Why the taxonomy of francolins and spurfowls (Galliformes, Phasianidae) needs revision: responses to Hustler (2021) and Hunter et al. (2021a,b)

References

• Benson CW. 1945. Notes on the birds of southern Abyssinia (part). Ibis 87: 366–400. https://doi.org/10.1111/j.1474-919X.1945.tb01370.x
   Google Scholar
• Benson CW, Benson FM. 1977. The birds of Malawi. Limbe: Montfort Press.
   Google Scholar
• Benson CW, Brooke RK, Dowsett RJ, Irwin MPS. 1971. The birds of Zambia. London: Collins.
   Google Scholar
• Bloomer P, Crowe TM. 1998. Francolin phylogenetics: molecular, morphobehavioral, and combined evidence. Molecular Phylogenetics and Evolution 9: 236–254. https://doi.org/10.1006/mpev.1997.0469
   PubMed Web of Science ®Google Scholar
• Bock WJ, Farrand J. 1980. The number of species and genera of Recent birds. American Museum Novitates 2703: 1–29.
   Google Scholar
• Bowie RCK, Fjeldså J. 2005. Genetic and morphological evidence for two species in the Udzungwa Forest Partridge Xenoperdix udzungwensis. Special Issue on the Eastern Arc Mts. Journal of East African Natural History 94: 191–201. https://doi.org/10.2982/0012-8317(2005)94[191:GAMEFT]2.0.CO;2
   Google Scholar
• Bowie RCK, Cohen C, Crowe TM. 2013. Ptilopachinae: a new subfamily of the Odontophoridae (Aves: Galliformes). Zootaxa 3670: 97–98. https://doi.org/10.11646/zootaxa.3670.1.11
   PubMed Web of Science ®Google Scholar
• Buck R, Flores-Rentería L. 2022. The eyngameon enigma. Plants (Basel) 11: 895. https://doi.org/10.3390/plants11070895
   PubMedGoogle Scholar
• Chapin JP. 1926. A new genus, Acentrortyx, proposed for Francolinus nahani Dubois. The Auk 43: 235. https://doi.org/10.2307/4074928
   Google Scholar
• Cicero C, Mason NA, Jiménez RA, Wait DR, Wang-Claypool CY, Bowie RCK. 2021. Integrative taxonomy and geographic sampling underlie successful species delimitation. Ornithology 138: ukab009. https://doi.org/10.1093/ornithology/ukab009
   Google Scholar
• Clancey PA. 1967. Gamebirds of southern Africa. Cape Town: Purnell and Sons.
   Google Scholar
• Cohen C, Wakeling JL, Mandiwana-Neudani TG, Sande E, Dranzoa C, Crowe TM, Bowie RCK. 2012. Phylogenetic affinities of evolutionarily enigmatic African galliforms: the Stone Partridge Ptilopachus petrosus and Nahan’s Francolin Francolinus nahani, and support for their sister relationship with New World quails. Ibis 154: 768–780. https://doi.org/10.1111/j.1474-919X.2012.01269.x
   Web of Science ®Google Scholar
• Conover HB. 1929. The races of the Bare-throated Francolin, Pternistes cranchi [sic]. The Auk 46: 344–347. https://doi.org/10.2307/4076110
   Google Scholar
• Cracraft J. 1997. Species concepts in systematics and conservation biology – an ornithological viewpoint. In: Claridge MF, Dawah HA, Wilson MR (eds). In: Species: the units of biodiversity. London: Chapman & Hall. pp. 325–339.
   Google Scholar
• Crowe TM. 1978. The evolution of guineafowl (Galliformes, Phasianidae, Numidinae): taxonomy, phylogeny, speciation and biogeography. Annals of the South African Museum 76: 43–136.
   Google Scholar
• Crowe TM. 1987. Species as individuals or classes: an ‘iconoclassificationist’s’ view. Biology and Philosophy 2: 167. https://doi.org/10.1007/BF00057960
   Google Scholar
• Crowe TM. 1988. Molecules vs morphology in phylogenetics: a non-controversy. Transactions of the Royal Society of South Africa 46: 317–334. https://doi.org/10.1080/00359198809520135
   Google Scholar
• Crowe TM. 2010. Phylogenetic affinities of enigmatic African galliforms: the Stone Partridge Ptilopachus petrosus and Latham’s and Nahan’s ‘Francolins’ Francolinus lathami and F. nahani. Cladistics 26: 206.
   Web of Science ®Google Scholar
• Crowe TM, Bloomer. 2017. A new approach to understanding subspecies can boost conservation. The Conversation, 2 February 2017. https://theconversation.com/a-new-approach-to-understanding-subspecies-can-boost-conservation-68364
   Google Scholar
• Crowe TM, Crowe AA. 1985. The genus Francolinus as a model for avian evolution and biogeography in Africa. In: Schuchmann KL (ed.), Proceedings of the International Symposium on African Vertebrates. Bonn: Museum Alexander Koenig. pp 207–231.
   Google Scholar
• Crowe TM, Little RM. 2004. Francolins, partridges and spurfowls: what’s in a name? Ostrich 75: 199–203. https://doi.org/10.2989/00306520409485445
   Web of Science ®Google Scholar
• Crowe TM, Keith GS, Brown LH. 1986. Galliformes. In: Urban E, Fry CH, Keith GS (eds), Birds of Africa, vol. 2. London: Academic Press. pp 24–75.
   Google Scholar
• Crowe TM, Harley EH, Jakutowicz MB, Komen J, Crowe AA. 1992. Phylogenetic, taxonomic and biogeographical implications of genetic, morphological, and behavioral variation in francolins (Phasianidae: Francolinus). The Auk 109: 24–42. https://doi.org/10.2307/4088264
   Web of Science ®Google Scholar
• Crowe TM, Essop MF, Allan DG, Brooke RK, Komen J. 1994. Overlooked units of comparative and conservation biology: a case study of a small African bustard, the Black Korhaan Eupodotis afra (Otididae) as a case study. Ibis 136: 166–175. https://doi.org/10.1111/j.1474-919X.1994.tb01081.x
   Web of Science ®Google Scholar
• Crowe TM, Bowie RCK, Bloomer P, Mandiwana T, Hedderson TAJ, Randi E et al. 2006. Phylogenetics, biogeography and classification of, and character evolution in gamebirds (Aves: Galliformes): effects of character exclusion, partitioning and missing data. Cladistics 22: 495–532. https://doi.org/10.1111/j.1096-0031.2006.00120.x
   PubMed Web of Science ®Google Scholar
• Crowe TM, Muasya M, Mandiwana-Neudani TG. 2016. The long struggle to understand species: from pre-Darwin to the present day. The Conversation, 6 November 2016. https://theconversation.com/the-long-struggle-to-understandspecies%20from-pre-darwin-to-the-present-day-65962
   Google Scholar
• Crowe TM, Little RM, Turner DA. 2019. Clarifying taxonomic, distributional and genetic information relating to subspecies of the Hildebrandt’s Spurfowl Pternistis hildebrandti and the Yellow-necked Spurfowl P. leucoscepus. Ostrich 90: 279–280. https://doi.org/10.2989/00306525.2019.1660431
   Web of Science ®Google Scholar
• Crowe TM, Mandiwana-Neudani TG, Donsker DB, Bowie RCK, Little RM. 2020. Resolving nomenclatural ‘confusion’ vis-à-vis Latham’s Francolin (Francolinus/Peliperdix/Afrocolinus lathami) and ‘Red-tailed’ francolins (Francolinus/Ortygornis/Peliperdix spp.). Ostrich 91: 134–136. https://doi.org/10.2989/00306525.2020.1723140
   Web of Science ®Google Scholar
• Crowe TM, Little RM, Mandiwana-Neudani TG, Bowie RCK. 2021. Francolin and spurfowl taxonomy revisited. African Birdlife 9: 8–10.
   Google Scholar
• del Hoyo J, Elliott A, Sargatal J. 1994. Handbook to the birds of the world, vol. 2. Barcelona, Spain: Lynx Edicions.
   Google Scholar
• Dinesen L, Lehmberg T, Svendsen JO, Hansen LA, Fjeldså J. 1994. A new genus and species of perdicine bird (Phasianidae, Perdicini) from Tanzania; a relict form with Indo-Malayan affinities. Ibis 136: 3–11. https://doi.org/10.1111/j.1474-919X.1994.tb08125.x
   Web of Science ®Google Scholar
• Dowsett RJ, Aspinwall DL, Dowsett-Lemaire F. 2008. The birds of Zambia: an atlas and handbook. Liége: Tauraco Press.
   Google Scholar
• Dowsett-Lemaire F, Dowsett RJ. 2006. The birds of Malawi: an atlas and handbook. Liége: Tauraco Press.
   Google Scholar
• Dowsett-Lemaire F, Dowsett RJ. 2014. The birds of Ghana: an atlas and handbook. Liège: Tauraco Press.
   Google Scholar
• Frankham R, Ballou JD, Dudash MR, Eldridge MDB, Fenster CB, Lacy RC et al. 2012. Implications of different species concepts for conserving biodiversity. Biological Conservation 153: 25–31. https://doi.org/10.1016/j.biocon.2012.04.034
   Web of Science ®Google Scholar
• Frost PGH. 1975. The systematic position of the Madagascan partridge Margaroperdix madagarensis (Scopoli). Bulletin of the British Ornithologists Club 95: 64–68.
   Google Scholar
• Hall BP. 1963. The Francolins, a study in speciation. Bulletin of the British Museum of Natural History 10: 105–204. https://www.biodiversitylibrary.org/page/2252779#page/181/mode/1up
   Google Scholar
• Harrison JA, Allan DG, Underhill LG, Herremans M, Tree AJ, Parker V et al. 1997. The atlas of southern African birds, vol. 1: Non-passerines. Johannesburg: BirdLife South Africa.
   Google Scholar
• Hunter N, Mills M, Cohen C. 2019. Elgon Francolin Scleroptila elgonensis should be treated as a species distinct from Moorland Francolin S. psilolaema. Bulletin of the African Bird Club 26: 92–100.
   Google Scholar
• Hunter N, Turner D, Borrow N, Finch B, Kennedy AS, Stevenson T et al. 2021a. Reflections concerning Spurfowl and Francolin species recommendations contained in Mandiwana-Neudani et al. (2019a and 2019b). Ostrich 92: 156–161. doi: 10.2989/00306525.2021.1890255
   Web of Science ®Google Scholar
• Hunter N, Turner D, Borrow N, Finch B, Kennedy AS, Stevenson T et al. 2021b. Francolin and spurfowl taxonomy revisited: Letter to the Editor. African Birdlife 9: 8–10.
   Google Scholar
• Hustler K. 2021. Taxonomic revision of francolins – reflections from a central African perspective. Ostrich 92: 324–328. https://doi.org/10.2989/00306525.2021.2005705
   Web of Science ®Google Scholar
• Irwin MPS. 1981. The birds of Zimbabwe. Harare: Quest.
   Google Scholar
• Johnsgard PA. 1988. The quails, partridges, and francolins of the world. Oxford: Oxford University Press.
   Google Scholar
• Kimball RT, Braun EL, Zwartjies PW, Crowe TM, Ligon JD. 1999. A molecular phylogeny of the pheasants and partridges suggests that these lineages are not monophyletic. Molecular Phylogenetics and Evolution 11: 38–54. https://doi.org/10.1006/mpev.1998.0562
   PubMed Web of Science ®Google Scholar
• Kimball RT, Hosner PA, Braun EL. 2021. A phylogenomic supermatrix of Galliformes (Landfowl) reveals biased branch lengths. Molecular Phylogenetics and Evolution 158. https://doi.org/10.1016/j.ympev.2021.107091
   Google Scholar
• Little R. 2021. Gamebirds of Africa – guineafowls, francolins, spurfowls, quails, sandgrouse & snipes. Johannesburg: Jacana Media.
   Google Scholar
• Little R, Crowe TM. 2011. Gamebirds of southern Africa (2nd edn). Cape Town: Random House Struik.
   Google Scholar
• Mackworth-Praed CW, Grant CHB. 1952. African handbook of birds, Series 1, volume 1: Birds of eastern and north-eastern Africa. London: Longmans.
   Google Scholar
• Mackworth-Praed CW, Grant CHB. 1962. African handbook of birds, Series 2, volume 2: Birds of the southern third of Africa. London: Longmans.
   Google Scholar
• Mackworth-Praed CW, Grant CHB. 1970. African handbook of birds, Series 3, volume 1: Birds of west central and western Africa. London: Longmans.
   Google Scholar
• Mandiwana-Neudani TG. 2013. Taxonomy, phylogeny and biogeography of francolins (‘Francolinus‘ spp.), Aves: order Galliformes, family Phasianidae. PhD thesis, University of Cape Town, South Africa.
   Google Scholar
• Mandiwana-Neudani TG, Kopuchian C, Louw G, Crowe TM. 2011. A study of gross morphological and histological syringeal features of true francolins (Galliformes: Francolinus, Scleroptila, Peliperdix and Dendroperdix spp.) and spurfowls (Pternistis spp.) in a phylogenetic context. Ostrich 82: 115–127. https://doi.org/10.2989/00306525.2011.603465
   Web of Science ®Google Scholar
• Mandiwana-Neudani TG, Bowie RCK, Hausberger M, Henry L, Crowe TM. 2014. Taxonomic and phylogenetic utility of variation in advertising calls of Francolins and Spurfowls (Galliformes: Phasianidae). African Zoology 49: 54–82. https://doi.org/10.1080/15627020.2014.11407620
   Web of Science ®Google Scholar
• Mandiwana-Neudani TG, Little RM, Crowe TM, Bowie RCK. 2019a. Taxonomy, phylogeny and biogeography of ‘true’ Francolins: Galliformes, Phasianidae, Phasianinae, Gallini; Francolinus, Ortygornis, Afrocolinus gen. nov., Peliperdix and Scleroptila spp. Ostrich 90: 191–221. https://doi.org/10.2989/00306525.2019.1632954
   Web of Science ®Google Scholar
• Mandiwana-Neudani TG, Little RM, Crowe TM, Bowie RCK. 2019b. Taxonomy, phylogeny and biogeography of African spurfowls, Galliformes, Phasianidae, Phasianinae, Coturnicini: Pternistis spp. Ostrich 90: 145–172. https://doi.org/10.2989/00306525.2019.1584925
   Web of Science ®Google Scholar
• Mandiwana-Neudani TG, Crowe TM, Little RM, Bowie RCK. 2021. The importance of adopting an integrative taxonomy framework in species delimitation: response to Hunter et al. (2021). Ostrich 92: 162–167. https://doi.org/10.2989/00306525.2021.1898170
   Web of Science ®Google Scholar
• Mayr E. 1963. Animal species and evolution. Cambridge: Harvard University Press. https://doi.org/10.4159/harvard.9780674865327
   Google Scholar
• McGowan PJK, Kirwan GM. 2020. Red-necked Francolin (Pternistis afer), version 1.0. In: del Hoyo J, Elliott A, Sargatal J, Christie DA, de Juana E (eds), Birds of the world. Ithaca: Cornell Laboratory of Ornithology.
   Google Scholar
• Milstein P le S, Wolff SW. 1987. The over-simplification of our ‘francolins’. South African Journal of Wildlife Research Suppl 1: 58–65.
   Google Scholar
• Moritz C. 1994. Defining ‘Evolutionary Significant Units’ for conservation. Trends in Ecology & Evolution 9: 373–375. https://doi.org/10.1016/0169-5347(94)90057-4
   PubMed Web of Science ®Google Scholar
• Ottenburghs J. 2019. Multispecies hybridization in birds. Avian Research 10: 20. https://doi.org/10.1186/s40657-019-0159-4
   Web of Science ®Google Scholar
• Peters JL. 1934. Checklist of birds of the world, vol. II. Cambridge: Harvard University Press.
   Google Scholar
• Randi E, Fusco G, Lorenzini R, Crowe TM. 1991. Phylogenetic relationships and rates of allozyme evolution within the Phasianidae. Biochemical Systematics and Ecology 19: 213–221. https://doi.org/10.1016/0305-1978(91)90005-K
   Web of Science ®Google Scholar
• Reichenow A. 1889. Uber einevogelsammlung aus ostafrika. Journal für Ornithologie Séries 4: 270.
   Google Scholar
• Roberts A. 1924. Synoptic checklist of the birds of South Africa. Annals of the Transvaal Museum 10: 89–195.
   Google Scholar
• Roberts A. 1947. A new Pternistis from Salisbury, S Rhodesia. Ostrich 18: 197. doi: 10.1080/00306525.1947.9639266
   Google Scholar
• Ruse M. 1975. Darwin’s debt to philosophy: an examination of the influence of the philosophical ideas of John F.W. Herschel and William Whewell on the development of Charles Darwin’s theory of evolution. Studies in History and Philosophy of Science Part A 6: 159–181. https://doi.org/10.1016/0039-3681(75)90019-9
   PubMed Web of Science ®Google Scholar
• Ruse M. 1979. Falsifiability, consilience, and systematics. Systematic Zoology 29: 530–536. https://doi.org/10.2307/2412565
   Google Scholar
• Sibley CG, Monroe BL Jr. 1990. Distribution and taxonomy of birds of the world. New Haven, Connecticut: Yale University Press.
   Google Scholar
• Snow DW. 1978. An atlas of speciation in African non-passerine birds. London: British Museum.
   Google Scholar
• Töpfer T, Podsiadlowski L, Gedeon K. 2014. Rediscovery of the Black-fronted Francolin Pternistis (castaneicollis) atrifrons (Conover, 1930) (Aves: Galliformes: Phasianidae) with notes on biology, taxonomy and conservation. Vertebrate Zoology 64: 261–271.
   Web of Science ®Google Scholar
• Turner D, Crowe TM. 2022. Comments on the taxonomy, common names and evolutionary relationships of some East African ‘Red-winged’ francolins currently recognised as Scleroptila g. gutturalis/lorti/archeri, Scleroptila s. shelleyi and S. s. uluensis. Scopus 42: 59–63.
   Google Scholar
• van Niekerk JH, Mandiwana-Neudani TG. 2018. The phylogeny of francolins (Francolinus, Dendroperdix, Peliperdix and Scleroptila) and spurfowls (Pternistis) based on chick plumage (Galliformes: Phasianidae). Avian Research 9: article 2. https://doi.org/10.1186/s40657-017-0093-2
   Google Scholar
• van Niekerk JH, Megía-Palma R, Forcina G. 2021. Call variation of Crested Francolin (Dendroperdix sephaena) across Africa defies the effects of spatial gradients. Ibis 163: 1409–1424. https://doi.org/10.1111/ibi.12945
   Web of Science ®Google Scholar
• Wang N, Kimball RT, Braun EL, Liang B, Zhang Z. 2013. Assessing phylogenetic relationships among Galliformes: a multigene phylogeny with extended taxon sampling. PLoS ONE 8: e:64312. doi: 10.1371/journal.pone.0064312
   Web of Science ®Google Scholar
• Ward RD. 2009. DNA barcode divergence among species and genera of birds and fishes. Molecular Ecology Resources 9: 1077–1085. https://doi.org/10.1111/j.1755-0998.2009.02541.x
   PubMed Web of Science ®Google Scholar
• Wilson EO, Brown WL. 1953. The subspecies concept and its taxonomic application. Systematic Biology 2: 97–111.
   Google Scholar
• Wolters HE. 1975–82. Die Vogelarten der Erde. Hamburg: Paul Parey.
   Google Scholar
• Zelnio K. 2012. Species concepts. Scientific American EvoEcoLab, 20 April 2012. https//:blogs.scientificamerican.com/evo-eco-lab/species-concepts.
   Google Scholar