Abstract
A new species, Nothobranchius sylvaticus sp. nov. is described from ephemeral swamps in open areas of the Gongoni Forest in the seasonal Mkurumudzi system in south-eastern coastal Kenya. The species occupies a unique ecological niche as it inhabits small seasonal wetland habitats situated in a relict forest ecosystem. Nothobranchius sylvaticus sp. nov. is distinguished from all other members of the genus by: iridescent light blue scales on its trunk extending to cover the entire caudal peduncle; dorsal portion of head grey with distinct red lobes along the supraorbital canal; posterior scale margins on the postorbital portion of the opercle creating two pronounced brown-red oblique bars; dorsal and anal fins golden, marked with dark grey dots and stripes proximally and medially, merging into a uniform dark grey zone distally; caudal fin dark red to red proximally, plain red medially and with a slender black distal band. Phylogenetic analysis of sequences of the mitochondrial ND2 and COI, and the nuclear Glyt, MyH6 and SNX33 genes, retrieve the new species as a phylogenetically distinct lineage, in the so-called Coastal Inland clade sister to subgenus Aphyobranchius Wildekamp, 1977 and subgenus Adiniops Myers, 1924.
Increasing impacts on land and water resources will continue to cause habitat degradation in the remnant forest ecosystem, exposing the ecologically vulnerable population of this newly described species to severe threats. Such changes may prevent the species from completing all stages of its annual life cycle in its ephemeral aquatic habitats, because these killifishes depend on a clay-rich substratum for embryonic development of the buried eggs through the dry season. This new species qualifies for a conservation status of Critically Endangered.
References
- Abell, R., Thieme, M.L., Revenga, C., Bryer, M., Kottelat, M., Bogutskaya, N., Coad, B., Mandrak, N., Contreras-Balderas, S., Bussing, W., Stiassny, M.L.J., Skelton, P., Allen, G.R., Unmack, P., Naseka, A., Ng, R., Sindorf, N., Robertson, J., Armijo, E., Higgins, J.V., Heibel, T.J., Wikramanayake, E., Olson, D., Loìpez, H.L., Reis, R.E., Lundberg, J.G., Sabaj Peìrez, M.H. & Petry, P. (2008) Freshwater ecoregions of the world: a new map of biogeographic units for freshwater biodiversity conservation. BioScience, 58 (5), 403–414. https://doi.org/10.1641/B580507
- Abuodha, J.O.Z. (2004) Geomorphological evolution of the southern coastal zone of Kenya, Journal of African Earth Sciences, 39, 517–525. https://doi.org/10.1016/j.jafrearsci.2004.07.012
- Atherton, R. (2003) The Fringing Reef Coasts of Eastern Africa – Present Processes in Their Long-term Context. Western Indian Ocean Journal of Marine Science, 2, 1–13. https://doi.org/10.4314/wiojms.v2i1.28424
- Barratt, C.D., Bwong, B.A., Onstein, R.E., Rosauer, D.F., Menegon, M., Doggart, N., Nagel, P., Kissling, W.D. & Loader, S.P. (2020) Environmental correlates of phylogenetic endemism in amphibians and the conservation of refugia in the Coastal Forests of Eastern Africa. Diversity and Distributions, 23, 875–887. https://doi.org/10.1111/ddi.12582
- Baur, H. & Leuenberger, C. (2011) Analysis of ratios in multivariate morphometry. Systematic Biology, 60 (6), 813–825. https://doi.org/10.1093/sysbio/syr061
- Boyden, P., Weil-Accardo, J., Deschamps, P., Oppo, D. & Rovere, A. (2021) Last interglacial sea-level proxies in East Africa and the Western Indian Ocean. Earth System Science Data, 13, 1633–1651. https://doi.org/10.5194/essd-13-1633-2021
- Bragança, P.H.N., Bellstedt, D.U., van der Merwe, P.D.W., Cotterill, F.P.D., Watters, B.R. & Chakona, A. (2022) Nothobranchius balamaensis (Cyprinodontiformes: Nothobranchiidae), a new species of annual killifish from northern Mozambique. Zootaxa, 5174 (5), 508–524. https://doi.org/10.11646/zootaxa.5174.5.2
- Butynski, T.M., Parker, I. & de Jong, Y.A. (2015) Historic and Current Distribution, Abundance, and Habitats of Roosevelt's Sable Antelope Hippotragus niger roosevelti (Heller, 1910) (Cerartiodactyla: Bovidae) in Kenya. Journal of African Earth Sciences, 104, 41–77. https://doi.org/10.2982/028.104.0107
- Bwong, B.A., Nyamache, J.O., Malonza, P.K., Wasonga, D.V., Ngwava, J.M., Barratt, C.D., Nagel, P. & Loader, S.P. (2017) Amphibian diversity in Shimba Hills National Reserve, Kenya: A comprehensive list of specimens and species. Journal of East African Natural History, 106, 19–46. https://doi.org/10.2982/028.106.0104
- Burgess, N., Butynski, T.M., Cordeiro, N.J., Doggart, N.H., Fjeldså, J., Howell, K.M., Kilahama, F.B., Loader, S.P., Lovett, J.C., Mbilinyi, B. & Menegon, M. (2007) The biological importance of the Eastern Arc Mountains of Tanzania and Kenya. Biological Conservation, 134, 209–231.
- Bytebier, B. (2001) Taita Hills Biodiversity Project Report. National Museums of Kenya, Nairobi, 47 pp.
- Cellerino, A., Valenzano, D.R. & Reichard, M. (2016) From the bush to the bench: the annual Nothobranchius fishes as a new model system in biology. Biological Reviews, 91, 511–533. https://doi.org/10.1111/brv.12183
- Chambers, J. (1984) A new Nothobranchius species (Teleostei, Cyprinodontiformes, Aplocheilidae) from the Sudan. Journal of Fish Biology, 25, 505–513. https://doi.org/10.1111/j.1095-8649.1984.tb04897.x
- Costa, W.J.E.M. (2018) Comparative morphology, phylogeny and classification of African seasonal killifishes of the tribe Nothobranchiini (Cyprinodontiformes: Aplocheilidae). Zoological Journal of the Linnean Society, 184 (1), 115–135. https://doi.org/10.1093/zoolinnean/zlx102
- Costa, W.J.E.M. (2019) Two new species of seasonal killifishes of the Nothobranchius melanospilus species complex from the East Africa biodiversity hotspot (Cyprinodontiformes: Aplocheilidae). Vertebrate Zoology, 69 (1), 73–82.
- Cui, R., Medeiros, T., Willemsen, D., Iasi, L.N., Collier, G.E., Graef, M., Reichard, M. & Valenzano, D.R. (2019) Relaxed selection limits lifespan by increasing mutation load. Cell, 178, 385–399. https://doi.org/10.1016/j.cell.2019.06.004
- Dagallier, L.M.J., Condamine, F.L. & Couvreur, T.L.P. (2024) Sequential diversification with Miocene extinction and Pliocene speciation linked to mountain uplift explains the diversity of the African rain forest clade Monodoreae (Annonaceae). Annals of Botany, 133, 677–696. https://doi.org/10.1093/aob/mcad130
- Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9 (8), 772. https://doi.org/10.1038/nmeth.2109
- de Rham, P. (1991) Safaris poissons au Kenya. Première partie. Aquarama, 120, 31–44.
- Dorn, A., Musilová, Z., Platzer, M., Reichwald, K. & Cellerino, A. (2014) The strange case of East African annual fishes: aridification correlates with diversification for a savannah aquatic group? BMC Evolutionary Biology, 14 (1), 210. https://doi.org/10.1186/s12862-014-0210-3
- Dytham, C. (2011) Choosing and using statistics: a biologist's guide. Third Edition. Wiley-Blackwell, Oxford, Chichester and Hoboken, 320 pp.
- Gereau, R.E., Cumberlidge, N., Hemp, C., Hochkirch, A., Jones, T., Kariuki, M., Lange, C.N., Loader, S.P., Malonza, P.K., Menegon, M., Ndang'ang'a, P.K., Rovero, F. & Shirk, P. (2016) Globally Threatened Biodiversity of the Eastern Arc Mountains and Coastal Forests of Kenya and Tanzania. Journal of East African Natural History, 105, 115–201. https://doi.org/10.2982/028.105.0104
- Githitho, A.N. (2021) Annotated checklist of the plants of the Arabuko-Sokoke Forest, Coastal Kenya. Journal of East African Natural History, 105, 13–74. https://doi.org/10.2982/028.110.0102
- Guindon, S. & Gascuel, O. (2003) A Simple, Fast, and Accurate Algorithm to Estimate Large Phylogenies by Maximum Likelihood. Systematic Biology, 52, 696–704. https://doi.org/10.1080/10635150390235520
- Hilgendorf, F.M. (1905) Fische von Deutsch und Englisch Ost-Afrika, gesammelt von Oskar Neumann, 1893–1895. Zoologische Jahrbücher Abteilung für Systematik, Geographie und Biologie der Tiere, 22 (4), 405–418.
- Hughes, R.H. & Hughes, J.S. (1992) A directory of African wetlands. IUCN, Gland and Cambridge and UNEP, Nairobi, 820 pp.
- IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1. 2nd Edition. IUCN, Gland and Cambridge, 32 pp.
- James, F.C. & McCulloch, C.E. (1990) Multivariate analysis in ecology and systematics: panacea or Pandora’s box? Annual Review of Ecology and Systematics, 21, 129–166. https://doi.org/10.1146/annurev.es.21.110190.001021
- Joordens, J.C., Feibel, C.S., Vonhof, H.B., Schulp, A.S. & Kroon, D. (2019) Relevance of the eastern African coastal forest for early hominin biogeography. Journal of Human Evolution, 131, 176–202. https://doi.org/10.1016/j.jhevol.2019.03.012
- Jubb, R.A. (1969) The Nothobranchius (Pisces, Cyprinodontidae) of southern Africa and a new species from Lake Chilwa, Malawi. Annals of the Cape Provincial Museums (Natural History), 8 (1), 1–9.
- Jubb, R.A. (1971) A new Nothobranchius (Pisces, Cyprinodontidae) from southeastern Rhodesia. Journal of the American Killifish Association, 8, 12–19.
- Jubb, R.A. (1981) Nothobranchius. T.F.H., Neptune City, 61 pp.
- Lévêque, C. (1997) Biodiversity dynamics and conservation: The freshwater fish of tropical Africa Cambridge University Press, Cambridge, 438 pp.
- Lönnberg, E. (1907) Fishes. In: Wissenschaftliche Ergebnisse der Schwedische Zoologische Expedition nach dem Kilimandjaro, dem Meru und den Umgebenden Massaisteppen 1905–11906 unter Leitung von Prof. Dr. Yngve Sjöstedt. K. Schwedischen Akademie der Wissenschaften, Uppsala, pp. 1–8.
- Lovett, J.C. & Wasser, S.K. (2008) Biogeography and ecology of the rain forests of East Africa. Cambridge University Press, Cambridge, 341 pp.
- Luke, Q. (2005) Annotated checklist of the plants of the Shimba Hills, Kwale District, Kenya. Journal of East African Natural History, 94, 5–121. https://doi.org/10.2982/0012- 8317(2005)94[5:ACOTPO]2.0.CO;2
- Malonza, P.K., Mulwa, D.M., Nyamache, J.O. & Jones, G. (2018) Biogeography of the Shimba Hills ecosystem herpetofauna in Kenya. Zoological Research, 39, 97–104. https://doi.org/10.24272/j.issn.2095-8137.2017.048
- Mayden, R.L. (1999) Consilience and a hierarchy of species concepts: advances toward closure on the species puzzle. Journal of Nematology, 31, 95–116.
- Mayden, R.L. (2002) On biological species, species concepts and individuation in the natural world. Fish and Fisheries, 3, 171–196. https://doi.org/10.1046/j.1467-2979.2002.00086.x
- McDonald, J.H. (2008) Handbook of biological statistics. Sparky House, Baltimore, Maryland, 287 pp.
- Miller, M.A., Pfeiffer, W., Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. 2010 Gateway Computing Environments Workshop (GCE). Institute for Electrical and Electronics Engineers, New Orleans, Louisiana, pp. 1–8. https://doi.org/10.1109/GCE.2010.5676129
- Minitab. (2010) [Statistical computer software]. Version 16.2.1. Minitab, Inc., State College.
- Myers, G.S. (1924) A new poeciliid fish from the Congo, with remarks on funduline genera. American Museum Novitates, 116, 1–9.
- Nagy, B. (2014) Nothobranchius milvertzi, a new species of killifish from the Lushiba Marsh in the Lake Mweru drainage, Zambia (Teleostei: Cyprinodontiformes: Nothobranchiidae). Ichthyological Exploration of Freshwaters, 24, 347–360.
- Nagy, B. (2015) Life history and reproduction of Nothobranchius fishes. Journal of the American Killifish Association, 47, 182–192. https://doi.org/10.5281/zenodo.4393325
- Nagy, B. (2018) Nothobranchius ditte, a new species of annual killifish from the Lake Mweru basin in Democratic Republic of Congo (Teleostei: Nothobranchiidae). Ichthyological Exploration of Freshwaters, 28, 115–134.
- Nagy, B. (2021) Nothobranchius elucens, a new species of seasonal killifish from the upper Nile drainage in Uganda (Cyprinodontiformes: Nothobranchiidae). Zootaxa, 4915 (1), 133–147. https://doi.org/10.11646/zootaxa.4915.1.10
- Nagy, B. & Watters, B.R. (2020) Threatened - the world is a dangerous place for seasonal fishes. Journal of the American Killifish Association, 53, 141–156.
- Nagy, B. & Watters, B.R. (2021) A review of the conservation status of seasonal Nothobranchius fishes (Teleostei: Cyprinodontiformes), a genus with a high level of threat, inhabiting ephemeral wetland habitats in Africa. Aquatic Conservation: Marine and Freshwater Ecosystems, 32 (1), 199–216. https://doi.org/10.1002/aqc.3741
- Nagy, B., Watters, B.R., van der Merwe, P.D.W., Cotterill, F.P.D. & Bellstedt, D.U. (2020) Review of the Nothobranchius ugandensis species group from the inland plateau of eastern Africa with descriptions of six new species (Teleostei: Nothobranchiidae). Ichthyological Exploration of Freshwaters, 30, 21–73. http://doi.org/10.23788/IEF-1129
- Nagy, B., Watters, B.R. & Raspopova, A.A. (2021) Nothobranchius nikiforovi, a new species of seasonal killifish from the lower Matandu drainage in south-eastern coastal Tanzania (Cyprinodontiformes: Nothobranchiidae). Zootaxa, 4950 (1), 103–122. https://doi.org/10.11646/zootaxa.4950.1.5
- Ng’oma, E., Valdesalici, S., Reichwald, K. & Cellerino A. (2013) Genetic and morphological studies of Nothobranchius (Cyprinodontiformes) from Malawi with description of Nothobranchius wattersi sp. nov. Journal of Fish Biology, 82, 165–188. https://doi.org/10.1111/jfb.12001
- Ngumbau, V.M., Luke, Q., Nyange, M., Wanga, V.O., Watuma, B.M., Mbuni, Yu.M., Munyao, J.N., Oulo, M.A., Mkala, E.M., Kipkoech, S., Itambo, M., Hu, G.-W. & Wang, Q.-F. (2020) An annotated checklist of the coastal forests of Kenya, East Africa. PhytoKeys, 147, 1–191. https://doi.org/10.3897/phytokeys.147.49602
- Nicholson, S.E. (2017) Climate and climatic variability of rainfall over eastern Africa, Reviews of Geophysics, 55, 590–635. https://doi.org/10.1002/2016RG000544
- Ogoma, M.O., Breckling, B., Reuter, H., Muchane, M. & Githiru, M. (2010) The birds of Gongoni Forest Reserve, South Coast, Kenya. Scopus, 30, 1–11.
- Peters, N. (1963) Zur Embryonalentwicklung bodenlaichender Zahnkarpfen. Die Aquarien und Terrarien Zeitschrift, 16, 201–204.
- Peters, W.C.H. (1868) Über eine neue Untergattung der Flederthiere so wie über neue Gattungen und Arten von Fischen. Monatsberichte der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 1868, 145–148.
- Poll, M. (1938) Poissons du Katanga (bassin du Congo) récoltés par le professeur Paul Brien. Revue de Zoologie et de Botanique Africaines, 30, 389–423.
- Quinn, G.P. & Keough, M.J. (2002) Experimental design and data analysis for biologists. University Press, Cambridge, 553 pp. https://doi.org/10.1017/CBO9780511806384
- Reichard, M. (2010) Nothobranchius kadleci (Cyprinodontiformes: Nothobranchiidae), a new species of annual killifish from central Mozambique. Zootaxa, 2332 (1), 49–60. https://doi.org/10.11646/zootaxa.2332.1.4
- Reichard, M. (2015) The evolutionary ecology of African annual fishes. In: Berois, N., García, G. & De Sá, R. (Eds.), Annual fishes: life history strategy, diversity, and evolution. CRC Press, Boca Raton, Florida, pp. 133–158.
- Rice, W.R. (1989) Analyzing tables of statistical tests. Evolution, 43, 223–225. https://doi.org/10.1111/j.1558-5646.1989.tb04220.x
- Robertson, R.J. & Luke, Q. (1993) Kenya Coastal Forests. Report to Director NMK. World Wide Fund for Nature, Nairobi. [unknown pagination]
- Rocha, J.L., Pinto, P.V., Siegismund, H.R., Meyer, M., van Vuuren, B.J., Veríssimo, L., Ferrand, N. & Godinho, R. (2022) African climate and geomorphology drive evolution and ghost introgression in sable antelope. Molecular Ecology, 31, 2968–2984. https://doi.org/10.1111/mec.16427
- Schedel, F.D.B., Friel, J.P. & Schliewen, U.K. (2014) Haplochromis vanheusdeni, a new haplochromine cichlid species from the Great Ruaha River drainage, Rufiji basin, Tanzania (Teleostei, Perciformes, Cichlidae). Spixiana, 37 (1), 135–149.
- Seegers, L. (1985) Prachtgrundkärpflinge, die Gattung Nothobranchius: Systematik, Vorkommen, Pflege und Zucht. Deutsche Killifisch Gemeinschaft Journal, Supplement No. 1, 1–48.
- Seegers, L. (1997) Killifishes of the world. Old world killis II. Aqualog, A.C.S., Mörfelden-Walldorf, 112 pp.
- Seegers L., De Vos, L. & Okeyo, D.O. (2003) Annotated Checklist of the Freshwater Fishes of Kenya (excluding the lacustrine haplochromines from Lake Victoria). Journal of East African Natural History, 92, 11–47. https://doi.org/10.2982/0012-8317(2003)92[11:ACOTFF]2.0.CO;2
- Sokal, R.R. & Rohlf, F.J. (1995) Biometry: the principles and practice of statistics in biological research. 3rd Edition. Freeman, New York, New York, 880 pp.
- Sokal, R.R. & Rohlf, F.J. (2009) Introduction to biostatistics. Second edition. Dover Publications, Mineola, 382 pp.
- Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313. https://doi.org/10.1093/bioinformatics/btu033
- Thieme, M.L., Abell, R., Stiassny, M.L.J., Skelton, P.H., Lehner, B., Teugels, G.G., Dinerstein, E., Kamdem-Toham, A., Burgess, N.D. & Olson, D. (2005) Freshwater ecoregions of Africa and Madagascar: a conservation assessment. Island Press, Washington, Covelo and London, 483 pp.
- Vanderplank, F.L. (1940) A study of Nothobranchius taeniopygus in its natural habitat. The Aquarist, 10, 247–249.
- van der Merwe, P.D.W., Cotterill, F.P.D., Kandziora, M., Watters, B.R., Nagy, B., Genade, T., Flügel, T. J., Svendsen, D.S. & Bellstedt, D.U. (2021) Genomic Fingerprints of Palaeogeographic History: The tempo and mode of Rift tectonics across tropical Africa has shaped the diversification of the killifish genus Nothobranchius (Teleostei: Cyprinodontiformes). Molecular Phylogenetics and Evolution, 158 (106988), 1–21. https://doi.org/10.1016/j.ympev.2020.106988
- Vinciguerra, D. (1897) Pesci raccolti dal Cap. V. Bottego, durante la sua seconda spedizione nelle regione dei Somali e dei Galla. Annali del Museo Civico di Storia Naturale Giacomo Doria, Genova, Series 2, 17, 343–364.
- Vinciguerra, D. (1927) Enumerazione de alcune specie di pesci della Somalia Italiana raccolte dal March.se Saverio Patrizi. Annali del Museo Civico di Storia Naturale Giacomo Doria, Genova, Series 3, 52, 246–25.
- Vrtílek, M., Žák, J., Pšenička, M. & Reichard, M. (2018) Extremely rapid maturation of a wild African annual fish. Current Biology, 28, 822–824. https://doi.org/10.1016/j.cub.2018.06.031
- Watters, B.R. (2009) The ecology and distribution of Nothobranchius fishes. Journal of the American Killifish Association, 42, 37–76.
- Watters, B.R. (2014) A classification of Nothobranchius fish habitats. Journal of the American Killifish Association, 47, 152–180.
- Watters, B.R., Wildekamp, R.H. & Shidlovskiy, K.M. (2014) Description and biogeography of Nothobranchius capriviensis, a new species of annual killifish from the Zambezi Region of Namibia (Cyprinodontiformes: Nothobranchiidae). Journal of the American Killifish Association, 47, 4–6.
- Watters, B.R., Wildekamp, R.H. & Cooper, B.J. (1998) Zwei neue Nothobranchius-Arten aus der Küstenebene Tansanias. Deutsche Killifisch Gemeinschaft Journal, 30, 52–63.
- Watters, B.R., Nagy, B., van der Merwe, P.D.W., Cotterill, F.P.D. & Bellstedt, D.U. (2019) Review of the Nothobranchius taeniopygus species group from central and western Tanzania with descriptions of five new species and redescription of Nothobranchius taeniopygus (Teleostei: Nothobranchiidae). Ichthyological Exploration of Freshwaters, 29, 239–278. http://doi.org/10.23788/IEF-1110
- Watters, B.R., Nagy, B., van der Merwe, P.D.W., Cotterill, F.P.D. & Bellstedt, D.U. (2020) Redescription of the seasonal killifish species Nothobranchius ocellatus and description of a related new species Nothobranchius matanduensis, from eastern Tanzania (Teleostei: Nothobranchiidae). Ichthyological Exploration of Freshwaters, 30, 151–178. https://doi.org/10.23788/IEF-1149
- White, F. (1983) The vegetation of Africa: a descriptive memoir to accompany the UNESCO/AETFAT/UNSO vegetation map of Africa. Natural Resources Research, 20, 1–356.
- Wildekamp, R.H. (1977) Nothobranchius lourensi spec. nov. und Nothobranchius janpapi spec. nov., zwei neue Rivulinen aus Ost Afrika. Das Aquarium, 11 (98), 326–331.
- Wildekamp, R.H. (1982) Die Nothobranchius Arten des Küstengebietes Kenias. Die Aquarien und Terrarien Zeitschrift, 35, 333–339.
- Wildekamp, R.H. (1992) A new Nothobranchius (Cyprinodontiformes: Aplocheilidae) from the Kenyan coastal plains, with a redefinition of the subgenus Aphyobranchius. Ichthyological Exploration of Freshwaters, 3, 187–192.
- Wildekamp, R.H. (2004) A world of killies. Atlas of the oviparous cyprinodontiform fishes of the world. Vol. IV. American Killifish Association, Elyria, 398 pp.
- Wildekamp, R.H. & Berkenkamp, H.O. (1979) Untersuchungen zur Identität von Nothobranchius neumanni (Hilgendorf, 1905) aus Tansania, mit der Beschreibung einer neue Art und einer Unterart aus dem Küstentiefland Kenias. Deutsche Killifisch Gemeinschaft Journal, 11, 65–75.
- Wildekamp, R.H., Shidlovskiy, K.M. & Watters, B.R. (2009) Systematics of the Nothobranchius melanospilus species group (Cyprinodontiformes: Nothobranchiidae) with description of two new species from Tanzania and Mozambique. Ichthyological Exploration of Freshwaters, 20, 237–254.
- Wildekamp, R.H., Watters, B.R. & Shidlovskiy, K.M. (2014) Review of the Nothobranchius neumanni species group with descriptions of three new species from Tanzania (Cyprinodontiformes: Nothobranchiidae). Journal of the American Killifish Association, 47 (1), 2–30.
- Wiley, E.O. (1978) The evolutionary species concept reconsidered. Systematic Zoology, 27, 17–26. https://doi.org/10.2307/2412809
- Zar, J.H. (2010) Biostatistical analysis. Fifth edition. Pearson/Prentice-Hall, Upper Saddle River, xii + 944 pp.
- Zuur, A.F., Ieno, E.N. & Smith, G.M. (2007) Analysing ecological data (statistics for biology and health). Springer, New York, New York, xxvi + 672 pp. https://doi.org/10.1007/978-0-387-45972-1