An integrative revision of the genus Isognathotermes (Termitidae: Cubitermitinae) with description of seven new species and four new subspecies
Abstract
Cubitermitinae species whose workers have enteric valves of the fungifaber and finitimus patterns were previously gathered into the genus Isognathotermes. Here, we revise this genus, combining the morphological and anatomical features of all castes, as well as the species’ chorological, ecological, and phylogenetic (mitogenome) characteristics in an integrative taxonomic approach. In addition to the genetic signature, the best morphological criteria for generic recognition are found in the workers’ EVA (enteric valve architecture) and the spinosity of the fore coxae of the workers, and the best criteria for specific recognition are found in (a) the valve and caecum of the workers, (b) the caecum, dimensions of the head, and curvature of the mandibles of the soldiers, and (c) the dimensions of the head and the relative dimensions of eyes and ocelli of the imagines. After revision, only eight of the 22 known taxa remain valid, 14 known taxa become junior synonyms, and one is considered as incertae sedis; six new morphologically recognizable species, one cryptic species and four new subspecies are described. The mitogenome of 12 out of the 15 valid species have been sequenced. The following described species are considered valid: I. bulbifrons (Sjöstedt, 1924), I. finitimus (Schmitz, 1916), I. fungifaber (Sjöstedt, 1896), I. minitabundus (Sjöstedt, 1913), I. planifrons (Sjöstedt, 1924), I. severus (Silvestri, 1914), I. ugandensis (Fuller, 1923) and I. zenkeri (Desneux, 1904). The following species are described as new: I. acristatus Josens & Deligne sp. nov., I. modicus Josens & Deligne sp. nov., I. magniplanifrons Josens & Deligne sp. nov., I. phallicaecalis Josens & Deligne sp. nov., I. phalloides Josens & Deligne sp. nov., I. rectimalatus Josens & Deligne sp. nov., and I. similifinitimus Josens & Deligne sp. nov. Four new subspecies of I. ugandensis are recognized: I. ugandensis burundii Josens & Deligne subsp. nov., I. ugandensis kenyae Josens & Deligne subsp. nov., I. ugandensis malawii Josens & Deligne subsp. nov., and I. ugandensis ugandensis Josens & Deligne subsp. nov. The following species are considered junior synonyms: I. antennalis (Sjöstedt, 1924), I. banksi (Emerson, 1928), I. comstocki (Emerson, 1928), I. fungifaber var. elongata (Sjöstedt, 1924), I. gaigei (Emerson, 1928), I. gibbifrons (Sjöstedt, 1924), I. heghi (Sjöstedt, 1924), I. kemneri (Emerson, 1928), I. loubetsiensis (Sjöstedt, 1924), I. modestior (Silvestri, 1914), I. schmidti (Emerson, 1928), I. silvestrii (Sjöstedt, 1925), I. speciosus (Sjöstedt, 1924), and I. subarquatus (Sjöstedt, 1926). Are considered as incertae sedis: I. bredoi (nomen nudum), I. congoensis (Emerson, 1928), an “I. aff. planifrons” (undescribed), and an unknown Isognathotermes from Malawi (undescribed). GenBank accessions: see Suppl. file 3 and ‘Material examined’ of each chapter.
References
Allio R., Schomaker-Bastos A., Romiguier J., Prosdocimi F., Nabholz B. & Delsuc F. 2020. MitoFinder: efficient automated large-scale extraction of mitogenomic data in target enrichment phylogenomics. Molecular Ecology Resources 20: 892–905. https://doi.org/10.1111/1755-0998.13160
Arora J., Buček A., Hellemans S., Beránková T., Romero Arias J., Fisher B.L., Clitheroe C. Brune A., Kinjo Y., Šobotník J. & Bourguignon T. 2023. Evidence of cospeciation between termites and their gut bacteria on a geological time scale. Proceedings of the Royal Society B 290: e20230619. https://doi.org/10.1098/rspb.2023.0619
Bouillon A. & Vincke P.P. 1971. Valvule entérique et révision du genre Cubitermes Wasmann. Cubitermes bugeserae sp. nov. (Isoptera, Termitidae). Revue de Zoologie et de Botanique africaines 84 (3–4): 269–280.
Bouillon A. & Vincke P.P. 1973. Hybridisme et génétique expérimentale chez les Termites (Isoptera). Revue de Zoologie et de Botanique africaines 87 (1): 31–50.
Bourguignon T., Lo N., Cameron S.L., Šobotník J., Hayashi Y., Shigenobu S., Watanabe D., Roisin Y., Miura T. & Evans T.A. 2015. The evolutionary history of termites as inferred from 66 mitochondrial genomes. Molecular Biology and Evolution 32: 406–421. https://doi.org/10.1093/molbev/msu308
Bourguignon T., Lo N., Šobotník J., Sillam-Dussès D., Roisin Y. & Evans T.A. 2016. Oceanic dispersal, vicariance and human introduction shaped the modern distribution of the termites Reticulitermes, Heterotermes and Coptotermes. Proceedings of the Royal Society B 283: e20160179. https://doi.org/10.1098/rspb.2016.0179
Bourguignon T., Lo N., Šobotnik J., Ho S.Y.W., Iqbal N., Coissac E., Lee M., Jendryka M.M., Sillam-Dusses D., Krizkova B., Roisin Y & Evans T. 2017. Mitochondrial phylogenomics resolves the global spread of higher termites, ecosystem engineers of the tropics. Molecular Biology and Evolution 34 (3): 589–597. https://doi.org/10.1093/molbev/msw253
Burgess N.D., D’Amico Hales J., Rickettsa T.H. & Dinerstein E. 2006. Factoring species, non-species values and threats into biodiversity prioritisation across the ecoregions of Africa and its islands. Biological Conservation 127: 383–401. https://doi.org/10.1016/j.biocon.2005.08.018
Cameron S.L., Lo N., Bourguignon T., Svenson G.J. & Evans T.A. 2012. A mitochondrial genome phylogeny of termites (Blattodea: Termitoidae): robust support for interfamilial relationships and molecular synapomorphies define major clades. Molecular Phylogenetics and Evolution 65: 163–173. https://doi.org/10.1016/j.ympev.2012.05.034
Chen S., Zhou Y., Chen Y. & Gu J. 2018. Fastp: an ultra-fast all-in-one FASTQ preprocessor. Bioinformatics 34: i884–i890. https://doi.org/10.1093/bioinformatics/bty560
Chernomor O., Von Haeseler A. & Minh B.Q. (2016). Terrace aware data structure for phylogenomic inference from supermatrices. Systematic Biology 65: 997–1008. https://doi.org/10.1093/sysbio/syw037
Chessel D., Dufour A.B. & Thioulouse J. 2004. The ade4 package-I: One-table methods. R News 4: 5–10.
Desneux J. 1904a. Notes termitologiques. Annales de la Société entomologique de Belgique 48 (3): 146–151.
Desneux J. 1904b. Isoptera, fam. Termitidae. In: Wytsman P. (ed.) Genera Insectorum. 25: 1–52.
Dowton M., Meiklejohn K., Cameron S.L. & Wallman J. 2014. A preliminary framework for DNA barcoding, incorporating the multispecies coalescent. Systematic Biology 63: 639–644. https://doi.org/10.1093/sysbio/syu028
Emerson A.E. 1928. Termites of the Belgian Congo and the Cameroon. Bulletin of the American Museum of Natural History 57 (7): 401–574.
Emerson A.E. 1938. Termite nests: a study of the phylogeny of behavior. Ecological Monographs 8: 247–284. https://doi.org/10.2307/1943251
Engel M.S., Grimaldi D.A. & Krishna K. 2009. Termites (Isoptera): their phylogeny, classification, and rise to ecological dominance. American Museum Novitates 3650: 1–27. https://doi.org/10.1206/651.1
Florencio D.F., Marins A., Rosa C.S., Cristaldo P.F., Araújo A.P.A., Silva I.R. & DeSouza O. 2013. Diet segregation between cohabiting builder and inquiline termite species. PLoS ONE 8 (6): e66535. https://doi.org/10.1371/journal.pone.0066535
Fuller C. 1923. Two new termites from Uganda. Annals and Magazine of Natural History (9) 11: 191–192. https://doi.org/10.1080/00222932308632837
Galtier N. 2019. Delineating species in the speciation continuum: A proposal. Evolutionary Applications 12 (4): 657–663. https://doi.org/10.1111/eva.12748
Gilbert M.T.P., Moore W., Melchior L. & Worebey M. 2007. DNA extraction from dry museum beetles without conferring external morphological damage. PLoS ONE 2: 1–4. https://doi.org/10.1371/journal.pone.0000272
Guindon S., Dufayard J.F., Lefort V., Anisimova M., Hordijk W. & Gascuel O. 2010. New algorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3.0. Systematic Biology 59: 307–321. https://doi.org/10.1093/sysbio/syq010
Han S.H. & Lepage M. 1991. Croissance des sociétés de Cubitermes fungifaber (Isoptera, Termitidae) en relation avec les dimensions de leurs nids. Annales des Sciences Naturelles, Zoologie et Biologie animale 12: 49–56.
Hardin G. 1960. The Competitive Exclusion Principle: An idea that took a century to be born has implications in ecology, economics and genetics. Science 131: 1292–1297. https://doi.org/10.1126/science.131.3409.1292
Harris W.V. 1958. Isoptera. In: De Witte G.F. (ed.) Exploration du Parc National de l’Upemba. I. Mission G.F. de Witte, en collaboration avec W. Adam, A. Janssens, L. van Meel et R. Verheyen (1946–1949) 52: 3–26. Institut des Parcs Nationaux du Congo Belge, Brussels.
Hellemans S., Marynowska M., Drouet T., Lepoint G., Fournier D., Calusinska M. & Roisin Y. 2019. Nest composition, stable isotope ratios and microbiota unravel the feeding behaviour of an inquiline termite. Oecologia 191: 541–553. https://doi.org/10.1007/s00442-019-04514-w
Hellemans S., Deligne J., Roisin Y. & Josens G. 2021. Phylogeny and revision of the ‘Cubitermes complex’ termites (Termitidae: Cubitermitinae). Systematic Entomology 46: 224–238. https://doi.org/10.1111/syen.12458
Hellemans S., Šobotník J., Lepoint G., Mihaljevič M., Roisin Y. & Bourguignon T. 2022a. Termite dispersal is influenced by their diet. Proceedings of the Royal Society B 289: e20220246. https://doi.org/10.1098/rspb.2022.0246
Hellemans S., Wang M., Hasegawa N., Šobotník J., Scheffrahn R.H. & Bourguignon T. 2022b. Using ultraconserved elements to reconstruct the termite tree of life. Molecular Phylogenetics and Evolution 173: e107520. https://doi.org/10.1016/j.ympev.2022.107520
Hoang D.T., Chernomor O., von Haeseler A., Minh B.Q. & Vinh L.S. 2018. UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35: 518–522. https://doi.org/10.1093/molbev/msx281
Holmgren N. 1912. Termitenstudien. 3. Systematik der Termiten. Die Familie Metatermitidae. Kungliga Svenska Vetenskapsakademiens Handlingar 48 (4): 1–166.
Johnson R.A. 1979. Configuration of the digestive tube as an aid to identification of worker Termitidae (Isoptera). Systematic Entomology 4: 31–38. https://doi.org/10.1111/j.1365-3113.1979.tb00609.x
Josens G. & Deligne J. 2019. Species groups in the genus Cubitermes (Isoptera: Termitidae) defined on the basis of enteric valve morphology. European Journal of Taxonomy 515: 1–72. https://doi.org/10.5852/ejt.2019.515
Josens G. & Makatia Wango S.P. 2019. Niche differentiation between two sympatric Cubitermes species (Isoptera, Termitidae, Cubitermitinae) revealed by stable C and N isotopes. Insects 10 (2): 38: 1–5. https://doi.org/10.3390/insects10020038
Kalyaanamoorthy S., Minh B.Q., Wong T.K.F., Von Haeseler A. & Jermiin L. S. 2017. ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods 14: 587–589. https://doi.org/10.1038/nmeth.4285
Katoh K. & Standley D.M. 2013. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30: 772–780. https://doi.org/10.1093/molbev/mst010
Klimov P.B., Skoracki M. & Bochkov A.V. 2019. Cox1 barcoding versus multilocus species delimitation: Validation of two mite species with contrasting effective population sizes. Parasites and Vectors 12: 8. https://doi.org/10.1186/s13071-018-3242-5
Krishna K., Grimaldi D.A., Krishna V. & Engel M.S. 2013. Treatise on the Isoptera of the World. Bulletin of the American Museum of Natural History 377 (1–7): 1–2704. https://doi.org/10.1206/377.1
Kück P. & Longo G. C. 2014. FASconCAT-G: extensive functions for multiple sequence alignment preparations concerning phylogenetic studies. Frontiers in Zoology 11: 81. https://doi.org/10.1186/s12983-014-0081-x
Lowry D.B. & Gould B.A. 2016. Speciation Continuum. In: Kliman R.M. (ed.) Encyclopaedia of Evolutionary Biology 4: 159–165. Elsevier. https://doi.org/10.1016/B978-0-12-800049-6.00080-9
Mathot G. 1964. Description d’une nouvelle espèce de Cubitermes du Congo (Isoptera, Termitinae). Cubitermes exiguus n. sp. In: Bouillon A. (ed.) Études sur les termites africains, un colloque international, Université Lovanium: 15–21. Édition de l’Université Léopoldville [= Kinshasa].
Mathur R.N. & Thapa R.S. 1962. A revised catalogue of Isoptera (white ants) of the Entomological Reference Collection at the Forest Research Institute. Dehra Dun. Indian Forest Leaflet (Entomology) 167: 1–122.
Minh B.Q., Schmidt H.A., Chernomor O., Schrempf D., Woodhams M.D., Von Haeseler A., Lanfear R. & Teeling E. 2020. IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution 37: 1530–1534. https://doi.org/10.1093/molbev/msaa015
Miura T., Roisin Y. & Matsumoto T. 2000. Molecular phylogeny and biogeography of the nasute termite genus Nasutitermes (Isoptera: Termitidae) in the Pacific tropics. Molecular Phylogenetics and Evolution 17: 1–10. https://doi.org/10.1006/mpev.2000.0790
Nishimaki T. & Sato K. 2019. An extension of the Kimura two-parameter model to the natural evolutionary process. Journal of Molecular Evolution 87: 60–67.https://doi.org/10.1007/s00239-018-9885-1
Noirot C. 1955. Recherches sur le polymorphisme des termites supérieurs (Termitidae). Annales des Sciences Naturelles, Zoologie (11) 17: 399–595.
Noirot C. & Kovoor J. 1958. Anatomie comparée du tube digestif des termites. 1 Sous-famille des “Termitinae”. Insectes sociaux 5: 439–471. https://doi.org/10.1007/BF02226857
Noirot C., Noirot-Thimothée C. & Han S.H. 1986. Migration and nest building in Cubitermes fungifaber (Isoptera, Termitidae). Insectes sociaux 33 (4): 361–374. https://doi.org/10.1007/BF02223945
Nurk S., Meleshko D., Korobeynikov A. & Pevzner P.A. 2017. metaSPAdes: a new versatile metagenomic assembler. Genome Research 27: 824–834. https://doi.org/10.1101/gr.213959.116
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.L., 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. BioScience 51 (11): 933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
Padial J.M., Miralles A., De la Riva I. & Vences M. 2010. The integrative future of taxonomy. Frontiers in Zoology 7: 1–14. https://doi.org/10.1186/1742-9994-7-16
Perdereau E., Dedeine F., Christidès J.-P., Dupont S. & Bagnères A.-G. 2011. Competition between invasive and indigenous species: an insular case study of subterranean termites. Biological Invasions 13: 1457–1470. https://doi.org/10.1007/s10530-010-9906-5
R Core Team 2018. R: a Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. Available from https://www.R-project.org/ [accessed 15 Sep. 2025].
Rice P., Longden L. & Bleasby A. 2000. EMBOSS: the European Molecular Biology Open Software Suite. Trends in Genetics 16: 276–277. https://doi.org/10.1016/S0168-9525(00)02024-2
Romero Arias J., Hellemans S., Kaymak E., Akama P.D., Bourguignon T., Roisin Y., Scheffrahn R.H. & Šobotník J. 2024. Mitochondrial phylogenetics position a new Afrotropical termite species into its own subfamily, the Engelitermitinae (Blattodea: Termitidae). Systematic Entomology 49: 72–83.https://doi.org/10.1111/syen.12607
Roy V., Demanche C., Livet A. & Harry M. 2006. Genetic differentiation in the soil-feeding termite Cubitermes sp. affinis subarquatus: occurrence of cryptic species revealed by nuclear and mitochondrial markers. BMC Evolutionary Biology 6: 102. https://doi.org/10.1186/1471-2148-6-102
Ruelle J.E. 1975. Type specimen of Isoptera in the National Collection of Insects, Pretoria. Entomology Memoir. Department of Agriculture and Technical Services (South Africa) 45: 1–22.
Ruelle J.E. 1992. The genus Cubitermes Wasmann (Isoptera: Termitidae: Termitinae): a review of its taxonomy, nomenclature and distribution in Africa. Journal of African Zoology 106: 499–502.
Sands W.A. 1965. A revision of the termite subfamily Nasutitermitinae (Isoptera, Termitidae) from the Ethiopian Region. Bulletin of the British Museum (Natural History) Entomology Supplement 4: 1–172. https://doi.org/10.5962/p.193228
Sands W.A. 1972. The soldierless termites of Africa (Isoptera, Termitidae). Bulletin of the British Museum (Natural History) Entomology Supplement 18: 1–244.
Sands W.A. 1998. The Identification of Worker Castes of Termite Genera from Soils of Africa and the Middle East. CAB International, Wallingford, U.K.
Schmitz H. 1916. Ein neuer Cubitermes vom belgischen Kongo. Tijdschrift voor Entomologie (Suppl.) 58: 119–124.
Servat É., Paturel J.E., Lubès-Niel H., Kouamé B., Masson J.M., Travaglio M. & Marieu B. 1999. De différents aspects de la variabilité de la pluviométrie en Afrique de l‘Ouest et Centrale non sahélienne. Regarding rainfall variability in non Sahelian Western and Central Africa. Revue des sciences de l’eau /Journal of Water Science 12 (2): 363–387. https://doi.org/10.7202/705356ar
Silvestri F. 1914. Contribuzione alla conoscenza dei Termitidi e Termitofili dell’Africa occidentale. I. Termitidi. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scoula Superiore d’Agricoltura, Portici 9: 1–146.
Sjöstedt Y. 1896. Termiten aus Kamerun. Entomologisk Tidskrift 17 (4): 297–298.
Sjöstedt Y. 1900. Monographie der Termiten Afrikas. Kungliga Svenska Vetenskapsakademiens Handlingar 34 (4): 1–236.
Sjöstedt Y. 1913. Über Termiten aus dem inneren Kongo, Rhodesia, und Deutsch-Ostafrika. Revue Zoologique africaine 2 (3): 354–391.
Sjöstedt Y. 1924a. Neue Termiten im Kongo-Museum (vorläufige Mitteilung). Revue Zoologique africaine 12 (1): 39–42.
Sjöstedt Y. 1924b. Neue Arten und Gattungen afrikanischer Termiten (vorläufige Mitteilung). Revue Zoologique africaine 12 (2): 253–257.
Sjöstedt Y. 1924c. Neue Afrikanische Termiten (vorläufige Mitteilung). Revue Zoologique africaine 12 (4): 490–494.
Sjöstedt Y. 1925. Neue Termiten aus Afrika und Madagaskar. Konowia 4 (1–2): 53–55.
Sjöstedt Y. 1926. Revision der Termiten Afrikas 3. Monographie. Kungliga Svenska Vetenskapsakademiens Handlingar (Ser.3) 3 (1): 1–419.
Smeathman H. 1781. Some accounts of the termites which are found in Africa and other hot climates. Philosophical Transactions of the Royal Society of London 71 (1): 139–192. https://doi.org/10.1098/rstl.1781.0033
Snyder T.E. 1949. Catalog of the termites (Isoptera) of the world. Smithsonian Miscellaneous Collections 112 (3953): 1–490.
Srivathsan A. & Meier R. 2012. On the inappropriate use of Kimura-2-parameter (K2P) divergences in the DNA-barcoding literature. Cladistics 28: 190–194. https://doi.org/10.1111/j.1096-0031.2011.00370.x
Suyama M., Torrents D. & Bork P. 2006. PAL2NAL: robust conversion of protein sequence alignments into the corresponding codon alignments. Nucleic Acids Research 34: W609–W612. https://doi.org/10.1093/nar/gkl315
Thomsen P.F., Elias S., Gilbert M.T.P., Haile J., Munch K., Kuzmina S., Froese D.G., Sher A., Holdaway R.N. & Willerslev E. 2009. Non-destructive sampling of ancient insect DNA. PLoS ONE 4 (4): e5048. https://doi.org/10.1371/journal.pone.0005048
Wang M., Buček A., Šobotník J., Sillam-Dussès D., Evans T.A., Roisin Y., Lo N. & Bourguignon T. 2019. Historical biogeography of the termite clade Rhinotermitinae (Blattodea: Isoptera). Molecular Phylogenetics and Evolution 132: 100–104. https://doi.org/10.1016/j.ympev.2018.11.005
Wang M., Hellemans S., Šobotník J., Arora J., Buček A., Sillam-Dussès D., Clitheroe C., Lu T., Lo N., Engel M.S., Roisin Y., Evans T.A. & Bourguignon T. 2022. Phylogeny, biogeography and classification of Teletisoptera (Blattaria: Isoptera). Systematic Entomology 47: 581–590. https://doi.org/10.1111/syen.12548
Wang M., Hellemans S., Buček A., Kanao T., Arora J., Clitheroe C., Rafanomezantsoa J.-J., Fisher B.L., Scheffrahn R., Sillam-Dussès D., Roisin Y., Šobotník J. & Bourguignon T. 2023. Neoisoptera repeatedly colonised Madagascar after the Middle Miocene climatic optimum. Ecography 2023: e06463. https://doi.org/10.1111/ecog.06463
Wango S.P. & Josens G. 2011. Comparison of nest shapes and densities of two sympatric species of Cubitermes (Isoptera: Termitidae: Termitinae) as clues for the study of their population dynamics. African Zoology 46: 156–168. https://doi.org/10.3377/004.046.0106
Wasmann E. 1906. Beispiele rezenter Artenbildung bei Ameisengästen und Termitengästen. Biologisches Central Blatt 26 (17–18): 565–580.
Wasmann E. 1911. Zur Kenntnis der Termiten und Termitengäste vom Belgischen Congo. Revue Zoologique africaine 1 (1–2): 91–117, 145–176.
Wikipedia. 2024a. Georg August Zenker.Available from https://de.wikipedia.org/wiki/Georg_August_Zenker [accessed 2024].
Wikipedia. 2024b. Karl Edvard Laman. Available from https://en.wikipedia.org/wiki/Karl_Edvard_Laman [accessed 2024].
Williams R.M.C. 1966. The East African termites of the genus Cubitermes (Isoptera: Termitidae). Transactions of the Royal Entomological Society of London 118 (4): 73–118. https://doi.org/10.1111/j.1365-2311.1966.tb00835.x
Wood T.G. & Lee K.E. 1971 Abundance of mounts and competition among colonies of some Australian termite species. Pedobiologia 11: 341–366. https://doi.org/10.1016/S0031-4056(23)00480-8
Wu L.W., Bourguignon T., Šobotník J., Wen P., Liang W.R. & Li H.F. 2018. Phylogenetic position of the enigmatic termite family Stylotermitidae (Insecta: Blattodea). Invertebrate Systematics 32: 1111–1117. https://doi.org/10.1071/IS17093
Copyright (c) 2025 Guy Josens, Jean Deligne, Myriam Harry, Virginie Roy, Pierre D. Akama, Tenon Coulibaly, Kanvaly Dosso, Georg Goergen, Michel Hasson, Patrick Kasangij a Kasangij, Carmel Kifukieto, Bruno Le Ru, Laura Estelle Yêyinou Loko, Abdoulaye Baila Ndiaye, Yves Roisin, Noémie Sion, Jan Šobotnik, Petr Stiblik, Soki Kue Di Kuenda, Saran Traoré, Manuela Viage, Solange Patricia Wango, Esra Kaymak, Thomas Bourguignon, Simon Hellemans
This work is licensed under a Creative Commons Attribution 4.0 International License.
Creative Commons Copyright Notices
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License (CC BY 4.0) that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are NOT PERMITTED to post their submitted work online (e.g., in institutional repositories or on personal websites) prior to publication, as it may lead to nomenclatural issues.
