Abstract
The Afrotropical endemic platystomatine genus Engistoneura Loew, 1873 is revised taxonomically, based on the study of name-bearing types of all described species and varieties, material housed in museums around the world and recently collected specimens from the rainforests of Central and West Africa. Following revisionary study, the genus now comprises 22 species, six previously described, namely: E. currani Steyskal, 1965 (Liberia), E. flavipennis Hendel, 1914 (Ghana), E. lugens (Fabricius, 1794) (Guinea and Sierra Leone), E. moerens (Fabricius, 1794) (Benin, Cameroon, Côte d’Ivoire, Equatorial Guinea, Ghana, Guinea, Nigeria and Togo), E. obscura Hendel, 1914 (Cameroon, Equatorial Guinea, Gabon and Republic of Congo) and E. parallela (Wiedemann, 1830) (Sierra Leone) and 16 species are described and figured as new to science, namely: E. ankasa, sp. nov. (Ghana), E. biseriata, sp. nov. (Côte d’Ivoire), E. circumfusus, sp. nov. (Guinea), E. deemingi, sp. nov. (Côte d’Ivoire, Nigeria and Togo), E. distincta, sp. nov. (Liberia and Sierra Leone), E. elvillah, sp. nov. (Guinea), E. hemifascia, sp. nov. (Côte d’Ivoire), E. hexafascia, sp. nov. (Sierra Leone), E. kachana, sp. nov. (Ghana), E. maya, sp. nov. (Guinea), E. mcalpinei, sp. nov. (Sierra Leone), E. nebula, sp. nov. (Sierra Leone), E. fatima, sp. nov. (Côte d’Ivoire and Sierra Leone), E. ghanensis, sp. nov. (Ghana), E. smithi, sp. nov. (Côte d’Ivoire and Ghana) and E. vicina, sp. nov. (Cameroon, Côte d’Ivoire, Equatorial Guinea, ?Gabon, Guinea and Liberia). The following nomenclatorial acts are proposed: E. catogastera (Bigot, 1891) syn. n. is sunk as a junior synonym of E. moerens; E. unilineata Bezzi, 1914 is removed from the genus Engistoneura and placed in new combination as Peltacanthina (Peltacanthina) unilineata (Bezzi, 1914) comb. n.; the variety names E. catogastera var. melanopleura Enderlein, 1924, syn. n., E. obscura var. duplicata Enderlein, 1924, syn. n. and E. obscura var. interrupa Enderlein, 1924, syn. n. are all sunk as junior synonyms of E. moerens and the variety name E. obscura var. simplex Enderlein, 1924, syn. n. is sunk as a junior synonym of E. obscura Hendel, 1914. Bezzi’s (1908) synonymy of E. albovaria (Walker, 1853) as a junior synonym of E. moerens (Fabricius, 1794) is here confirmed and upheld, based on the examination of the name-bearing types of both species. Lectotypes are designated for the following 18 taxa: E. albolineata Bezzi, 1908, E. bicolor Bezzi, 1908, E. catogastera var. melanopleura Enderlein, 1924, E. cohaesa Speiser, 1911, E. flavipennis Hendel, 1914, E. guttata Bezzi, 1908, E. obscura Hendel, 1914, E. obscura var. duplicata Enderlein, 1924, E. obscura var. interrupta Enderlein, 1924, E. obscura var. simplex Enderlein, 1924, E. octodecim Speiser, 1911, E. surniipennis Speiser, 1911, E. unilineata Bezzi, 1914, Megaglossa catogastera Bigot, 1891, Musca lugens Fabricius, 1794, Mu. moerens Fabricius, 1794, Ortalis parallela Wiedemann, 1830 and Trypeta albovaria Walker, 1853. An identification key to the 22 species is provided and the distributions of the species are mapped and biogeography of the genus is discussed in relation to the Riverine Barriers Hypothesis, the Refuge Hypothesis, forest fragmentation and elevational considerations. Although the limited distribution of some species appear to be bounded by rivers, examination of collective patterns indicate that forest refugia rather than rivers appear to have driven diversification and speciation in the genus.
References
Aduse-Poku, K., van Bergen, E., Sáfián, S., Collins, S.C., Etchangienne, R.S., Herrera-Alsina, H., Brakefield, P.M., Brattström, O., Lohman, D.J. & Wahlberg, N. (2022) Miocene climate and habitat change drove diversification in Bicyclus, Africa’s largest radiation of satyrine butterflies. Systematic Biology, 71, 570–588. https://doi.org/10.1093/sysbio/syab066
Aleixo, A. (2006) Historical diversification of floodplain forest specialist species in the Amazon: a case study with two species of the avian genus Xiphorhynchus (Aves: Dendrocolaptidae). Biological Journal of the Linnean Society, 89, 383–395. https://doi.org/10.1111/j.1095-8312.2006.00703.x
Allen, K.E., Greenbaum, E., Hime, P.M., Tapondjou, N.W.P., Sterkhova, V.V., Kusamba, C., Rödel, M.-O., Penner, J., Townsend Peterson, A. & Brown, R.M. (2021) Rivers, not refugia, drove diversification in arboreal sub-Saharan African snakes. Ecology and Evolution, 11, 6133–6152. https://doi.org/10.1002/ece3.7429
Anthony, N.M., Johnson-Bawe, M., Jeffery, K., Clifford, S.L., Abernethy, K.A., Tutin, C.E., Lahm, S.A., White, L.J.T., Utley, J.F., Wickings, E.J. & Bruford, M.W. (2007) The role of Pleistocene refugia and rivers in shaping gorilla genetic diversity in Central Africa. Proceedings of the National Academy of Sciences, 104, 20432–20436. https://doi.org/10.1073/pnas.0704816105Arillo, A. & Ortuño, V.M. (2005) Catalogue of fossil insect species described from Dominican amber (Miocene). Stuttgarter Beiträge zur Naturkunde, Serie B, 352, 1–68.
Ayres, J.M.C. & Clutton-Brock, T.H. (1992) River boundaries and species range size in Amazonian primates. American Naturalist, 140, 531–537. https://doi.org/10.1086/285427
Bates, H.W. (1863) The naturalist on the river Amazons. A record of adventures, habits of animals, sketches of Brazilian and Indian life, and aspects of nature under the Equator, during eleven years of travel. London: John Murray, Albemarle Street.
Bates, J.M., Haffer, J. & Grismer, E. (2004) Avian mitochondrial DNA sequence divergence across a headwater stream of the Rio Tapajós, a major Amazonian River. Journal of Ornithology, 145, 199–205. https://doi.org/10.1007/s10336-004-0039-4
Bezzi, M. (1908a) Diagnoses d’espèces nouvelles de Diptères d’Afrique. Bulletin et Annales de la Société entomologique de belgique, 52, 374–388. [https://www.biodiversitylibrary.org/item/47476]
Bezzi, M. (1908b) Ditteri eritrei raccolti dal Dott. Andreini e dal Prof. Tellini. Parte seconda. Diptera cyclorrhapha. Bollettino Societa Entomologia Italiana, 39 (1907), 3–199.
Bezzi, M. (1914) Ditteri raccolti dal Prof. F. Silvestri durante il suo viaggio in Africa del 1912–13. Bollettino del Laboratorio di zoologia generale e agraria della R. Scuola superiore d’agricoltura, 8, 279–308.
Bigot, J.M.F. (1891) Voyage de M. Ch. Alluaud dans le territoire d’Assinie. 8e mémoire (Afrique occidentale) en juillet et août 1886. Diptères. Annales de la Société entomologique de France, 60, 365–386. [https://www.biodiversitylibrary.org/item/101300#page/371/mode/1up]
Boadi, S., Nsor, C.A., Yakubu, D.H., Acquah, E. & Antobre, O.O. (2017) Conventional and indigenous biodiversity conservation approach: a comparative study of Jachie Sacred Grove and Nkrabea Forest Reserve. International Journal of Forestry Research, 2017, 1–8. https://doi.org/10.1155/2017/1721024
Bossart, J.L. & Antwi, J.B. (2016) Limited erosion of genetic and species diversity from small forest patches: sacred forest groves in an Afrotropical biodiversity hotspot have high conservation value for butterflies. Biological Conservation, 198, 122–134. https://doi.org/10.1016/j.biocon.2016.03.029
Bossart, J.L., Opuni-Frimpong, E., Kuudaar, S. & Nkrumah, E. (2006) Richness, abundance, and complementarity of fruit-feeding butterfly species in relict sacred forests and forest reserves of Ghana. In: Hawksworth, D.L. & Bull, A.T. (Eds), Arthropod diversity and conservation. Topics in biodiversity and conservation. Volume 1. Springer, Dordrecht. https://doi.org/10.1007/978-1-4020-5204-0_20
Cockerell, T.D.A. (1921) An ortalid fly in British amber. Entomologist, 54, 30.
Coetzee, J. (1993) African flora since the terminal Jurassic. New Haven: Yale University Press.
Cumming, J.M. & Wood, D.M. (2017) 3. Adult morphology and terminology. In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds), Manual of Afrotropical Diptera. Volume 1. Introductory chapters and keys to Diptera families. Suricata 4. South African National Biodiversity Institute, Pretoria, pp. 89–133.
Enderlein, G. (1912) Zur Kenntnis orientalischer Ortalinen und Loxoneurinen. Zoologische Jahrbücher, 33, 363–378.
Enderlein, G. (1924) Beiträge zur Kenntnis der Platystominen. Mitteilungen aus dem Zoologischen Museum in Berlin, 11, 99–153.
Ernst, M., Rödel, M.O. & Blom, M.P.K. (2025) Towards a comprehensive view on evolutionary refugia in West African rainforests. Frontiers of Biogeography, 18, e139537. https://doi.org/10.21425/fob.18.139537
Evenhuis, N.L. (1997) Family Platystomatidae. Catalog of the fossil flies of the world (Insecta: Diptera) website. Available from: https://hbs.bishopmuseum.org/fossilcat/fossplatystom.html (Version 2.0, 17 February 1997) (accessed 8 June 2025).
Fabricius, J.C. (1794) Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species adjectis synonimis, locis, obervationibus, descriptionibus. C.G. Proft, Hafniae [= Copenhagen], 6 pp. https://doi.org/https://archive.org/details/entomologiasy-st04fabr/page/n3/mode/2up
Fabricius, J.C. (1805) Systema antliatorum secundum ordines, genera, species, adiectis synonymis, locis, observationibus, descriptionibus. Reichard, Brunsvigae. [https://www.biodiversitylibrary.org/item/53432#page/9/mode/1up]
Fallén, C.F. (1810) Specim. entomolog. novam Diptera disponendi methodum exhibens. Berlingianis, Lund. [https://www.biodiversitylibrary.org/item/188591#page/1/mode/1up]
Gascon, C., Malcolm, J.R., Patton, J.L., da Silva, M.N.F., Bogart, J.P., Lougheed, S.C., Peres, C.A., Neckel, S. & Boag, P.T. (2000) Riverine barriers and the geographic distribution of Amazonian species. Proceedings of the National Academy of Sciences of the United States of America, 97, 13672–13677. [https://www.pnas.org/doi/pdf/10.1073/pnas.230136397]
Goloboff, P.A. & Morales, M.E. (2023) TNT version 1.6, with a graphical interface for MacOS and Linux, including new routines in parallel. Cladistics, 39, 144–153. https://doi.org/10.1111/cla.12524
Haffer, J.R. (1969) Speciation in Amazonian forest birds: most species probably originated in forest refuges during dry climatic periods. Science, 165, 131–137. [https://www.science.org/doi/abs/10.1126/science.165.3889.131]
Haffer, J.R. (1974) Avian speciation in tropical South America. Publications of the Nuttall Ornithological Club, 14, 1–390. https://doi.org/10.1080/01650529009360816
Haffer, J.R. (1982) General aspects of the refuge theory. In: Prance, G.T., (Ed,), Biological diversification in the tropics. Columbia University Press, pp. 6–24.
Haffer, J.R. (2008) Hypotheses to explain the origin of species in Amazonia. Brazilian Journal of Biology, 68, 917–947. https://doi.org/10.1590/s1519-69842008000500003
Hall, J.B. & Swain, M.D. (1981) Distribution and ecology of vascular plants in a tropical rainforest: forest vegetation of Ghana. W. Junk Publishers, The Hague.
Hall, J.P.W. & Harvey, D.J. (2002) The phylogeography of Amazonia revisited: new evidence from riodinid butterflies. Evolution, 56, 1489–1497. https://doi.org/10.1111/j.0014-3820.2002.tb01460.x
Hawthorne, W.D. (1988) The flora and vegetation of Ghana’s forests. In: Ghana forest inventory proceedings. Overseas Development Agency/Ghana Forestry Dept, Accra, pp. 8–13.
Hawthorne, W.D. & Abu-Juam, M. (1995) Forest protection in Ghana with particular reference to vegetation and plant species. IUCN, Gland, Switzerland.
Hendel, F. (1914a) Diptera. Fam. Muscaridae, Subfam. Platystominae. Genera Insectorum, 157, 1–179 + 15 plates. [https://www.biodiversitylibrary.org/item/223809#page/165/mode/1up]
Hendel, F. (1914b) Die Arten der Platystominen. Abhandlungen der Kaiserlich-Königlichen Zoologisch-botanischen Gesellschaft in Wien, 8, 1–409 + 4 plates. [https://www.biodiversitylibrary.org/item/28240#page/31/mode/1up]
Hennig, W. (1965) Die Acalyptratae des Baltischen Bernsteins und ihre Bedeutung für die Erforschung der phylogenetischen Entwicklung dieser Dipteren-Gruppe. Stuttgarter Beiträge zur Naturkunde, 145, 1–215. [https://www.biodiversitylibrary.org/part/241980]
Hugueny, B. & Lèvêque, C. (1994) Freshwater fish zoogeography in West Africa: faunal similarities between river basins. Environmental Biology of Fishes, 39, 365–380.
International Commission on Zoological Nomenclature (1925) Opinion 82. Suspension of rules for Musca Linnaeus, 1758A, type of M. domestica. Smithsonian miscellaneous collections, 73, 1–7. [https://www.biodiversitylibrary.org/page/8911029]
Jacobs, B.F. (2004) Palaeobotanical studies from tropical Africa: relevance to the evolution of forest, woodland and savannah biomes. Philosophical Transactions of the Royal Society of London. B. Biological Sciences, 359, 1573–83. https://doi.org/10.1098/rstb.2004.1533
Kennis, J., Nicolas, V., Hulselmans, J., Katuala, P.G.B., Wendelen, W., Verheyen, E., Dudu, A.M. & Leirs, H. (2011) The impact of the Congo River and its tributaries on the rodent genus Praomys: speciation origin or range expansion limit? Zoological Journal of the Linnean Society, 163, 983–1002. https://doi.org/10.1111/j.1096-3642.2011.00733.x
Kirk-Spriggs, A.H. (2017) 1. Introduction and brief history of Afrotropical dipterology. In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds), Manual of Afrotropical Diptera. Volume 1. Introductory chapters and keys to Diptera families. Suricata 4. Pretoria: South African National Biodiversity Institute, pp. 1–67.
Kirk-Spriggs, A.H. & Muller, B.S. (2017) 9. Biogeography of Diptera. In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds), Manual of Afrotropical Diptera. Volume 1. Introductory chapters and keys to Diptera families. Suricata 4. Pretoria: South African National Biodiversity Institute, pp. 203–238.
Lampert, K.P., Rand, A.S., Mueller, U.G. & Ryan, M.J. (2003) Fine-scale genetic pattern and evidence for sex-biased dispersal in the tungara frog, Physalaemus pustulosus. Molecular Ecology, 12, 3325–3334. https://doi.org/10.1046/j.1365-294X.2003.02016.x
Larsen, T.B. (2006) The Ghana butterfly fauna and its contribution to the objectives of the protected areas system. Wildlife Division Support Project (WDSP) Report No. 63. [https://abdb-africa.org/library/bibliography/2006%20Ghana%20Report%20complete%201.5MB.pdf]
László, G.M. & Volynkin, A.V. (2023) Review of the genus Neomardara Hering, 1926 with descriptions of two new species and updated information on the distribution of the genus (Lepidoptera: Erebidae: Lymantriinae). Ecologica Montenegrina, 68, 103–126. https://doi.org/10.37828/em.2023.68.11
Lebbie, A.R. & Freudenberger, M.S. (1996) Sacred groves in Africa: forest patches in transition. In: Schelhas, J. & Greenberg, R. (Eds), Forest patches in tropical landscapes. Island Press, Washington, pp. 300–326.
Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum caracteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. L. Salvii, Holmiae [= Stockholm]. [https://www.biodiversitylibrary.org/item/10277#page/1/mode/1up]
Loew, H. (1873) Monographs of the Diptera of North America. Part III. Smithsonian Miscellaneous Collections, 11 (No. 256), vii + 1–351 + xiii + 4 plates. [https://repository.si.edu/handle/10088/23107]
Maley, J. (1996) The African rainforest – main characteristics of changes in vegetation and climate from the Upper Cretaceous to the Quaternary. Proceedings of the Royal Society of Edinburgh, 104b, 31–73. https://doi.org/10.1017/S0269727000006114
Massa, B. (2023) Biodiversity hotspot in the Sangha Trinational Protected Area revealed through Orthoptera Tettigoniidae. Zootaxa, 5331, 1–67. https://doi.org/10.11646/zootaxa.5331.1.1
Mayr, E. (1942) Systematics and the origin of species from the viewpoint of a zoologist. Columbia University Press, New York, NY.
Mayr, E. & O’Hara, R.J. (1986) The biogeographic evidence supporting the Pleistocene forest refuge hypothesis. Evolution, 40, 55–67. https://doi.org/10.2307/2408603
McAlpine, D.K. (1994) Review of the species of Achias (Diptera: Platystomatidae). Invertebrate Taxonomy, 8, 117–281.
Meigen, J.W. (1803) Versuch einer neuen Gattungs-Eintheilung der europäischen zweiflügligen Insekten. Magazin für Insektenkunde, 2, 259–281. [https://www.biodiversitylibrary.org/item/137736#page/4/mode/1up]
Meunier, F. (1908) Sur quelques diptères (Muscinae, Ortalinae, Helomyzinae) du copal récent de Zanzibar. Annales de la Société scientifique de Bruxelles, 32, 252–257. [https://www.biodiversitylibrary.org/part/198315]
Micheels, A., Eronen, J. & Mosbrugger, V. (2009) The Late Miocene climate response to a modern Sahara Desert. Global and Planetary Change, 67, 193–204. https://doi.org/10.1016/j.gloplacha.2009.02.005
Moretto, P., Cosson, B., Takano, H., Basquin, P., Bordat, P., Boucher, S., Bouyer, T., Danflous, S., Dérozier, V., Eitscheberger, U. & Juhel, P. (2021) Un refuge forestier menacé: la forêt d’altitude à Parinari du Mont Tonkoui en Côte d’Ivoire: évaluation de la biodiversité entomologique, botanique et ornithologique, valeurs de conservation du site. Catharsius La Revue, Hors-série, 1, 1–123.
Morley, R.J. & Richards, K. (1993) Graminae cuticle – a key indicator of late Cenozoic climatic-change in the Niger Delta. Review of Palaeobotany and Palynology, 77, 119–127. [https://www.sciencedirect.com/science/article/abs/pii/0034666793900608]
Murienne, J., Benavides, L.R., Prendini, L., Hormiga, G. & Giribet, G. (2013) Forest refugia in Western and Central Africa as ‘museums’ of Mesozoic biodiversity. Biology Letters, 23, 9(1): 20120932. https://doi.org/10.1098/rsbl.2012.0932
Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A.B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853–858. https://doi.org/10.1038/35002501
NASA (2023) IMERG grand average climatology (11 km resolution): Average annual precipitation (mm/year) June 2000–May 2023 [Spatial data]. NASA. Available from: https://gpm.nasa.gov/data/imerg/precipitation-climatology (accessed 18 June 2025).
Nicolas, V., Missoup, A.D., Denys, C., Peterhans, J.K., Katuala, P., Couloux, A. & Colyn, M. (2011) The roles of rivers and Pleistocene refugia in shaping genetic diversity in Praomys misonnei in tropical Africa. Journal of Biogeography, 38, 191–207. https://doi.org/10.1111/j.1365-2699.2010.02399.x
Ntiamoa-Baidu, T. (2001) Indigenous versus introduced biodiversity conservation strategies: the case of protected area systems in Ghana. In: Weber, W., White, L.J.T., Vedder, A. & Naughton-Treves, L. (Eds), African rain forest ecology and conservation. Yale University Press, New Haven & London, pp. 385–394.
Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D’amico, J.A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H., Kura, Y., Lamoreux, J.F., Wettengel, W.W., Hedao, P. & Kassem, K.R. (2001) Terrestrial ecoregions of the world: a new map of life on Earth: a new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. BioScience, 51, 933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
Patton, J.L. & Silva, M.F. (2005) The history of Amazonian mammals: mechanisms and timing of diversification. In: Bermingham, E., Dick, C.W. & Moritz, C. (Eds,), Tropical rainforests: past, present and future. The University of Chicago Press, pp. 107–126.
Pellegrino, K.C.M., Rodrigues, M.I., Waite, A.N., Morando, M., Yassuda, Y.Y. & Sites, J.W. (2005) Phylogeography and species limits in the Gymnodactylus darwinii complex (Gekkonidae, Squamata): genetic structure coincides with river systems in the Brazilian Atlantic Forest. Biological Journal of the Linnean Society, 85, 13–26. https://doi.org/10.1111/j.1095-8312.2005.00472.x
Plana, V. (2004) Mechanisms and tempo of evolution in the African Guineo-Congolian rainforest. Philosophical Transactions of the Royal Society of London. B. Biological Sciences, 359, 1585–1594. https://doi.org/10.1098/rstb.2004.1535
Pont, A.C., Chainey, J.E. & Evenhuis, N.L. (2024) The types of Diptera (Insecta) described by J.-M.-F. Bigot. Privately published, Hilo, Hawaiʽi, ii–x + 1–658 pp. [https://archive.org/details/bigot-diptera-types/page/n13/mode/2up?view=theatre]
Prance, G.T. (1982) Biological diversification in the tropics: Proceedings of the Fifth International Symposium of the Association for Tropical Biology, held at Macuto Beach, Caracas, Venezuela, February 8–13, 1979. Columbia University Press, New York, NY.
Querouil, S., Verheyen, E., Dillen, M. & Colyn, M. (2003) Patterns of diversification in two African forest shrews: Sylvisorex johnstoni and Sylvisorex ollula (Soricidae, Insectivora) in relation to paleo-environmental changes. Molecular Phylogenetics and Evolution, 28, 24–37. [https://www.sciencedirect.com/science/article/abs/pii/S1055790303000277]
Ribas, C.C., Aleixo, A., Nogueira, A.C.R., Miyaki, C.Y. & Cracraft, J. (2012) A palaeobiogeographic model for biotic diversification within Amazonia over the past three million years. Proceedings of the Royal Society B, 279, 681–689. https://doi.org/10.1098/RSPB.2011.1120
Richardson, J.E., Pennington, R.T., Pennington, T.D. & Hollingsworth, P.M. (2001) Rapid diversification of a species-rich genus of Neotropical rain forest trees. Science, 293, 2242–2245. https://doi.org/10.1126/science.1061421
Runge, J. (2022) Chapter 15. The Congo River, Central Africa. In: Gupta, A. (Ed.) Large rivers. Geomorphology and management. Second Edition. John Wiley & Sons, pp. 433–456. https://doi.org/10.1002/9781119412632.ch15
Sæther, O.A. & Ekrem, T. (2023) Biogeography of Afrotropical Chironomidae (Diptera) with special reference to Gondwanaland. Cimbebasia, 19, 123–139.
Senut, B., Pickford, M. & Segalen, L. (2009) Neogene desertification of Africa. Comptes Rendus Geoscience, 341, 591–602. https://doi.org/10.1016/j.crte.2009.03.008
Sick, H. (1967) Rios e enchentes na Amazônia como obstáculo para a avifauna. Atas do Simpósio Sobre a Biota Amazônica, 5, 495–520.
Sinthumule, N.I. (2024) Sacred forests as repositories of local biodiversity in Africa: a systematic review. Forest Science and Technology, 20, 337–348. https://doi.org/10.1080/21580103.2024.2397522
Speiser, P. (1910) 5. Cyclorhapha, Aschiza. In: Sjöstedt, Y. (Ed.) Wissenschaftliche Ergebnisse der schwedischen zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen, Deutsch-Ostafrikas 1905–1906 unter Leitung von Prof. D. Yngve Sjöstedt. P. 2. Band. [Abteilung] 10. P. Palmquist, Stockholm, pp. 113–198.
Speiser, P. (1911) Zur Kenntnis aussereuropäischer Dipteren. Jahrbücher des Nassauischen Vereins für Naturkunde, 64, 237–261. [https://archive.org/details/jahrbcherdesna64nass]
Steyskal, G.C. (1965a) Records and descriptions of African Otitidae and Platystomatidae (Diptera). Annals of the Natal Museum, 18, 171–178. [https://archive.org/details/aja03040798_767]
Steyskal, G.C. (1965b) Family Sciomyzidae (Tetanoceridae). In: Stone, A., Sabrosky, C.W., Wirth, W.W., Foote, R.H. & Coulson, J.R., A catalog of the Diptera of America north of Mexico. United States Department of Agriculture, Agricultural Handbook 276, pp. 685–695.
Steyskal, G.C. (1980) 43. Family Platystomatidae. In: Crosskey, R.W. (Ed.) Catalogue of the Diptera of the Afrotropical Region. British Museum (Natural History) London, pp. 563–574.
Taberer, T.R. (2024) A new species of Delorhachis Karsch, 1896 from Nouabalé-Ndoki National Park, Republic of Congo. Journal of Natural History, 58, 639–644. https://doi.org/10.1080/00222933.2024.2354483
Takano, H. (2024) Descriptions of three new species of Lycaenidae (Lepidoptera: Papilionoidea) from Parc National de Nouabalé-Ndoki, Republic of Congo with additional notes on little-known species. Zootaxa, 5496, 214–230. https://doi.org/10.11646/zootaxa.5496.2.4
Takano, H. & László, G.M. (2024) Descriptions of new Hypotrabala Holland, 1893 (Lepidoptera: Lasiocampidae: Lasiocampinae: Selenepherini) in the collections of the African Natural History Research Trust, with notes on allied genera and the description of a new genus. Ecologica Montenegrina, 72, 19–63. https://doi.org/10.37828/em.2024.72.4
Telfer, P.T., Souquière, S., Clifford, S.L., Abernethy, K.A., Bruford, M.W., Disotell, T.R., Sterner, K.N., Roques, P., Marx, P.A. & Wickings, E.J. (2003) Molecular evidence for deep phylogenetic divergence in Mandrillus sphinx. Molecular Ecology, 12, 2019–2024. https://doi.org/10.1046/j.1365-294x.2003.01877.x
UNEP-WCMC & IUCN (2025) Protected Planet: The World Database on Protected Areas (WDPA) and World Database on Other Effective Area-based Conservation Measures (WD-OECM) [Spatial data]. UNEP-WCMC and IUCN. Available from: www.protectedplanet.net (accessed 18 June 2025)
UNESCO (2003) The importance of sacred natural sites for biodiversity conservation. In: Lee, K. & Schaaf, T. (Eds), Proceedings of the international workshop, UNESCO (Division of Ecological Sciences) Paris.
USDA (n.d.) World 30m DEM [Spatial data]. USDA Foreign Agricultural Service. Available from: https://geo.fas.usda.gov/arcgis1/services/G_Elevation/WorldElevation_30m/ImageServer (accessed 18 June 2025)
Vallinoto, M., Araripe, J., do Rego, P.S., Tagliaro, C.H., Sampaio, I. & Schneider, H. (2006) Tocantins River as an effective barrier to gene flow in Saguinus niger populations. Genetic and Molecular Biology, 29, 215–219. https://doi.org/10.1590/S1415-47572006000200005
Vanzolini, P.E. (1973) Paleoclimates, relief, and species multiplication in equatorial forests. In: Meggers, B.J., Ayensu, E.S. & Duckworth, W.D. (Eds), Tropical forest ecosystems in Africa and South America: a comparative review. Smithsonian Press, Washington D.C., pp. 255–258.
Vanzolini, P.E. & Williams, E.E. (1970) South American anoles: geographic differentiation and evolution of the Anolis chrysolepis species group (Sauria, Iguanidae). Arquivos de Zoologia, 19, 291–298. https://doi.org/10.11606/issn.2176-7793.v19i3-4p125-298
Voelker, G., Marks, B.D., Kahindo, C., A’genonga, U., Bapeamoni, F., Duffie, L.E., Huntley, J.W., Mulotwa, E., Rosenbaum, S.A. & Light, J.E. (2013) River barriers and cryptic biodiversity in an evolutionary museum. Ecology and Evolution, 3, 536–545. https://doi.org/10.1002/ece3.482
Walker, F. (1853) Diptera (part IV). In: Saunders, W.W. (Ed.), Insecta saundersiana: or characters of undescribed insects in the collection of William Wilson Saunders, Esq., F.R.S., F.L.S., &c. 1. Van Voorst, London, pp. 253–474 + plates 7–8. [https://www.biodiversitylibrary.org/bibliography/5112]
Wallace, A.R. (1852) On the monkeys of the Amazon. Proceedings of the Zoological Society of London, 20, 107–110. [https://digitalcommons.wku.edu/dlps_fac_arw/3/]
Weir, J.T. (2006) Divergent timing and patterns of species accumulation in lowland and highland Neotropical birds. Evolution, 60, 842–855. https://doi.org/10.1111/j.0014-3820.2006.tb01161.x
Westwood (1840) Synopsis of the genera of British insects as appendix of Westwood: an introduction to the modern classification of insects. Longman, Orme, Brown, Green & Longmans, London.
Whittington, A.E. & Kirk-Spriggs, A.H. (2021) 70. Platystomatidae (signal flies). In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds), Manual of Afrotropical Diptera. Volume 3. Brachycera–Cyclorrhapha, excluding Calyptratae. Suricata 8. Pretoria: South African National Biodiversity Institute, pp. 1619–1667.
Wiedemann, C.R.W. (1830) Aussereuropäische zweiflügelige Insekten. Als Fortsetzung des Meigenschen Werks. Zweiter Theil. Schulz, Hamm. [https://www.biodiversitylibrary.org/item/88495]
Wiley, E.O. (1981) Phylogenetics: the theory and practice of phylogenetic systematics. New York: Wiley.
WWF & The Nature Conservancy. (2001) Terrestrial Ecoregions of the World (TEOW) v2 [Shapefile]. World Wildlife Fund. Available from: https://www.worldwildlife.org/publications/terrestrial-ecoregions-of-the-world (accessed 18 June 2025).
Zachos, J.C., Dickens, G.R. & Zeebe, R.E. (2008) An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics. Nature, 451, 279–283. https://doi.org/10.1038/nature06588
Zachos, J., Pagani, M., Sloan, L., Thomas, E. & Billups, K. (2001) Trends, rhythms, and aberrations in global climate 65 Ma to present. Science, 292, 686–693. [https://www.science.org/doi/10.1126/science.1059412]
Zhang, Z., Nisancioglu, K.H., Flatoy, F., Bentsen, M., Bethke, I. & Wang, H. (2011) Tropical seaways played a more important role than high latitude seaways in Cenozoic cooling. Climate of the Past, 7, 801–813. https://doi.org/10.5194/cp-7-801-2011