Family CACATUIDAE
Compiler and date details
R. Schodde & I.J. Mason, CSIRO Australian National Wildlife Collection, Canberra, Australia
Introduction
[After Schodde (1997), but updated following Joseph et al. (2012)]
Cacatuidae (cockatoos) comprise about 19–21 species in six genera; fourteen species in all six genera occur in Australia. The Australian fossil record, of mainly extant species, is limited to the Holocene-Pleistocene of southern Victoria and south-eastern South Australia, except for recently recorded material of 'Cacatua' from the early-mid Miocene in north-western Queensland (Boles 1993). The family is centred in Australia, where four of the six genera are endemic; beyond, it ranges through islands in the south-west Pacific to the Philippines and Solomon Islands.
Cockatoos are gregarious birds that feed on the ground or in trees on seeds won from hard fruit by the cutting action of the lower mandible or the scooping action of the maxilla, according to species. They clump-roost, drink by scooping, and bathe only (?) in rain; their voices are characteristically loud and hoarse screeches and whistles. Nests, defended in undefended territories, are of unconstructed beds in hollows or holes; eggs are ellipsoidal, plain-white and slightly glossy, and are incubated by sexes according to subfamily, q.v.; young are altricial, nidicolous, blind at hatching, covered with long yellow down (grey in Probosciger, pink in Eolophus), and they are fed by pump-regurgitation.
Family-group Systematics
In the Catalogue, cockatoos are split from other parrots as a family, for two reasons. Firstly, given that parrot-like birds form a single order (Psittaciformes), their primary lineages are better treated as families than infra-familial taxa wherever they are not so closely linked phylogenetically as to make this unrealistic. Secondly, biochemical and chromosomal studies (Adams et al. 1984; Christidis et al. 1991a, 1991b; Joshua & Parker 1993) have now built on the morphological and behavioural assessments of Beddard & Parsons (1893); Beddard (1898); Thompson (1899); Mudge (1902); Boetticher (1943, 1959); Verheyen (1956); Glenny (1957, 1959); Brereton & Immelmann (1962); Smith (1975) and Homberger (1980, 1991) to single out the cockatoos as a particularly distinctive assemblage among the Psittaciformes, well distant from other parrots. Condon (1975), Wolters (1975–1982), Homberger (1991) and Christidis et al. (1991a) took this view, discounting the idiosyncratic approach of Mathews (1913) and his subsequent revisions which split the cockatoos into further families. The most widespread alternative arrangement follows Fürbringer (1888) in combining cockatoos with all other parrots in a single family, Psittacidae, e.g. Stresemann (1927–1934); Peters (1937); Boetticher (1943, 1959); Mayr & Amadon (1951); Smith (1975); Homberger (1980); Sibley et al. (1988) and Sibley & Ahlquist (1990).
Within Cacatuidae four principal lineages (but cf. Homberger 1991) were recognised by Schodde (1997) as follows:
Microglossinae Bonaparte, 1853 (palm cockatoos)—large, heavily built, slatey-black cockatoos with massive, permanently open bills, coloured facial skin, rather long and square plain tails, finely plumed erectile but uncoloured crests, feathered ceres and little sexual dimorphism; they also head-scratch under the wing, clump and allopreen (?), and only the female broods—young are uniquely grey-downed; diploid karyotype 2n = 74, with four pairs of acrocentric and four pairs of telocentric autosomal macrochromosomes, and Z-chromosome ?metacentric. The single species is shared by Cape York Peninsula and New Guinea, in the genus Probosciger Kuhl, 1820.
Calyptorhynchinae Bonaparte, 1853 (black cockatoos)—large but slender black cockatoos with long and colour-banded tails, uncoloured recumbent crests, naked ceres, and moderate sexual dimorphism; they also head-scratch under the wing, neither clump nor allopreen, and only the female broods, fed by the male; diploid karyotype 2n = 78, with eight pairs of telocentric autosomal macrochromosomes, and Z-chrosomome submetacentric. Their five species comprise one genus, Calyptorhynchus Desmarest, 1826, endemic to Australia.
Cacatuinae G.R. Gray, 1840 (white cockatoos)—stocky white, pink or grey cockatoos with short and usually plain tails, coloured and often up-curved erectile crests, feathered nares, and slight sexual dimorphism (Callocephalon Lesson, 1830 excepted); they also head-scratch under the wing, clump and allopreen, and both sexes brood without support feeding; diploid karyotype 2n = 82 or 76, with 8 pairs of telocentric autosomal macrochromosomes or two pairs acrocentric, and Z-chromosome metacentric (Callocephalon Lesson, 1857 not known). Comprising three genera, the group is centred in Australia where two genera are endemic: Callocephalon Lesson, 1837 and Eolophus Bonaparte, 1854. Callocephalon is included in this lineage based on data in Adams et al. (1984) contra Condon (1975), but its position is problematic, as is that of Eolophus (Homberger 1991). Callocephalon, in particular, may comprise a separate subfamial group (Homberger loc. cit.).
Nymphicinae Bonaparte, 1857 (cockatiel)—diminutive grey cockatoos with slender, attenuate parrot-like wings and tails, up-curved crests, naked ceres, and sexually dimorphic faces and patterned tails; both sexes also head-scratch over the wing, allopreen and brood without support feeding; diploid karyotype 2n = 72, with three pairs of acrocentric, one pair of metacentric and three pairs of telocentric autosomal macrochromosomes, and Z-chromosome submetacentric. The single species is endemic to Australia, in the genus Nymphicus Wagler, 1832. Despite past dispute, there is now an overwhelming consensus of morphological, karyotypic, behavioural and biochemical evidence to show that the cockatiel forms a root lineage among the Cacatuidae (Salvadori 1891; Mudge 1902; Wolters 1975–1982; Adams et al. 1984 and references therein; Christidis et al. 1991b; Birt et al. 1992; Joshua & Parker 1993). Its relationships within the cockatoos, however, are still unclear.
These four lineages were treated as subfamilies by Schodde (1997), after Condon (1975) and data in Adams et al. (1984), Christidis et al. (1991a) and Joshua & Parker (1993), cf. Homberger (1991). However, fowwlowing Joseph et al. (2012), the Microglossinae are reduced in status to a tribe within subfamily Cacatuinae, together with Cacatuini.
Genus-group Systematics
Calyptorhynchus Desmarest, 1826—Two subgenera of black cockatoos are distinguishable. Although recognised by Peters (1937) and Wolters (1975–1982), they have never been diagnosed comprehensively; only their external morphological traits are summarized here. In the red-tailed nominotypical subgenus, adult males are plain black with a plain vermillion band in the tail, and females and/or immatures are banded and spotted yellowish with a yellowish red to yellow band in the tail barred black; ear coverts and periophthalmic skin are plain (or spotted) and black in both sexes. Calyptorhynchus lathami (Temminck, 1807) is a member of this group, as pointed out by Ford (1980). In subgenus Zanda Mathews 1913, the contour plumage is edged yellowish or whitish, and the tail has a broad pale yellow or white band flecked variably with dusky speckling in both sexes; ear coverts are yellow or white (duller in males) and periophthalmic skin is sexually dimorphic: flesh-pink in males, black in females; males also have paler feet. That the two subgenera merit generic separation is a likelihood that needs further study, cf. Adams et al. (1984, Figs 3 and 4).
Cacatua Vieillot, 1817—Limits and lineages in the white cockatoos are not well resolved. Included here in Cacatua Vieillot, 1817 are those species with prevailingly white or pinkish white plumage, frequently coloured crests, prevailingly telocentric karyotype (cf. Christidis et al. 1991b; Joshua & Parker 1993), no palatine processes on the maxillaries (D.G. Homberger, pers. comm.), a single left carotid artery and yellow-downed nestlings. This circumscription is conventional except for the exclusion of the Galah, Eolophus roseicapillus (Vieillot, 1817), q.v. Within Cacatua, three principal species-groups were identified by Adams et al. (1984) (cf. Homberger 1991)—they are treated as subgenera here:
(1) nominotypical Cacatua which includes the broad-winged, black-billed, plain-colour-crested 'white cockatoo' group;
(2) subgenus Licmetis Wagler, 1832 which includes the slender-winged, white-billed, vestigially crested corellas; and
(3) subgenus Lophochroa Bonaparte, 1857, comprising the broad-winged, white-billed, multi-colour-crested Pink Cockatoo, Cacatua leadbeateri Vigors, 1831, which may be of disparate affinity (Homberger 1991).
Eolophus Bonaparte, 1854—The position of the Galah is controversial because morphological evidence favouring monogeneric status (Holyoak 1970; Forshaw 1981; Homberger & Schodde in Schodde 1989) is in conflict with protein evidence grouping it with the corellas in Cacatua Vieillot, 1817 (Adams et al. 1984). The last study, and Boles (1993), presume that the protein evidence—or its computation by the distance-Wagner algorithm—is necessarily correct phylogenetically. Inspection of the protein data indicates that the Galah shares almost as many apparent allelic synapomorphies with the Cockatiel (Nymphicus Wagler, 1832) as with the corellas. Moreover, judged by allelic expression across both calyptorhynchine and cacatuine lineages, there seems to be some homoplasy in the loci screened. The protein-based phylogeny thus needs corroboration. For the Galah, this is not forthcoming from morphology in traits which extend from plumage colour pattern and the structure of the auditory region of the skull (Holyoak 1970) to pink-downed nestlings (Forshaw 1981), form of the wing (Miller 1915), paired carotid arteries (Glenny 1955, in Homberger 1991), karyotype of 2n = 76, with two pairs of acrocentric macrochromosomes (Christidis et al. 1991b; Joshua & Parker 1993), and broadened palate with erect and terminally hooked palatine processes on the maxillaries. These processes, first reported by Homberger & Schodde in Schodde (1989) contra Boles (1993), are missing from all other Cacatuidae except Nymphicus and Callocephalon Lesson, 1837, being vestigial in the latter. Together with the forward attachment of the M. genioglossus in the lower mandible and the absence of fine paired caudo-medial processes on the maxillary shelf fronting the ventral surface of the naso-frontal hinge, they indicate that the oral myology of the Galah is different from that of Cacatua, including the corellas (D.G. Homberger & R. Schodde, pers. comm.). Courtney (1993) recorded ten traits in juvenile begging and external morphology that tied the Galah into Cacatua, but two of them (white = pigmentless contour plumage, colour of down) are misreported and the others can be interpreted as symplesiomorphies in the cacatuine lineage which may include Callocephalon as well. Symplesiomorphic too, it may argued, are the oral osteology and myology of the Galah. Until such questions are answered, cladogenesis among the cacatuine cockatoos will remain unresolved; in the interim, the relatively distinctive Galah is kept in Eolophus Bonaparte, 1854 pending clarification of its position.
Species-group Systematics
Probosciger aterrimus (Gmelin, 1788)—Geographical variation in size, apparently chequered in pattern, is understood from series of specimens that are still too meagre, leaving the limits of subspecies around the Arafura Sea in doubt. The earlier arrangements of five subspecies proposed by Peters (1937) and Mayr (1937) may be more realistic than those put forward by Mees (1957, 1982) and Forshaw (1978). Accordingly, a conservative approach is taken here in accepting as distinct the zoogeographically coherent populations on Cape York Peninsula and Trans-Fly region of New Guinea; these are intermediate in size between the small nominotypical aterrimus on the Aru Islands and the huge P. a. goliath (Kuhl, 1820) throughout rainforested southern New Guinea.
Calyptorhynchus banksii (Latham, 1790)—Ford's thorough revision (1980) leaves several questions unresolved: what the nature of intergradation among northern populations around the eastern head of the Gulf of Carpentaria, and are the similar-looking isolates across inland Australia, between the Western Australian wheatbelt and central Darling River, New South Wales, monophyletic or convergent from different ancestral stocks? In the Catalogue, the similarities among inland populations are taken at face value, as for Cacatua sanguinea gymnopis Sclater, 1871 (q.v.), and treated as representing a single fragmented form. The nexus in northern populations at the head of the Gulf of Carpentaria, moreover, is treated conservatively as secondary, as in so many other Torresian species (Macdonald 1969; Ford 1987). The status of the morphologically and ecologically distinctive southeast Australian isolate, C. b. graptogyne Schodde, Saunders & Homberger, 1989, also needs further clarification.
Calyptorhynchus funereus (Shaw, 1794)—Despite Saunders' (1979) detailed revision, documentation of infra-specific differentiation is incomplete. Zones of intergradation between large eastern and small southern populations on the south-east mainland have not been defined; Courtney (1986) suggested that the two forms are specifically distinct. The isolated population in Tasmania, moreover, may be inversely sexually dimorphic in size. Accordingly, subspecies recognised here are provisional.
Calyptorhynchus latirostris Carnaby, 1948—When Saunders (1979) showed that the two white-tailed black cockatoos in south-western Australia (baudinii Lear, 1832 and latirostris Carnaby, 1848) were better treated as specifically distinct, he combined the short-billed form, latirostris, with the eastern yellow-tailed allospecies, C. funereus (Shaw, 1794). Biochemical evidence, however, indicates that the two white-tailed forms are one another's closest relatives (Adams et al. 1984), a relationship consistent with their colour and finer feet with straighter and slenderer claws. Accordingly, latirostris is treated here as a separate species (also Sibley & Monroe 1990). Mechanisms for its speciation are canvassed in Schodde & Mason (1991).
Cacatua galerita (Latham, 1790)—The last revision of Australian populations (Forshaw 1968) is followed here, pending a more complete analysis of variation in body size, bill form and length and breadth of the crest. If eastern C. g. galerita (Latham, 1790) and north-western C. g. fitzroyi (Mathews, 1912) prove to be subspecifically distinct, their nexus will probably lie along the Carpentaria Barrier around the eastern head of the Gulf of Carpentaria, cf. Macdonald (1969) and Ford (1987).
Cacatua pastinator (Gould, 1841)—Given the likelihood that C. sanguinea Gould, 1843 has only recently invaded the range of pastinator Gould, 1841 from the north, their treatment by Ford (1985) as distinct species is followed contingent on their continuing sympatry. Evidence of contact older than the last several decades is based on one unsatisfactory sight record. The alternative treatment of pastinator and sanguinea as conspecific by Schodde et al. (1979) was also expressly provisional for the same reason, a point commonly misquoted by omission in citations.
Cacatua sanguinea Gould, 1843—Following Schodde et al. (1979), the two isolated but similar-looking eastern and western populations across inland Australia are combined subspecifically under C. s. gymnopis Sclater, 1871. Ford (1985) separated them but his data show that they are barely different morphologically; their present geographical separation may be no older than post-glacial times and transient. In discounting relictual intergradation between gymnopis and nominotypical sanguinea in the southern Kimberley, Ford (loc. cit.) did not consider the primary diagnostic character (Schodde et al. loc. cit.): residual red feathering around lores and eyes. The issue needs clarification, cf. Ford (1987).
Eolophus roseicapillus (Vieillot, 1817)—Nowhere in literature are the three forms of the Galah well diagnosed. In summary, western E. r. roseicapillus (Vieillot, 1817) has grey crustose periophthalmic skin, a pale pink crown grading into a deeper pink collar, and pale grey rump; eastern E. r. albiceps Schodde, 1989 has warty red periophthalmic skin, a pinkish white crown sharply demarcated from a deeper pink collar, and white-grey rump; and northern E. r. kuhli (Mathews, 1912) has the periophthalmic skin of albiceps, the pinker crown of nominotypical roseicapillus (particularly north-westwards) and is smaller and paler than both. Nominotypical roseicapillus and albiceps intergrade on a broad front through central Australia, details of which have yet to be documented. Since the name roseicapilla Vieillot, 1817 was shown to apply to the western form (Schodde 1989), there has been a tendency to apply the name howei Mathews, 1917 to the eastern form in its stead. The holotype of howei, nevertheless, is from the zone of intergradation between the two (type locality: between Claravale and Ruby Gap, about 100 km northeast of Alice Springs) and its traits intergrade. Accordingly, it is treated here as unidentifiable, i.e. as incertae sedis.
Nymphicus hollandicus (Kerr, 1792)—A modern revision of geographical variation is lacking.
Diagnosis
Stocky, usually large, erectile-crested parrots with large hooked and cered bills, and prevailingly white, black or grey plumage without green or blue tones conferred by 'dyck texture'; body feathering coarse, in diffuse tracts, powder downs present in lumbar region and otherwise scattered; aftershafts present, downy; uropygial gland small to vestigial. Feet zygodactylous; tarsi short with fine reticulate scaling, toes longer, reticulately scaled, holding food to bill. Sexes similar (Cacatua, Probosciger, Eolophus) or dissimilar (other genera). Wings long and broadly rounded: 10 primaries and 10 diastataxic secondaries moulting erratically; tail broadly rounded (except Nymphicus): 12 rectrices moulting erratically. Nares holorhinal and impervious, nasal septum imperforate; desmognathous palate, with or without vestigial vomer; maxillaries articulating with deep, bilaterally compressed and twisted palatines, naso-frontal hinge jointed by a diarthrosis or ligamentis syndesmosis; lachrymals extended in processes to reach zygomatic processes, encircling orbit and bridging the temporal fossae (Nymphicus (sometimes) and Calyptorhynchus subg. Zanda excepted); basipterygoid processes absent; cervical vertebrae (13-14); sternum entire, only spina externa present, clavicles ankylosed in furcula. Musculus expansor secundariorum and biceps slip absent, M. tensor patagium brevis with wristward slip, posterior M. deltoideus larger than anterior; pelvic muscle formula AXY, M. ambiens present; deep plantar tendons Type I. Carotid arteries paired in hypophyseal canal or single through coalescence. Syrinx bronchial, the first rings straight, cartilaginous and separated by membrane. Bare orbital skin around eyes often enlarged and coloured; tongue muscular, tactile, grooved, moved by hyoid apparatus with extensive median foramen in entoglossum; crop present; gall bladder present; no caeca. Diploid karyotype of 72–82 chromosomes, with 7–8 pairs of prevailingly telocentric macrochromosomes.
General References
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Boetticher, H. von 1943. Gedanken über die systematische Stellung einiger Papageien. Zoologischer Anzeiger 143: 191-200
Boles, W.E. 1993. A new cockatoo (Psittaciformes: Cacatuidae) from the Tertiary of Riversleigh, northwestern Queensland, and an evaluation of rostral characters in the systematics of parrots. Ibis 135: 8-18
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History of changes
| Published | As part of group | Action Date | Action Type | Compiler(s) |
|---|---|---|---|---|
| 10-Nov-2020 | 14-Oct-2013 | MODIFIED | ||
| 12-Feb-2010 | (import) |