Taxonomy, phylogeny and biogeography of African spurfowls (Galliformes, Phasianidae, Coturnicinae, Pternistis spp.)

During much of the 20th Century, partridge/quail-like, Afro-Asian phasianine birds referred to commonly as African spurfowls, francolins and/or partridges had a tortuous taxonomic history. Because of striking autapomorphic differences in plumage, vocalizations and ecology in some of these taxa, as many as nine genera and nearly 200 clinal and/or idiosyncratic subspecies, embedded within a polyphyletic Perdicinae, were recognized. In 1963, two clades, 28 ‘francolin’ and ‘spurfowl’ species (fisante in Afrikaans) and 13 Afro-Asiatic ‘francolins’ and ‘partridges’ (patryse in Afrikaans), were combined into a single genus (Francolinus) – the largest within the Galliformes – comprising about 100 subspecies. Furthermore, fisante and patryse were partitioned into several unnamed “Groups” and four “Unplaced” species. Here, we use morphological, behavioural and DNA evidence to produce: a comprehensive revision of the taxonomy and phylogeny of the fisante clade; a stable classification system for tis component taxa; and hypotheses vis-à-vis eco-biogeographical processes that promoted their speciation and cladogenesis. We shift Francolinus spp. sensu stricto (members of the Spotted Group) and the Unplaced ‘Francolinus’ gularis from the fisante clade to the patryse [discussed in Mandiwana-Neudani et al., in review]. An Unplaced fisant, ‘F.’ nahani, is linked with Ptilopachus petrosus (another African endemic ‘partridge‘) within the Odontophoridae (New World ‘Quails‘). We recognize 25 species of fisante (hereafter spurfowls), including seven with subspecies. They comprise 34 terminal taxa placed within a single genus, Pternistis, sister to Ammoperdix and Perdicula spp., Coturnix ‘quails’ and Alectoris ‘partridges‘, within the now monophyletic Coturnicinae. Only one of four putative Groups of spurfowls, the Bare-throated Group, is monophyletic. The other three Groups (Montane, Scaly and Vermiculated) are para- or polyphyletic. Several species pairs of spurfowls, most notably P. afer and cranchii, hybridize in para/sympatry. One Bare-throated spurfowl, P. rufopictus, may be the product of stabilized hybridization between P. afer and/or cranchii and P. leucoscepus.


Introduction
During much of the 20 th Century, there was little consensus relating to the taxonomy and phylogeny 43 of Afro-Asian quail and partridge-like galliforms within the Phasianidae, variously commonly known 44 as francolins, spurfowls and/or partridges. As many as nine genera [1] and nearly 200 clinal and/or 45 idiosyncratic subspecies [2], embedded within a polyphyletic Perdicinae [3] were recognized. In 1963, 46 Mrs B.P. 'Pat' Hall comprehensively revised the taxonomy of many of these taxa [2]. She argued 47 convincingly that they should be combined within a single 'mega-genus', Francolinus, comprising 41 48 species -the largest genus in the order Galliformes and the twelfth largest in Aves [4]. Thirty-six of 49 these species are endemic to Africa, five to Asia. Hall also synonymized many subspecies, reducing 50 the nearly 200 to just over 100 [2]. 51 However, literally immediately after this 'lumping/ synonymizing' exercise, Hall divided 52 "francolins" into two, unnamed, putatively monophyletic major clades, comprised of eight (also 53 unnamed and putatively monophyletic) "Groups" and four "Unplaced" species [2]. The major clades 3 54 of francolins correspond to what Afrikaans-speakers commonly refer to as fisante ('pheasants') and 55 patryse ('partridges') [5,6]. We deal with the patryse elsewhere [Mandiwana-Neudani et al., in 56 review]. Hall's fisante (hereafter spurfowls) included an Asiatic Spotted Group (incorporating the 57 nominate species F. francolinus and congeners), four other Groups (Vermiculated, Montane, Scaly and 58 Bare-throated) and two Unplaced species (nahani and gularis) [2]. Morphologically, these taxa 59 generally have: plain or plain-vermiculated back-plumage; brown/black/red tarsi with long -60 sometimes multiple -spurs; emit raucous, grating vocalizations; and roost/perch in large bushes or 61 trees [6]. Members of one Group of spurfowls (the Bare-throated), differ from the others in having 62 bare, brightly coloured skin around the eye and/or on the throat [2]. 63 Within the spurfowls, Hall [2] recognized 28 species, which are generally sexually monomorphic, 64 with males (and females of some taxa) of most species having at least a single (often two), long tarsal 65 spurs. The species differ markedly in plumage, ecology, behaviour and distributional patterns [7,8,9].  'Francolin' [21] should also be removed from the spurfowl clade and placed outside francolins sensu 91 Hall as sister to another phylogenetically enigmatic African 'partridge', Ptilopachus petrosus [22]. 92 These now congeneric Ptilopachus spp. Morpho-behavioural characters 161 The basic body plan of study skins was divided into discrete sections (Fig 1) and scored for 162 variation in colour and patterning: 33 organismal characters reflecting assessment of 163 plumage/integument colour/pattern, measurements of study skins, and vocal characters (

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For within-group molecular analyses of spurfowls, 51 putative terminal taxa were studied (Table 3)

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Morpho-behavioural characters 236 Character information for morpho-behavioural characters are presented in Table 6. 237 Table 6. Morpho-behavioural character scores matrix used for the phylogenetic analysis of spurfowls.
Character reference numbers (see Table 4). The 'total evidence' parsimony analysis based on 5149 characters (33 organismal and 5116 DNA 240 bases) and 33 terminal taxa produced two equally parsimonious trees of length 2124, the strict 241 consensus of which is presented as (Fig 2). Pternistis camerunensis is sister to nobilis. It is endemic to, and locally distributed within dense 324 undergrowth and edges of forests on the south-eastern slopes of Mt. Cameroon, between 850 and 2100 325 m above sea level (Fig 3) [54]. The Mt Cameroon Spurfowl is a small (male ~593 g., female ~509 g.), 326 sexually dimorphic spurfowl, and has an orange-red bill, red eye-ring, and orange-red tarsi with 1-2 327 spurs, only in males. The male has a dark brown crown and nape. Its throat is grey-buff with the belly and is locally common in dense undergrowth, forest edge and moist bamboo thickets (Fig 3). north-western southern Africa (Fig 4) [54]. The closest CYTB taxon is its sister-species, P. schuetti, at 3.4% sequence divergence. The closest CYTB taxon to schuetti is griseostriatus at 2.7% sequence divergence. Ghana through the central Togo and central Benin to south-western Nigeria (Fig 4). 460 The Ahanta Spurfowl is a medium-sized spurfowl (males +-608 g., females +-487 g.) and has an The closest CYTB taxon to ahantensis is P. squamatus at 4.2% sequence divergence.

470
Pternistis griseostriatus is a small spurfowl (males 265-430 g., females 213-350 g. [54]) endemic to 471 vestigial patches of forest in the Angolan western escarpment (Fig 4).   (Fig 6). Heuglin's Spurfowl has a yellow-orange black bill, small yellow eye-patch, yellow-orange 552 tarsi with 1-2 (upper longer), in males only. It is monotypic and sexually monomorphic species (Fig 6) Pternistis bicalcaratus comprises three sexually monomorphic subspecies (Fig 6). northern Cameroon and southern Chad (Fig 6). 563 The Zimbabwe, Mozambique into eastern South Africa (Fig 8)  Zambia, western Tanzania, Uganda and Lake Victoria shores (Fig 8). In the nominate P. a. afer, confined to south-western Angola (Fig 8) 692 and afer have been described where these three forms are para/sympatric, but they lack the 693 morphological cohesion necessary for recognition. These occur mainly in two hybrid zones between