Family COLUMBIDAE

Phaps (Phaps) chalcoptera

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Family COLUMBIDAE

Compiler and date details

R. Schodde & I.J. Mason, CSIRO Australian National Wildlife Collection, Canberra, Australia

Introduction

The family Columbidae (pigeons) comprises about 300 to 310 species in about 40 to 45 genera; 25 species in 12 genera occur naturally in Australia and its territories, and three introduced species are established. Two of the indigenous species are now extinct. The Australian fossil record is mid-Miocene from the Lake Eyre basin, and Pleistocene-Holocene from southern Western Australia east to the Nullarbor Plain, south-eastern South Australia, the Lake Eyre basin, south-eastern and north-eastern Queensland, western New South Wales, southern Victoria, Tasmania, and Norfolk and Lord Howe islands; the latter series includes often extant species of the genera Phaps, Geophaps, Leucosarcia, Ptilinopus and Columba. Pigeons are cosmopolitan and widespread in warmer parts of the world, with centres of diversity in Africa-Madagascar, the Neotropics, and particularly Australo-Papuasia where about 76 species in some 21 genera are centred.

Pigeons are solitary or colonial birds that feed either terrestrially or arboreally on seeds and fruit. On the ground their gait is a deliberate walk often with teetering movements of the head; their flight is strong and heavy with some gliding and clattering but no soaring; and they drink by sucking or 'pumping'. Nests are flimsy unlined platforms of twigs; eggs of one or two per clutch are plain-white, dull-glossy and ovoid; young are altricial and nidicolous, blind at hatching with sparse pale buff to russet hair-like down on upper parts; and they are fed by regurgitation on crop-milk produced by parents.

Family-group Systematics

Columbidae are a well-defined monophyletic group with no close relatives except the dodos and solitaires (Raphidae). Infra-familial lineages are poorly understood and their relationships are in a state of flux (see summaries in Sibley & Ahlquist 1972, 1990). The four principal assemblages in Australia, circumscribed conservatively here as tribes because their higher group relationships are unclear, are:

Columbini Leach, 1820 (ordinary pigeons and doves)—tree- and ground-feeding, fruit- and seed-eaters, with medium to long tails, thickly ossified maxillaries, slender palatines with compressed shelf, both caeca and uropygial gland, and no gall bladder; they are also diastataxic. Diverse in the Old and New World, they are poorly represented in Australia with few species in two genera, one of which is introduced.

Macropygiini Bonaparte, 1854 (long-tailed pigeons)—arboreal fruit- and seed-eaters with long tapered tails, much rich brown and/or grey in plumage, a palate as in Columbini, both caeca and uropygial gland, and no gall bladder; all are diastataxic. They are centred with three genera in the Malesian-Melanesian archipelagos, only one of which, Macropygia Swainson, 1837, reaches Australia.

Phabini Bonaparte, 1853 (Australo-Papuan bronzewings)—ground-feeding seed-eaters, usually with short, rounded tails, and eutaxic wings (Miller 1915) that are often glossed green- or purplish-bronze; bronzewings also have a palate as in Columbini, vestigial caeca and small uropygial gland, a gall bladder, and rather slender naked tarsi. There is a marked trend to narrowing of the sternum with reduced notching (W.E. Boles, pers. comm.). Apart from the possible inclusion of African Oena Swainson, 1837 and Turtur Boddaert, 1783, the group is centred in Australia-New Guinea where all nine remaining genera occur; seven are present in Australia, five of which are endemic.

Lopholaimini Bonaparte, 1853 (Australo-Papuan fruit-pigeons)—rainforest-inhabiting arboreal fruit-eaters often with much green in plumage, rather short tails, reduced maxillaries, flattened palatines with extended shelf, and gall bladder but no caeca or (usually) uropygial gland; they are also diastataxic and have thick and extensively feathered tarsi. Their seven or eight genera are centred in the archipelagos of the south-western Pacific with an outlier, Alectroenas G.R. Gray, 1840, in Malagasy; four genera are present in Australia and its territories, with one endemic. Here, Lopholaimini Bonaparte, 1853 is used for this group in preference to Alectroenadini Bonaparte, 1853 and Chrysoenini Bonaparte, 1853 (cf. Bock 1994).

For reviews of familial and infra-familial classification, see Garrod (1874), Gadow (1893), Salvadori (1893), Martin (1904), Peters (1937), Verheyen (1957), Harrison (1960), Sibley & Ahlquist (1972, 1990), Wolters (1975–1982) and Goodwin (1983). Here the genera are arranged in their tribes, as circumscribed above, which commence with Columbini Leach, 1820, after the sequence in Goodwin (1983) and Sibley & Monroe (1990).

Genus-group Systematics

Columba Linnaeus, 1758—Included here in subgenus Columba are the rock pigeons of the species-group Oenas Linnaeus, 1758 of Johnston (1962) and the wood pigeons of his subgroup Palumbus Linnaeus, 1758, after Goodwin (1959). These revisions and those of Boetticher (1954), Corbin (1968), Wolters (1975–1982) and Goodwin (1983) all recognise the Janthina (=Janthoenas Reichenbach, 1853) group as separate, although not as a subgenus as here; this distinctive group comprises six dark, dorsally iridescent, arboreal species occurring around the western Pacific rim.

Macropygia Swainson, 1837—Wolters (1975–1982) indicated the need for a second subgenus to contain the small non-Australian species with short bills and bifurcate breast feathers: M. nigrirostris Salvadori, 1876, M. mackinlayi Ramsay, 1878 and M. ruficeps (Temminck, 1835), which are allopatric and are often treated as a superspecies, as by White & Bruce (1986).

Geophaps G.R. Gray, 1842—Generic limits are currently unsettled by conflicting arrangements; that of Frith (1982) is followed here because it is argued most comprehensively. For alternative arrangements separating Lophophaps Reichenbach, 1853 generically, see Peters (1937) and Wolters (1975–1982); for combining Geophaps with Petrophassa Gould, 1841, see Goodwin (1967b, 1983) and Condon (1975), cf. also Cracraft (1986); and for combining Ocyphaps G.R. Gray, 1842 with Geophaps, see Schodde (1982, Appendix 14) and Sibley & Monroe (1990).

Leucosarcia Gould, 1843—For relationships of this genus, see Goodwin (1967b) and Frith (1982, p. 284).

Ocyphaps G.R. Gray, 1842—However likely it is that this genus is linked closely to Geophaps G.R. Gray, 1842 via G. plumifera Gould, 1842 (Goodwin 1967b; Schodde 1982, Appendix 14), proper documentation of monophyly is needed before the two are combined.

Lopholaimus Gould, 1841—For relationships of this isolated genus, see Peters (1938), Goodwin (1967b) and Frith (1977).

Ducula Hodgson, 1836—Of the seven species-groups defined in Ducula by Goodwin (1960), the prevailingly milky-white islet-inhabiting members of the Myristicivora-group are particularly distinct, see Cadow (1933), Stresemann (1940–1941, p. 56) and Goodwin (1967b). Accordingly, Myristicovora Reichenbach, 1853 is treated here as a subgenus; it was recognised generically by Salvadori (1893), Mathews (1927) and Wolters (1975–1982). Differences among other species-groups are less clear-cut such that recognition of further subgenera is difficult (cf. Goodwin 1960) and at this stage premature, pace Wolters (loc. cit.).

Hemiphaga Bonaparte, 1854—For relationships, see Peters (1938).

Ptilinopus Swainson, 1825—Infra-generic groups and relationships are unsettled despite recent revision. Here only two subgenera are accepted for Australian species: nominotypical Ptilinopus which includes the smaller, short-tailed 'typical' fruit-doves; and Megaloprepia Reichenbach, 1853 which includes eleven species of large and long-tailed Malesian-Papuasian fruit-doves with pale grey heads and large patches or bands of deep red, dark green or black on the breast. Among them are Cain's (1954) subgenera: Leucotreron Bonaparte, 1854 and Ramphiculus Bonaparte, 1854. This concept of Megaloprepia has been defined phyletically by Goodwin (1983, Fig. D13) and taxonomically by Wolters (1975–1982).

Species-group Systematics

Macropygia amboinensis (Linnaeus, 1766)—Implicit in specific limits proposed by Mayr (1944), Goodwin (1967b) and Frith (1982)—and adopted here—is that M. phasianella (Temminck, 1821) of Australia, M. amboinensis (Linnaeus, 1766) of Papuasia-Sulawesi, and M. magna Wallace, 1864 of the Banda Arc and eastern Lesser Sundas are closely related allotaxa and best treated as members of one polytypic species, pace White & Bruce (1986) and Sibley & Monroe (1990). This interpretation is reinforced by the recently described isolate on Cape York Peninsula (Schodde 1989) which is morphologically transitional between eastern Australian phasianella and New Guinean amboinensis in a geographically intermediate region. Subspecific differentiation between central- and north-eastern Australian populations north to the Cairns-Cooktown region requires documentation.

Chalcophaps indica (Linnaeus, 1758)—As stressed by Goodwin (1983), populations of this Malesian-Australasian species fall into two well-marked groups on male plumage: grey-crowned, white-browed and almost plain-shouldered forms in Malesia and brown-headed and extensively white-shouldered forms in Australia. According to White & Bruce (1986), the two groups replace one another abruptly in Wallacea; but R.E. Johnstone (pers. comm.) reports intergradation between them through the Lesser Sundas. Pending clarification of interaction there, they are combined in one species here following current convention. The Christmas Island population is of the Malesian form.

Geopelia humeralis (Temminck, 1821)—Southern New Guinean populations may be subspecifically distinct from northern Australian G. h. inexpectata Mathews, 1912—see Frith (1982) cf. Mees (1982); they may be significantly smaller in size, but confirmation is needed. If so, northern Torres Strait populations on Boigu, Saibai, Dauan and Deliverance islands are probably of the form, G. h. gregalis Bangs & Peters, 1926.

Geopelia placida Gould, 1844—Although Johnstone (1992) recognised no subspecies in Australia, other revisers distinguish three (Condon 1975; Frith 1982); neither arrangement appears satisfactory. Northern and eastern populations grade slightly larger and more intensely grey from north-west to south-east (data in Frith loc. cit.; Johnstone 1992), the shift being clinal (Goodwin in Hall 1974). Accordingly, no subspecies are separated in those populations; Frith (loc. cit.) could not distinguish any either. However, all revisers except Johnstone (loc. cit.) agree that the isolated form in the Pilbara is subspecifically distinct (Mayr 1951; Mees 1961; Condon loc. cit.; Frith loc. cit.), a conclusion corroborated by specimens checked for this volume. Irrespective of local chequer-board variation well analysed by Johnstone (loc. cit.), the Pilbara populations are paler than those from anywhere else, buffier grey dorsally, and have a disproportionally pinker suffusion ventrally. Whether these differences in pigmentation are ecophenotypic, or the erratic similarities between De Grey and southern Kimberley enclaves expressions of past secondary intergradation, are questions that need to be resolved before the conventional view of the Pilbara form is changed.

Given infraspecific differentiation in Australia and probably New Guinea (Rand 1938; Frith 1982), and similar differentiation within at least one of the other major forms of the Geopelia striata complex in Malesia (Peters 1937; White & Bruce 1986), all three principal taxa of the complex are treated here as allospecies after Harrison (1969); they are Malesian striata Linnaeus, 1766, Lesser Sundan maugeus Temminck, 1809 (correct original spelling), and Australo-Papuan placida Gould, 1844. The rationale for this arrangement is explained in the Introduction (Taxonomic Scope) to this volume. In further clarification, placida differs from maugeus or striata or both in its hind-neck barring, ventral barring that is open and restricted to across the whole upper breast, extent of vinaceous cast to the ventrum, reduced bare periorbital skin with whitish post-orbital stripe, chestnut-rufous on the underwing that is dark, immaculate and restricted to the underwing coverts, and more pointed wing in which the penultimate primary is longest.

Geophaps plumifera Gould, 1842—Although combining major regional white- (plumifera Gould, 1842) and cinnamon- (ferruginea Gould, 1865) ventrumed forms in one species, the two current revisions differ in their interpretation of subspecies: Crome et al. (1980) recognised three subspecies and Johnstone (1981a) none, the latter inferring that intergradation between plumifera and ferruginea in the south Kimberley Division and west centralian ranges is primary. Variation in ventrum and iris colour reported by Johnstone (loc. cit.) in these intergrading populations is nevertheless discombinant, indicating that intergradation is secondary; accordingly, ferruginea and plumifera are treated as subspecies here. Among white-ventrumed forms, Kimberley-Victoria River populations differ from those to the east in their greyer dorsa, whiter bellies, contrastingly deeper rufous breast bands and yellower irides, consistent with their subspecific recognition by Crome et al. (1980).

Phaps chalcoptera (Latham, 1790)—Geographical variation is clinal, from darker and larger across southern Australia to slightly paler and smaller in the north, and sandier dorsally in inland regions—see Frith (1982).

Ducula bicolor (Scopoli, 1786)—The taxonomy of this complex is unsettled by two unsatisfactorily resolved questions:

(1) are Columba bicolor Scopoli, 1786 and Carpophaga spilorrhoa G.R. Gray, 1858 conspecific; and

(2) are populations around the fringes of the Arafura Sea, including coastal northern Australia, consubspecific?

Johnstone's (1981b) revision is followed here because it is the only modern review based on a comprehensive analysis of specimens. Sight records of different members of the complex co-occurring or replacing one another on islands in northern Wallacea and Sulawesi should be discounted as evidence for speciation until breeding sympatry is demonstrated cf. White & Bruce (1986). The sedentary, consistently white population in the Kimberley Division is not distinguished from other Australian populations here because those on the north coast of Queensland are often as white and just as large: wings 229–247 mm in 19 males, 222–237 mm in 14 females in collections in Australian museums (ANWC, QM, NMV, SAMA) not examined by Bruce (1989). A variable grey wash in white plumage is characteristic of northern Australian-southern New Guinean populations as a whole (=spilorrhoa G.R. Gray, 1858), and chequer-board variation there seems taxonomically trivial; the explanation given by Bruce (1989) for this variation is sound.

Ptilinopus cinctus (Temminck, 1809)—Despite the current tendency to recognise the grey-and-white population in Arnhem Land specifically (e.g. Wolters 1975–1982; Goodwin 1983; Sibley & Monroe 1990), it is retained as a subspecies of P. cinctus (Temminck, 1809) here because the only detailed character analysis available—still that of Hartert (1904)—treats it as such.

Ptilinopus regina Swainson, 1825—Specific circumscription is uncertain because of the unsettled status of extra-limital forms, notably of the P. r. xanthogaster (Wagler, 1827) group, see implications in Hartert (1904), Peters (1937), and White & Bruce (1986) cf. Johnstone (1981b) and Goodwin (1983). For relationships, see Ripley & Birkhead (1942) and Cain (1954).

Excluded Taxa

Vagrant Species

CAVS:0769
COLUMBIDAE: Ducula concinna (Wallace, 1865) [Elegant Imperial-pigeon; vagrant to NT. Sight record from urban Darwin during 1993, accepted by RAOU Records Appraisal Committee (case no. 181). Elsewhere widespread on islets in Flores and Banda Seas east to Tanimbar and Aru Ils.] — Stanger, M., Clayton, M., Schodde, R., Wombey, J. & Mason, I. 1998. CSIRO List of Australian Verebrates: A Reference with Conservation Status. Collingwood : CSIRO Publishing iii 124 pp. [97]; Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp. [16, 75]

CAVS:9932
COLUMBIDAE: Ptilinopus iozonus G.R. Gray, 1858 [Orange-bellied Fruit-Dove; vagrant to Tores Strait Islands] — Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp. [75]

COLUMBIDAE: Streptopelia orientalis Latham, 1790

CAVS:8056
COLUMBIDAE: Streptopelia tranquebarica humilis (Temminck, 1824) [Single occurrences at Christmas Island]

CAVS:8055
COLUMBIDAE: Streptopelia tranquebarica (Hermann, 1804)

Diagnosis

Plump, densely plumaged, soft-skinned birds in grey, green and brown tones often barred or marked on tail, wing and neck; heads small, bills weak and plover-like with hardened dertrum and fleshly operculum over nostrils; body feathering diffuse, with broad shafts and dense down producing powder downs; aftershafts and down tracts vestigial or absent; uropygial gland naked, small to rudimentary or absent. Feet often pink-red, anisodactylous, the three forward toes and reversed hallux with long basal phalanges; tarsi short, the acrotarsia scutellate and the sides and plantar tarsi reticulate with hexagonal scales. Sexes alike to slightly dimorphic. Wings broad, short to long: 11 primaries, the outer often distinctively emarginate, moulting in staggered descending sequence, 10–15 eutaxic or mostly diastataxic secondaries; tail rounded, short to long: 12–14 rectrices (16 in Goura, 20 in Otidiphaps), moulting in staggered centrifugal sequence. Nares schizorhinal and impervious, nasal septum perforate; schizognathous palate, with slender vomer, lachrymals fused with ectethmoids in a broad passerine-like plate; basipterygoid processes present, small; cervical vertebrae 14–15; sternum either two-notched or one-notched in some Australian phabinine pigeons, both spina externa and broad interna present. Musculus expansor secundariorium consistently present, if reduced, M. tensoris patagium brevis extensive, and biceps slip usually present and independent; pelvic muscle formula usually complete A(B)XY+, usually with M. ambiens (except Ptilinopus, Goura); deep plantar tendons Type 1. Carotid arteries paired. Syrinx tracheo-bronchial, with extrinsic muscles asymmetric due to crop. Tongue small, narrow and cartilaginous; crop present, two-lobed, producing crop-milk from lining; gizzard heavily muscled, less so in fruit-eaters; gall bladder absent except in Australo-Papuasian fruit-pigeons; non-functional caeca small to rudimentary. Diploid karyotype of 76–80 chromosomes, with five pairs of macrochromosomes.