAU (TAUI) 318991.
Material examined: Suppl. materials
= Rhinia rugosa Bigot, 1888: 591. Type locality: Sierra Leone.
= Stomorhina mitis Curran, 1931: 18. Type locality: South Africa, Natal [KwaZulu-Natal].
= Rhinia hyphena Séguy, 1958: 188. Type locality: Guinea.
Afrotropical: Democratic Republic of Congo, Ghana, Guinea (today could be Guinea, Guinea-Bissau or Equatorial Guinea), Kenya, Lesotho*, Malawi, Mozambique, Namibia, Nigeria, Sierra Leone, Sudan (reported as Anglo-Egyptian Sudan), South Africa (Fig.
Preferred environment: coast scarp forest, grassland, scrub and wooded grassland area, near streams, on the margin of dune forest, sewage-seepage area, bushveld and in picnic area. In Zimbabwe, the species was collected indoors and in Namibia, it was collected at a damaged termite mound and is apparently restricted to the Mesic Savannah Biome (
Material examined: Suppl. materials
= Cosmina Robineau-Desvoidy, 1830: 423. Type species: Cosmina fuscipennis Robineau-Desvoidy, 1830, by subsequent designation of Townsend (1916: 6).
= Seseromya Rondani, 1863: 32. Type species: Musca punctulata Wiedemann, 1819 (= Cosmina fuscipennis Robineau-Desvoidy, 1830), by original designation.
= Synamphoneura Bigot, 1887: xiv. Type species: Synamphoneura cuprina Bigot, 1887 (= Idia limbipennis Macquart, 1848), by original designation.
= Idiopsis Brauer and Bergenstamm, 1889:153. Type species: Idiopsis prasina Brauer and Bergenstamm, 1889, by monotypy.
= Eusynamphoneura Townsend, 1917: 189. Type species: Idia seriepunctata Loew, 1852 (= Dictya aenea Fabricius, 1805), by original designation.
= Synamphoneuropsis Townsend, 1917: 199. Type species: Synamphoneuropsis viridis Townsend, 1917, by original designation.
= Dictya aenea Fabricius, 1805: 328. Type locality: "Guinea" Krieger, [Ghana, Teshi. Remarks: type-serie in ZMUC.
= Idia seriepunctata Loew, 1852: 660 [1864:32]. Type locality: Mozambique, Inhambane.
= Cosmina despressa Karsch, 1888: 377. Type locality: Tanganyika [Tanzania], Usambara.
= Cosmina punctulata var. microps Malloch, 1926: 518. Type locality: Gold Coast [Ghana], N. territories [North East Region], Yapi.
Afrotropical: Burkina Faso, Cameroon, Democratic Republic of Congo, Côte d'Ivoire*, Kenya*, Liberia, Malawi, Mali, Mozambique, Namibia*, Nigeria, South Africa (Fig.
Preferred environment: forest, open woodlands areas, woodland savannah, grasslands, coastal grassland, open mixed grassland, mixed dune woodland and bushveld near to a river (close to Nylstroom in Limpopo Province). In Namibia, the species was associated with the Kwanso River floodplain, in Kenya to Ngorowa Gorge/Stream and Lukenya cliffs/bushveld and in Malawi, to the Acacia woodland. Recorded elevations: 24–1219 m a.s.l. Seasonality: highest abundance in November and March and lowest between April and October (absent in July). Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: sweeping net and Malaise trap. Hand net in Mozambique and Malaise trap in Namibia. Illustrations and photographs: male habitus as in Fig.
Material examined: Suppl. materials
= Cosmina fuscipennis Robineau-Desvoide, 1830: 423. Type locality: Cap de le bonne-Espérance [Western Cape, South Africa]. Remarks: type-series in MNHN, destroyed, not in remnants of Robineau-Desvoidy's collection.
= Musca punctulata Wiedemann, 1819: 21. Type locality: Cape of Good Hope [Western Cape, South Africa]. Remarks: type-series in ZMUC.
= Cosmina cuprina Bigot, 1860: 539. Type locality: Madagascar.
= Cosmina aethiopissa Séguy, 1958: 176. Type locality: Kenya.
Afrotropical: Botswana, ?Kenya, Madagascar, Mozambique, Namibia, Oman, Seychelles, South Africa (Fig.
Preferred environment: Indigenous dune forest; bushveld and dune vegetation forest; open woodland areas, sandy woodland savannah areas; Karoo and valley Acacia woodland, succulent Karoo garden on hot N slope, Acacia and Ziziphus Mill. veld; Worcester Macchia, coastal Macchia and sandy area; Fynbos, mesic mountain Fynbos on sandstone; rocky slopes at road cutting, stream edge or bushes; rocky hillside areas with vegetation or sandy areas below; rocky outcrops; open mixed grassland, riverine bush and grass, plains; mountain slope overlooking sea on vegetated cliff; and Caravan Park and surrounding area, dry stream bed near staff houses, orchards and grasslands. This species also was reported for Arid-Savannah and Succulent Karoo Biomes in Namibia (
Type material examined: M. punctulata: 1 ? // HT Cape Good Hope (South Africa) / June 1817 // Mus. Westerm // [ZMUC 00025139]; 1 ? // Type // Mus. Westerm // [ZMUC 00025140].
Material examined: Suppl. materials
= Cosmina gracilis Curran, 1927: 2. Type locality: South Africa, Barberton. Remarks: type-serie in SANC at ARC.
Preferred environment: dry scrub forest, mixed bushveld-grass, sand and broad-leafed deciduous forest and Rhus and Acacia savannah. In Namibia, associated with the Mesic Savannah, Arid Savannah, Karoo and Desert Biomes (
Type material examined: C. gracilis: 1 ? HT 1 ? AT // Barberton / May 17 1914 / (H. K. Munro).
Material examined: Suppl. materials
= Cosmina margaritae Peris, 1952a: 229. Type locality: Nyasaland [Malawi], Cholo. Remarks: type-series in NHMUK.
Afrotropical: ?Botswana, Democratic Republic of Congo, ?Kenya, Malawi, Mozambique*, Namibia, ?Senegal, South Africa (Fig.
Preferred environment: Arid and Mesic Savannah Biome in Namibia (
Type material examined: C. margaritae: 1 ? Holotype // Nyasaland / Cholo. / R. C. Wood // Pres. by / Com. Inst. Ent. / BM. 1950-323 // Cosmina / margaritae / tipo n. sp. / Dr. S-V. Peris det. // [NHNUK 010579920]. 1 ? // Paratype // Nyasaland / Cholo. / R. C. Wood // Pres. by / Com. Inst. Ent. / BM. 1950-323 // Cosmina / margaritae / paratipo n. sp. / Dr. S-V. Peris det. // [NHNUK 010579921].
Material examined: Suppl. materials
= Cosmina thabaniella Lehrer, 2010: 26. Type locality: South Africa, Natal [KwaZulu-Natal], Uvongo, South Coast.
Afrotropical: South Africa.
No specimens examined for South Africa, based on
= Cosmina undulata: Malloch, 1926: 518. Type locality: S. Nigeria [Nigeria], Ibadan. Remarks: HT in NHMUK.
Afrotropical: ?Benin, Botswana, Burkina Faso, Cameroon, Côte d'Ivoire, Democratic Republic of Congo, Eritrea*, Ethiopia*, Malawi, Namibia, ?Niger, Nigeria, South Africa* (Fig.
Preferred environment: Mesic Savannah Biome in Nambia (
Type material examined: C. undulata: 1 ? // Holo-type // Nigeria: / Ibadan. / 2.viii.1923. / A.W.J. Pomeroy // Pres. by / Imp. Bur. Ent. / Brit. Mus. / 192x-94 // Cosmina / undulata / Type / Det / J.R. Malloch // [NHMUK 010579923].
Material examined: Suppl. materials
= Eurhyncomyia Malloch, 1926: 513. Type species: Xysta obtusa Bigot, 1891 = Eurhyncomyia diversicolor (Bigot, 1888) by original designation.
= Rhyncomya diversicolor Bigot, 1888: 595. Type locality: Somalis [Somalia]. Remarks: LT in NHMUK, designated by Dear and Pont in the collection.
= Rhyncomyia bigoti Villeneuve, 1913: 155. Type locality: South Africa, Natal [KwaZulu-Natal], Port Natal [Durban].
= Eurhyncomyia thoracica Curran, 1931: 21. Type locality: South Africa, Natal [KwaZulu-Natal], Port Shepstone.
Preferred environment: broad-leafed, broad-leafed deciduous, coastal and sand forest. In Namibia, it is associated with a broken veld at the base of small hill. Recorded elevations: 77 m a.s.l. Seasonality: present almost all year-round, peaking in December, lower abundance in January, May, June and August and absent in September. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: Malaise trap. In Mozambique, Malaise trap. Illustrations and photographs: male habitus as in Fig.
Type material examined: R. diversicolor: 1 ? // Lecto-type // Somalis // Brauer / Wien. CVIII / (No. 87) // Lectotype ? / Rhyncomyia / diversicolor / Bigot / Designated by / Dear and Pont. // BMNH (E) / #231121 // [NHMUK 010579922].
Material examined: Suppl. materials
= Rhyncomyia metzi Zumpt, 1981: 487 (see taxonomic notes). Type locality: South Africa, Natal Zululand [KwaZulu-Natal], Umfalozi Game Park. Remarks: HT and PTs in NMSA.
Preferred environment: no data. Recorded elevations: no data. Seasonality: recorded only in September. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: unknown. Illustrations and photographs: male habitus as in Fig.
Taxonomic notes: the HT and PTs of R. metzi were examined at the NMSA and their supra-squamal ridge is setulose in the posterior half, a diagnostic character that separates Eurhyncomyia (setulose) from Rhyncomya (bare). Another characteristic of Eurhyncomyia is that the aristal hairs are long and pubescent, the longest hairs slightly exceeding half the width of the post-pedicel, just as R. metzi, whereas in Rhyncomya, it is either bare or the hairs rarely exceed the width of the basal arista (
Type material examined: R. metzi: 1 ? // HOLOTYPUS // SOUTH AFRICA, Natal / Zululand, Umfalozi / Game Park, 2831Bd / 21-VII-1973, ME Irwin // slide no. 30 // Rhyncomya ? / metzi Zumpt / det. Zumpt 80 // [NMSA-DIP 074954] // (N.M. Type No. 2437). 6 ?? // PARATYPE // SOUTH AFRICA, Natal / Zululand, Umfalozi / Game Park, 2831Bd / 21-VII-1973, ME Irwin // Rhyncomya ? / metzi Zumpt / det. Zumpt 80 // [NMSA-DIP [NMSA DIP 019991, 061680, 074915, 074918, 074952, 0749523] // (N.M. Type No. 2437).
Material examined: Suppl. material
= Isomyia Walker, 1859: 134. Type species: Musca delectans Walker, 1859 by original designation and monotypy.
= Strongyloneura Bigot, 1886: xiv. Type species: Strongyloneura prasina Bigot, 1887 (Oriental genus) by monotypy.
= Thelychaeta Brauer and Bergenstamm, 1891: 390. Type species: Thelychaeta chalybea Brauer and Bergenstamm, 1891 = Isomyia viridaurea (Wiedemann, 1819) by monotypy.
= Apollenia Bezzi, 1911: 79. Type species: Pollenia nudiuscula Bigot, 1911 = Phumosia nudiuscula (Bigot, 1888) [missidentification = Curtonevra tristis Bigot, 1888] by original designation.
= Anna Malloch, 1926: 520. Type species: Anna calliphoroides Malloch, 1926 by original designation
= Gerschia Lehrer, 1970: 30. Type species: Isomyia eos Zumpt, 1958 by original designation.
= Strongyloneura cuthbertsoni Curran, 1938: 2. Type locality: S. Rhodesia [Zimbabwe], Vumba Mts.
Preferred environment: woodland savannah, woodland, dune, sand and broad-leafed deciduous forest. Recorded elevations: 6–1095 m a.s.l. Seasonality: highest abundance in October and December, lower numbers or absent the rest of the year. Behaviour and ecology: specimens were swept from Asparagus L. sp. in Sileza Natural Reserve (KwaZulu-Natal). In Zimbabwe,
Material exmained: Suppl. material
= Strongyloneura darwini Curran, 1938: 3. Type locality: S. Rhodesia [Zimbabwe].
Preferred environment: Ficus forest. Apparently restricted to the Arid Savannah Biome in Namibia (
Material examined: Suppl. materials
= Somomyia deserti Karsch, 1888: 378. Type locality: [Tanzania].
= Thelychaeta versipellis Villeneuve, 1917: 344. Type locality: Congo Belge [Democratic Republic of Congo], Kilimbi and Sankisia; Nyasaland [Malawi], Mt. Mlanje; Mozambique; South Africa. Remarks: PT in SAMC.
Afrotropical: ?Botswana, Burundi, Democratic Republic of Congo, Malawi, Mozambique, Namibia, South Africa (Fig.
Preferred environment: dune vegetation and beach environment. Apparently restricted to the Arid and Mesic Savannah Biome in Namibia (
Type material examined: T. versipellis: 1? / Marley / Kloof Natal/ 2-1915 // 359 // Thelychaeta / versipellis Ville. // Paratype // [SAMC DIP A011140]. Remarks: with head glued on a card.
Material examined: Suppl. materials
= Thelychaeta distinguenda Villeneuve, 1917: 352. Type locality: Congo Belge [Democratic Republic of Congo], Elisabethville [Lubumbashi] and Kundelungu; l'Afrique Orientale anglaise [Kenya], Nairobi.
Preferred environment: sand, forest and broad-leafed deciduous forest and woodland. Vegetated stream-bed in Togo. Recorded elevations: no data. Seasonality: highest abundance between November and January, rest of the year generally absent or only one specimen per month. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: Malaise traps. In Togo, Malaise trap. Illustrations and photographs: male habitus as in Fig.
Material examined: Suppl. materials
= Idiopsis buccata Bezzi, 1911: 73. Type locality: South Africa, Pretoria.
= Thelychaeta dubiosa Villeneuve, 1917: 350. Type locality: Belgian Congo [Democratic Republic of Congo], Urwald Moera; British East Africa [Kenya], Wa-Taita Boura District.
= Thelychaeta claripennis Villeneuve, 1917: 350. Type locality: Nyasaland [Malawi]; Southern Rhodesia [Zimbabwe], Salisbury [Harare].
= Strongyloneura sheppardi Curran, 1938: 3. Type locality: Southern Rhodesia [Zimbabwe], Balla-Balla.
= Apollenia nasica Séguy, 1949: 131. Type locality: Kenya, Nairobi.
= Apollenia promula Séguy, 1949: 133. Type locality: Guinea, Nimba, Keoulenta.
Afrotropical: Cameroon, Democratic Republic of Congo, Equatorial Guinea, Ghana, Guinea, Côte d’Ivoire, Kenya, Liberia, Malawi, Namibia, Nigeria, Rwanda, Senegal, Sierra Leone, South Africa (Fig.
Preferred environment: broad-leafed deciduous woodland. In Democratic Republic of Congo associated with bush paths and village environments; and in Togo, with remnant forest patches and vegetated stream-beds.
Material examined: Suppl. materials
= Isomyia eos Zumpt, 1958: 45. Type locality Southern Rhodesia [Zimbabwe], Farfell Farm, Melsetter District. Remarks: PT in NMSA.
Preferred environment: broad-leafed deciduous woodland. Recorded elevations: no data. Seasonality: low numbers during March, November and December. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: Malaise trap. Illustrations and photographs: male habitus as in Fig.
Type material examined: I. eos: 1 ? // Grampirks / Inyamadzi River / Melsetter District / Dept. Agric. S. Rhodesia / 3/6/1939 // slide no 11 // Paratype // Isomyia / eos n.sp. / Zumpt 1956 // [NMSA DIP 019909].
Material examined: Suppl. materials
= Isomyia innia Lehrer, 2009: 21. Type locality: South Africa, Inn on Robber's Pass, 1,400 m.
Afrotropical: South Africa.
No specimens examined for South Africa, based on
= Thelychaeta longicauda Villeneuve, 1917: 350. Type locality: South Africa, East London [Eastern Cape]; N. W. Tanganika [Decomocratic Republic of Congo]. Remarks: PT in SAMC.
Afrotropical: Burundi, Democratic Republic of Congo, Kenya, Malawi, South Africa (Fig.
Preferred environment: savannah/grassland, mist-belt mixed forest, grass and forest edges and river stream. In Malawi, in grassland and forest edges. Recorded elevations: 1100–1350 m a.s.l. In Burundi, between 1800 and 2000 m a.s.l. (
Type material examined: T. longicauda: 1 ? // Paratype // E. London / Lightfoot / July 1914 // S.A.M // Thelychaeta / longicauda / Det. Villeneuve // [SAMC DIP A011144].
Material examined: Suppl. materials
= Thelychaeta natalensis Villeneuve, 1917: 347. Type locality: South Africa, Natal [KwaZulu-Natal].
Preferred environment: forest, closed woodlands, montane forest, mist-belt forest in the margin, dune forest, forest margins, grass, grasslands, montane grasslands, Little Berg Summits Themeda grassland, riverine bush Montane slopes, riverine vegetation (Tugela River), straddling Mahai River, riverside in open road and environments with little anthropogenic intervention (Simes cottage, caravan park environments and garden areas). In Namibia, the Mesic Savannah Biome (
Material examined: Suppl. material
= Thelychaeta oculosa Villeneuve, 1917: 342.Type locality: South Africa and North-West Rhodesia [Zambia], Chilanga R. Remarks: STs in NHMUK and PT in SAMC.
Preferred environment: in Zimbabwe,
Type material examined: T. oculosa: 1 ? SYN-TYPE // 24/9/13 / Chilanga R. / W. Lusaka; N.W.R. / (R.C.W.) / in house // R.C. Wood // Pres. by / Com. Inst. Ent. / B.M. 1956-102 // [NHMUK 010580057]. 1 ? SYN-TYPE // 25/09/13 M / Chilanga / N.W. Rhodesia / (R.C.W) / 4030 / in house // Pres. by / Com. Inst. Ent. / B.M. 1956-102 // Thelychaeta / oculosa Villen / Villeneuve det. // [NHMUK 010580055]. T. oculosa: 1 ? Para-Type // Bulawayo / 10.ix.1910 / E.C. Chubb / caught in house // S. Afri. Mus. // 119 // Thelychaeta / oculosa / Type Villeneuve // [SAMC DIP A011149].
Material examined: Suppl. materials
= Thelychaeta pubera Villeneuve, 1917: 340. Type locality: British East Africa [Kenya], Eldoret; South Africa, Kloof [Durban]; North-West Tanganyka [Tanzania]; Uganda, Entebbe.
= Thelychaeta jactatrix Villeneuve, 1917: 343. Type locality: Belgian Congo [Democratic Republic of Congo], Stanleyville [Kisangani].
= Thelychaeta villeneuvei Curran, 1927: 3. Type locality: Democratic Republic of Congo [Belgian Congo], Stanleyville [Kisangani].
= Strongyloneura cupreithorax Curran, 1931: 1. Type locality: South Africa, Transvaal [Gauteng], Pretoria, Barberton.
Preferred environment: no data. Recorded elevations: 1450–1981 m a.s.l. Seasonality: present almost all year, peaking in September, absent in April, May, June and August with low numbers in other months. Behaviour and ecology:
Type material examined: T. pubera: 1 ? LT // Kloff / 16.xii.14 / H.W. Bell-Marley // S.A.M. // Type // Lectotype // Thelychaeta / pubera n. sp / Det. Villeneuve // Isomyia / pubera Ville. / Vid. Zumpt 56 // slide no 39 // Slide SAM: 33B10 // [SAMC DIP A011150].
Material examined: Suppl. material
= Isomyia transvaalensis Zumpt and Stimie, 1965: 8. Type locality: South Africa, Transvaal [North-West], Rustenburg. Remarks: HT in NMSA.
Preferred environment: no data. Recorded elevations: no data. Seasonality: only one specimen collected in December. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: unknown. Illustrations and photographs: male habitus as in Fig.
Type material examined: I. transvaalensis: 1 ? Rustenburg / Transvaal / 3.xii.61. // HOLOTYPUS // Isomyia / transvaalensis / Z[um]pt & Stimie // [NMSA-DIP 019760].
Material examined: Suppl. material
= Curtonevra tristis Bigot, 1888: 613. Type locality: South Africa, Port-Natal [Durban]. Remarks: HT in NHMUK and LT in SAMC.
= Apollenia psophis Séguy, 1933: 74. Type locality: Mozambique, Macequece.
Afrotropical: Angola*, Botswana, Cameroon*, Democratic Republic of Congo, Ethiopia, ?Ghana, Kenya, Liberia, Lesotho, Mozambique, Namibia, Rwanda, South Africa (Fig.
Preferred environment: montane environments (grassland and bush, meadow, Podocarpus L'Hér ex Pers sp. forest margins and slopes), savannahs (Acacia and dry open), forests (Mbhome, pine and swamp), areas associated with forests (creeks, gorges, margins, stream, slopes, trail, roads and burnt-out forests), bush (bushveld, coastal, dry bushveld, in front beach, riverine; mixed Acacia woodland and mixed woodland), grasslands, amongst others. Additionally, caravan park, grotto near farm, thornveld camp ground. In Mozambique, it was associated with gallery forest. In Namibia, associated with the Mesic Savannah and the Succulent Karoo Biomes (
Type material examined: C. tristis: 1 ? // Holo / type // Curtoneura / tristis Bigot / Port-Natal / ex. Bigot Coll: / B.M.1960-539. // Genus / Apollenia / Bezzi / det. Villen. // BMNH(E) # 231136 // [NHMUK 010832105]. 1 ? Durban / Natal // Pres. by / Com. Imp. Bur. Ent. // 1917-94 // Thelychaeta / Dr Villeneuve det. / tristis / sec type. Bigot // 861 // [NHMUK 010832122]. 1 ? // Marley / Jan. 1915 / P. Port Shepstone // Apollenia / tristis Bigot. / Lec Type // S.A. Museum. // [SAMC DIP A011159]. 1 ? / K/Kloof / Marley / 1-15 // S.A. Museum // Apollenia / tristis /sec.type Bigot / det. Villeneuve // [SAMC DIP A015196].
Material examined: Suppl. materials
= Pseudorhyncomyia Peris, 1952: 58. Type species: Rhyncomyia braunsi Villeneuve, 1920, by original designation.
= Rhyncomyia braunsi Villeneuve, 1920: 158. Type locality: South Africa, Cape Province [Western Cape], Willowmore. Remarks: LT in NMSA, designated by Zumpt (1958: 124).
Preferred environment: no data. Recorded elevations: no data. Seasonality: present in low numbers, more abundant in May and absent most of the year. In Namibia, low numbers in February (
Type material examined: P. braunsi: 1 ? // Cape Province / Willowmore / 5.V.1920 / Brauns // Lectotype // [NMSA-DIP 19835].
Material examined: Suppl. materials
= Beria Robineau-Desvoidy, 1830: 418. Type species: Beria inflata Robineau-Desvoidy, 1830, by monotypy.
= Rhyncomya Robineau-Desvoidy, 1830: 424. Type species: Musca felina Fabricius, 1794, by monotypy.
= Trichometallea Townsend, 1917: 194. Type species: Trichometallea pollinosa Townsend, 1917, by original designation.
= Rhynchomyiopsis Townsend, 1917: 195. Type species: Rhynchomyiopsis indica Townsend, 1917 (= Rhyncomya townsendi James, 1977), by original designation.
= Doljia Suster, 1953: 769. Type species: Doljia viridicauda Šuster, 1953 (= Rhyncomya speciosa (Loew, 1844)), by monotypy.
= Sokotra Lehrer, 1970: 32. Type species: Rhyncomyia varifrons Beckerm 1910, by original designation.
= Idia bicolor Macquart, 1844: 124. Type locality: South Africa, Cape of Good Hope [Western Cape]. Remarks: HT in NHMUK.
= Rhyncomya crinicauda Villeneuve, 1927: 22. Type locality: South Africa, Cape of Good Hope [Western Cape]. Remarks: HT in SAMC.
Preferred environment: sand dunes. In Namibia, the Arid Savannah Biome (
Type material examined: I. bicolor: 1 ? // Holo-type // Idia / bicolor / n. sp. // S. Bicolor / Idia . iv. Macq / C. B. Sp. / J. Bigot // [NHMUK 010832205]. B. crinicaudata: 1 ? // Stellenbush / 9.86 // Holotypus // Beria / crinicaudata / n. sp. // crinicaudata / type Villen. // slide no. 6 // Slide / SAM: / 33B12 // [SAM-DIP A011181].
Material examined: Suppl. material
= Rhyncomya botswanae Zumpt, 1974: 129. Type locality: Botswana, Moremi Reserve. Remarks: HT and PT in NHMUK.
Preferred environment: Gordonia Plains Shrubland, Senegalia mellifera (M. Vahl) Seigler and Ebinger on red sand. Recorded elevations: 1035 m a.s.l. Seasonality: a single specimen collected between September and November. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: Malaise trap. Illustrations and photography: male habitus as in Fig.
Type material examined: R. botswanae: 1 ? // Holo-type // Botswanhoa (B11) / Moremi Reserve, / 19º23'S, 23º33'E / 18-20.iv.1972 // Southern / African Exp. / B.M.1972-1 // slide no. 38 // Rhyncomya ? / botswanae n.sp / Zumpt 1973 // Slide prep.: / 010194475 // [NHMUK 010832201]. R. botswanae: 1 ? // Para-type // Botswana (B7) / Kuke Pan, / 20º56'S, 22º25'E / 14-15.iv.1972 // Southern / African Exp. / B.M.1972-1 // slide no. 82 // Rhyncomya ? / botswanae n. sp / Zumpt 1973 // Slide prep.: / 010194473 // [NHMUK 010832200]
Material examined: Suppl. materials
= Rhyncomya buccalis Villeneuve, 1927: 24. Type locality: Congo Belge [Democratic Republic of Congo], Mufungwa; South Africa.
Afrotropical: Democratic Republic of Congo, Kenya, Malawi, Mozambique, South Africa, Tanzania*, Uganda and Zambia*.
No specimens examined for South Africa, based on
Material examined: Suppl. material
= Tachina cassotis Walker, 1849: 761. Type locality: Sierra Leone. Remarks: HT in NHMUK.
Afrotropical: Angola*, Benin*, Cameroon, Democratic Republic of Congo, Ethiopia, Gambia, Kenya, Mali, Malawi, Mozambique, Namibia, Nigeria, Sierra Leone, South Africa (Fig.
Preferred environment: Acacia savannah and Acacia veld, sand forest and broad-leafed deciduous forest, mixed woodland, broad-leafed deciduous woodland and woodland near stream and camp. In Benin, to lowland gallery forest and stream-bed, remnant forest and agricultural plots; Cameroon to degraded savannah forest; Democratic Republic of Congo to bush paths and village environs; Kenya to rocks and river margins; in Namibia, to degraded sand forest, cultivated plots and Kwando River floodplain; eSwatini to Panata Ranch and Bushveld; Togo to vegetated stream-bed; and Zambia to Central Zambezian Miombo Woodlands. In Namibia, also associated with the Arid and Mesic Savannah Biome (
Type material examined: T. cassotis: 1? // Holo-type // Sierra Leone. / W. Africa. / Reud D.T. Morgan. // HOLOTYPE ? / Tachina / cassotis Walker / 1849, List Dipt. / Brit. Mus., 4:761 // Tachinia / Type / cassotis / Walk // ? Nomia sp / det. G.R. Else. 1974 (Hymenoptera) // [NHMUK 010832158].
Material examined: Suppl. materials
= Rhyncomya pollinosa Curran, 1931: 20. Type locality: South Africa, Transvaal [Gauteng], Pretoria. Remarks: nec. R. pollinosa Townsend 1917. Remarks: HT in SANC.
= Rhyncomya currani Zumpt, 1958: 144. Type locality: South Africa, Transvaal [Gauteng], Pretoria. Remarks: new replacement name for R. pollinosa Curran, 1931 by Zumpt in 1958; HT and AT in SANC at ARC.
Preferred environment: no data. Recorded elevations: no data. Seasonality: uncommon, present only in September. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: unknown. Illustrations and photographs: male habitus as in Fig.
Type material examined: R. pollinosa: 1 ? HT // Pretoria / 21.9.15 / H.K. Munro // Rhyncomya / Type / pollinosa ? / Curran. / No. // 1901 was ST258 // TYPH01901 // Rhyncomya currani Zumpt, 1958 / nom. nov.
Material examined: Suppl. material
= Rhyncomya dasyops Bezzi, 1908: 382. Type locality: Zaire [Democratic Republic of Congo], Haut-Congo [Oriental Province].
= Beria proxima Séguy, 1926: 12. Type locality: French Congo [Republic of Congo]. Remarks:
= ?Rhyncomya rugosa Séguy, 1926: 12. Type locality: Mozambique, Vallée du Pungoué. Remarks:
= Rhyncomya nigropilosa Villeneuve, 1927: 19. Type locality: Rhodesia mérdionale [Zimbabwe], Salisbury [Harare]; Chilanga [Zambia]; Nigeria septetrionale [Nigeria]; Uganda Prot. [Uganda], Mt. Elgon, 3,700–3,900 ft.; Congo Belge [Democratic Republic of Congo], Elisabethville [Lubumbashi]; Abyssinie [Ethiopia]; région du Cap [South Africa, Western Cape].
= Metallea pseudoinflata Peris, 1951: 240. Type locality: Anglo-Egipsian Sudan [Sudan], W. Darfur, S. Jebel Murrs, Kallikitting, 4,4450 ft.
Afrotropical: Angola, Benin, Botswana, Burundi*, Cameroon, Democratic Republic of Congo, Ethiopia, Gambia, Ghana, Kenya, Malawi, Mozambique, Namibia, Nigeria, South Africa (Fig.
Preferred environment: in Namibia, restricted to the Arid and Mesic Savannah Biomes (
Material examined: Suppl. materials
= Rhyncomya depressifrons Villeneuve, 1927: 24. Type locality: South Africa, Natal [KwaZulu-Natal], Estcourt. Remarks: STs in NHMUK.
Preferred environment: no data. Recorded elevations: no data. Seasonality: only present from December to March. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: unknown. Illustrations and photography: male habitus as in Fig.
Type material examined: R. depressifrons: 1 ? // SYN-TYPE // Natal / Estcourt / 5.ii.1913 / R.C. Wroughton // Slide no. 61 // Rhyncomya ? / depressifrons / Villeneuve // Slide prep.: / 010194477 // [NHMUK 010832207]. 1 ? // SYN-TYPE // Natal / Estcourt / 5.ii.1913 / R.C. Wroughton //Pres. by / Com. Inst. Ent. / BM. 1955-504 // [NHMUK 010832208]. R. depressifrons: 1 ? // SYN-TYPE // Natal / Estcourt / 26.i.1913 / R.C. Wroughton // Pres. by / Com. Inst. Ent. / B.M.1955-504 // depressifrons / typ. Villen. // [NHMUK 010832209].
Material examined: Suppl. material
= Rhyncomya disclusa Villeneuve, 1927: 24. Type locality: South Africa, Natal [KwaZulu-Natal], Willow Grange. Remarks: STs in NHMUK.
Preferred environment: no data. Recorded elevations: no data. Seasonality: only present in March, September, October and December. Behaviour and ecology: reported as collected on flowers of Acacia Mill. and visiting flowers of onions. Life cycle and developmental stages: unknown. Collection methods: unknown. Illustrations and photographs: male habitus as in Fig.
Type material examined: R. disclusa: 1 ? // SYN-TYPE // Natal / Willow Grange / Mooi River / R.C. Wroughton / 18.iii.1913 // Pres. by / Imp. Inst. Ent. / Brit. Mus. / 1933-14 // [NHMUK 010832216]. R. disclusa: 1 ? // SYN-TYPE // Natal / Willow Grange / Mooi River / R.C. Wroughton // slide no. 43 // [NHMUK 010832217]. R. disclusa: 1 ? // SYN-TYPE // Natal / Willow Grange / Mooi River / R.C. Wroughton // ?llBrook / 22.x.13 // Pres. by / Com. Inst. Ent. / B.M. 1955-504. // Rhyncomyia / Dr. Villeneuve det. / disclusa / Typ. Villen. // [NHMUK 010832218].
Material examined: Suppl. material
= Rhyncomya discrepans Villeneuve, 1927: 22. Type locality: South Africa, Gt. Wint-hock, Tulbach, 3,600 ft.; and Cap [Eastern Cape], Algoa-bay. Remarks: LT in SAMC; the specimens are labelled as "lectotypus", apparently designated by Zumpt, with no data about the date.
Preferred environment: in Namibia, the edge of the Nama-Karoo Biome (
Type material examined: R. discrepans: 1 ? Gt.Wint-hoek Tulbagh 3600 ft // April1916 R.M.L // discrepans Typ. Villen. // semihirta Typ. Villen. // Lectotypus // slide no 52 // Slide SAM 33B13 [SAMC DIP A011192].
Material examined: Suppl. materials
= Rhyncomya forcipata Villeneuve, 1927: 17. Type locality: Northern Rhodesia [Zambia]; South Africa, Transvaal [Gauteng], Pretoria; and Zaire [Democratic Republic of Congo]. Remarks: ST in NHMUK.
Afrotropical: Botswana, Democratic Republic of Congo, Gambia*, Kenya, Malawi, Mozambique, Namibia, Nigeria*, South Africa (Fig.
Preferred environment: on red sand and directly associated with Acacia sp., Boscia Lam. ex J.St.-Hil. sp. and Rhus lancea L.fil savannah. Additionally, in broad-leafed deciduous woodland and forest, dry scrub (Ficus) riverine and sand forest, mixed Bushveld-grass and Kathu Bushveld to the Olifants River near Balule riparian woodland and Olifantshoek Plains Thornveld., Gordonia Plains Schrubland and Savanah Biome to Mesinda. In Kenya, the dry Acacia savannah, rocks and river margins. In Namibia, the indigenous and degraded sand forest, Miombo and Mopane Woodlands, Kwando River and open savannah floodplain and cultivated plots. Additionally, reported as restricted to Arid and Mesic Savannah Biome in Namibia (
Type material examined: R. forcipata: 1 ? // SYN-TYPE // Mid' / Luangara Koller / N.E. Rhodesia / Aug. 28.1910 / S.A. Neave // Pres. by / Com. Inst. Ent. / B.M. 1955-504 // Rhyncomyia / forcipata / Typ. Villen. // [NHMUK 010832190].
Material examined: Suppl. materials
= Rhyncomya fovealis Bezzi, 1908: 188. Type locality: South Africa, Port Nolloth, Klein Namaland.
Afrotropical: South Africa.
No specimens examined for South Africa, based on
= Rhyncomya hessei Zumpt, 1958: 150. Type locality: South Africa, C. P. [Cape Province - Western Cape], Dikbome, Merweville Koup. Remarks: PTs and HT in SAMC and PTs in NMSA.
Preferred environment: flowers at the roadsides, rocky hillsides and sandy and rocky ridges with succulents. In Namibia, virtually restricted to the semi-arid region, Nama-Karoo and Succulent Karoo Biomes (
Type material examined: R. hessei: 1 ? // HOLOTYPUS // Dikbome / Merweville / Koup / C.P. // Mus. Expd. / Oct. 1952 // slide no. 94 // // Rhyncomyia ? / hessei n.sp. / Det. Zumpt '56 // Slide / SAM: / 33B9 // [SAMC DIP A011196]. 2 ?? // PARATYPE // Wallekraal / Namaqualand // Mus., Expd. / Oct. 1950. // Rhyncomyia ? / hessei n.sp. / Det. Zumpt 1956 // [SAMC DIP A015139, A015140]. 1 ? // PARATYPE // Wallekraal / Namaqualand // Rhyncomyia ? / hessei n.sp. / Det. Zumpt 1956 [NMASA-DIP 19954]. 1 ? // PARATYPE // Wallekraal / Namaqualand // Mus., Expd. / Oct. 1950. // slide no. 92 // Rhyncomyia ? / hessei n.sp. / Det. Zumpt 1956 [SAMC DIP A011197]. 1 ? // PARATYPE // Vogelfontein / P. Albert Div. // A. J. Hesse / Mar.-Apr. 1929 // ? term. 4 // Rhyncomyia / hessei n. sp. / Zumpt 1956 // [SAM-DIP A011198].
Material examined: Suppl. materials
= Beria inflata Robineau-Desvoidy, 1830: 418. Type locality: Cape [South Africa].
= Rhyncomya elegantula Villeneuve, 1927: 20. Type locality: Congo Belge [Democratic Republic of Congo], Kashiobwe.
Afrotropical: Democratic Republic of Congo, Mozambique* and South Africa.
No specimens examined, based on
Material examined: Suppl. material
= Rhyncomya interclusa Villeneuve, 1920: 160. Type locality: South Africa, Western Cape [Capland], Willowmore. Remarks: PT in NMSA and PLT (see
Preferred environment: mesic Mountain Fynbos to plains. In Namibia, apparently restricted to the Desert Biome (
Type material examined: R. interclusa: 1 ? // PARA- / LECTO- / TYPE / Capland / Willowmore, Mai 5 1920 / Dr. Brauns // Pres By / Com Inst Ent / B M 1953-354 // Rhyncomyia / interclusa / Villen. // Paralectotype ? / See Zumpt, 1958 / Explor Parq Nati. / Albert Miss. G.F. / de Witte, 92:156 // [NHMUK 010832194]. 1 ? // Capland / Willowmore / März 1926 / Dr. H. Brauns // PARATYPE // Rhyncomyia ? / interclusa Vill. / det. Zumpt // [NMSA-DIP 19961].
Material examined: Suppl. materials
= Rhyncomyia maculata Macquart, 1846: 322. Type locality: Cape [South Africa].
Preferred environment: no data. Recorded elevations: 479–747 m a.s.l. Seasonality: present August-October, December-January and May, rest of the year absent. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: hand net with hilltopping. Illustrations and photographs: male habitus as in Fig.
Material examined: Suppl. material
= Rhyncomya messoria Villeneuve, 1927: 25. Type locality: Congo Belge [Democratic Republic of Congo], Kilwa; South Africa, Cape of Good Hope [Western Cape]. Remarks: HT in RMCA.
= Beria erula Séguy, 1933: 68. Type locality: Oriental Portuguese East Africa [Mozambique], Moulima.
Afrotropical: Botswana, Democratic Republic of Congo, Kenya, Lesotho, Mozambique, Namibia, South Africa (Fig.
Preferred environment: in Namibia, the Arid and Mesic Savannah and Desert Biomes (
Type material examined: R. messoria: 1 ? // HOLOTYPUS // COLL. MUS. CONGO / Kilwa / (Dr.: J. Bequaert.) // Rhyncomyia / messoria / Typ. Villen. // Kilwa / 28- XII.11 // [RMCA ENT 000012156].
Material examined: Suppl. materials
= Rhyncomya minutalis Villeneuve, 1927: 22. Type locality: Cape [South Africa].
Preferred environment: Acacia Karoo thicket, succulent Karoo, Karoo vegetation, Tanqua succulent Karoo; rocky sandy in Fynbos area, sandy bank and Olifantshoek Plains Thornveld; Savannah Biome and Macchia with Proteas. More or less restricted to the hyper-arid zone in Namibia (
Material examined: Suppl. materials
= Rhyncomyia nana Peris, 1951: 240. Type locality: South Africa, Transvaal [?Gauteng], Plaas Kopernijn. Remarks: HT and PT in NHMUK.
Preferred environment: coastal grassland, open Acacia woodland with thick grass-cover, adjoining thicker valley bush and dry forest; subtropical grassland and open lala-palm woodland. Recorded elevations: 35–830 m a.s.l. Seasonality: low numbers in April, July, November and December, absent the rest of the year. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: sweeping hand net and with Malaise trap. In Mozambique, by sweeping hand net. Illustrations and photographs: male habitus as in Fig.
Type material examined: R. nana: 1 ? // Holo- / type // Plaas Kopesrijn / J.J. Rust Tvl / 19.vii.1924 // Rhyncomyia ? nana n. sp S-V. Peris det. / Tipo // [NHMUK 010832203]. 1 ? // Para-type // Durban. / F. Muir. / 1905-313. // Sharp Coll. / 1905-313 // Rhyncomyia / nana n. sp / S-V. Peris det. / Paratipo // [NHMUK 010832204].
Material examined: Suppl. materials
= Rhyncomya paradoxa Zumpt, 1958: 159. Type locality: South Africa, Cape Province [Eastern Cape], Resolution, Albany District. Remarks: HT in NMSA, PTs in SAMC and SANC at ARC.
Preferred environment: no data. Recorded elevations: 958 m a.s.l. Seasonality: low numbers, present only in February, March, June and December. Behaviour and ecology: two males were collected from a nest of Dasyproctus braunsii and Dasyproctus ?nificanidis (Hymenoptera). Life cycle and developmental stages: unknown. Collection methods: unknown. Illustrations and photography: male habitus as in Fig.
Type material examined: R. paradoxa: 1 ? //Holotype // Resolution / Albany District / 21.ii.1928 / A. Walton // slide no. 88 // [NMSA-DIP 20008] - NMSA Type 2077. 1 ? // Garies / Namaqualand // Museum Staff /June 1930 // slide no. 89 // PARATYPE // Rhyncomyia ? / paradoxa n.sp. / Zumpt, 1956 // Slide number / SAM 4A9 // [SAM-DIP A011222]. 1 ? // Uitenhage / De Hoek / 1x.3.19 / H.K. Munro // PARATYPE // slide no.60 // Rhyncomyia ? / paradoxa n. sp / det. Zumpt 56 // SANC-Pretoria / Database No. / DIPT00302 // "Slide not available in the collection".
Material examined: Suppl. material
= Rhyncomya paratristis Zumpt and Stimie, 1965: 9. Type locality: South Africa, Zululand Natal [KwaZulu-Natal], Ndumu Reserve, Ingwavuma District. Remarks: HT and PTs in NMSA.
Preferred environment: broad-leafed deciduous woodland, sand forest and broad-leafed deciduous forest, dry scrub forest, grassy floodplain and at the margin of dune forest. Recorded elevations: 10–98 m a.s.l. Seasonality: abundant, present in June and November, peaking in December. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: Malaise traps. Illustrations and photographs: male habitus as in Fig.
Type material examined: R. paratristis: 2 ?? // HOLOTYPE // Ndumu Reserve / Ingwavuma dist. / Zululand, Natal / South Africa / 1-10.xii.63 // collectors / B. & P. Stuckenberg // Rhyncomya ? / paratristis / Zumpt & Stimie // [NMSA-DIP 074955, 074956]. 2 ?? // PARATYPE // Ndumu Reserve / Ingwavuma dist. / Zululand, Natal / South Africa / 1-10.xii.63 // collectors / B. & P. Stuckenberg // Rhyncomya ? / paratristis / Zumpt & Stimie // [NMSA-DIP 061694, 074957].
Material examined: Suppl. material
= Rhyncomya peraequa Villeneuve, 1929: 186. Type locality: South Africa, Transvaal [Mpumalanga], Barberton. Remarks: HT in SAMC.
Preferred environment: all Namibian biomes, except desert (
Type material examined: R. peraequa: 1 ? // Type // Barberton / Transvaal / H Edwards // Dec / 1911 // Type // Rhyncomyia / peraequa / Type Villen //[SAMC DIP A011223].
Material examined: Suppl. materials
= Rhyncomya pruinosa Villeneuve, 1922: 65. Type locality: Kenya, Zaire [Democratic Republic of Congo], Nyasaland [Malawi]; Anglo-Egipsian Sudan [Sudan].
Afrotropical: Angola, Botswana, Cameroon*, Democratic Republic of Congo, Ethiopia*, Gambia, Kenya, Malawi, Mauritania*, Mozambique, Namibia, Nigeria, Senegal*, South Africa (Fig.
Preferred environment: Dry, sand and broad-leafed deciduous forest and Acacia savannah. In Cameroon, in degraded savannah forest. In Namibia, in Miombo and Mopane Woodlands, Arid and Mesic Savannah and Nama-Karoo Biomes (
Material examined: Suppl. materials
= Rhyncomya soyauxi Karsch, 1886: 262. Type locality: Pongo-Andongo [Angola]. Remarks: HT in ZMHB.
= Rhyncomya pictifacies Bigot, 1888: 595. Type locality: Cape [South Africa].
= Rhynchomyia isaea Séguy, 1933: 69. Type locality: Cameroon.
= Rhynchomyia proterva Séguy, 1938: 378. Type locality: Kenya, Mt. Elgon.
Afrotropical: Angola, Botswana, Cameroon, Democratic Republic of Congo, Kenya, Malawi, Namibia, Rwanda*, South Africa (Fig.
Preferred environment: wild environments such as Acacia veld (dry mixed bush, savannah and woodland), forests (indigenous Afromontane, broad-leafed deciduous woodland, Ficus forest, sand and red sand), grasslands (grassy floodplain, mixed, grass and Kathu), savannah, Kalahari thornveld and rural and urban environments, such as the Albany Museum grounds, camp site areas and sewage-seepage areas. In Kenya, Kenyan dry forest; in Malawi, forest edge, margins and grasslands; in Namibia, the Kwando River floodplain, Miombo and mopane woodlands and open savannah floodplain. Almost all Namibian biomes, except the Hyper-Arid Desert and Succulent Karoo Biomes (
Type material examined: R. soyauxi: 1 ? // Typus // Pungo-Andongo / Leg. V. Homeyer // 11013 // Rhynch. / ?Soyeauxi / K.* // Rhyncomyia ? / soyauxi Karsch / C = pictifacies Bigot) // [ZMHB].
Material examined: Suppl. materials
= Rhyncomya stannocuprea Speiser, 1910: 150. Type locality: Tanganyika [Tanzania], Meru Kilimandjaro.
= Rhyncomyia stannocuprea spp. abyssinica Peris, 1951: 244. Type locality: Abyssinia [Ethiopia], Gatelo Amaizu.
Preferred environment: Acacia savannah, rocky hills with Acacia veld and grassland and forest edges. In Malawi, forest edges and grasslands; in Namibia the Kwando River floodplain. Recorded elevations: 1200–1500 m a.s.l. Seasonality: low numbers year-round, peaking in September, absent in March and May to July. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: Malaise trap. In Nambia, Malaise trap. Illustrations and photographs: male habitus as in Fig.
Material examined: Suppl. materials
= Rhyncomya trispina Villeneuve, 1929: 62. Type locality: Southern Rhodesia [Zimbabwe], Bulawayo. Remarks: HT in NHMUK.
Preferred environment: broad-leafed deciduous woodland and dry scrub forest and to Acacia thornveld, Kwando river floodplain and the Miombo and mopane woodlands. In Namibia, degraded sand forest and cultivated plots.
Type material examined: R. trispina: 1 ? // Holo- / type // Bulawayo / S. Rhodesia / 25.ix.1923 / Coll. R. Stevenson // Pres. by / Com. Inst. Ent. / B.M. 1955-504 // Rhyncomyia / trispina / type Villen. // [NHMUK 010832199].
Material examined: Suppl. materials
= Rhynchomyia tristis Séguy, 1933: 67. Type locality: Mozambique, Zambèze [Zambezi], N. Chupanga [Nova-Chupanga]. Remarks: HT and PT in MNHN.
Afrotropical: Botswana, Chad, Namibia, Nigeria, Mozambique, South Africa* (Fig.
Preferred environment: in Tanzania, riverine in dry forest. In Namibia, the Arid Savannah, Mesic Savannah and Nama-Karoo Biomes (
Type material examined: R. tristis: 1 ? // TYPE // MUSEUM PARIS / Zambèze / N. Chupanga / J. SURCOUF 1926 // Octobre 27 // Rhyncomyia / tristis Séguy / TYPE / E. Séguy det. 1925 // slice no. 16 // Prépar. microsc. / nº 3344. 1 ? // PARATYPE // MUSEUM PARIS / Zambèze / N. Chupanga / J. SURCOUF 1926 // Janvier 28 // Rhyncomyia ? / tristis Séguy / vide Zumpt 75.
Material examined: Suppl. materials
= Rhyncomya viduella Villeneuve, 1927: 18 (see taxonomic notes). Type locality: South Africa, Transvaal [Mpumalanga], Barberton. Remarks: HT in SAMC; nec Rhynchomya cassotis (Walker, 1849) sensu
Preferred environment: hillside with flowers. Recorded elevations: 495 m a.s.l. Seasonality: low numbers in September, October and December. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: sweeping from Asparagus sp. (Asparagaceae). Illustrations and photographs: male habitus as in Fig.
Taxonomic notes: Rhyncomya viduella stat. rev. is reinstated as a valid species. Previously listed as a synonym of R. cassotis (Fig. 37) by
Type material examined: R. viduella: 1 ? // Barberton / Transvaal /H Edwards // Dec 1911 // Rhyncomyia / viduella / ? Type // [SAMC DIP A011183].
Material examined: Suppl. material
= Stegosoma Loew, 1863: 15. Type species: Stegosoma vinculatum Loew, 1863, by monotypy.
= Stegosoma bowdeni Peris, 1951: 239. Type locality: Golden Coast [Ghana], Mampong. Remarks: HT and PT in NHMUK and PTs in NMSA.
Preferred environment: in Togo, vegetated stream-bed and lowland evergreen secondary forest. Recorded elevations: no data. Seasonality: recorded in February, October and December. Behaviour and ecology:
Type material examined: S. bowdeni: 1 ? // Holotype // GOLD COAST / Mampong / (Ashanti) / 12.iv.1947 / J. Bowden // Stegosoma / bowdeni / n.sp. / S-V. Peris 1948 // [NHMUK]. 5 ?? // GOLD COAST / Mampong / (Ashanti) / 12.iv.1947 / J. Bowden / 451/47 // Ovipositing in / opened mount /of macrotermes // NB Par-of / onqunalantes: / presumably rank as / Paratypes J.B. // Stegosoma / bowdeni / n.sp. / S-V. Peris 1948 // [NMSA DIP 019516].
Material examined: Suppl. materials
= Stegosoma vinculatum Loew, 1863: 15. Type locality: South Africa, Orange Free State [Free State], Bloemfontein.
Afrotropical: Benin, Botswana, Democratic Republic of Congo, Ghana, Kenya, Malawi, Mali, Mozambique, Namibia, Nigeria, South Africa (Fig.
Preferred environment: Acacia savannah and mixed bushveld-grass. In Namibia, degraded sand forest and cultivated plots; apparently restricted to the Arid and Mesic Savannah Biomes (
Material examined: Suppl. materials
= Rhynchomyia wellmani Lichtwardt, 1908: 338. Type locality: Angola, Benguella.
Afrotropical: Angola, Cameroon, Central African Republic*, Democratic Republic of Congo, Equatorial Guinea, Ghana, Kenya, Liberia, Nigeria, Sierra Leone, South Africa (Fig.
Preferred environment: in Democratic Republic of Congo, in lowland evergreen swamp forest. Recorded elevations: no data. Seasonality: recorded in January, February and April, absent the rest of the year. Behaviour and ecology: collected at Microtermes Wasmann nest in Kenya and ovipositing in opened mound of Macrotermes Holmgren in Ghana. Life cycle and developmental stages: unknown. Collection methods: in Democratic Republic of Congo, with Malaise trap. Illustrations and photographs: male habitus as in Fig.
Material examined: Suppl. materials
= Thoracites Brauer and Bergenstamm, 1891: 363. Type species: Musca abdominalis Fabricius, 1805, by original designation.
= Thoracites cingulatus Bezzi, 1914: 290. Type locality: Senegal, Theis.
Preferred environment: no data. Recorded elevations: no data. Seasonality: three specimens recorded in December. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: unknown. Illustrations and photographs: female habitus as in fig. 29a in
Material examined: Suppl. material
= Thoracites kirkspriggsi Kurahashi, 2001: 146. Type locality: Namibia, Rundo District.
Preferred environment: in Namibia, apparently restricted to the Arid and Mesic Savannah Biomes (Kurahashi 2001;
Material examined: Suppl. materials
= Ochromyia petersiana Loew, 1852: 660. Type locality: South Africa, Zululand [KwaZulu-Natal], Mtubatuba.
= Thoracites neglectus Zumpt, 1972: 49. Type locality: South Africa, Zululand [KwaZulu-Natal], Mtubatuba. Remarks: HT in NMSA.
Preferred environment: broad-leafed deciduous woodland forest, grassy floodplain and sand and forest. Recorded elevations: 77–98 m a.s.l. Seasonality: abundant species present in May, November and December (peaking in December). Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: Malaise trap. Illustrations and photographs: male habitus as in Fig.
Type material examined: T. neglectus: 1 ? // Holotypus // Mtubatuba / Zululand / May 1941 / H.K. Munro // slide no. 95 // Thoracites / neglectus n. sp. / Zumpt 1972 // [NMSA-DIP 019669].
Material examined: Suppl. material
= Thoracites sarcophagoides Kurahashi, 2001: 155. Type locality: Namibia, West Caprivi PK, Kwando River.
Preferred environment: Gordonia Plains Schrubland, Senegalia mellifera (as Acacia) on red sand in the Savannah Biome. Recorded elevations: 1035 m a.s.l. Seasonality: only one specimen between September and November. In Namibia, recorded year-round, most abundant from December to February and September (Kurahashi 2001,
Material examined: Suppl. material
= Trichoberia Townsend, 1933: 439. Type species: Trichoberia rufopilosa Townsend, 1933 (= Rhyncomyia lanata Villeneuve, 1920), by original designation.
= Trichoberia kamita Lehrer, 2007: 13. Type Locality: South Africa, Natal [KwaZulu-Natal], St. Lucia Park.
Afrotropical: South Africa.
No specimens examined for South Africa, based on
= Rhyncomya lanata Villeneuve, 1920: 162. Type locality: Congo Belge [Democratic Republic of Congo].
= Trichoberia rufopilosa Townsend, 1933: 440. Type locality: Guinea.
Preferred environment: forest and open woodland areas. Recorded elevations: no data. Seasonality: three males between September and October. Behaviour and ecology: unknown. Life cycle and developmental stages: unknown. Collection methods: unknown. Illustrations and photographs: male habitus as in Fig.
Material examined: Suppl. material
= Zumba Peris, 1951: 239. Type species: Zumba rhinoidea Peris, 1951, by original designation.
= Rhyncomyia antennalis Villeneuve, 1929: 185. Type locality: South West Africa [Namibia]. Remarks: LT in SAMC designated by Zumpt 1958.
= Pseudorhyncomyia deserticola Zumpt and Argo, 1978: 35. Type locality: South West Africa [Namibia], Gobabeb. Remarks: HT in NMSA.
Preferred environment: Gordonia Plains Shrubland, Olifantshoek Thornveld plains, Senegalia mellifera on red sand and the Savannah Biome. All the Namibian Biomes, but mainly present in the Desert and Succulent Karoo Biomes showing preferences for the hyper-arid regions (
Type material examined: R. antennalis: 1 ? // Mafa / Feb. 1923 // S.W. Africa / Mus. Exped. // Rhyncomyia / antennalis / Typ. Villen. // Zumba ? / antennalis Vill. / det. Zumpt 56 // LECTO-TYPE / designated / Zumpt 1958 // [SAMC-DIP A011283]; 1 ? // Mafa / Feb. 1923 // S.W. Africa / Mus. Exped. // Type series // [SAMC-DIP A015172]. P. deserticola: 1 ? // S.W. Africa: Namib / Desert, Welwitschia / “Forest” nr Gobabeb / 3-X-1967 E. S. Ross / A. R. Stephen // slide no. 6 // Pseudorhyncomyia ? / deserticola n. sp. / Zumpt & Argo 1976 // HOLOTYPUS // [NMSA-DIP 19837].
Material examined: Suppl. materials
= Zumba rhinoidea Peris, 1951: 239. Type locality: N. Rhodesia [Zambia], Mozabuka.
Afrotropical: South Africa and Zambia.
No specimens examined for South Africa, based on
This review showed that the life cycles of the Rhiniinae are more diverse than previously thought (
Seasonal abundance of South African Rhiniinae species, based on the material reviewed for this study and with available data. Abbreviations used in the table: J: January; F: February; M: March; A: April; M: May; J: June; J: July; A: August; S: September; O: October; N: November; D: December; T: Total.
Species | J | F | M | A | M | J | J | A | S | O | N | D | T |
Cosmina aenea | 5 | 2 | 14 | 1 | 1 | 3 | 0 | 1 | 5 | 1 | 14 | 5 | 52 |
C. fuscipennis | 7 | 1 | 9 | 0 | 1 | 0 | 2 | 8 | 41 | 64 | 12 | 7 | 152 |
C. gracilis | 2 | 0 | 2 | 2 | 1 | 0 | 0 | 0 | 18 | 6 | 9 | 3 | 43 |
C. margaritae | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 3 | 4 |
C. undulata | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 3 |
Eurhyncomyia diversicolor | 1 | 4 | 5 | 5 | 1 | 1 | 6 | 2 | 0 | 5 | 3 | 24 | 57 |
E. metzi | 0 | 0 | 0 | 0 | 0 | 0 | 7 | 0 | 1 | 0 | 0 | 0 | 8 |
Fainia albitarsis | 13 | 14 | 1 | 2 | 7 | 5 | 7 | 6 | 1 | 4 | 16 | 22 | 98 |
F. elongata | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
Isomyia cuthbertsoni | 0 | 0 | 4 | 0 | 0 | 0 | 0 | 0 | 0 | 11 | 2 | 15 | 32 |
I. darwini | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 4 | 1 | 7 |
I. deserti | 2 | 2 | 2 | 0 | 2 | 1 | 0 | 0 | 0 | 0 | 0 | 2 | 11 |
I. distinguenda | 3 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 8 | 8 | 23 |
I. dubiosa | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 3 | 1 | 0 | 4 |
I. eos | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 3 |
I. longicauda | 3 | 0 | 1 | 0 | 0 | 1 | 3 | 2 | 1 | 0 | 1 | 1 | 13 |
I. natalensis | 17 | 48 | 42 | 23 | 0 | 0 | 0 | 5 | 25 | 17 | 35 | 26 | 238 |
I. oculosa | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 3 | 0 | 3 |
I. pubera | 2 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 5 | 1 | 1 | 1 | 13 |
I. transvaalensis | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 |
I. tristis | 39 | 13 | 62 | 43 | 11 | 3 | 2 | 5 | 34 | 21 | 26 | 35 | 294 |
Pseudorhyncomyia braunsi | 0 | 1 | 0 | 2 | 5 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 9 |
Rhinia. apicalis | 5 | 10 | 10 | 32 | 5 | 6 | 8 | 4 | 7 | 9 | 20 | 43 | 159 |
Rh. coxendix | 5 | 2 | 2 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 12 |
Rh. nigricornis | 3 | 1 | 6 | 6 | 1 | 0 | 0 | 1 | 0 | 0 | 2 | 10 | 30 |
Rhyncomya bicolor | 1 | 1 | 2 | 0 | 0 | 0 | 0 | 0 | 1 | 2 | 0 | 0 | 7 |
R. botswanae | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 |
R. cassotis | 2 | 3 | 10 | 5 | 4 | 7 | 1 | 4 | 3 | 4 | 6 | 12 | 61 |
R. currani | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 4 | 0 | 0 | 0 | 4 |
R. dasyops | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 6 |
R. depressifrons | 2 | 2 | 2 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 0 | 8 |
R. disclusa | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 5 | 4 | 0 | 4 | 14 |
R. discrepans | 0 | 0 | 0 | 6 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 7 |
R. forcipata | 0 | 10 | 6 | 4 | 3 | 1 | 0 | 0 | 18 | 9 | 32 | 54 | 137 |
R. hessei | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 6 | 14 | 7 | 0 | 0 | 28 |
R. interclusa | 3 | 1 | 2 | 1 | 2 | 0 | 0 | 0 | 3 | 9 | 6 | 3 | 30 |
R. maculata | 1 | 0 | 0 | 0 | 3 | 0 | 0 | 1 | 3 | 4 | 0 | 3 | 15 |
R. messoria | 2 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 2 | 0 | 8 | 14 |
R. minutalis | 4 | 16 | 6 | 6 | 2 | 0 | 0 | 1 | 9 | 23 | 6 | 5 | 78 |
R. nana | 0 | 0 | 4 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 3 | 10 |
R. paradoxa | 0 | 3 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 6 |
R. paratristis | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 10 | 111 | 122 |
R. peraequa | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 1 |
R. pruinosa | 0 | 0 | 0 | 1 | 0 | 1 | 2 | 1 | 1 | 0 | 3 | 6 | 15 |
R. soyauxi | 14 | 16 | 11 | 26 | 8 | 2 | 7 | 4 | 73 | 8 | 31 | 54 | 254 |
R. stannocuprea | 3 | 1 | 0 | 3 | 0 | 0 | 0 | 2 | 7 | 1 | 1 | 2 | 20 |
R. trispina | 0 | 0 | 0 | 2 | 1 | 0 | 1 | 0 | 0 | 0 | 5 | 0 | 9 |
R. tristis | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 |
R. viduella | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 5 | 0 | 1 | 7 |
Stegosoma bowdeni | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 0 | 1 | 4 |
St. vinculatum | 5 | 7 | 18 | 2 | 3 | 6 | 0 | 0 | 0 | 7 | 6 | 15 | 69 |
St. wellmani | 2 | 1 | 1 | 2 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 6 |
Stomorhina apta | 4 | 0 | 0 | 2 | 0 | 0 | 0 | 0 | 2 | 0 | 2 | 2 | 12 |
S. armatipes | 1 | 1 | 2 | 1 | 0 | 1 | 0 | 0 | 3 | 1 | 2 | 0 | 12 |
S. chapini | 6 | 0 | 1 | 6 | 2 | 1 | 1 | 1 | 0 | 0 | 5 | 4 | 27 |
S. cribrata | 5 | 0 | 4 | 3 | 0 | 1 | 0 | 1 | 7 | 1 | 3 | 15 | 40 |
S. guttata | 2 | 1 | 3 | 11 | 0 | 3 | 0 | 3 | 17 | 16 | 2 | 6 | 64 |
S. lunata | 40 | 50 | 24 | 11 | 7 | 4 | 2 | 27 | 34 | 26 | 28 | 18 | 271 |
S. malobana | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 |
S. rugosa | 13 | 31 | 12 | 10 | 7 | 10 | 8 | 3 | 2 | 8 | 19 | 28 | 151 |
Thoracites cingulatus | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 3 | 3 |
Th. kirkspriggsi | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 |
Th. petersiana | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 20 | 57 | 78 |
Th. sarcophagoides | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 |
Trichoberia lanata | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 3 | 0 | 3 |
Zumba antennalis | 2 | 3 | 2 | 0 | 0 | 0 | 0 | 0 | 4 | 2 | 2 | 0 | 15 |
Total | 222 | 251 | 275 | 222 | 81 | 60 | 61 | 88 | 354 | 285 | 356 | 628 | 2883 |
Biological information compiled for the Afrotropical Rhiniinae from the material reviewed (Appendices I and II) and literature. Abbreviations used in the table: W: wasp association; T: Termite association; A: Ant association; F: collected or observed on flowers; S: females observed laying eggs on soil or immature stages associated with soil; AB: collected in animal burrows; H: collected or observed hovering; S: collected or observed swarming; FD: collected on fresh dung of different animals; O: Association with Orthoptera oothecae. Notes: †oviparous, †† Unilarviparous.
Species | W | T | A | F | S | AB | H | Sw | FD | O |
Cosmina aenea | ||||||||||
C. fuscipennis | x | x | ||||||||
C. gracilis† | x | |||||||||
C. margaritae† | ||||||||||
C. undulata | ||||||||||
C. thabaniella | ||||||||||
Eurhyncomyia diversicolor | ||||||||||
E. metzi | ||||||||||
Fainia albitarsis† | x | x | x | |||||||
F. elongata | ||||||||||
Isomyia cuthbertsoni | x | |||||||||
I. darwini | x | |||||||||
I. deserti | x | |||||||||
I. distinguenda | ||||||||||
I. dubiosa | x | |||||||||
I. eos | ||||||||||
I. innnia | ||||||||||
I. longicauda | ||||||||||
I. natalensis | x | |||||||||
I. oculosa | x | |||||||||
I. pubera | x | |||||||||
I. transvaalensis | ||||||||||
I. tristis | x | x | x | |||||||
Pseudorhyncomyia braunsi | x | x | ||||||||
Rhinia apicalis† | x | x | x | x | x | x | ||||
Rh. coxendix† | x | |||||||||
Rh. nigricornis | x | x | ||||||||
Rhyncomya bicolor | ||||||||||
R. botswana | x | |||||||||
R. buccalis | ||||||||||
R. cassotis | x | |||||||||
R. currani | ||||||||||
R. dasyops | x | |||||||||
R. depressifrons | ||||||||||
R. disclusa | x | |||||||||
R. discrepans | ||||||||||
R. forcipata | x | x | ||||||||
R. fovealis | ||||||||||
R. hessei | ||||||||||
R. inflata | ||||||||||
R. interclusa | x | |||||||||
R. maculata | ||||||||||
R. messoria | ||||||||||
R. minutalis | x | |||||||||
R. nana | x | |||||||||
R. paradoxa | x | |||||||||
R. paratristis | ||||||||||
R. peraequa | ||||||||||
R. pruinosa† | x | x | x | |||||||
R. soyauxi | x | x | x | |||||||
R. stannocuprea | ||||||||||
R. trispina | x | |||||||||
R. tristis | ||||||||||
R. viduella | x | |||||||||
Stegosoma bowdeni | x | |||||||||
St. vinculatum† | x | x | x | x | x | |||||
St. wellmani | x | |||||||||
Stomorhina apta | ||||||||||
S. armatipes† | x | x | x | |||||||
S. chapini | ||||||||||
S. cribrata† | x | x | x | x | x | |||||
S. guttata | x | x | ||||||||
S. lunata† | x | x | x | x | ||||||
S. rugosa | x | x | x | |||||||
S. malobana†† | x | |||||||||
Thoracites kirkspriggsi | ||||||||||
Th. petersina | ||||||||||
Th. cingulatus | ||||||||||
Th. sarcophagoides | ||||||||||
Trichoberia kamita | ||||||||||
T. lanata | ||||||||||
Zumba antennalis†† | ||||||||||
Z. rhinoides | x |
Data on the ecology and natural history of several rhiniid species also improved. All species of Pseudorhyncomyia and Stegosoma, several species of Stomorhina and a few species of Rhyncomya show some level of association with termites (Table
Similarly, three Rhinia species (R. apicalis, R. coxendix and R. nigricornis) showed different associations with wasps (specifically Bembix melanopa in Cuthbertson 1938 and Cerceris yngvei on material examined), as both adults and larvae were collected inside nests and females were observed attending nests. This suggests an ecological relationship between Rhinia and these wasps, the nature of which remains unknown. Four other species (F. albitarsis, R. apicalis, R. pruinosa and S. cribrata) showed a relationship with soft, turned or humus-rich soil, as several females were reported ovipositing in soil (Table
The abundance of adult Rhiniinae varies seasonally (Table
Seventy-three species of Rhiniinae belonging to twelve genera are now known for South Africa, nine of which are new records (Cosmina undulata, Isomyia cuthbertsoni, Rhyncomya botswanae, R. trisis, Stomorhina apta, S. malobana, Thoracites kirkspriggsi, Th. sarcophagoides and Trichoberia lanata). Of the 73 species, 66 were examined (identifications, re-identifications and corroborations) and the other seven were assessed exclusively on literature records.
The 73 species represent almost 50% of the Rhiniinae fauna known for the Afrotropical Region (approx. 150 species sensu
Our review confirms that South Africa has the highest species richness of Rhiniinae, followed by India with 57 species (
At the tribe level, Cosminini is the most diverse in South Africa with 60 species in nine genera, with Rhyncomya the largest genus of the subfamily with 27 species that represent 54% of the 50 known Afrotropical Rhyncomya species sensu
Specimens here studied have been sampled using a variety of techniques, each of them offering different advantages and targets. The most common technique is Malaise trap, followed by hand netting, pan trapping (23 yellow, eight blue, six white, two brown and one orange) and pitfall trapping (Table
Collection methods reported for the Afrotropical Rhiniinae examined (Appendices I and II). Abbreviations used in the table: sweeping with hand net (Sb: Sweeping on bait); Pan traps (Br: Brown pan; Bu: Blue pan; Y: Yellow pan; W: White pan; Mc: McPhail trap with Nu-Lure); Light trap (B: black light; MV-B: Mercury Vapour and black light; UV: ultraviolet light); Traps baited with organic matter (R: Rotten fish; Ba: Banana; Ff: Hanging traps with fermenting fruit; Dm: dead millipedes).
Species | Malaise trap | Hand net | Pan traps | Pitfall trap | Light trap | Organic baits |
Cosmina aenea | x | x | ||||
C. fuscipennis | x | x | x (Y, W) | x (Ba) | ||
C. gracilis | x | x | x (Y, Bu) | x | x, x (UV) | x (Ff) |
C. margaritae | x | x (Y, Bu) | x | |||
C. undulata | x | |||||
C. thabaniella | ||||||
Eurhyncomyia diversicolor | x | |||||
E. metzi | ||||||
Fainia albitarsis | x | x | x | |||
F. elongata | x | x | x (R) | |||
Isomyia cuthbertsoni | x | x | ||||
I. darwini | x | x | x (UV) | |||
I. deserti | x | x | x (Y) | |||
I. distinguenda | X | |||||
I. dubiosa | x | x (MV-B) | ||||
I. eos | x | |||||
I. innnia | ||||||
I. longicauda | x | x | ||||
I. natalensis | x | x | x (Y) | x (MV) | ||
I. oculosa | ||||||
I. pubera | x | |||||
I. transvaalensis | ||||||
I. tristis | x | x, (Sb) | x (Y, W) | x | x (MV) | |
Pseudorhyncomyia braunsi | x (UV) | |||||
Rhinia apicalis† | X, x (UV) | x | x (Y) | x | ||
Rh. coxendix† | x | x | ||||
Rh. nigricornis | x | x (R) | ||||
Rhyncomya bicolor | x | |||||
R. botswana | x | |||||
R. buccalis | ||||||
R. cassotis | x | x | x (Y) | x | x (MV) | |
R. currani | ||||||
R. dasyops | x | x | ||||
R. depressifrons | ||||||
R. disclusa | ||||||
R. discrepans | X | |||||
R. forcipata | x, x (MV) | X | x | x, x (UV, MV-B) | ||
R. fovealis | ||||||
R. hessei | x | x (Y) | x | x (UV) | ||
R. inflata | ||||||
R. interclusa | x | x (Y) | ||||
R. maculata | ||||||
R. messoria | x | X | x (Y) | x | ||
R. minutalis | x | X | x (Y) | x | ||
R. nana | x | X | ||||
R. paradoxa | ||||||
R. paratristis | x | |||||
R. peraequa | x | x (Y) | x | x (UV) | ||
R. pruinosa | x | X | x (Y) | x | x, x (UV) | |
R. soyauxi | x, x (Ex) | X | x (Bu, Y, W, Mc) | x | x, x (UV, MV-B) | |
R. stannocuprea | x | |||||
R. trispina | x | x | x (Bu, Y) | x | x (UV) | x (Ff) |
R. tristis | x | x | x (Y) | x | x (UV) | |
R. viduella | x | |||||
Stegosoma bowdeni | ||||||
St. vinculatum | x | x | x | |||
St. wellmani | x | |||||
Stomorhina apta | x | |||||
S. armatipes | x | x | ||||
S. chapini | x | |||||
S. cribrata | x | |||||
S. guttata | x | x | x (Y) | x | ||
S. lunata | x | x | x (Y) | x | x (B) | |
S. rugosa | x | x (Bu) | x | |||
S. malobana | x | x | ||||
Thoracites kirkspriggsi | x | x | x (Bu, Y, W) | x (Dm) | ||
Th. petersina | x | |||||
Th. cingulatus | ||||||
Th. sarcophagoides | ||||||
Trichoberia kamita | ||||||
T. lanata | ||||||
Zumba antennalis | x | x | x (Br, Bu, Y, W) | x | x (UV) | |
Z. rhinoides |
Compared with other African countries, South Africa has had a broad sampling effort as indicated by the almost 3,000 collected specimens. However, sampling has been uneven. KwaZulu-Natal has been extensively sampled and showed the largest number of collected individuals and diversity of species, while the Free State, Northern Cape and North West Provinces have the lowest, showing that more collection effort is necessary in the central areas of the country. This is clearly evident in most of the distribution maps for each species and was very well illustrated in a trapping and mapping study of the common and economically significant calliphorids Lucilia sericata (Meigen, 1826) and Lucilia cuprina (Wiedemann, 1830) (
Overall, although knowledge on the Diptera of South Africa is vast, collection expeditions and research have never focused specifically on Rhiniinae. In this context, thorough revisions, based on specimens housed in entomological collections, such as the one presented here, are very useful for gathering information that would otherwise be scattered and lost across different institutions. Our main findings and contributions include nine new records for South Africa, one new combination, one reinstated species and significant information on the life habits and ecology of the group, all of which form a base for productive future expeditions and studies focused on Rhiniinae. In particular, future studies should focus on the taxonomic value of the immature stages, exploring the ecological association of some species of Rhiniinae with termites, ants, wasps and soils rich in organic matter and the phylogeny of the family. Promoting and training local taxonomists on Diptera would be important to increase our knowledge of this complex group.
This study would not have matured without the help of numerous institutions and individuals. We especially thank Terence Bellingan (AMGS); Ashley Kirk-Spriggs and Burgert Muller (BMSA); Cinta Quirce (CIBIO-UA); Natasha Govender (DMSA); Emmanuel Delfosse (MNHN); Pierfilippo Cerretti (MZSUR); Daniel Whitmore and Nigel Wyatt (NHMUK); Igor Muratov, John Midgley, Kirstin Williams, Linda Davies and Tricia Pillay (NMSA); Kurt Jordaens (RMCA); Aisha Mayekiso and Simon van Noort (SAMC); Kurt Jordaens and Robert Copeland (PINDIP-project in Kenya); Vivienne Uys (SANC); Concepción Ornosa, Eduardo Ruiz Piña, María Angeles Vazquez, Raimundo Outerelo (UCME); Mike Mostovski and Netta Dorchin (SMNHTAU (TAUI)); Knut Rognes (UiS); Torsten Dikow (USNM-SM); Bernhard Schurian, Eliana Buenaventura, Jenny Pohl, Joachim Ziegler and Sven Marotzke (ZMHB); and Arn Rytter Jensen and Thomas Pape (ZMUC). We also thank J. Camilo Azpúrua for his invaluable comments and linguistic input and to the reviewers/editor of this manuscript, Thomas Pape, Krzysztof Szpila, Daniel Whitmore, Pierfilippo Cerretti and Kurt Jordaens. This work was partially funded by the H2020 Research and Innovation Staff Exchange Programme of the European Commission (RISE), project 645636: ‘Insect-plant relationships: insights into biodiversity and new applications’ (FlyHigh); by the International mobility Grant of the University of Alicante; and the Bøje Benzon's Foundation Grant from the Natural History Museum of Denmark.
Arianna Thomas-Cabianca: designed and developed the research project, revised all the specimens, wrote the entire text, generated all the high resolution pictures, figures and tables.
The co-authors provided funds, made suggestions on the scope of the research and critically reviewed and edited the text for its improvement.