Taxonomy, phylogeny and biogeography of African spurfowls Galliformes, Phasianidae, Phasianinae, Coturnicini: Pternistis spp.

Abstract

Afro-Asiatic perdicine galliform birds, commonly and inconsistently referred to as francolins, spurfowls and partridges, have contentious taxonomic and phylogenetic histories. In a widely followed monograph, Hall combined two putative monophyletic, but taxonomically unnamed, clades comprising 28 perdicine species known as ‘francolins’ or fisante in South Africa and 13 additional quail-like species (partridges or patryse) into a single genus, Francolinus, which was the largest genus within the Galliformes. Furthermore, she partitioned fisante + patryse into eight, also formally unnamed, putative monophyletic ‘Groups’ and speculated on the phylogenetic affinities of four ‘Unplaced’ species. We investigate fisante using combined morphological, vocalisation and DNA-based evidence and produce a comprehensive revision of fisante taxonomy and phylogeny, a stable classification system and common terminology, and hypotheses vis-à-vis eco-biogeographical processes that promoted their speciation and cladogenesis. Three of Hall’s four Groups of fisante sensu stricto (Montane, Scaly and Vermiculated) are para- or polyphyletic evolutionary grades. Only her Bare-throated Group emerges as monophyletic. We recommend the recognition of only one genus, Pternistis, and the use of ‘spurfowl’ as its collective common name. The proposed new system recognises 25 species, elevating two of Hall’s subspecies (schuetti and cranchii) to species level and reduces the number of subspecies taxa from 59 to 16. Several species pairs of spurfowls, most notably P. afer and P. cranchii, hybridise in para/sympatry. At least one Bare-throated spurfowl, P. rufopictus, may be the product of stabilised hybridisation between P. afer and/or P. cranchii and P. leucoscepus, and hybridisation between proto-taxa in the Montane and Scaly grades may undermine nodal support for basal spurfowl clades.

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Ostrich
Journal of African Ornithology
ISSN: 0030-6525 (Print) 1727-947X (Online) Journal homepage: https://www.tandfonline.com/loi/tost20
Taxonomy, phylogeny and biogeography of African
spurfowls Galliformes, Phasianidae, Phasianinae,
Coturnicini: Pternistis spp.
Tshifhiwa G Mandiwana-Neudani, Robin M Little, Timothy M Crowe & Rauri
CK Bowie
To cite this article: Tshifhiwa G Mandiwana-Neudani, Robin M Little, Timothy M Crowe & Rauri
CK Bowie (2019) Taxonomy, phylogeny and biogeography of African spurfowls Galliformes,
Phasianidae, Phasianinae, Coturnicini: Pternistis spp., Ostrich, 90:2, 145-172
To link to this article: https://doi.org/10.2989/00306525.2019.1584925
Published online: 04 Jun 2019.
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Ostrich 2019, 90(2): 145–172
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OSTRICH
ISSN 0030–6525 EISSN 1727-947X
https://doi.org/10.2989/00306525.2019.1584925
Ostrich is co-published by NISC (Pty) Ltd and Informa UK Limited (trading as Taylor & Francis Group)
Taxonomy, phylogeny and biogeography of African spurfowls Galliformes,
Phasianidae, Phasianinae, Coturnicini: Pternistis spp.
Tshifhiwa G Mandiwana-Neudani1,2 , Robin M Little2* , Timothy M Crowe2 and Rauri CK Bowie2,3
1 Department of Biodiversity, University of Limpopo, Sovenga, South Africa
2 FitzPatrick Institute of African Ornithology, DST-NRF Centre of Excellence, Department of Biological Sciences, University of
Cape Town, Cape Town, South Africa
3 Museum of Vertebrate Zoology and Department of Integrative Biology, University of California, Berkeley, CA 94720-3160, USA
* Corresponding author, email: rob.little@uct.ac.za
Afro-Asiatic perdicine galliform birds, commonly and inconsistently referred to as francolins, spurfowls and
partridges, have contentious taxonomic and phylogenetic histories. In a widely followed monograph, Hall
combined two putative monophyletic, but taxonomically unnamed, clades comprising 28 perdicine species known
as ‘francolins’ or fisante in South Africa and 13 additional quail-like species (partridges or patryse) into a single
genus, Francolinus, which was the largest genus within the Galliformes. Furthermore, she partitioned fisante +
patryse into eight, also formally unnamed, putative monophyletic ‘Groups’ and speculated on the phylogenetic
affinities of four ‘Unplaced’ species. We investigate fisante using combined morphological, vocalisation and
DNA-based evidence and produce a comprehensive revision of fisante taxonomy and phylogeny, a stable
classification system and common terminology, and hypotheses vis-à-vis eco-biogeographical processes that
promoted their speciation and cladogenesis. Three of Hall’s four Groups of fisante sensu stricto (Montane,
Scaly and Vermiculated) are para- or polyphyletic evolutionary grades. Only her Bare-throated Group emerges
as monophyletic. We recommend the recognition of only one genus, Pternistis, and the use of ‘spurfowl’ as its
collective common name. The proposed new system recognises 25 species, elevating two of Hall’s subspecies
(schuetti and cranchii) to species level and reduces the number of subspecies taxa from 59 to 16. Several species
pairs of spurfowls, most notably P. afer and P. cranchii, hybridise in para/sympatry. At least one Bare-throated
spurfowl, P. rufopictus, may be the product of stabilised hybridisation between P. afer and/or P. cranchii and
P. leucoscepus, and hybridisation between proto-taxa in the Montane and Scaly grades may undermine nodal
support for basal spurfowl clades.
Taxonomie, phylogénie et biogéographie des Oiseaux Galliformes africains à éperons apparentant
aux Phasianidae, Phasianinae, Coturnicini: Pternistis spp.
Les oiseaux galliformes «Perdicidae» afro-asiatiques couramment et inconsistamment appelés francolins, faisons
et perdrix ont une histoire taxonomique et phylogénétique litigieuse. Dans une monographie largement suivie,
Hall associe deux clades supposés monophylétiques, mais non identifiables sur le plan taxonomique, comprenant
28 espèces de Perdicidae connues sous le nom de «francolins» ou fisante « faisons » en Afrique du Sud et
13 autres espèces ressemblant à des cailles (perdrix ou patryse) en un seul le genre, Francolinus, qui était le plus
grand genre des Galliformes. En outre, elle a divisé fisante + patryse en huit groupes, également appelés «groupes»
monophylétiques supposés, et a spéculé sur les affinités phylogénétiques de quatre espèces «non placées».
Nous étudions fisante en combinant des preuves morphologiques, vocales et basées sur l’ADN et produisons
une révision complète de la taxonomie et de la phylogénie des fisante, un système de classification stable et
une terminologie commune, ainsi que des hypothèses sur les processus éco-biogéographiques favorisant leur
spéciation et leur cladogénèse. Trois des quatre groupes de fisante sensu stricto de Hall (Francolin du cap, écaillé
et Criard) sont des grades évolutifs para- ou polyphylétiques. Seul le groupe à gorge nue se révèle monophylétique.
Nous recommandons la reconnaissance d’un seul genre, Pternistis, et l’utilisation de «spurfowl» comme nom
commun anglais. Le nouveau système proposé reconnaît 25 espèces, élève deux espèces de la sous-espèce de Hall
(schuetti et cranchii) au niveau de l’espèce et réduit le nombre de taxons de la sous-espèce de 59 à 16. Plusieurs
couples d’espèces de francolin, notamment P. afer et P. cranchii, se croisent en para / sympatrie. Au moins un
francolin à gorge nue, P. rufopictus, peut être le produit d’une hybridation stabilisée entre P. afer et / ou P. cranchii
et P. leucoscepus, et une hybridation entre des proto-taxons des grades Francolin du cap et F. écaillé peut nuire au
support nodal pour les clades basiques des francolins.
Keywords: biogeography, gamebird, phylogeny, Pternistis, spurfowl, taxonomy
Published online 04 Jun 2019

Mandiwana-Neudani, Little, Crowe and Bowie
146
Stable, evidence-based, philosophically sound taxonomies
and phylogenies and plausible biogeographical scenarios
are essential for comparative and conservation biology
(Cracraft 1997; Ladle and Whittaker 2011; Pellens
and Grandcolas 2016). During much of the twentieth
century, Afro-Asiatic francolins, spurfowls and several
other partridge-like perdicine galliforms had conten-
tious taxonomic histories paralleled by the inconsistent
use of common names. Because of striking autapomor-
phic differences in plumage, vocalisations and ecology,
a plethora of small, even monotypic, genera were
described. For example, Roberts (1924) recognised seven
genera of ‘francolins’ and ‘partridges’ within southern
Africa alone: Dendroperdix, Ortygornis, Scleroptila,
Peliperdix, Chapinortyx, Chaetopus and Pternistis. Other
generic epithets have been used inconsistently to group
Afro-Asian francolins, spurfowls and partridges. Peters
(1934) and Mackworth-Praed and Grant (1952, 1962,
1970) recognised two genera for the same species taxa.
They used Francolinus for the bulk of species taxa that
have feathered throats, and restricted Pternistis only to
those that have unfeathered throats with brightly coloured
skin, also distinguishing them with the common name
spurfowls. Alternatively, Wolters (1975–82) recognised six
genera: Francolinus, Ortygornis, Dendroperdix, Peliperdix,
Scleroptila and Pternistis, using Francolinus-through-
Scleroptila to group smaller, short-single-spurred taxa with
quail-like dorsal plumage, and employed Pternistis more
‘generously’ to incorporate larger, long-multiple-spurred
taxa with streaked and/or vermiculated backparts, irrespec-
tive of throat plumage. With regards to common names,
Wolters (1975–82) also referred to bare-throated Pternistis
spp. as ‘spurfowls’ and to those with feather-throated taxa
as ‘francolins’. Also confusing is the genus Ortygornis,
which Roberts (1924) used for the African Coqui Partridge
O. coqui, Wolters (1975–82) for the Asiatic Grey Francolin
pondicerianus, and Darwin (1871) for the Asiatic Swamp
Francolin gularis. Chapin (1926) further proposed that
Nahan’s Francolin Francolinus nahani be removed from
Francolinus and placed in a monotypic genus Acentrortyx
because it lacks tarsal spurs. Complicating matters further,
geographically variable species were split into more than
160 clinal and idiosyncratic subspecies (Peters 1934).
Finally, all these taxa were embedded within a highly
polyphyletic Perdicinae subfamily within the Phasianinae
(Kimball et al. 1999; Crowe et al. 2006; Wang et al. 2017).
The history of the use of common names for these taxa is
outlined in detail by Crowe and Little (2004).
In her seminal monograph dealing with some of this
‘taxonomic turbulence’, Hall (1963) lumped 41 species
of Afro-Asiatic francolins, spurfowls and partridges into a
single, monophyletic genus, Francolinus, the largest genus
within the Galliformes, and one of the largest genera in Aves
(Bock and Farrand 1980). She proposed that they should
all bear the common name ‘francolin’. Nevertheless, Hall
(1963) partitioned francolins into two major, taxonomically
unnamed, putatively monophyletic clades, comprising eight
also putatively monophyletic and taxonomically unnamed
‘Groups’ of species and speculated on the phylogenetic
affinities of four ‘Unplaced’ species, including Acentrortyx
nahani. Her two clades correspond closely with what
southern African Griqua peoples and Afrikaans-speaking
wingshooters and farmers call fisante (her Spotted,
Bare-throated, Montane, Scaly and Vermiculated Groups,
plus the ‘unplaced’ gularis and nahani) and patryse (Scaly,
Red-tailed and Red-winged Groups and the ‘unplaced’
pondicerianus and lathami) (Milstein and Wolff 1987). Hall
(1963) also recognised 100 Francolinus subspecies – 59 for
fisante and 41 for patryse (Table 1). Many of these taxa are
endemic or near endemic to well-established Afrotropical
avian biogeographical zones (Crowe and Crowe 1982,
1985) and thus could be useful biogeographical indicators or
model species for conservation planning and action. Hall’s
(1963) species-level classification is still employed in the
IOC World Bird List (Gill and Donsker 2018).
Crowe and Crowe’s (1985) phylogenetic analysis of
morpho-behavioural characters questioned, but did not
refute, the monophyletic status of the genus Francolinus
as defined in Hall (1963). Nevertheless, it supported the
monophyly of at least four of Hall’s groups (Montane,
Red-tailed, Red-winged and Spotted), whereas the other
groups (Bare-throated, Scaly, Striated and Vermiculated)
were para- or polyphyletic. Analyses of mitochondrial
DNA (mtDNA) restriction fragment length polymorphisms
and morpho-behavioural characters (Crowe et al. 1992)
recommended that F. streptophorus be moved from the
Striated to the Red-winged Group, but still retained a
monophyletic Francolinus.
Bloomer and Crowe’s (1998) expanded phylogenetic
analyses of Francolinus sensu Hall (1963) to include
mtDNA cytochrome b sequences and morpho-behavioural
characters. Although analyses of the two data partitions
separately produced relatively poorly resolved cladograms,
analysis of both partitions combined produced a
well-resolved cladogram that identified two clades. One
clade comprised a subset of fisante (Hall’s Montane, Scaly
and Vermiculated Groups) minus the Spotted Group and
F. gularis. The second clade comprised the patryse plus
the Spotted francolins and gularis. The combined-evidence
cladogram also challenged the monophyly of Francolinus
sensu Hall (1963) and suggested that fisante sensu stricto
link phylogenetically with Alectoris partridges and Old World
quails Coturnix spp. None of these studies decisively placed
the enigmatic Francolinus/Acentrortyx nahani taxonomically
or phylogenetically.
Crowe et al. (2006) investigated all putative Francolinus
spp. using a broader range of mtDNA, nuclear DNA (nDNA)
and morpho-behavioural characters and demonstrated
decisively that Francolinus sensu Hall (1963) is not
monophyletic. Fisante sensu stricto (Hall’s Montane, Scaly
and Vermiculated Groups) were confirmed to link with Old
World ‘true’ quails (Coturnix spp.), Alectoris ‘partridges’
and several other Old World perdicines that are sister to
Ammoperdix ‘partridges’ and Perdicula ‘quails’. Patryse link
with Gallus and Bambusicola spp. Crowe et al. (2006) placed
nahani phylogenetically distant from francolins as sister to
another enigmatic African perdicine, Ptilopachus petrosus,
within an expanded Odontophoridae (Cohen et al. 2012).
Introduction

Ostrich 2019, 90(2): 145–172
147
Group English name Species and subspecies
Clade 1
Spotted Black Francolin Francolinus francolinus (Linnaeus, 1766)
francolinus (caucasicus, sarudyni, billypaynei); arabistanicus Zarudny and Härms, 1913; henrici
Bonaparte, 1856 (festinus, bogdanovi); asiae Bonaparte, 1856 (parkerae); melanotus Hume, 1888
Painted Francolin Francolinus pictus (Jardine and Selby, 1828)
pictus; pallidus (Grey, 1831); watsoni Legge, 1880
Bare-throated Yellow-necked Francolin Francolinus leucoscepus (Gray, GR, 1867)
leucoscepus; infuscatus (Cabanis, 1868) (holtemulleri, muhamed-ben-abdullah, keniensis,
kilimensis, tokora, oldowai)
PTERNISTIS Wagler, 1832)
Grey-breasted Francolin Francolinus rufopictus (Reichenow, 1887)
PTERNISTIS
Swainson’s Francolin Francolinus swainsonii (Smith, A, 1836)
swainsonii; lundazi (White,1947); damarensis (Roberts, 1931); gilli (Roberts, 1932)
PTERNISTIS
Red-necked Francolin Francolinus afer (Statius Muller, 1776)
PTERNISTIS
‘Black and white’ taxa: afer (palliditectus); castaneiventer (Gunning and Roberts, 1911) (krebsi);
notatus (Roberts, 1924); lehmanni (Roberts, 1931); humboldtii (Peters, 1854) (swynnertoni);
melanogaster (Neumann, 1898) (loangwae); leucopareus Fischer and Reichenow, 1884
‘Vermiculated’ taxa: cranchii (Leach, 1818) (punctulata, nyanzae); intercedens (Reichenow,
1909); harterti (Reichenow, 1909)
PTERNISTIS
Montane Swierstra’s Francolin Francolinus swierstrai (Roberts, 1929)
PTERNISTIS
Mount Cameroon Francolin Francolinus camerunensis Alexander, 1909
PTERNISTIS
Handsome Francolin Francolinus nobilis (Reichenow, 1908)
nobilis; chapini (Mackworth-Praed and Grant, 1934)
PTERNISTIS
Jackson’s Francolin Francolinus jacksoni Ogilvie-Grant, 1891
jacksoni (gurae); pollenorum (Meinertzhagen, 1937)
PTERNISTIS
Chestnut-naped Francolin Francolinus castaneicollis Salvadori, 1888
castaneicollis (bottegi, gofanus); ogoensis Mackworth-Praed, 1920; kaffanus (Grant and
Mackworth-Praed, 1934) (patrizii); atrifrons Conover, 1930
PTERNISTIS
Erckel’s Francolin Francolinus erckelii (Rüppell, 1835)
erckelii; pentoni Mackworth-Praed, 1920
PTERNISTIS
Djibouti Francolin Francolinus ochropectus Dorst and Jouanin, 1952
PTERNISTIS
Scaly Scaly Francolin Francolinus squamatus Cassin, 1857
squamatus (whitei); schuetti Cabanis, 1880 (tetraoninus, zappeyi, dowashanus); maranensis
Mearns, 1910 (kapitensis, keniensis, chyuluensis); usambarae Conover, 1928; uzungwensis
Bangs and Loveridge, 1931; doni (Benson, 1939)
PTERNISTIS
Ahanta Francolin Francolinus ahantensis Temminck, 1854
ahantensis; hopkinsoni (Bannerman, 1934)
PTERNISTIS
Grey-striped Francolin Francolinus griseostriatus Ogilvie-Grant, 1890
PTERNISTIS
Vermiculated Double-spurred Francolin Francolinus bicalcaratus (Linnaeus, 1766)
bicalcaratus; ayesha Hartert, 1917; adamauae Neumann, 1915; ogilvie-granti Bannerman, 1922
(molunduensis); thornei Ogilvie-Grant, 1902
PTERNISTIS
Clapperton’s Francolin Francolinus clappertoni Children and Vigors, 1826
clappertoni (tschadensis); gedgii Ogilvie-Grant, 1891 (cavei); heuglini Neumann, 1907; sharpii
Ogilvie-Grant, 1892 (testis); konigseggi von Madarász, 1914; nigrosquamatus Neumann, 1902
PTERNISTIS
Heuglin’s Francolin Francolinus icterorhynchus Heuglin, 1863
icterorhynchus (grisecens); dybowskii Oustalet, 1892 (emini, ugandensis)
PTERNISTIS
Harwood’s Francolin Francolinus harwoodi Blundell and Lovat, 1899
PTERNISTIS
Table 1: Francolinus Stephens 1819 – clades, groups, species and subspecies recognised by Hall (1963). Synonymised subspecies are in
parentheses, pre-Hall (1963) alternative generic epithets are below in upper-case letters

Mandiwana-Neudani, Little, Crowe and Bowie
148
Group English name Species and subspecies
Vermiculated Hildebrandt’s Francolin Francolinus hildebrandti Cabanais, 1878
hildebrandti (helleri); altumi Fischeri and Reichenow, 1884; johnstoni Shelley, 1894 (grotei, lindi)
PTERNISTIS
Natal Francolin Francolinus natalensis Smith, A, 1834
natalensis (thamnobium); neavei Mackworth-Praed, 1920
PTERNISTIS, CHAETOPUS (Swainson, 1837)
Hartlaub’s Francolin Francolinus hartlaubi Bocage, 1869
(bradeldi, crypticus, ovambensis)
PTERNISTIS, CHAPINORTYX Roberts, 1928
Cape Francolin Francolinus capensis (Gmelin, 1789)
PTERNISTIS, CHAETOPUS
Red-billed Francolin Francolinus adspersus Waterhouse, 1838
(Kalahari)
PTERNISTIS, CHAETOPUS
Unplaced Swamp Francolin Francolinus gularis (Temminck, 1815)
ORTYGORNIS (Reichenbach, 1853)
Nahan’s Francolin Francolinus nahani Dubois, 1905
ACENTRORTYX Chapin, 1928
Clade 2
Striated Crested Francolin Francolinus sephaena (Smith, A, 1836)
sephaena (zuluensis); spilogaster Salvadori, 1888; rovuma Gray, GR, 1867 (kirkii); grantii Hartlaub,
1865 (schoanus, ochrogaster, delutescens, jubaensis); zambesiae Mackworth-Praed, 1920
(thompsoni, chobiensis, mababiensis)
DENDROPERDIX Roberts, 1924
Ring-necked Francolin Francolinus streptophora Ogilvie-Grant, 1891
SCLEROPTILA Blyth, 1849
Red-tailed Coqui Francolin Francolinus coqui (Smith, 1836)
coqui (stuhlmanni, campbelli, lynesi); vernayi (Roberts, 1932); hoeschianus (Stresemann, 1937);
angolensis (Rothschild, 1902); kasiacus (White, 1945); ruahdae (Monard, 1934); hubbardi (Ogilvie-
Grant, 1895); thikae (Grant and Mackworth-Praed, 1934); maharao (Sclater, 1927); spinetorum
Bates, 1928
ORTYGORNIS, PELIPERDIX
White-throated Francolin Francolinus albogularis Hartlaub, 1854
albogularis; buckleyi (Ogilvie-Grant, 1892) (gambagae); meinertzhageni (White, 1944); dewittei
(Chapin, 1937)
PELIPERDIX
Schlegel’s Francolin Francolinus schlegelii Heuglin, 1863
PELIPERDIX
Red-winged Shelley’s Francolin Francolinus shelleyi Ogilvie-Grant, 1890
shelleyi (trothae, sequestris); whytei Neumann, 1908; uluensis Ogilvie-Grant, 1892 (macarthuri)
SCLEROPTILA
Grey-winged Francolin Francolinus africanus (Latham, 1790)
SCLEROPTILA
Orange River Francolin Francolinus levaillantoides (Smith, A, 1836)
North: gutturalis (Rüppell, 1835) (eritreae); archeri (Sclater, 1927) (stantoni, friedmanni); lorti
(Sharpe, 1897)
South: levaillantoides (gariepensis, ludwigi); jugularis (Büttikofe, 1889) (cunenensis, stresemanni);
kalaharica (Roberts, 1932) (langi); pallidior (Neumann, 1908) (wattii)
SCLEROPTILA
Red-winged Francolin Francolinus levaillantii (Valenciennes, 1825)
levaillantii; crawshayi Ogilvie-Grant, 1896; kikuyuensis Ogilvie-Grant, 1897 (mulemae, adolriederii,
benguellensis, clayi, mombolensis)
SCLEROPTILA
Finsch’s Francolin Francolinus nschi Bocage, 1881
SCLEROPTILA
Moorland Francolin Francolinus psilolaemus Gray, 1867
psilolaemus; theresae (Meinertzhagen, 1937); elgonensis Ogilvie-Grant, 1891; ellenbecki Erlanger,
1905 (fricki)
SCLEROPTILA
Unplaced Grey Francolin Francolinus pondicerianus (Gmelin, 1789)
pondicerianus; mecranensis Sarudny and Härms, 1913; ceylonensis Whistler, 1941; interpositus
Hartert, 1917 (paganus, titar, prepositus)
ORTYGORNIS
Latham’s Francolin Francolinus lathami Hartlaub, 1854
lathami; schubotzi Reichenow, 1915
PELIPERDIX
Table 1: (cont.)

Ostrich 2019, 90(2): 145–172
149
These relationships have been corroborated by several
additional studies, including large multi-locus phylogenies
(Hosner et al. 2016; Cai et al. 2017; Wang et al. 2017).
Crowe et al. (2006) and Little (2016a) followed Little
and Crowe (2011) and Crowe and Little (2004) and
recommended that the now fisante sensu stricto be known
commonly as ‘spurfowls’ and the patryse sensu lato as
‘francolins’. Spurfowls occupy bushy and wooded habitats,
regularly take refuge and may roost in trees, prefer to run
rather than flush to escape danger, have raucous, grating
or cackling, pheasant-like calls, tend not to respond to
play-back of their advertisement calls, weigh >400 g, have
streaked or vermiculated dorsal plumage and black, orange
or red tarsi armed with two prominent spurs. Spurfowl
chicks have a crown surmounted by a broad dark patch.
Many species are capable of withstanding responsible
wingshooting and few appear in the IUCN Red List of
Threatened Species. ‘True’ francolins occupy grassland
habitats grading into bush and thicket, are ground-dwelling/
roosting, sit tight when disturbed flushing (often in coveys)
only when pressed, have musical, whistling calls, vocally
respond to their play-back (van Nierkerk 2010) and have
yellow tarsi armed with a short, single spur. Francolin chicks
have a crown surmounted by a narrow dark patch, bordered
by several dark and light stripes. Only the Grey-winged
Francolin Francolinus/Scleroptila afra is wingshot sustain-
ably and populations of several species are threatened
by agricultural practices (e.g. overgrazing and frequent
‘remedial’ burning of grasslands).
Here, we revise the taxonomy, phylogenetics and
biogeography of fisante sensu stricto (Hall’s Vermiculated,
Montane, Scaly and Bare-throated Groups) through
the investigation of further expanded morphological
(Mandiwana-Neudani et al. 2011), mtDNA and nDNA
sequence data, and characters derived from vocalisations
(Mandiwana-Neudani et al. 2014).
Materials and methods
Taxon sampling
The specimens studied morphologically include 88
putative African spurfowl (fisante) species and subspecies
(Appendix 1), and cover all putative taxa examined and
recognised by Hall (1963) at the then British Museum
of Natural History (now The Natural History Museum at
Tring, UK), supplemented by a broader array of material
from other major natural history museums mentioned in
the Acknowledgements.
Taxa from which DNA sequence data were obtained are
annotated, with four mitochondrial markers: Cytochrome
b (CYTB; 1 143 base pairs), Control region (CR; 820 bp),
NADH dehydrogenase subunit 2 (ND2; 1 041 bp) and 12S
rRNA (12S; 706 bp); and three nuclear DNA markers:
Ovomucoid G (OVO-G; 449 bp), Glyceraldehyde-3-
phosphodehydrogenase (GAPDH; 361 bp) and Trans
Globulin Growth Factor Beta2 intron-5 (TGFB; 596 bp)
(Appendices 2 and 3).
Taxonomic approach
Taxonomy involves the discovery, description, naming and
classification of taxa at all levels. Generally, it focuses on
the phylogenetically terminal components of biodiversity,
species and subspecies (Dayrat 2005). Great emphasis has
been placed on the relative merits of different operational
‘species concepts’ (Zachos 2016). We employ a species
concept that emphasises their ontological status and
processes by which species arise (de Queiroz 2007).
Species
Empirically and conceptually, we employ the Consilience
Species Concept (Crowe et al. 1994; Bowie et al. 2016;
Crowe et al. 2016) that requires evidence from combined,
multiple, consilient, independent data sets to discover and
diagnose taxa based on the reinforcing convergence of
complementary characters. In short, species are:
(1) reciprocally monophyletic groups of specimens that
are qualitatively similar in terms of suites of diagnostic
and consilient morpho-vocalisation and DNA-sourced
characters;
(2) morphologically, genetically, behaviourally and often
ecologically distinct entities discernible from evolutionary
near relatives; and
(3) geographically ‘meaningfully’ distributed, e.g. in relation
to past/present vegetation types and/or topography and
well-established African biogeographical provinces/
regions (Crowe and Crowe 1982, 1985; Linder et al.
2014).
Our goal is to identify evolutionarily independent species
lineages buffered from the homogenising effects of tokogeny
(Wheeler and Platnick 2000) and confounding effects of
horizontal gene flow and incomplete lineage sorting (Pruett
and Winker 2010; Zachos 2016). This is important for
spurfowls since ‘hybridisation’ between them is reportedly
common and widespread (Hall 1963; McCarthy 2006).
Subspecies
Subspecies are groups of populations delineated by
geographically steep, congruent clinal variation in multiple
characters where their distributional ranges meet (Crowe
1978; Winker 2010). The ‘suture’ zones of parapatry
are characterised by morphologically intermediate and
genetically heterogeneous individuals with ‘shuffled’,
non-diagnosing characters that appear to reflect
interbreeding between largely allo- and parapatric popula-
tions. Thus, our goal for subspecies is for them to reflect
phylogeographic structure and genealogy characterised
by consilient, potentially ecologically adaptive anatomical
and behavioural differences. Subspecies in one clade may
have geographically similar distributions to those in other
clades comprised of genetically diagnosable Evolutionarily
Significant Units (Casacci et al. 2014) or even full species
(Crowe and Crowe 1985). Putative ‘francolin’ subspecies
deemed to be clinal or local variants are not supported.
In practice within this study, morphologically, behaviour-
ally and genetically distinct taxa were evaluated as putative
species if there was little morphological evidence of inter-
taxon interbreeding and molecular genetic divergence
from their sister taxon in unweighted, uncorrected, overall
molecular sequence divergence of mitochondrial CYTB
(Swofford 2002) exceeded 1.5%. By way of comparison,
well-marked subspecies of Africa’s most widespread and
well-studied galliform species, the Helmeted Guineafowl

Mandiwana-Neudani, Little, Crowe and Bowie
150
Numida meleagris (Crowe 1978), are 1% to 1.4% CYTB
divergent (van Alphen-Stahl 2005).
Morphology
The spurfowl basic body plan was divided into discrete
sections (Figure 1) and variation in patterning and
colouration was assessed for morphological characters
relating to plumage/integument colour/pattern (Table 2).
Measurements to the nearest 1 mm of the bill from the
cere, tail, tarsus and spur length were taken using a
Vernier calliper, the flattened wing was measured using a
wing-rule, and gap-coded states were assigned following
Archie (1985).
Vocalisations
Character information for vocalisations was extracted from
Mandiwana-Neudani et al. (2014).
Molecular samples and primers
Sample information for putative spurfowl taxa for which
DNA sequences were obtained are presented in Table 3.
Primers used in sequencing are listed in Tables 4 and 5.
The 1 143 bp CYTB was sequenced for all taxa included
in this study, whereas data for the other markers are
missing for some taxa. Contrary to earlier work (Crowe
et al. 1992; Bloomer and Crowe 1998), which focused on
few species, all putative species and most subspecies
attributed to African spurfowls were included (Appendix 1).
Some 72% of specimens sequenced in this study derived
from DNA extractions of toe-pad scrapes of museum
skins. As a result, only CYTB was sequenced for both
fresh and historical tissues, and the other six markers
were sequenced for species for which fresh tissues were
available. Due to the fragmented nature of the historical
sourced DNA, the CYTB gene for the toe-pads was
sequenced in multiple fragments (six for each sample)
using spurfowl-specific primers (Table 5).
C
HN
LN
B
UPTC UNTC
BE
BR
G
T
SOH
W
DORSAL VENTRAL LATERAL
Figure 1: Spurfowl body parts scored when generating plumage
characters. C = crown, HN = hind neck, LN = lower neck, B = back,
UPTC = upper tail coverts, T = throat, G = gorget, BR = breast, BE =
belly, UNTC = under tail coverts, SOH = side of head, W = wing
Character Character scores
1 Crown margins unmargined = 0; grey = 1; buff = 2; grey brown = 3
2 Nares black = 1; chestnut = 2; grey brown = 3; buff or white = 4; white = 5
3 Hind neck patterning unpatterned = 0; mottled = 1; streaked = 2
4 Hind neck base colour grey brown = 1; grey black = 2; grey chestnut = 3; rufous brown = 4; black = 5
5 Hind neck margins unmargined = 0; grey = 1; buff = 2; grey brown = 3
6 Lower neck patterning streaked = 1; mottled = 2; barred = 3
7–10 Back plumage plain = 0; streaked = 1 (7); mottled = 1 (8); vermiculated = 1 (9); barred = 1 (10)
11–13 Upper tail coverts plain = 0; barred = 1 (11); vermiculated = 1 (12); streaked = 1 (13)
14 Throat feathered = 1; yellow skin = 2; orange skin = 3; red skin = 4
15–18 Under tail coverts plain = 0; barred = 1 (15); streaked = 1 (16); vermiculated = 1 (17); mottled = 1 (18)
19 Bare skin around eye none = 0; red = 1; yellow = 2
20 Leg colour yellow = 1; red = 2; orange red = 3; orange = 4; olive green = 5; orange yellow = 6; black = 7
21 Number of spurs one = 1; two = 2
22 Wing length (mm) males <160 = 1; 160–180 = 2; 181–200 = 3; >200 = 4
23 Culmen length / Wing length (mm) <0.16 = 1; 0.17–0.2 = 2; >0.2 = 3
24 Tail length / Wing length (mm) <0.54 = 1; >0.54 = 2
25 Sexual dimorphism (plumage) absent = 0; present = 1
26 Sexual dimorphism (wing length) ♀ >0.9 of ♂ = 0; ♀ <0.9 of ♂ = 1
27 Vocalisation strophe duration (s) <0.3 = 1; 0.3–0.6 = 2; >0.6 = 3
28 Number of elements one = 1; two = 2; >2 = 3
29 Inter-element interval absent or indistinct = 1; distinct = 2
30 Cackle trill absent = 0; present = 1
31–32 Strophe character tonal = 0 (31); trill = 1 (32)
33 Ko-waaark advertisement call absent = 0; present = 1
Table 2: Thirty-three morpho-vocalisation characters with scoring criteria used for the phylogenetic analyses of spurfowls

Ostrich 2019, 90(2): 145–172
151
Taxa Sample number Origin Date collected Sample type
Bare-throated Group
F. afer PFIAO 108 Tudor East, Watervalboven 2004 Liver
F. a. benguellensis AMNH 267682 Mombola Toe pad
F. a. harterti AMNH 541485 Russisi River Toe pad
F. a. nudicollis BM 1903.10.14.91 E. Transvaal 1903 Toe pad
F. a. böhmi BM 1932.5.10.214 S. Tanganyika 1932 Toe pad
F. a. cunenensis TM 28584 Cunene River 1957 Toe pad
F. humboldtii, swynnertoni TM 20341 Selindu, Mabsettler 1935 Toe pad
F. cranchii cranchii BM 1953.54.56 Mwinilunga, Northern Rhodesia 1953 Toe pad
F. c. itigi AMNH 202502 Poona Singida Toe pad
F. c. intercedens AMNH 416180 Tukuyu Toe pad
F. c. nyanzae AMNH211906 Buhumbiro Toe pad
F. swainsonii TMC 40 Marico River 2004 Liver
F. s. lundazi SAM 2055756a Deka 1969 Toe pad
F. s. chobiensis SAM 2003501 Victoria Falls 1904 Toe pad
F. rufopictus AMNH 202503 Gagayo, Muranza Toe pad
F. leucoscepus PFIAO 109 Kenya 2004 Heart
F. l. infuscatus AMNH 419169 Tana River, Kenya Toe pad
F. l. muhamed-ben-abdullah AMNH 541581 Toe pad
Montane Group
F. erckelii AMNH 541471 Badaltino, Shoa Toe pad
F. erckelii AMNH DOT11039 Ethiopia Liver
F. ochropectus FNHM 1971-1072 Djbouti Toe pad
F. castaneicollis GB Toe pad
F. c. bottegi AMNH541435 Rafissa, Abyssinia Toe pad
F. c. ogoensis AMNH541426 Lower Sheikh Toe pad
F. jacksoni AMNH261929 East slope, Mt Kenya Toe pad
F. nobilis AMNH1759 West Ruwenzori Toe pad
F. camerunensis TMC 42 Mount Cameroon Liver
F. swierstrai AMNH 419126 Angola Toe pad
F. swierstrai TMC 67 Angola, 14.49° S 13.23° E 2010 Blood
Scaly Group
F. ahantensis AMNH 541409 Near York Pass, Sierra Leone Toe pad
F. squamatus PFIAO 117 Tot. DNA
F. s. maranensis AMNH 541407 Kilimanjaro district Toe pad
F. s. schuetti AMNH 763912 Tshibati, DR Congo Toe pad
F. griseostriatus AMNH 541411 Ndalla Tanda Toe pad
Vermiculated Group
F. bicalcaratus TM 14682 Gold Coast, Hinterland 1901 Toe pad
F. b. ayesha AMNH 541250 Forest of Mamora Toe pad
F. b. thornei AMNH 541280 Kavene, Sierra Leone Toe pad
F. b. adamauae AMNH 704359 Cameroon Toe pad
F. clappertoni AMNH 541305 Takoukout, Cameroon Toe pad
F. clappertoni TMC 68 Cameroon 2005 Breast muscle
F. c. sharpie AMNH 541324 Adarte Toe pad
F. c. nigrosquamatus AMNH 541341 S. Ethiopia Toe pad
F. icterorhynchus AMNH 156922 Fanadji Toe pad
F. hildebrandti GB Blood
F. h. altumi AMNH 551345 Gilgil River Toe pad
F. h. fischeri AMNH 261945 N. Tanganyika Territory Toe pad
F. h. johnstoni AMNH 347277 Mafinga Mt, Northern Rhodesia Toe pad
F. h. helleri AMNH 207771 Neng Toe pad
F. natalensis TMC 120 Marico River, South Africa 2004 Liver
F. hartlaubi TMC 121 Namibia 2006 Breast muscle
F. h. crypticus AMNH 703654 Erungo Plateau Toe pad
F. capensis PFIAO 229 Kakamas, South Africa Heart
F. adspersus PFIAO 206A Liver
F. harwoodi BM 1927.11.5.18 1927 Toe pad
Table 3: Sample information for putative spurfowl taxa for which DNA sequences were generated. AMNH = American Museum of Natural
History, FHHM = French Natural History Museum, TM = Transvaal Museum/Ditsong National Museum of Natural History, BM = British
Museum/Natural History Museum at Tring, SAM = Iziko Museums of Cape Town (Natural History), FIAO = FitzPatrick Institute of
African Ornithology, TMC = Timothy M Crowe, University of Cape Town, South Africa, GB = GenBank. Generic terminology follows that
generated in this study

Mandiwana-Neudani, Little, Crowe and Bowie
152
Phylogenetic approach
Taxa were placed phylogenetically following the principle
of apomorphic character consilience to reflect sequen-
tially more inclusive reciprocally monophyletic groupings
(Hennig 1950, 1966). Qualitative morpho-behavioural
characters (morphology, behaviour and life history) were
analysed in combination with DNA sequence characters, in
a ‘total evidence’ phylogenetic analysis (Kluge 1998; Rieppel
2009). This approach was chosen because combined data
sets may show clade support and resolution that is ‘hidden’
by separate analysis of character partitions. For instance,
when data are concatenated, different classes of charac-
ters that evolve at somewhat different rates may ‘kick in’ at
different depths of phylogeny (Nixon and Carpenter 1996;
Givnish and Systma 2000; Nylander et al. 2004; Wortley
and Scotland 2006), i.e. at deep, shallow and intermediate
nodes. Moreover, previous phylogenetic studies of galliforms
(Crowe et al. 2006) and ‘francolins’ sensu Hall (1963)
Primer name Primer sequence (5′ to 3′) Reference
Cytochrome b
L14578 cta gga atc atc cta gcc cta ga JG Groth (pers. comm.)
MH15364 act cta cta ggg ttt ggc c P Beresford (pers. comm.)
ML15347 atc aca aac cta ttc tc P Beresford (pers. comm.)
H15915 aac gca gtc atc tcc ggt tta caa gac Edwards and Wilson (1990)
Control region
PHDL agg act acg gct tga aaa gc Fumihito et al. (1995)
PH-H521 tta tgt gct tga ccg agg aac cag EA Scott (pers. comm.)
PH-L400 att tat tga tcg tcc acc tca cg EA Scott (pers. comm.)
PHDH cat ctt ggc atc ttc agt gcc Fumihito et al. (1995)
12S rRNA
L1267 aaa gca tgg cac tga ag(atc) tg Moum et al. (1994)
H2294 gtg cac ctt ccg gta cac ttac c O Haddrath (S Pereira pers. comm.)
NADH dehydrogenase subunit 2
L5216 gcc cat acc ccr aaa atg Sorenson et al. (1999)
H6313 ctc tta ttt aag gct ttg aag gc Sorenson et al. (1999)
Ovomucoid G
OVO-G Forward caa gac ata cgg caa caa rtg Armstrong et al. (2001)
OVO-G Reverse ggc tta aag tga gag tcc crt t Armstrong et al. (2001)
GAPDH intron-11
GapdL890 acc ttt aat gcg ggt gct ggc att gc Friesen et al. (1997)
GapdH950 cat caa gtc cac aac acg gtt gct gta Friesen et al. (1997)
Tran Globulin Growth Factor Beta2 intron-5
TGFb2-5F ttg tta ccc tcc tac aga ctt gag tc Primmer et al. (2002)
TGFb2-6R gac gca ggc agc aat tat cc Primmer et al. (2002)
Table 4: DNA markers sequenced and primers used for PCR amplification and sequencing of preserved tissues
Primer name Primer sequence (5′ to 3′) Reference
Cytochrome b
Spurfowl-specific primers
L14851 (General) cct act tag gat cat tcg ccc t Kornegay et al. (1993)
Pt-H195 ttt cgr cat gtg tgg gta cgg ag R Moyle and T Mandiwana-Neudani
Pt-H194 cat gtr tgg gct acg gag g R Bowie
MH15145 aag aat gag gcg cca ttt gc P Beresford
Pt-L143 gcc tca tta ccc aaa tcc tca c R Moyle and T Mandiwana-Neudani
Pt-H361 gtg gct att agt gtg agg ag R Moyle and T Mandiwana-Neudani
Pt-L330 tat act atg gct cct acc tgt ac R Bowie
Pt-H645 ggg tgg aat ggg att ttg tca gag R Moyle and T Mandiwana-Neudani
Pt-L633 ggc tca aac aac cca cta ggc R Moyle and T Mandiwana-Neudani
Pt-H901 agg aag ggg att agg agt agg at R Moyle and T Mandiwana-Neudani
L2-2312 cat tcc acg aat cag gct c R Bowie
H15696 aat agg aag tat cat tcg ggt ttg atg Edwards et al. (1991)
Pt-L851alt cct att tgc cta cgc cat cct ac R Bowie
Pt-H1050 gat gct gtt tgg ccg atg R Bowie
Pt-L961 cga acc ata aca ttc cca c R Moyle and T Mandiwana-Neudani
Pt-L961alt ctc atc cta ctc cta atc ccc R Bowie
HB20 (General) ttg gtt cac aag acc aat gtt J Feinstein (pers. comm.)
Table 5: DNA markers sequenced and primers used for PCR amplification and sequencing of museum toe pads

Ostrich 2019, 90(2): 145–172
153
(Bloomer and Crowe 1998) have demonstrated improved
phylogenetic resolution and nodal support when a ‘total
evidence’ strategy is employed.
Parsimony was employed as the optimality criterion for
the combined DNA and morpho-behavioural character
analyses to minimise ad hoc assumptions vis-à-vis
character evolution (Farris 1983). Indeed, the meta-analysis
of more than 500 articles using model- and parsimony-
based methods found strongly supported topological
incongruence in only two of the studies examined (Rindal
and Brower 2011). Model-based approaches were also not
employed since they may not cater adequately for morpho-
vocalisation characters.
All parsimony-based phylogenetic analyses were
conducted using TNT (Goloboff et al. 2008). The search
strategy employed was the default Ratchet Island Hopper
option: 200 iterations per replication; one tree to hold per
iteration; four characters to sample, amb-poly, and random
constraint level 10. When multiple, equally parsimonious
cladograms persisted, a strict consensus cladogram was
constructed. The extent to which each non-terminal node is
supported by character data was determined by using the
‘jackknife’ option (Farris et al. 1996; Källersjö et al. 1998)
using the following strategy: 1 000 replications, branch-
swapping switched on, random addition of five sequences
per replicate, and ~37% of the characters deleted per
jackknife replicate.
The combined African spurfowl data matrix was rooted on
its sister-species Perdicula asiatica and Ammoperdix heyi
(Crowe et al. 2006; Cai et al. 2017). For inter-taxon genetic
distances, uncorrected pairwise distances were calculated
in PAUP* 4.0b10 (Swofford 2002) and were transformed
into percentages. Both absolute and cladistic CYTB nearest
taxa were identified to obtain both minimum and ‘cladistics-
dependent’ estimates of genetic relationship to allow
speculation on non-dichotomous phylogenetic structure.
Distributional range maps
Another challenging and indispensable aspect in the
analyses was to produce maps showing the distribu-
tional ranges of the various spurfowl taxa ultimately
recognised. Step 1 in developing the range maps for each
taxon that emerged used the Atlas of Speciation in African
Non-passerine Birds (Snow 1978), since it presents the
best distribution ranges of species produced from the point
localities of the specimens collected. This was supple-
mented in Step 2 by consulting the Atlas of Southern
African Birds (Harrison et al. 1997), which was helpful in
filling distribution gaps for southern African species. Step
3 involved using Hall’s (1963) inferred distributions to
complete the ranges of species and subspecies recognised.
Results
Morpho-vocalisation characters
Morpho-vocalisation character state entries for spurfowl
taxa recognised are presented in Table 6.
Genetic distance
Percentage unweighted, uncorrected mitochondrial
cytochrome b sequence divergences between Hall’s (1963)
fisante sensu stricto and all the species recognised from
our analyses (i.e. previous subspecies) are presented
in Table 7.
Taxonomy
The implementation of the Concilience Species Concept
and its attendant criteria for recognising species increases
the number of fisante sensu stricto species from 23 to 25
and reduces the number of subspecies taxa from 59 to 16
(Appendix 4). Two of Hall’s (1963) subspecies (schuetti and
cranchii) are elevated to species level. We recommend that
all taxa be referred to commonly as spurfowls (Crowe and
Little 2004).
Hartlaub’s spurfowl
Hartlaub’s Spurfowl Pternistis hartlaubi is the smallest
spurfowl, which occupies dense, mixed grass–shrub
cover on boulder-strewn slopes and rocky outcrops in hilly
and mountainous regions within granite and sandstone
substrate semi-desert open savanna (Komen 1987). It is
confined to central and northern Namibia, particularly on the
Namibian escarpment and extreme south-western Angola
(Little 2016a).
The upper mandible is horn coloured and the lower
yellowish. The male has a dark grey-brownish crown, a
pronounced white eyestripe, offset by a black line below
and chestnut ear coverts. The back is grey, faintly streaked
and barred with brown. The belly is pale grey, heavily
streaked with brown. The black and white under-tail coverts
are conspicuous in flight and in courtship display. The adult
female has an orange-brown eyestripe, and a grey-brown
head, cheeks, chin and belly. The back is grey-brown with
strong vermiculations (Komen 1987).
It is markedly distinct from other African spurfowls
in organismal (Hall 1963; Komen 1987) and molecular
biology (Table 7). It differs from all other spurfowls in that
it is socially monogamous throughout the year (Komen
1987). Other spurfowls in southern Africa are monoga-
mous only during the courtship phase of reproduction (van
Niekerk et al. 2009; van Niekerk 2011) with only the female
providing care for offspring (van Niekerk 2017). It also has
markedly sexually dimorphic plumage and size-dimorphism
(♂ 245–290 g, ♀ 210–240 g; Little 2016a), a dispropor-
tionately long bill (Komen 1987), and yellow (normally
black or red/orange-red in spurfowl) tarsi with virtually no
tarsal spurs, only tiny bumps (Hall 1963). It has vocalisa-
tions markedly different from, but that still link with, other
spurfowls (van Niekerk 2013; Mandiwana-Neudani et al.
2014); it demarcates and defends its territory year-round
using a combination of antiphonal duet calling initiated
by the hen; displays, rather than using overt aggression
(Komen 1987); and seems not to require standing/flowing
water for drinking (Komen 1987).
Putative subspecies populations in southern Angola
(nominate ‘hartlaubi ’) are somewhat smaller in body size
than those in Namibia. Those in the Kaokoveld and Erongo
(‘crypticus’) are paler than those in the Waterberg and
Otavi (‘bradfieldi ’) in the east. We regard these differences
as clinal variation. The two specimens from Erongo and
the Waterberg were 0.4% CYTB divergent. Therefore, we
recognise no subspecies for this spurfowl.

Mandiwana-Neudani, Little, Crowe and Bowie
154
Taxon Character reference number (see Table 2)
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33
Francolinus hartlaubi 011101110011011000012132103320110
F. camerunensis 01120201101101100012221110222010 0
F. nobilis 011201100000010000122311002220111
F. erckelii 021301001000010110022412013321010
F. swierstrai 010501100000010000022322103321010
F. castaneicollis 010301101000010100122412013321010
F. c. atrifrons 012121101000010100122412013321010
F. ochropectus 021301001000010110022412012220100
F. jacksoni 01030110100001010002241200??????0
F. squamatus 030102011011010001032212003320110
F. s. schuetti 031102011011010100032212 003320110
F. ahantensis 04010110100101011004221200222011 0
F. griseostriatus 030101101110010000031112003110010
F. bicalcaratus 011401101011010100052221001210010
F. b. ayesha 011401101011010100052221001210010
F. b. adamauae 011202101011010100052221001210010
F. icterorhynchus 011201001111011000262221001210010
F. clappertoni 011401100011010100122221002210010
F. c. sharpii 010101100011010100122221002210010
F. harwoodi 011401100011010100122222?12210010
F. h. hildebrandti 112211001111011010022212101320010
F. h. fischeri 11221100111101101002221210132001 0
F. natalensis 0301010011110110 00021212002320110
F. adspersus 010203000110011000221212003321110
F. capensis 232121100010010100022412003320100
F. leucoscepus 342131101011020010172321003220011
F. l. infuscatus 012101001011020010172321003220011
F. rufopictus 030101101001130100172421002220011
F. afer afer 352101100000040100122321002220111
F. a. cranchii 010101101001040110122321002220111
F. a. humboldtii 342131100000040000122321002220111
F. swainsonii 33213110100114101017132100322011 1
Table 6: Morpho-vocalisation character score matrix used for the phylogenetic analysis of spurfowls
Taxon 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25
1F. hartlaubi
2F. swierstai 10.8
3F. camerunensis 10.8 7.8
4F. nobilis 11.2 9.2 7.4
5F. jacksoni 10.8 6.6 8.6 9.7
6F. castaneicollis 9.9 7.3 7.6 8.4 7.5
7F. erckelii 8.9 6.4 7.5 8.0 6.6 4.4
8F. ochropectus 8.5 7.7 7.8 8.5 7.3 4.5 2.6
9F. squamatus 10.0 6.9 8.4 8.7 5.8 6.6 6.7 7.1
10 F. schuetti 7.8 5.3 7.9 8.0 6.8 6.2 4.8 6.3 3.7
11 F. ahantensis 8.9 7.4 8.1 8.2 7.7 7.3 5.7 7.6 4.2 4.2
12 F. griseostriatus 10.0 6.6 8.0 8.7 5.0 6.0 5.6 5.9 4.2 2.7 5.1
13 F. hildebrandti 10.0 7.0 8.7 9.4 6.8 7.1 7.6 8.0 5.4 4.2 4.8 5.6
14 F. natalensis 10.1 6.9 8.3 8.8 5.9 6.6 6.8 7.2 5.0 5.0 6.4 4.8 3.1
15 F. adspersus 10.2 6.9 8.6 9.5 6.0 7.0 7.0 6.9 5.5 5.6 7.9 4.7 5.0 4.2
16 F. capensis 10.2 6.5 8.2 9.1 5.6 6.5 6.6 7.3 5.0 4.8 7.7 4.5 4.6 4.0 3.8
17 F. icterorhyncus 11.1 8.2 9.3 9.2 8.0 8.3 7.4 7.3 7.3 8.2 6.0 6.3 7.5 7.1 7.3 7.3
18 F. bicalcaratus 10.3 6.7 8.4 8.6 6.8 6.8 6.3 6.6 5.9 5.7 6.4 5.0 6.7 6.2 6.4 6.1 3.3
19 F. harwoodi 9.4 6.6 8.3 9.0 5.6 6.2 5.9 6.8 4.7 5.1 6.8 4.7 6.0 4.8 5.7 5.2 6.8 5.7
20 F. clappertoni 9.4 6.1 8.0 8.7 5.4 6.0 5.4 6.3 4.5 4.6 6.4 4.4 5.9 4.4 5.2 5.2 6.4 5.4 1.4
21 F. leucoscepus 9.8 7.5 8.5 9.4 6.6 6.8 6.8 7.1 5.8 5.8 7.1 6.1 6.4 6.0 6.9 6.4 7.5 6.6 5.2 5.4
22 F. swainsonii 9.5 7.2 7.5 8.7 6.1 6.4 6.0 6.5 5.6 5.1 8.2 5.8 6.3 5.6 5.7 5.9 7.4 6.6 5.0 4.6 4.5
23 F. cranchii 9.2 7.0 8.2 8.3 5.9 6.4 6.0 6.5 5.2 4.7 6.2 5.7 5.4 5.3 5.8 5.4 7.4 6.6 4.7 4.7 3.7 3.9
24 F. rufopictus 9.2 7.4 7.6 8.0 6.2 6.0 6.2 6.6 5.2 4.5 5.8 5.5 5.6 5.5 5.5 5.2 7.5 6.6 4.9 4.9 4.1 3.9 1.7
25 F. afer 9.7 7.3 8.0 8.0 6.5 7.2 6.6 7.3 5.4 5.1 7.2 6.2 6.3 5.8 6.6 6.0 7.0 7.3 5.4 5.3 4.4 4.9 1.6 2.4
Table 7: Percentage unweighted, uncorrected, overall, ‘P’ molecular sequence divergence of mitochondrial cytochrome b between Hall’s
(1963) fisante ‘Francolinus’ sensu stricto and the upgraded subspecies

Ostrich 2019, 90(2): 145–172
155
Pternistis hartlaubi is not closely genetically related to
other spurfowls, diverging by between 7.8% to 11.2% at
CYTB from other taxa (Table 7). The closest CYTB taxon is
the Scaly schuetti at 7.8% sequence divergence (Table 7).
Its closest Vermiculated spurfowl (where Hall [1963]
placed it) is harwoodi at 9.4%. Given its striking morpho-
behavioural and genetic divergence, a case might be made
to taxonomically elevate Hartlaub’s Spurfowl to generic
status. Should this be supported, the generic epithet
Chapinortyx (Roberts 1928) is available.
Montane spurfowls
Swierstrai’s Spurfowl Pternistis swierstrai is an endemic
of Angola confined to undergrowth within patches and
edges of relict evergreen forest in the highlands of western
Angola, Mountains Moco and Soque, the Bailundu
Highlands and Mombolo Plateau along the escarpment,
with isolates on the Chela Escarpment, Tundavala (Huila
District) and Cariango (Cuanza Sul District) (Figure 2) (Little
2016a). It ventures into grass- and bracken-covered slopes
and gullies (Little 2016a).
It is a small spurfowl (both sexes 375–565 g) (Little
2016a) with an orange-red bill, a yellow ear-patch, yellow
eye-ring on males (blue in females), red tarsi with one spur
only in males. It is weakly sexually dimorphic in plumage.
Both sexes have a conspicuous white eye-stripe and
throat, and brown back plumage (irregularly blotched in the
female). The male’s black breast contrasts with the white
throat, whereas the belly feathers have broad buff central
streaks with blackish margins. The belly plumage of the
female is white, barred/blotched with dark brown (Hall
1963).
Pternistis swierstrai is an early diverging lineage of
spurfowl, diverging by 5.3% to 9.2% at CYTB and by 10.8%
from P. hartlaubi. Like hartlaubi, its closest CYTB taxon is
schuetti at 5.3% sequence divergence (Table 7). Its closest
Montane taxon is erckelii at 6.4%.
Mount Cameroon Spurfowl Pternistis camerunensis is
endemic to dense undergrowth and edges of forests on the
south-eastern slopes of Mt Cameroon, between 850 and
2 100 m above sea level (Figure 2) (Little 2016a). It is a
small (♂ ~593 g, ♀ ~509 g) sexually dimorphic spurfowl
with an orange-red bill, red eye-ring, and orange-red
tarsi with 1–2 spurs only in males. The male has a dark
brown crown and nape. Its throat is grey-buff with the belly
feathers chestnut with grey edges. Its upper tail coverts
and primaries are grey brown, and wing coverts and the
lower neck are deep maroon, with light grey scalloping
on the lower neck. Its back is rich dark brown, excluding
the lower neck. The belly and lower neck are plain grey
with some black feather centres and shaft streaks. The
chest and belly plumage of the female is mottled and
1
2
23
4
5
6
7
8
9
A
A
1 Pternistis camerunensis
2 P. erckelii
3 P. ochropectus
4 P. castaneicollis castaneicollis
5 P. c. atrifrons
6 P. jacksoni
7 P. nobilis
8 P. swierstrai
9 P. hartlaubi
A ‘Arid Corridor’
0°
0°
10° N
20° N
10° S
10° E10° W 20° E 30° E 40° E 50° E
20° S
30° S
1000 km
MONTANE AND
HARTLAUB’S SPURFOWLS
Figure 2: Geographical distribution of Hartlaub’s Spurfowl, Montane spurfowls and the ‘Arid Corridor’. Arrows draw attention to
phylogenetically sequential cladogenesis

Mandiwana-Neudani, Little, Crowe and Bowie
156
vermiculated with black, dark brown and buff with some
off-white U- to V-patterning on the belly and lower neck
(Hall 1963).
Pternistis camerunensis is sister (and closest) to the East
African montane P. nobilis, from which it diverges by 7.4%
at CYTB (Table 7).
Handsome Spurfowl Pternistis nobilis is endemic to
the highland Ruwenzori and Kivu forests in the Albertine
Rift and mountains in far western Democratic Republic of
Congo (hereafter DR Congo), south-western Uganda, and
borders of Rwanda and Burundi (Little 2016a), and is locally
common in dense undergrowth, forest edge and moist
bamboo thickets (Figure 2). It is medium-sized and sexually
monomorphic (♂ averaging 877 g, ♀ 635 g) (Little 2016a).
It has a red bill, eye-ring and tarsi with 1–2 spurs only in
the male. It has a grey-brown head, primaries and rump,
and a buff throat. It is dark maroon overall, particularly on
the wings and back, with light grey scalloping on the lower
neck. The remainder of the belly feathers are chestnut with
narrow grey or whitish edges or scallops (Hall 1963).
We regard the subspecies ‘chapini ’ from the Ruwenzori
Mountains as a localised variant since it differs only by
having somewhat narrower greyish edges to the belly
feathers (Hall 1963).
Pternistis nobilis is sister (and CYTB closest) to the
Mount Cameroon endemic P. camerunensis, from which it
diverges by 7.4% at CYTB (Table 7).
Erckel’s Spurfowl Pternistis erckelii is the most northerly
distributed Montane spurfowl and the largest African
spurfowl (♂ 1 050–1 590 g, one ♀ 1 136 g) (Little 2016a).
It occurs in giant heath, forest scrub remnants and edges
above 2 000 m, extending relatively continuously up to
3 000 m, from the vicinity of Addis Ababa in the massif of
central and northern Ethiopia northwards to southern Eritrea
(Figure 2). Unlike other Montane spurfowls, it will venture
out of forest into adjacent heath and grassland. It has
a black bill and yellowish tarsi with two spurs only in the
male. It is sexually monomorphic and has a black forehead
and eyestripe, chestnut crown, grey ear coverts and white
throat. Its lower neck is grey like the breast, but with greyish
brown margins and a thin central buff streak, whereas the
breast feathers have central greyish black streaks. Belly
feathers have a broad buff central streak constricted in the
middle and expanded distally into a tear-drop, margined
with rufous (Hall 1963). The somewhat greyer putative
subspecies, ‘pentoni ’, isolated in the Red Sea Hills at
Erkowit, is not recognised.
The montane Ethiopian and Eritrean P. erckelii is sister
(and closest) to its geographical neighbour the Montane
Djibouti P. ochropectus, diverging by 2.6% at CYTB. These
two species are sister to the remaining Ethiopian montane
endemic spurfowl, P. castaneicollis, diverging by 4.4% and
4.5%, respectively (Table 7).
Djibouti Spurfowl Pternistis ochropectus is large (one
♂ 809 g, one ♀ 605 g) (Little 2016a) and endemic to the
evergreen juniper forest mostly above 1 200 m on the
Plateau du Day of Djibouti (Figure 2). It has a black bill
with the lower mandible yellowish and yellow tarsi with two
spurs only in the male. The belly feathers of P. ochropectus,
P. erckelii and P. castaneicollis are similar, but P. erckelii
and P. castaneicollis are more heavily marked with brown
on the back and breast. The belly feathers of ochropectus
have a broad buff central streak constricted in the middle
and expanded distally into a tear-drop, margined by a
greyish black U-shaped streak (Hall 1963; Little 2016a).
Pternistis ochropectus is sister (and closest) to P. erckelii
(2.6% at CYTB) (Table 7).
Chestnut-naped Spurfowl Pternistis castaneicollis is a
large spurfowl (♂ 915–1 200 g, ♀ 550–650 g) (Little 2016a)
restricted to montane heath moorlands, juniper forests and
forest edge/scrub above 2 800 m. It extends broadly in
montane ‘islands’ along the mountain ranges of central and
southern Ethiopia on both sides of the Rift Valley to Somalia
in the north-west, and to the Kenyan border in the south
(Figure 2) (Little 2016a). It is morphologically geographically
variable and most similar to P. erckelii (Little 2016a). It has a
red bill, yellow ear-patch, yellowish eye-ring in males (blue in
the female) and orange-red legs with two equally long spurs
only in the male. It is sexually monomorphic in plumage,
but females are smaller. It has less black on the face than
P. erckelii and P. ochropectus. Its belly feathers have a
broad buff central streak, constricted in the middle and
expanded distally into a tear-drop, margined with rufous. Its
eastern Ethiopian populations have an extensive double-U-
patterning on the back with wing coverts and breast clearly
defined in black and white, with some ochre and chestnut,
grading to mainly white on the belly (Hall 1963).
Pternistis castaneicollis is sister (and closest) to P. erckelii
and P. ochropectus, from which it diverges by 4.4% and
4.5% at CYTB, respectively (Table 7). The putative subspe-
cies ‘bottegi ’ and ‘ogoensis’ are morphologically clinal
variants that differ from nominate canstaneicollis by 0.2%
and 0.6%, respectively (Mandiwana-Neudani 2012). Other
putative subspecies ‘gofanus’ and ‘kaffanus’ are clinally
less well-defined and U-patterned (Hall 1963). Moreover,
their CYTB divergence from nominate castaneicollis is
0.2% to 0.7%.
The subspecies atrifrons, for which we had no DNA
sequence data, is confined to the Mega Mountains of
southern Ethiopia (Figure 2). Peters (1934) and Töpfer et
al. (2014) recognised it as a full species and it is 1.2% to
1.3% CYTB divergent from P. c. castaneicollis, from which it
differs by having the throat and belly cream instead of white
and reduced or absent chestnut colouration and U-patterning
on the neck and flanks. Despite these genetic and morpho-
logical differences, atrifrons has similar vocalisations, habits
and habitat use to other forms of P. castaneicollis (Hall 1963;
Little 2016a). Hence, although populations of this taxon
are highly endangered, in terms of our stated criteria, its
elevation to full species is not supported.
Jackson’s Spurfowl Pternistis jacksoni occurs between
2 200 and 3 700 m (Little 2016a), primarily in forests,
forest edges, moorlands, bamboo patches and within the
Aberdares and Mt Kenya, Mau Escarpment and Cherangani
Mountains in Podocarpus, Juniperus and other Afro-alpine
forests of western and central Kenya, extending margin-
ally into Uganda (Figure 2). It is large (1 130–1 160 g),
with females slightly smaller (Little 2016a). It has a red
bill, yellow-orange ear-patch and eye-ring, and tarsi with
1–2 spurs only in the male. Its throat is buff and the lower
neck greyish with the proximal part of the lower neck
similarly patterned to the remainder of the belly. Lower neck

Ostrich 2019, 90(2): 145–172
157
feathers are chestnut coloured and edged with buff to white,
variable among individuals (Hall 1963).
The subspecies ‘pollenorum’ from Mt Kenya is not
supported because it is only ‘somewhat darker’ (Hall 1963)
than other forms of P. jacksoni.
Pternistis jacksoni is sister to the balance of spurfowls. It
is deeply divergent from most spurfowl lineages, differing
by >5% at CYTB from other spurfowl (Table 7). Its closest
taxon is Scaly griseostriatus at 5.0% sequence divergence
(Table 7). The closest Montane taxon is swierstrai at 6.6%.
Scaly spurfowls
Scaly Spurfowl Pternistis squamatus occurs in forests of
south-eastern Nigeria, extending east into the DR Congo
and up to 3 000 m (on Mt Elgon) in Uganda/Kenya
(Figure 3) (Little 2016a). This small spurfowl has a red bill,
orange-red tarsi with 1–2 spurs in males only (Little 2016a).
There is no size dimorphism (♂ 372–565 g, ♀ 377–515 g)
and plumage, with U-patterned vermiculated upperparts,
less so in males. It is the least distinctly patterned scaly
taxon. The brown upperparts are indistinctly vermiculated
faint grey with each feather having a blackish centre tinged
maroon, and the upper back has faint buff U-patterning.
The scaly underparts are brown with ill-defined dark shaft
streaks (Hall 1963).
Pternistis squamatus is sister (and closest) to fellow
Scaly P. schuetti, from which it diverges by 3.7% at CYTB
(Table 7).
Schuett’s Spurfowl Pternistis schuetti occurs in
eastern DR Congo extending east to Uganda, Ethiopia,
Kenya, Tanzania and Malawi (Little 2016a) (Figure 3). It
resembles P. squamatus, but is less vermiculated overall,
with the scaly pattern on the lower neck less clearly
defined, each feather has a deep red-brown centre (Hall
1963). Populations west of the Rift Valley in Kenya south
towards Kilimanjaro, Monduli and Mt Meru in north-eastern
Tanzania (Hall 1963) become clinally increasingly darker
and greyer (more readily seen in males) and tend to have
less white on the belly. Poorly sampled, isolated popula-
tions to the south ‘usumbarae’, ‘uzungwensis’ and ‘doni ’
are clinal or idiosyncratic variants of P. schuetti but may
warrant subspecific status should they exhibit significant
genetic divergence.
Pternistis schuetti is sister to P. squamatus, from which
it diverges by 3.7% at CYTB. However, its closest taxon is
P. griseostriatus at 2.7% sequence divergence (Table 7).
We elevate schuetii to full species status (Appendix 4).
Pternistis s. maranensis (1.2% divergent from nominate
schuetti; Mandiwana-Neudani 2012) occurs further east
on Mt Kilimanjaro (up to 2 000 m), Monduli, Mt Meru
and in the Chyulu Hills (Figure 3). It is darker and less
patterned than schuetti (Hall 1963). About 240 km south-
east of Kilimanjaro, birds (‘usambarae’) from the Usambara
Mountains (Hall 1963) have the area around their eyes and
cheeks freckled with black and white instead of uniform
brown. Another isolated population from the forests on the
11
2
3
3
5
2
4
SCALY SPURFOWLS
1 Pternistis ahantensis
2 P. squamatus
3 P. schuetti schuetti
4 P. s. maranensis
5 P. griseostriatus
0°
0°
10° N
20° N
10° S
10° E10° W 20° E 30° E 40° E 50° E
20° S
30° S
1000 km
Figure 3: Geographical distribution of Scaly spurfowls

Mandiwana-Neudani, Little, Crowe and Bowie
158
Vipya Plateau between 900–2 800 m (‘doni ’) in Malawi
have more reddish-brown upper and underparts with some
white streaking on the underparts (Hall 1963). These
populations, for now, are included within nominate schuetti
(Appendix 4).
Ahanta Spurfowl Pternistis ahantensis occurs within
gallery and secondary coastal lowland West African
forests in three disjunct populations west of the Niger
River: southern Senegambia and northern Guinea-Bissau;
southern Guinea, Sierra Leone and western Liberia; north-
eastern Côte d’Ivoire and Ghana through central Togo and
central Benin to south-western Nigeria (Figure 3).
Pternistis ahantensis is medium-sized (♂ ~608 g,
♀ ~487 g) and has an orange bill with a black base and
yellow-orange tarsi with 1–2 spurs only in the male (Little
2016a). It is the most patterned Scaly spurfowl, with breast
and flank feathers having paler edges and darker centres.
The feathers on its upperparts are vermiculated (distinct
on the lower neck, indistinct on the back) with blackish
centres and a reddish-brown shaft-streaking, those on the
lower neck have some white U-patterning. The underparts
are dark-brown chestnut with white and darker brown
U-patterning (Hall 1963). The isolated western populations
‘hopkinsoni ’ (for which we had no CYTB data) are paler
overall (Hall 1963) than those in the east and probably do
not warrant taxonomic status.
Pternistis ahantensis is sister (and closest) to
P. squamatus and P. schuetti, diverging from both species
by 4.2% at CYTB (Table 7).
Grey-striped Spurfowl Pternistis griseostriatus is a small
(♂ 265–430 g, ♀ 213–350 g) (Little 2016a) endemic to
vestigial patches of forest in the Angolan western escarp-
ment (Figure 3). It has a black bill with a red base (lower
mandible orange-red) and its tarsi are orange-red with a
single spur in the male. It is sexually monomorphic, and
its lower neck feathers and wing coverts are chestnut
and broadly edged and vermiculated with grey, similar to
P. squamatus and P. ahantensis, but paler. However, the
underparts are plain, and the breast and flank feathers are
chestnut, edged with greyish or creamy buff (Hall 1963).
Pternistis griseostriatus is sister to four species of southern
Vermiculated spurfowls: P. hildebrandti, P. natalensis,
P. adspersus and P. capensis; CYTB divergent from these
taxa by 4.5% to 5.6% (Table 7). However, its closest CYTB
taxon is P. schuetti at 2.7% sequence divergence (Table 7).
Vermiculated spurfowls
Hildebrandt’s Spurfowl Pternistis hildebrandti is medium-
sized (two ♂ 600 and 645 g, two ♀ 430 and 480 g) (Little
2016a). Its distribution extends from central and western
Kenya, south-eastern DR Congo, north-eastern Zambia,
Tanzania, Malawi and northern Mozambique (Figure 4)
and comprises two subspecies, hildebrandti and fischeri,
with the former being sexually dimorphic. It has a reddish
mandible and brown culmen with a yellow base, a yellow
ear-patch and eye-ring, red tarsi with 1–2 spurs in both
sexes. The dorsal plumage of males resembles that of
P. icterorhynchus. It is greyish brown with vermicula-
tions, the hind and lower neck are streaked black with
white margins, and the belly plumage has marked black
blotching. Females have similar back plumage to males,
but (especially in P. h. fischeri) differ markedly in having
orange-brown underparts (Hall 1963).
Pternistis h. fischeri (Hall 1963) (0.5% sequence
divergent from hildebrandti) from southern Malawi,
Mozambique and south-western Tanzania (Figure 4) differs
from hildebrandti in that females have an unpatterned
nape, hind neck and breast, in contrast to an orange-brown
abdomen. Birds from Kenya, altumi (Hall 1963), do not
warrant taxonomic recognition because their plumage is
intermediate between nominate hildebrandti and fischeri.
All other putative forms of hildebrandti are >2% divergent
from natalensis, its sister-species. In the Luangwa Valley,
the presence of specimens with intermediate plumage
suggests that P. hildebrandti may interbreed (or have
interbred) with P. natalensis (Hall 1963). Its closest CYTB
taxon is the southern Vermiculated natalensis at 3.1%
sequence divergence (Table 7).
Natal Spurfowl Pternistis natalensis is medium-sized
(♂ 415–723 g, ♀ 370–482 g) (Little 2016a) occurring
across south-eastern Africa, from Zambia, Zimbabwe,
inland Mozambique, eastern Botswana, Swaziland and
north-eastern South Africa (Figure 4). It occurs in thick
riverine bush but will venture into dry lowveld savanna
and adjacent grasslands (Little and Crowe 2011). It has an
orange bill with a dull greenish base, and orange tarsi with
a single spur only in the male (Little 2016a). It is sexually
monomorphic, but some populations ‘neavei ’ from southern
Zambia and western Mozambique are slightly dimorphic.
The hindneck is mottled black and white, the back is highly
vermiculated in greyish-brown and black, with white and
buff markings. The belly is buff with the breast to mid-belly
being heavily patterned in black and buff U-patterning
is concentrated on the breast with the extreme lower
abdomen having no or few marks (Hall 1963).
Pternistis natalensis is sister to P. hildebrandti from
which it diverges by 3.1% at CYTB (Table 7). The closest
CYTB ‘taxon’ to P. natalensis is southern Vermiculated
P. hildebrandti fischeri (a possible product of hybridisation)
at 0.5% sequence divergence (Mandiwana-Neudani 2012).
Red-billed Spurfowl Pternistis adspersus is smallish
(♂ 340–635 g, ♀ 340–549 g) (Little and Crowe 2011; Little
2016a) and occurs in dense bush, mixed woodland and low
scrub thickets interspersed with open ground, mostly on
Kalahari sands along watercourses in Namibia, Botswana,
southern Angola and south-western Zambia (Figure 4).
It is monotypic with an orange-red bill and tarsi, yellow
ear-patch and eye-ring. Males have a single spur. The
upperparts are finely vermiculated, and the underparts have
distinct narrow black and white bars, variable on the lower
neck (Hall 1963).
Pternistis adspersus is sister (and closest) to P. capensis,
from which it diverges by 3.8% at CYTB (Table 7).
Cape Spurfowl Pternistis capensis is the largest
Vermiculated spurfowl (♂ 870–1 000 g, ♀ 640–900 g).
It is endemic to thick cover and rocky river valleys in the
Fynbos Biome of south-western South Africa, with isolated
populations extending deep into the Succulent Karoo Biome
and lower stretches of the Orange River (Figure 4) (Little
and Crowe 2011). It has a brown upper mandible (lower
red), and orange-red tarsi with one spur (females) and
sometimes two (males). It has distinctive uniform brown and

Ostrich 2019, 90(2): 145–172
159
white double V- or U-shaped patterning on the back, breast
and belly, while the throat has irregular black flecking. The
breast and belly feathers have broad white shaft streaks
(Hall 1963; Little and Crowe 2011).
Pternistis capensis is sister (and closest) to its fellow
southern Vermiculated P. adspersus, from which it diverges
by 3.8% at CYTB (Table 7).
Heuglin’s Spurfowl Pternistis icterorhynchus is medium-
sized (♂ 504–588 g, ♀ 200–462 g) (Little 2016a) and occurs
in grasslands, open woodlands and adjacent agricultural
lands in the Central African Republic, northern DR Congo,
extending east to South Sudan and Uganda (Figure 5). It
has a yellow-orange black bill, small yellow eye-patch,
yellow-orange tarsi with 1–2 in males only. It is monotypic
and sexually monomorphic (Figure 5), with a chestnut
crown, brown back diagnosed by having less V-shaped
patterning on the lower neck and more vermiculations on
the back than other vermiculated taxa. Its underparts are
buff, heavily marked with dark brownish-back V-shaped
markings (Hall 1963).
Pternistis icterorhynchus is sister (and closest) to its
northern Vermiculated sister-taxon, P. bicalcaratus, at 3.3%
sequence divergence at CYTB (Table 7).
Double-spurred Spurfowl Pternistis bicalcaratus
comprises three sexually monomorphic subspecies
(Figure 5). All are similarly patterned above and below,
differing in the degree of colouration and vermiculation, and
the size of the arrow-shaped buff marks in the centre of the
belly feathers (Hall 1963). They occur in dry grasslands,
open savanna, palm groves and cultivated areas of West
Africa from Senegal east to northern Cameroon and
southern Chad (Little 2016a) (Figure 5).
The nominate form, bicalcaratus, is medium-sized
(♂ ~507 g, ♀ ~381 g) and has a greenish-black bill and
greenish tarsi with 1–2 spurs, much shorter in females. It
has no bare facial skin, a pale rufous crown, and a white
eyestripe. It has rufous-chestnut on the lower neck and
the remaining upperparts are vermiculated with V-shaped
patterning. It has buff heavily streaked underparts with
small black and chestnut arrow-shaped buff marks on most
belly feathers (Hall 1963).
The more heavily patterned ayesha (from Morocco, not
mapped) is similar (1.0% CYTB divergent) to bicalcaratus
but is faintly vermiculated and slightly more rufous on the
lower neck, with small arrow-shaped buff marks on the belly
feathers (Hall 1963). The darkest form is adamauae (1.7%
CYTB divergent) with very little rufous on the lower neck,
and the underparts are more buff with extremely reduced
chestnut and larger arrow-shaped buff marks along the
belly feather shafts (Hall 1963).
Pternistis bicalcaratus is sister (and closest) to fellow
northern Vermiculated P. icterorhynchus, diverging by 3.3%
at CYTB (Table 7).
Clapperton’s Spurfowl Pternistis clappertoni comprises
two widespread subspecies extending up to 2 300 m in
semi-arid grassland and bushy savanna and adjacent
1
2
3
4
5
5
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0°
10° N
20° N
10° S
10° E10° W 20° E 30° E 40° E 50° E
20° S
30° S
1000 km
VERMICULATED
SPURFOWLS – SOUTH
1 Pternistis hildebrandti
hildebrandti
2 P. h. fischeri
3 P. natalensis
4 P. adspersus
5 P. capensis
Figure 4: Geographical distribution of Vermiculated spurfowls (South)

Mandiwana-Neudani, Little, Crowe and Bowie
160
cultivations across north-central Africa from far eastern
Mali, central Niger, far north-eastern Nigeria, Chad,
southern Sudan, South Sudan, north-eastern Uganda and
western Ethiopia (Little 2016a) (Figure 5). It also occurs in
the Nile and Blue Nile river valleys (Hall 1963).
It is medium-sized (♂ 450–604 g, ♀ 300–530 g) (Little
2016a) with a black bill with a red base and red tarsi with
1–2 spurs in males only. The bare skin around the eye
distinguishes it from P. bicalcaratus, P. icterorhynchus
and P. castaneicollis. The brown upperparts of the
nominate clappertoni have U-shaped patterning similar
to P. icterorhynchus, but more orange brown and vary
geographically in the degree of vermiculation and
U-patterning. It has a white throat and the neck is buff below
with black to brownish marks.
Pternistis c. sharpii (1.4% CYTB divergent from
clappertoni) has streakier marks on the belly than those in
clappertoni having a more buffy white background below
with the breast being similarly U-patterned extending onto
the back (Hall 1963). A specimen collected at ‘Ngeem’ at
Lake Chad (possibly Nguigmi), the type of Francolinus
‘tschadensis’, is possibly a hybrid between P. clappertoni
and P. icterorhynchus (Hall 1963).
Pternistis clappertoni is sister (and closest) to northern
Vermiculated P. harwoodi, differing by 1.4% sequence
divergence at CYTB (Table 7). Despite the limited
sequenced divergence between these two taxa, we retain
P. clappertoni and P. harwoodi as distinct species due to
their morphological differences. The next closest taxon is
southern Vermiculated P. natalensis, jumping to 4.4%
divergence.
Harwood’s Spurfowl Pternistis harwoodi is a medium-
sized (one ♂ 545 g, one ♀ 446 g) (Little 2016a) poorly
known species occurring in Typha reedbeds, scrub, thicket
and adjacent cultivations along the gorges of the Jemmu
Valley, the Blue Nile and its tributaries of East Africa, and
the highlands of central Ethiopia (Figure 5). It is markedly
distinct morphologically from other northern Vermiculated
spurfowls (Hall 1963). It has a red bill with a black tip, bare
red eye-ring and tarsi with 1–2 spurs in males only. It most
closely resembles P. natalensis, which lacks the bare red
facial skin but has more defined U-patterning on the nape,
with similar U-patterning on the underparts. The upperparts
of the male that we examined were grey speckled and finely
barred with blackish and buff above. The lack of a white
eyestripe sets it apart from other Vermiculated spurfowls.
The hind and lower neck, sides of face, and throat are
speckled with black and white. It has irregular double-
V-shaped patterning on its underparts, which tends to be
scattered on the lower extreme of the buff belly.
The closest CYTB taxon to P. harwoodi is northern
Vermiculated P. clappertoni at 1.4% sequence divergence
(Table 7). However, since it is so strikingly different morpho-
logically, it was not relegated to subspecies status. The next
C
12
3
6
45
0°
0°
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20° N
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10° E10° W 20° E 30° E 40° E 50° E
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VERMICULATED
SPURFOWLS – NORTH
1
2
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C
Pternistis bicalcaratus
bicalcaratus
P. b. adamauae
P. clappertoni clappertoni
P. c. sharpii
P. harwoodi
P. icterorhynchus
Mega Lake Chad
Figure 5: Geographical distribution of Vermiculated spurfowls (North)

Ostrich 2019, 90(2): 145–172
161
closest taxa are Scaly P. squamatus, P. griseostriatus and
Bare-throated P. cranchii at 4.7%.
Bare-throated spurfowls
Yellow-necked Spurfowl Pternistis leucoscepus is a
medium-sized (♂ 615–896 g, ♀ 400–615 g), markedly
dimorphic spurfowl (Little 2016a). It is the most morpho-
logically and ecologically differentiated Bare-throated
species and comprises two subspecies, leucoscepus and
infuscatus (Figure 6). It occurs in arid-acacia savanna and
subdesert scrub in eastern Africa (most of Kenya, north-
eastern Uganda, south-eastern South Sudan and northern
Tanzania), extending north and east through Ethiopia and
Somalia into the Horn of Africa (Hall 1963; Little 2016a)
(Figure 6).
Both subspecies (Hall 1963; Little 2016a) have black
bills with a red base, bare red skin around the eye, bare
yellow throat skin, and black tarsi with 1–2 spurs in the
males. The upper back plumage is dark brown with white
shaft streaks and the underparts are streaked with white
and chestnut with narrow white edges and a triangular white
patch at the tip, tapering up the shaft. The primaries have a
conspicuous white patch visible during flight. The northern
subspecies P. l. infuscatus, at 0.9% sequence divergence
from P. l. leucoscepus, differs in having more chestnut than
white on the underparts in contrast with the dominant white
over chestnut in P. l. leucoscepus.
Pternistis leucoscepus is sister to a clade of four
Bare-throated spurfowls: P. swainsonii, P. cranchii,
P. rufipictus and P. afer. Its CYTB divergence from these
species is 3.5% to 3.7% (Table 7). Its closest CYTB taxon is
P. cranchii at 3.7% divergence (Table 7).
Grey-breasted Spurfowl Pternistis rufopictus is
a monotypic medium-large spurfowl (♂ 779–964 g,
♀ 400–666 g) (Hall 1963; Little 2016a)) distributed in dry
savanna, thickets and plains from the south-eastern
shores of Lake Victoria to the Wembere River in north-
western Tanzania (Figure 6). It is narrowly sympatric with
P. leucoscepus in the southern part of its range (Hall 1963).
It has a red bill, orange-pink throat skin, bare red skin
around the eye, and brown tarsi with 1–2 spurs in males
only. The eye-stripe and sides of the face are black and
white with a white chin stripe on either side of the bare
throat. Its upper back plumage is grey-brown with dark
vermiculations and dark shaft streaks, grading posteriorly
to black, white and chestnut streaking. The wing coverts
and feathers on the back are edged with rufous chestnut.
The breast is grey with black shaft streaks and the belly is
streaked black and white. The belly feathers have narrow
central black streaks separated from rufous chestnut
margins by broad buff to white streaks. It is similar to the
cranchii-type taxa in western Tanzania except for the white
chin stripes (as in P. humboldtii), and no vermiculations
(Hall 1963).
Figure 6: Geographical distribution of Bare-throated spurfowls (part 1)
4
1
2
3
X
0°
0°
10° N
20° N
10° S
10° E10° W 20° E 30° E 40° E 50° E
20° S
30° S
1000 km
BARE-THROATED
SPURFOWLS (PART 1)
Pternistis leucoscepus
leucoscepus
P. l. infuscatus
P. rufopictus
P. swainsonii
P. afer/P.swainson ii hybrid
1
2
3
4
X

Mandiwana-Neudani, Little, Crowe and Bowie
162
Pternistis rufopictus is sister to Bare-throated P. afer
from which it diverges by 2.4% at CYTB (Table 7). Its
closest CYTB taxon is Bare-throated P. cranchii at 1.7%
sequence divergence (Table 7). It is 4.1% divergent from
P. leucoscepus.
Swainson’s Spurfowl Pternistis swainsonii is monotypic,
medium-sized (♂ 400–875 g, ♀ 340–750 g) (Little and
Crowe 2011) distributed across south-western Africa from
northern Namibia, eastern Botswana, Zimbabwe, southern
and eastern Zambia, southwards to north-eastern South
Africa (Figure 6). It frequents acacia/mopane savanna and
tall grassland and is especially partial to cultivated lands
where there is suitable cover. Its range and numbers have
increased in recent decades in the south-eastern parts of
its distribution due to agriculture-related alteration of the
environment (Little 2016a).
It has a black upper mandible (lower dull orange), bare
red throat skin and black tarsi, normally with a single spur
in the male (Little and Crowe 2011). Its upperparts are grey
brown with faint dark shaft-streaking. The underparts are
similar but with a grey wash on the breast and chestnut
streaking lower down. Specimens from southern Zimbabwe
and northern South Africa have blackish mottling on the
abdomen. The feathers have a narrow central greyish black
streak separated from greyish chestnut margins by broad
buff grey vermiculated streaks (Hall 1963).
Pternistis swainsonii is sister to three Bare-throated
spurfowl species: P. cranchii, P. rufopictus and P. afer from
which it diverges by 3.9% to 4.9% at CYTB (Table 7). The
closest CYTB taxon to P. swainsonii is P. cranchii at 3.9%
sequence divergence (Table 7).
Hall’s (1963) ‘Red-necked’ Spurfowl ‘Francolinus afer’ is
the most widespread and morphologically and genetically
variable ‘species’ of Bare-throated spurfowl. It has a
complex geographical distribution and occurs in relatively
mesic evergreen forest edges, and woodland in central
Africa and Kenya, extending southwards through, Zambia,
Malawi, Tanzania, south-western Angola, north-western
Namibia, eastern Zimbabwe, Mozambique into eastern
South Africa (Figure 7) (Little and Crowe 2011).
Hall (1963) lumped all taxa ascribed to this spurfowl into
one species, afer, with two polytypic, ‘diverse blocks’ of
subspecies; the ‘black and white’ afer and ‘vermiculated’
cranchii. We elevate cranchii to full species status
(Figure 7; Appendix 4). Both species are medium-sized
(♂ 480–1 000 g, ♀ 370–690 g) and have a red bill, throat
skin and tarsi with 1–2 spurs in males only (Little and
Crowe 2011).
Cranch’s Spurfowl Pternistis cranchii (Hall 1963;
Little 2016a) includes highly geographically variable, but
not taxonomically distinct, populations from southern
DR Congo, northern Angola, northern Zambia, western
Tanzania, Uganda and Lake Victoria shores (Figure 7). It
is characterised by having no white on the head or black
on the abdomen. The underparts are heavily and finely
vermiculated with grey with heavy chestnut-brown streaking
1
3
1
2
4
4
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H2
0°
0°
10° N
20° N
10° S
10° E10° W 20° E 30° E 40° E 50° E
20° S
30° S
1000 km
BARE-THROATED
SPURFOWLS (PART 2)
1 Pternistis cranchii
2 P. a. humboldtii
3 P. afer afer
4 P. a. castaneiventer
H1 P. cranchii/P. afer hybrids
H2 P. cranchii/P. humboldtii
hybrids
Figure 7: Geographical distribution of Bare-throated spurfowls (part 2)

Ostrich 2019, 90(2): 145–172
163
on the abdomen. Its belly feathers have buff central streaks
vermiculated with blackish grey and margined with broad
chestnut (the degree of chestnut colour varies geographi-
cally) and a black and grey facial pattern. Populations from
the Ruzizi Valley, north of Lake Tanganyika, (‘harterti ’ 0.6%
CYTB divergent from P. cranchii; Mandiwana-Neudani
2012) are darker overall and the streaking on the abdomen
is maroon rather than chestnut (Hall 1963).
In our phylogenetic analyses (Figure 8), P. cranchii is
sister to P. afer and P. rufopictus, and its CYTB divergence
from these species is 1.6% and 1.7%, respectively. The
CYTB divergences between ‘core’ P. cranchii and various
putative subspecies and ‘hybrid’ forms (e.g. ‘cunenensis’,
‘nyanzae’, ‘harterti ’, ‘intercedens’, ‘benguellensis’, ‘itigi ’ and
‘bohmi ’) are 0.5% to 0.8% (Mandiwana-Neudani 2012).
Red-necked Spurfowl P. afer sensu stricto, in marked
contrast to P. cranchii, has unvermiculated upperparts (Hall
1963; Little and Crowe 2011; Little 2016a), and is strongly
patterned black and white on the face and underparts.
Underpart feathers have broadly streaked greyish black
central streaks with buff margins, particularly in the
nominate subspecies afer, or have thin greyish black central
streaks separating the long buff parallel streaks margined
with black or sometimes maroon in south-eastern South
African specimens. In the nominate P. a. afer, confined to
south-western Angola (Figure 7), the face is white, and the
underparts are streaked broadly with black and white, with
black centres and white margins.
The closest CYTB taxon to nominate afer is P. cranchii
at 1.6% sequence divergence (Table 7). Since it is 2.7%
96
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80
77
68
55
80
78
100
100
100
80
77
52
92
93
97
96 98
90
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63
Pternistis hartlaubi
Ammoperdix & Perdicula spp.
Pternistis camerunensis
Pternistis nobilis
Pternistis swierstrai
Pternistis erckelii
Pternistis ochropectus
Pternistis castaneicollis
castaneicollis
P. c. atrifrons
Pternistis jacksoni
Pternistis squamatus
Pternistis schuetti schuetti
P. s. maranensis
Pternistis ahantensis
Pternistis griseostriatus
Pternistis hildebrandti
hildebrandti
P. h. fischeri
Pternistis natalensis
Pternistis adspersus
Pternistis capensis
Pternistis icterorhynchus
P. b. adamauae
P. b. ayesha
Pternistis bicalcaratus
bicalcaratus
Pternistis harwoodi
Pternistis clappertoni
clappertoni
Pternistis c. sharpii
P. l. infuscatus
Pternistis leucoscepus
leucoscepus
Pternistis swainsonii
Pternistis cranchii
Pternistis rufopictus
Pternistis afer afer
P. a. castaneiventer
P. a. humboldtii
Figure 8: Strict consensus parsimony tree for spurfowls constructed from two most parsimonious trees. Numbers mapped above nodes
are jackknife support values. MS = Montane spurfowls, SCS = Scaly spurfowls, SVS = Southern Vermiculated spurfowls, NVS = Northern
Vermiculated spurfowls and BTS = Bare-throated spurfowls

Mandiwana-Neudani, Little, Crowe and Bowie
164
divergent from the wholly black-faced South African
P. a. castaneiventer, which has maroon streaking on its
underparts, it may warrant full species status. Pternistis a.
castaneiventer is 3.6% CYTB divergent from core P. cranchii
(Mandiwana-Neudani 2012). Its closest ‘cranchii-type’ CYTB
taxon is the P. c. ‘cunenensis’ at 2.7% sequence divergence
(Mandiwana-Neudani 2012).
Pternistis a. humboldtii ranges from southern Kenya
and Tanzania south to Mozambique (Figure 7). It is 1.3%
divergent from P. a. castaneiventer and 2.4% to 3.4%
divergent from P. cranchii sensu lato (Mandiwana-Neudani
2012), has a black face with a white jaw-beard and black
belly patch. Feathers on the breast belly are mainly grey
with black shaft streaks, which contrast with the abdomen to
form a black patch, and the flanks, which are streaked black
and white. Birds from coastal Kenya have a white face and
black and white eyestripe. Birds from northern Tanzania
southwards to Malawi and south-eastern Zambia have a
wholly black face (Hall 1963).
There are two broad ‘hybrid zones’ between P. cranchii
and P. afer (H1 and H2 in Figure 7). H2 stretches from
Kondoa Dodoma in central Tanzania through central
Malawi into the Luangwa Valley. Hybrids have well-defined
streaks on the abdomen and varying amounts of chestnut
and black-and-white depending on relative proximity to the
respective parental forms. They exhbit relatively low within-
locality morphological variation (Hall 1963), but do not
warrant taxonomic recognition. H1 in northern and central
Angola is characterised by morphologically much less
structured intermediate populations (Hall 1963).
Phylogenetics
The ‘total evidence’ parsimony analysis based on
5 149 characters (33 morpho-vocalisation and 5 116
DNA bases) and 33 terminal taxa produced two equally
parsimonious trees of length 2 124, the strict consensus
of which is presented in Figure 8. Only one of Hall’s
(1963) spurfowl species groups, the phylogenetically
terminal Bare-throated Group, emerged as monophyletic
and the others are paraphyletic. The Montane, Scaly and
Vermiculated (minus P. hartlaubi) Groups are possibly
‘Grades’ sensu Huxley (1959), since their component
species are often each other’s CYTB nearest neighbours
(Table 7). Therefore, we recognise only one monophyletic
genus for the African spurfowls: Pternistis.
Phylogenetics and biogeography
The basal spurfowl
Pternistis hartlaubi, one of Hall’s (1963) Vermiculated
taxa, is the basal African spurfowl (Figure 8). It occurs in
north-central Namibia and south-western Angola in rocky
outcrops within desertic biotopes and is highly divergent
from other spurfowls in virtually all aspects of its biology,
perhaps warranting recognition as a monotypic genus, for
which Chapinortyx (Roberts 1928) has priority.
Montane spurfowls
Hall’s (1963) Montane spurfowls follow on phylogenetically
sequentially and paraphyletically in the strict-consensus
cladogram from P. hartlaubi (Figure 8). They are
confined to island-like montane forests distributed across
sub-Saharan Africa from the highlands of Angola, to
Mt Cameroon in west-central Africa, extending eastwards
to the Ruwenzori highlands and Kivu forests in the Albertine
Rift and mountains in far western DR Congo, south-
western Uganda, Kenya, Ethiopia, Somalia, Eritrea and
Djibouti (Figure 2). They form two, monophyletic clades:
camerunensis + nobilis and erckelii + ochropectus +
castaneicollis ‘linked’ to P. hartlaubi by P. swierstrai and to
each other by P. nobilis. Pternistis jacksoni falls between this
paraphyletic assemblage and the Scaly spurfowls further
along the cladogram.
Montane spurfowls are the morphologically least homo-
geneous and within-group genetically divergent (Table 7)
spurfowls (Hall 1963). The two isolated, sexually dimorphic
western species (P. camerunensis and P. swierstrai) are
the least heavily spurred and the smallest species, and
most closely resemble the basal, and also strongly sexually
dimorphic P. hartlaubi. The central African species, P. nobilis,
is intermediate in body mass between the two small western
species and the three larger species in the north-east.
Generally, Montane spurfowls differ from one another
primarily in their belly plumage, particularly on the mid-
and lower belly. There is no diagnostic ‘Group’ morpho-
logical character other than that the males have the crown,
lower back, primaries and tail plain brown or red-brown.
Females of the relatively small, moderately sexually
dimorphic species (P. camerunensis and P. swierstrai) have
vermiculated primaries, lower back and tail. Variation in
some characters follows geographically clinal trends, with
the birds of the extreme north-east being the largest and
most heavily spurred with dark bills, yellowish tarsi, no bare
skin round the eyes, with the sexes alike (Hall 1963).
Scaly spurfowls
Moving further along the cladogram, the paraphyletic Scaly
spurfowls comprise three allopatric species (P. squamatus,
P. ahantensis and P. griseostriatus). The fourth species,
P. schuetti, is parapatric with P. squamatus (Figure 3) (Hall
1963). The species (P. squamatus, P. ahantensis and
P. schuetti) form a monophyletic assemblage with P. griseo-
striatus acting as a ‘link’ to the Vermiculated species. Scaly
spurfowls have the plainest plumage (Hall 1963), with the
least patterning and no strong colour. They are charac-
terised by having ‘scaly’ underparts, and inhabit vestigial
patches of montane and lowland forest, secondary and
riverine forests, forest edges and clearing/cultivation therein
of West Africa eastwards to the Sudan and north-eastern
Tanzania, and Central Africa and the Benguela district of
north-western southern Africa (Figure 3) (Little 2016a).
Compared with other spurfowls, these taxa are poorly
diagnosed in terms of plumage pattern and colouration
(Little 2016a). All taxa have unpatterned faces, whitish
throats and brown upperparts, some with faint vermicula-
tions. The underparts are brown or creamy-buff with very
narrow darker edges, providing the characteristic ‘scaly’
appearance. There is no marked plumage dimorphism, with
the exception that females tend to be more vermiculated
than males (Hall 1963).
With regards to CYTB genetic divergence, Scaly spurfowls
have anomalously low (~3% vs >8%) values compared
with some Montane and Vermiculated spurfowls (Table 7).

Ostrich 2019, 90(2): 145–172
165
Vermiculated spurfowls
Thereafter, come the also paraphyletic woodland, savanna,
scrub and bush dwelling Vermiculated spurfowls, divided
into northern and southern grades. Vermiculated taxa are
the most widely distributed spurfowl assemblage within
Africa. They occur almost continuously from Senegal to
Eritrea southwards to Namibia and South Africa (Figures
4 and 5). There is an isolated population (ayesha) of
P. bicalcaratus in Morocco, making it one of the few
sub-Saharan bird species with natural populations north
and south of the Sahara (Moreau 1966). Northern taxa
frequent grasslands and cultivation within woodlands and
acacia savanna and steppe. South of the equator, they
frequent thick bush on hillsides and riparian watercourses.
Bare-throated spurfowls
The monophyletic Bare-throated spurfowls are largely allo/
parapatric (Figures 6 and 7) and ecologically segregated
meta-populations, extending from Ethiopia and Eritrea in
north-east Africa, westwards through Kenya, Tanzania,
Sudan and Uganda to the Congo and Gabon, and south
through Angola, northern Namibia, Botswana, Zimbabwe,
and Mozambique to South Africa. Species inhabit a
range of biotopes sorting by rainfall regimes from desertic
grasslands, through open woodland savanna/bush. Some
taxa are tied to bush adjacent to water.
Bare-throated spurfowls are sexually monomorphic in
plumage, although females of some species are slightly
vermiculated, with a body mass ranging from 340–950
g (Hall 1963; Little 2016a) with males being much larger
(van Niekerk 2009, 2017). They are distinguished from
other spurfowls by having bare skin on the throat and a
patch around the eye and plain dark upperparts without
pale vermiculations. Their tarsi are black, red, orange or
brown with spurs well-developed in males only. They have
a long robust lower spur and, in some taxa (P. leucoscepus
and P. rufopictus), often a shorter blunt upper spur, less
prevalent in P. afer and rare in P. swainsonii (Hall 1963).
Discussion
Origin of African spurfowls and ‘groups’
The results presented here demonstrate that African
spurfowls represent an ancient and ongoing biogeograph-
ical, morphological, behavioural and ecological radiation
within the African continent. The existence of a subspecies
of P. bicalcaratus (ayesha; 1% divergent from the nominate
subspecies) in Morocco, which is exceptional amongst
Afrotropical birds (Moreau 1966), demonstrates relatively
recent biogeographic connectivity between North and
sub-Saharan Africa.
African spurfowls are sister to Ammoperdix heyi (native
range from Egypt and Israel east to southern Arabia) and
Perdicula asiatica (native range India, Nepal, Bangladesh,
Pakistan and Sri Lanka) (Crowe et al. 2006), both of which
are arid-zone taxa (Madge and McGowan 2002). Moreover,
the phylogenetically most basal and highly peculiar
African spurfowl, P. hartlaubi, is also a desert-associated
bird (Komen 1987; Little and Crowe 2011). Therefore,
spurfowls are probably derived from an arid-adapted taxon
that dispersed from the Middle East or Asia into Africa
30–40 mybp, during an intercontinental arid era (Crowe
et al. 2006). Hall (1963) also suggested an Asiatic origin.
Within Africa, north-to-south dispersal of proto-hartlaubi
may have been facilitated by an ‘Arid Corridor’ (Figure 2)
that opened (and closed) connecting/disconnecting the
north-east arid Horn of Africa to arid Namibia and the Karoo
in the south-west (Balinsky 1962; Irwin et al. 1962; Vernon
1999; Bobe 2006).
Montane and Scaly spurfowls
The Montane and Scaly spurfowls follow on from P. hartlaubi
paraphyletically (Figure 8). They probably result from
invasions (depicted by lines in Figure 2) by proto-hartlaubi
of, and diversification within, forested biotopes where they
predominated thereafter during subsequent wetter eras. The
marked CYTB divergences between the basal P. hartlaubi,
P. swierstrai, P. camerunensis and P. nobilis (Table 7)
suggest that this radiation was relatively ancient. Thereafter,
when forests subsequently contracted geographically during
renewed dry eras, proto-Montane spurfowls became isolated
in relictual, island-like patches of montane forest (e.g. Crowe
and Crowe 1982; Fjeldså and Bowie 2008; Voelker et al.
2010). This scenario suggests that the two earliest emergent
taxa in this radiation were the relatively basal, most isolated,
western Montane taxa (P. camerunensis and P. swierstrai),
being geographically most proximal to the hill/mountain-
dwelling P. hartlaubi and similarly relatively small, sexually
dimorphic, and poorly spurred.
The Handsome Spurfowl P. nobilis is geographically
intermediate (Figure 2) between western and north-eastern
African Montane taxa and is sister to P. camerunensis. It
is phylogenetically ‘intermediate’ between the western taxa
and those in the north-east. It is also of intermediate body
mass between the two species assemblages (Little 2016a).
The divergence of Scaly from Montane spurfowls is
probably more a consequence of further ecologically
opportunistic speciation in secondary and riverine forests
during multiple expansions and contractions of lowland
forests separated by intervening savanna/steppe, hence
the relatively close genetic propinquity between Montane
P. jacksoni and Scaly P. griseostriatus, and Montane
P. swierstrai and Scaly P. squamatus. The anomalously
low CYTB molecular divergence values between Scaly taxa
and P. hartlaubi and Montane taxa (Table 7) may indicate
ancient hybridisation between these taxa, hence the poor
cladistic nodal support in their respective sections (Figure 8).
Although the core ranges of P. ahantensis and
P. squamatus closely coincide with the current distribution
of present-day lowland forest, the existence of peripheral,
island-like isolates suggests a much broader continuous
distribution in more widespread forest during wetter eras.
Indeed, the primordial ‘scaly’ spurfowl may have been a
single species distributed continuously from West Africa
eastwards to the East African coast and south to Angola,
with an initial vicariance event producing P. griseostriatus.
The second major forest vicariance event and physical
barrier of the Niger River may have split P. ahantensis from
P. squamatus.
Furthermore, vicariant speciation within proto-squamatus
may have promoted the divergence of P. schuetti in paleo-
forest isolates in the east (Figure 4) during drier eras, as

Mandiwana-Neudani, Little, Crowe and Bowie
166
it seems to have done within Latham’s Forest Francolin
Afrocolinus lathami (Mandiwana-Neudani 2012) and
Plumed Guineafowl Guttera plumifera (Crowe and Crowe
1985). Finally, Hall (1963) noted that P. squamatus extends
its range to higher altitudes on mountains uninhabited by
Montane spurfowls, suggesting that competition might also
have restricted its range.
Vermiculated and Bare-throated spurfowls
Moving into relatively open arid-steppe, savanna, woodland
and bush biotopes, the vicariant speciation of Vermiculated
and Bare-throated taxa within pockets of these biotopes
was probably promoted by physical barriers (lakes, rivers
and valleys), other geomorphological events and the
expansion and contraction of forest biotopes (Crowe and
Crowe 1982; Cotterill 2003; Fjeldså and Bowie 2008).
For example, the southern Vermiculated P. hildebrandti
and P. natalensis occupy similar habitats and are separated
by the valleys of the Shire and Luangwa Rivers (Hall 1963).
Within the northern Vermiculated taxa, Lake Chad (Figure 5)
probably played a similar role in speciation between proto-
bicalcaratus and proto-icterorhynchus + clappertoni (Crowe
and Crowe 1982). These latter two spurfowls perhaps
diverged in broad stretches of arid (P. clappertoni) and
mesic (P. icterorhynchus) savanna/grassland. Riverine
forest along the Nile River and in Kenya/Uganda could also
have separated proto-icterorhynchus in the north from proto-
hildebrandti in the south (Hall 1963).
The initial divergence of Vermiculated taxa probably
occurred in central/southern Africa with the proto-
southern taxa radiating within the region into xeric western
(adspersus + capensis) and mesic eastern (hildebrandti
+ natalensis) clades. Northern taxa may be a result of
‘invasion’ from the south via the ‘Arid Corridor’ (Figure 2).
In sharp contrast, the northward dispersal of P. bicalcaratus
from West Africa into Morocco was probably via a relatively
recent corridor of savanna biotope that subsequently
reverted to the western Sahara.
With regards to Bare-throated taxa, proto-leucoscepus
originated in arid biotopes in the north and subsequently
dispersed southwards, once again via the ‘Arid corridor’,
with proto-cranchii/afer/swainsonii biogeographically
insinuating themselves within the savanna- and bush-living
southern Vermiculated taxa.
Ecological speciation due to competition may also have
contributed to speciation in Vermiculated taxa. Those
north of the equator are birds of grasslands and cultiva-
tion in woodland, savanna and steppe and their evolution
may have been influenced by competition with francolins.
However, south of the equator, these habitats are occupied
by Bare-throated taxa, and southern Vermiculated taxa are
relegated to thickets on rocky hillsides and along rivers.
Relevance of the ‘realm’ of tokogeny
There is also evidence that tokogenetic processes, hybrid-
isation and the resulting horizontal gene flow may have
played and still play significant roles in the evolution of
Pternistis species and subspecies. Interbreeding is most
apparent between and among the Vermiculated and
Bare-throated taxa, which may continue to ‘hybridise’
where they come into contact in nature or anthropogenically
induced sympatry. For example, where P. cranchii and
P. afer hybridise along the ‘Arid corridor’ and especially in
eastern Zambia west of the Luangwa River, south to 13°30′
S and in the Eastern Province plateau in Lundazi, hybrid
forms show remarkably high within-locality morphological
homogeneity, forming microgeographic ‘races’. However,
where P. cranchii and P. afer hybridise in southern Angola
and northern Namibia, there is no such localised morpho-
logical homogeneity. Indeed, Roberts (1947) described
a ‘new species’ of spurfowl, P. cooperi, from near Harare,
Zimbabwe (Figure 7), which turned out to be a hybrid
between P. cranchii and P. swainsonii, probably due to
range expansion by P. swainsonii into P. cranchii habitat
that was transformed by agriculture (Little and Crowe 2011).
McCarthy (2006) also reports a range of spurfowl hybrids
in the wild, mainly within and between Vermiculated and
Bare-throated taxa: afer × leucoscepus; afer × swainsonii;
bicalcaratus × erckelii; castaneicollis × erckelii; hildebrandti
× natalensis; leucoscepus × rufopictus; natalensis ×
swainsonii; adspersus × natalensis (van Niekerk 2009);
and adspersus × swainsonii reported by Little (2016b,
2016c). Van Niekerk (2009) reported hybridisation between
Red-billed P. adspersus and Natal P. natalensis spurfowls in
captivity. More recently, D Engelbrect (University of Limpopo)
published a photograph of an alleged hybrid between a
Swainson’s Spurfowl P. swainsonii and Crested Francolin
Ortygornis sephaena taken in the Borakalao National Park
near Polokwane, Limpopo, South Africa, where the mixed
pair were observed making advertisement calls side by side
(http://www.zestforbirds.co.za/hybridfrancolin1.html).
Perhaps the most intriguing taxon in this regard is
P. rufopictus, which Hall (1963) speculated might have
resulted from stabilised hybridisation because it is
‘diagnosed’ by a combination of characters of the other
Bare-throated taxa (e.g. orange, rather than red or yellow
facial skin) and ‘hybrid’ (vermiculated, chestnut, white
and black) plumage. Genetically, it is ~4% divergent
from P. leucoscepus, <2% from P. cranchii and hybrids,
and 2.4% to 3.2% from P. afer. Phylogenetically, it ‘links’
P. cranchii with P. afer. Vocally, it sounds very similar to
P. leucoscepus except that its call is ‘faster’. The strophes
of P. leucoscepus and P. rufopictus are both high-pitched,
with an element of screeching and more protracted trilling
(Mandiwana-Neudani et al. 2014). Nevertheless, its
specific status seems appropriate since it seems to exist
partially sympatric with P. leucoscepus and P. afer without
unfettered hybridisation. Its putative hybrid origins remain to
be tested using genomic data.
Acknowledgements — We acknowledge Pat Hall’s contributions
as a pioneering systematist/biogeographer and as a generous
mentor to TMC. We thank the curators and collection managers at
the American Museum of Natural History (New York), The Natural
History Museum at Tring, Humboldt-Museum (Berlin), the National
Museum of Natural History – Smithsonian Institution (Washington,
DC), the Ditsong National Museum of Natural History (Ditsong
Museums of South Africa), the Iziko Museum of Natural History,
the Natural History Museum of Zimbabwe (Bulawayo), the Field
Museum of Natural History (Chicago), Carnegie Museum of Natural
History (Pittsburgh), Academy of Natural Sciences of Philadelphia,
Natural History Museum of Los Angeles County, Museum of
Comparative Zoology (Harvard University), and the Royal Museum

Ostrich 2019, 90(2): 145–172
167
for Central Africa (Turvuren) for access to specimens under their
care. We thank Robert Moyle for help with designing primers
and Julie Feinstein for helping with the sequencing of toe-pads
subsampled from museum skins. This project was funded by
the South African Department of Science and Technology (DST)
and the National Research Foundation (NRF) through their
South African Biosystematics Initiative and Centre of Excellence
Programmes (grant number UID: 40470), and the African Gamebird
Research, Education and Development Trust. Amy Bruce kindly
assisted with constructing the figures.
ORCID
Tshifhiwa Mandiwana-Neudani https://orcid.org/0000-0001-7276-
5983
Robin Little https://orcid.org/0000-0003-3167-182X
Timothy Crowe https://orcid.org/0000-0002-6430-8085
Rauri Bowie https://orcid.org/0000-0001-8328-6021
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Associate Editor: Graeme Oatley

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Appendix 1: Putative spurfowl taxa (n = 88) sensu Hall (1963) examined morphologically. Those from which molecular marker data were
obtained are annotated with the markers concerned
Bare-throated Group
Francolinus leucoscepus leucoscepus (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH), infuscatus (CYTB), muhamed-ben-abdullah (CYTB),
holtemulleri, kilimensis, tokora
F. rufopictus (CYTB)
F. afer
‘Black and white’: afer (CYTB,12S), palliditectus, castaneiventer (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH), notatus, lehmanni,
nudicollis (CYTB), krebsi, humboldtii, swynnertoni (CYTB), melanogaster, loangwae, leucopareus
‘Vermiculated’: cranchii (CYTB), intercedens (CYTB), harterti (CYTB), benguellensis (CYTB), itigi (CYTB), bohmi (CYTB), nyanzae (CYTB),
cunenensis (CYTB)
F. swainsonii swainsonii (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH), chobiensis (CYTB), damaraensis, lundazi (CYTB)
Montane Group
F. swierstrai
F. camerunensis (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH)
F. nobilis nobilis (CYTB), chapini
F. jacksoni jacksoni (CYTB), gurae
F. castaneicollis castaneicollis (CYTB), atrifrons, bottega (CYTB), ogoensis (CYTB), gofanus, kaffanus
F. erckelii erckelii (CYTB, ND2, CR), pentoni
F. ochropectus (CYTB)
Scaly Group
F. squamatus squamatus (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH), maranensis (CYTB), schuetti (CYTB), tetraoninus, zappeyi,
usambarae, uzungwensis
F. ahantensis ahantensis, hopkinsoni
F. griseostriatus (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH)
Vermiculated Group
F. hartlaubi hartlaubi (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH), crypticus (CYTB), bradfieldi
F. adspersus adspersus (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH), kalahari
F. capensis (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH)
F. bicalcaratus bicalcaratus (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH), adamauae (CYTB), ayesha (CYTB), thornei (CYTB),
ogilviegranti
F. clappertoni clappertoni (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH), nigrosquamatus (CYTB), sharpii (CYTB), cavei, gedgii, heuglini,
konigseggi
F. icterorhynchus icterorhynchus (CYTB), grisescens, emini, ugandensis, dybowskii
F. harwoodi (CYTB)
F. hildebrandti hildebrandti (CYTB, CR), altumi (CYTB), fischeri (CYTB), helleri (CYTB), johnstoni (CYTB)
F. natalensis natalensis (CYTB, ND2, CR, 12S, OVOG, TGFB, GADPH), thamnobium, neavei

Ostrich 2019, 90(2): 145–172
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Appendix 2: Sources of mitochondrial molecular marker data for 49 (CYTB), 13 (12S), 13 (CR) and 14 (ND2) spurfowl taxa
Hall (1963) taxon Markers and GenBank accession numbers
CYTB 12S Control region ND2
F. hartlaubi hartlaubi FR691618 FR691555 FR716656 FR691572
F. h. crypticus FR691619
F. adspersus adspersus FR691623 DQ832113 FR691381 DQ768276
F. afer afer FR694158 DQ832111 DQ834533 FR691579
F. a. benguellensis FR694159
F. a. bohmi FR694162
F. cranchii cranchii FR694164
F. afer cunenensis FR694160
F. a. harterti FR694161
F. a. intercedens FR694165
F. a. itigi FR694166
F. a. nudicollis FR694163
F. a. nyanzae FR694167
F. a. castaneiventer AM236908 DQ832111 DQ834533 DQ768280
F. a. swynnertoni FR694168
F. bicalcaratus bicalcaratus FR691624 FR691551 FR691370 FR691578
F. b. adamauae FR691626
F. b. ayesha FR691625
F. b. thornei FR691627
F. camerunensis FR691591 FR691552 FR691382 FR691577
F. capensis AM236909 DQ832112 DQ834534 DQ768282
F. castaneicollis castaneicollis AM236903
F. c. bottegi FR691629
F. c. ogoensis FR691628
F. clappertoni clappertoni FR691602 FR716655 FR691383 FR691576
F. c. nigrosquamatus FR691604
F. c. sharpii FR691603
F. erckelii FR691589 FR691553 FR691575
F. griseostriatus AM236905 FR691554 FR691384 DQ768284
F. harwoodi FR691600
F. hildebrandti hildebrandti FR691595 FR691385
F. h. altumi FR691597
F. h. fischeri FR691598
F. h. helleri FR691599
F. h. johnstoni FR691596
F. icterorhynchus icterorhynchus FR691601
F. jacksoni FR691594
F. leucoscepus leucoscepus AM236906 FR691556 FR691387 DQ768283
F. l. infuscatus FR691587
F. l. muhamed-ben-abdullah FR691586
F. natalensis AM236911 FR691557 DQ834536 DQ768285
F. nobilis FR691592
F. ochropectus FR691590
F. rufopictus FR691588
F. squamatus squamatus AM236904 DQ832109 FR691388 DQ768286
F. s. maranensis FR691630
F. s. schuetti FR691631
F. swainsonii swainsonii AM236907 DQ832110 DQ834532 DQ768287
F. s. chobiensis FR694170
F. s. lundazi FR694169
F. swierstrai FR691593

Mandiwana-Neudani, Little, Crowe and Bowie
172
Appendix 3: Sources of nuclear molecular marker sequences for spurfowl taxa
Hall (1963) taxon Markers and GenBank accession numbers
OVOG TGFB GAPDH
Francolinus hartlaubi FR691692 FR694129 FR694095
F. camerunensis FR691694 FR694124 FR694090
F. squamatus DQ832088 FR694133 FR694099
F. griseostriatus DQ832089 FR694128 FR694094
F. natalensis FR694132 FR694098 FR694098
F. adspersus DQ832095 FR694122 FR694087
F. capensis DQ832093 FR694125 FR694091
F. bicalcaratus FR691690 FR694103 FR694089
F. clappertoni FR691693 FR694126 FR694092
F. leucoscepus FR694131 FR694097 FR694097
F. swainsonii DQ832091 FR694134 FR694100
F. afer DQ832092 FR694123 FR694088
Appendix 4: Revised classification and English common names for Pternistis spurfowls based on multiple lines of evidence presented in this
study. Order: Galliformes; Family: Phasianidae; Subfamily: Phasianinae; Tribe: Coturnicini
Taxon English common name
Genus: Pternistis
P. hartlaubi Hartlaub’s Spurfowl
P. camerunensis Mount Cameroon Spurfowl
P. nobilis Handsome Spurfowl
P. swierstrai Swierstra’s Spurfowl
P. erckelii Erckel’s Spurfowl
P. ochropectus Djibouti Spurfowl
P. castaneicollis castaneicollis Chestnut-naped Spurfowl
P. c. atrifrons Black-fronted Spurfowl
P. jacksoni Jackson’s Spurfowl
P. squamatus Scaly Spurfowl
P. schuetti schuetti * Schuett’s Spurfowl
P. s. maranensis
P. ahantensis Ahanta Spurfowl
P. griseostriatus Grey-striped Spurfowl
P. hildebrandti hildebrandti Hildebrandt’s Spurfowl
P. h. fischeri
P. natalensis Natal Spurfowl
P. adspersus Red-billed Spurfowl
P. capensis Cape Spurfowl
P. icterorhynchus Heuglin’s Spurfowl
P. bicalcaratus bicalcaratus Double-spurred Spurfowl
P. b. ayesha Moroccan Spurfowl
P. b. adamauae
P. harwoodi Harwood’s Spurfowl
P. clappertoni clappertoni Clapperton’s Spurfowl
P. c. sharpii
P. leucoscepus leucoscepus Yellow-necked Spurfowl
P. l. infuscatus
P. swainsonii Swainson’s Spurfowl
P. cranchii * Cranch’s Spurfowl
P. rufopictus Grey-breasted Spurfowl
P. afer afer Red-necked Spurfowl
P. a. castaneiventer
P. a. humboldtii
* Subspecies elevated to species