Family CUCULIDAE
Compiler and date details
R. Schodde & I.J. Mason, CSIRO Australian National Wildlife Collection, Canberra, Australia
Introduction
Cuculidae (true cuckoos and koels) comprise about 46–54 species in about 13–17 genera, excluding phaenicophaeids (malcohas) and New World coccyzids, crotophagids and neomorphids; 12 species in five genera are accepted here as occurring in Australia and its territories. The Australian fossil record is of extant genera from Quaternary deposits across southern Australia. The family is centred today in the Palaeotropics, ranging from Eurasia, Africa and Madagascar to the Malesian archipelagos and Australasia.
Rather solitary and usually unobtrusive and secretive, cuckoos are arboreal perchers that feed on small invertebrates and fruit by perch-and-pounce and scrambling among shrubbery, woodland and forest. Many species are migratory; flight is graceful and often swift on usually pointed wings, with or without undulations. Males call monotonously by day and often at night from song posts during breeding; mating systems—monogamous, polygamous or promiscuous—are not well understood. Breeding system parasitic, the female depositing her eggs in the nests of other birds (passerines) usually at the rate of one per nest; host specialization is highly developed, involving egg and nestling mimicry and hereditary lines (gentes) within species adapted to particular host taxa; reproductive success is facilitated by timing of laying to coincide with the host's, by shorter incubation period allowing cuckoo to hatch first, and by removal of host's eggs by breeding female cuckoo; young are reared by hosts.
Family-group Systematics
Both the limits and composition of cuculiform families are in a state of flux, see the conflicting treatments among Peters (1940), Mayr & Amadon (1951), Wetmore (1960), Wolters (1975–1982) and Sibley et al. (1988), Sibley & Ahlquist (1990) and Sibley & Monroe (1990). Because of significant morphological and behavioural differences additional to the large genomic distances found among the principal lineages by Sibley et al. (loc. cit.) and Sibley & Ahlquist (loc. cit.), circumscription of Cuculidae Leach, 1820 itself is limited here to the Cuculinae of Peters (loc. cit.), cf. Wolters (loc. cit.) and Sibley & Monroe (loc. cit.). Their 'hawk'-like form, smooth-textured plumage, structure of sternum and pelvic girdle, pelvic musculature, tracheo-bronchial syrinx, arboreal niche and brood parasitism together set them apart from other groups of Cuculiformes. The Afro-Eurasian Clamator Kaup, 1829 complex is nevertheless aberrant in its cresting and the structure of its sternum, pelvis and pelvic muscle formula, and may not belong in the family (also Wolters loc. cit.). Infra-familial groups are also unclear, Wolters (loc. cit.) having separated the koels as well at family level; their definition is avoided here.
Genus-group Sytematics
Cuculus Linnaeus, 1758—Cuculus pallidus (Latham, 1802) is kept in Cuculus Linnaeus 1758, as is conventional, but recognised subgenerically because of its syringeal structure, perforate atlas and relatively plain ventrum. Wolters (1975–1982) distinguished it generically while Mayr (1964) interpreted it as a link between Cuculus and Cacomantis S. Müller, 1843. Nevertheless, it has the cuculine, not cacomantine traits of multiple wing bars, 11 secondaries and three pairs of lumbar foveae, and it exhibits vestigial cuculine ventral barring.
Cacomantis S. Müller, 1843—Although Condon (1975) and White & Bruce (1986) combined Cacomantis in Cuculus Linnaeus, 1758, following Mayr (1964), no substantiating case has been documented. The members of Cacomantis are distinguished by simple broad wing bar, plain rufous ventrum (excluding paedomorphic C. sonneratii (Latham, 1790)), 10 secondaries and two pairs of lumbar foveae. Other traits, including patterns of moult, have been discussed by Stresemann & Stresemann (1961) and Devillers (1976–1980) when recognising Cacomantis, as is conventional. Within Cacomantis, there are two species-groups which are treated here as subgenera: nominotypical Cacomantis with coarsely mottled immature plumage that is retained into adulthood in C. sonneratii (Latham, 1790), and subgenus Vidgenia Mathews, 1918 in which juveniles are comparatively plain; Vidgenia is Australo-Papuasian-centred.
Chalcites Lesson, 1830—The recent trend towards combining the duller Australo-Asian bronze-cuckoos (Chalcites Lesson, 1830) with the brighter Afro-Asian glossy-cuckoos (Chrysococcyx Boie, 1826) follows Berger (1955) who had limited material of the Australo-Asian members for his anatomical review. Even revisers adopting this arrangement since have viewed these two groups as phylogenetically distinct regional entities (Friedmann 1968; Marchant 1972); accordingly, they are separated generically here, as was traditional (Shelley 1891; Hartert & Stresemann 1925; Peters 1940; Stresemann & Stresemann 1961; and Wolters 1975–1982). The morphological traits distinguishing the two groups are summarized by Marchant (loc. cit.: 225), to which can be added the consistently broad simple wing bar in Australo-Asian Chalcites, in contrast to frequently multiple wing bars in Afro-Asian Chrysococcyx. However divergent, Chalcites osculans (Gould, 1847) is considered a member of the Australo-Asian group by consensus (Friedmann loc. cit.; Wolters loc. cit.; cf. Marchant loc. cit.; Harrison 1973); it has the wing pattern of that group but is here distinguished subgenerically after Wolters (loc. cit.), given its distinctness in the group (Stresemann & Stresemann loc. cit.; Marchant loc. cit.). Despite recent attention (Friedmann loc. cit.; Marchant loc. cit.; Harrison loc. cit.), phylogenetic relations in the bronze- and glossy-cuckoos are still far from resolved.
Eudynamys Vigors & Horsfield, 1827—Mayr's (1944a) broad definition is followed but Urodynamis Salvadori, 1880 is kept separate subgenerically (cf. Stresemann & Stresemann 1961) pending clarification of the phylogeny of the koels, including New Guinean Caliechthrus Cabanis & Heine, 1863.
Species-group Systematics
Cuculus saturatus Hodgson, 1843—According to material in Australian museums, Oriental Cuckoos reaching Australia on migration are all of the large northern Palaearctic subspecies horsfieldi Moore, 1857 = optatus Gould, 1845 according to measurements by Roselaar in Cramp (1985); cf. Vaurie (1965); Condon (1975).
Cacomantis flabelliformis (Latham, 1802)—The case made by Mason (1982) for reviving flabelliformis Latham, 1802 as the senior synonym for this species is followed here for the additional reason that the junior name pyrrophanus Vieillot, 1817 is a source of nomenclatural confusion in Australian ornithology: it has been used both for this species by Condon (1975) and for the Brush Cuckoo Cacomantis variolosus (Vigors & Horsfield, 1827) by the RAOU Checklist Committee (1926). For summary of other nomenclatural complications involving pyrrophanus Vieillot, 1817, see Peters (1940: 27, footnote), Amadon (1942), and Mason (loc. cit.).
Cacomantis variolosus (Vigors & Horsfield, 1827)—Hartert's (1925) demonstration of two Australian subspecies—one eastern, the other paler and smaller in the north — is followed because the cases made against it (Peters 1940: 26, footnote; Condon 1975; White & Bruce 1986) were not substantiated. The prior name tymbonomus S. Müller, 1843 (type locality: Timor) has usually been presumed to supersede dumetorum Gould, 1845 for northern Australian populations (Junge 1937; Peters 1940; Condon 1975); and it later misled White & Bruce (1986) to claim that Australian populations reach Timor on migration. The type of tymbonomus (examined!) nevertheless belongs to the endemic long-tailed, rufous-marked form recorded from Timor by Mayr (1944b); accordingly, Cuculus variolosus whitei Bruce, 1986 is its synonym, and dumetorum Gould, 1845 the valid senior name for the northern Australian subspecies.
Chalcites lucidus (Gmelin, 1788)—Gill's (1983) finding of no consistent differentiation between breeding Australian and New Zealand populations is not confirmed by material in Australian museums. The Australian form differs in its combination of variable purplish wash over the crown and mantle, reduced white flecking on frons, and proportionally narrower bill. Accordingly, Mayr's (1932) revision is followed, as is conventional.
Chalcites minutillus (Gould, 1859)—The recent revisions of the C. minutillus (Gould, 1859)-C. russatus (Gould, 1868) complex by Parker (1981) and Ford (1981) leave the systematics of this difficult group in conflict. Parker separated minutillus and russatus specifically, based on evidence of sympatry in Borneo and the uncertainty that the forms there are respectively conspecific with these taxa in Australia. Ford, however, combined them in one species, based on complete introgressive intergradation between minutillus and russatus in eastern Queensland (also Ford 1987). His in-depth statistical analysis, nevertheless, did not discriminate southern migratory (barnardi Mathews, 1912) from northern sedentary material in north-eastern Queensland, the centre of presumed genetic mixing. Accordingly, cues for species limits have here been drawn instead from the earlier overview of Hartert & Stresemann (1925) which, corroborated by Peters (1940), Friedmann (1968) and Wolters (1975–1982), treats minutillus and russatus as conspecific, pace Condon (1975), White & Bruce (1986) and Sibley & Monroe (1990). One point of agreement in Parker's (loc. cit.) and Ford's (loc. cit.) reviews is their separation of three infra-specific forms in Australia; these are accepted here. The name malayanus Raffles, 1822, used as the senior synonym for this complex in earlier literature, has been found by Parker (loc. cit.) to apply instead to South-East Asian Chalcites xanthorhynchus xanthorhynchus (Horsfield, 1821).
Eudynamys orientalis (Linnaeus, 1766)—Since Hartert (1903) combined all forms of the Eudynamys scolopacea (Linnaeus, 1758) complex in one species, the status of component members have been in dispute, see Peters (1940), Stresemann (1940–1941), Rand (1941), Mayr (1944b), Wolters (1975–1982), White & Bruce (1986) and Sibley & Monroe (1990). Here the three species arrangement of the last two revisers is followed, except for including the sedentary forms of the Moluccas and eastern Lesser Sundas in the Australo-Papuan allospecies; its senior name is therefore orientalis Linnaeus, 1766 (type locality: Ambon, Moluccas). Australo-Papuasian forms are distinguished from other members of the scolopacea group in South-East Asia and the Greater Sundas by their cinnamon-buff-skinned nestlings, females with black crowns and coarser white dorsal spotting, salmon-pink-pigmented eggs, and hosts which are primarily large honeyeaters and oriolids instead of corvids. That the group extends to the Moluccas and eastern Lesser Sundas is also zoogeographically logical. Australo-Papuan populations are not split into further species here, cf. Rand (1941) and White & Bruce (1986), because of the likelihood that the sympatry recorded among their forms in southern New Guinea, Moluccas and Lesser Sundas is effected by non-breeding migrants from Australia.
Excluded Taxa
- Vagrant Species
CUCULIDAE: Centropus (Polophilus) bengalensis sarasinorum Stresemann, 1912 [Lesser Coucal]
CUCULIDAE: Cuculus saturatus Blythe, 1843
CAVS:9937
CUCULIDAE: Eudynamys scolopaceus (Linnaeus, 1758) [Asian Koel; vagrant on Christmas Island] — Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp. [163]CAVS:9949
CUCULIDAE: Hierococcyx sparverioides (Vigors, 1832) [Large Hawk-Cuckoo; vagrant on Christmas Island] — Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp. [163]
Diagnosis
Small to large, slender birds with barred tails and small to stout, decurved bills with swollen rounded nostrils and vestigial rictal bristles; body feathering soft, sleek and sometimes glossed brilliantly green or blue-black, close in varied tracts but without apterium between dorsal-cervical and interscapular tracts; no downs; aftershafts absent (vestigial?); uropygial gland well developed, naked. Feet relatively weak, adapted for arboreal perching, zygodactylous with outer toe permanently reversed; tarsi rather coarsely scutellate, toes with short claws; hypotarsus with two closed canals. Sexes dimorphic or similar; males usually larger than females. Wings rounded to commonly pointed: 10 primaries plus remicle and 9 to 12 eutaxic secondaries moulting erratically in 'transilient' mode, alula 4-feathered; tail short to long and graduated: 10 rectrices. Nares holorhinal and impervious, nasal septum imperforate; desmognathous palate with vestigial, incomplete vomer, palatines with narrowly rounded shelf, maxillary processes swollen and straight, lachrymals moderately enlarged to vestigial, sometimes almost reaching jugal bar, free from thin, variably enlarged ectethmoids, uncinate bone consistently(?) present; basipterygoid processes vestigial; interorbital septum with two or three principal perforations; atlas notched or perforated; cervical vertebrae 14 (13 in Clamator); sternum double-notched to almost entire on either side, both spina interna and externa present (fused in Clamator) or only spina externa (Eudynamys, Scythrops), furcula with or without prolonged hypocleideum; fossae flanking preacetabular processes of pelvic girdle narrow to wide, shelf of postacetabular processes square to rounded, not overlapping ischium dorsally, lateral perforations along either side of iliac crest obsolete to pronounced. Musculus expansor secundariorum 'cuculine'; pelvic muscle formula ABXY to AXY (ABEXY in Clamator), M. ambiens present, deep plantar tendons Type 1. Carotid arteries paired. Syrinx tracheo-bronchial. Tongue small, cartilaginous; no crop; caeca present, rather long. Diploid karyotype of 78–80 chromosomes, with 6 pairs of macrochromosomes.
General References
Amadon, D. 1942. Birds collected during the Whitney South Sea Expedition. L Notes on some non-passerine genera, 2. American Museum Novitates 1176: 1-21
Baker, E.C.S. 1942. Cuckoo Problems. London : H.F. & G. Witherby 207 pp.
Becking, J.H. 1981. Notes on the breeding of Indian cuckoos. Journal of the Bombay Natural History Society 78: 201-231
Beddard, F.E. 1898. The Structure and Classification of Birds. London : Longmans, Green xx 548 pp.
Berger, A.J. 1955. On the anatomy and relationships of Glossy Cuckoos of the genera Chrysococcyx, Lampromorpha, and Chalcites. Proceedings of the United States National Museum 103: 585-597
Berger, A.J. 1960. Some anatomical characters of the Cuculidae and the Musophagidae. Wilson Bulletin 72: 60-104
Brooker, M.G. & Brooker, L.C. 1989. Cuckoo hosts in Australia. Australian Zoological Reviews 2: 1-67
Brush, A.H. & Witt, H.H. 1983. Intraordinal relationships of the Pelecaniformes and Cuculiformes: electrophoresis of feather keratins. Ibis 125: 181-199
Devillers, P. 1976–1980. Project de Nomenclature française des Oiseaux du Monde. Le Gerfaut 66: 153-168, 391-421; 67: 171-200, 337-365, 469-489; 68: 129-136, 233-240, 703-720; 70: 121-146
Devillers, P. 1977. Projet de Nomenclature française des Oiseaux du Monde 5. Trogonides aux Picides. Le Gerfaut 67: 469-489
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Ford, J. 1987. Hybrid zones in Australian birds. The Emu 87: 158-178
Friedmann, H. 1968. The evolutionary history of the avian genus Chrysococcyx. Memoirs of the Queensland Museum 265: 1-137
Gill, B.J. 1983. Morphology and migration of Chrysococcyx lucidus, an Australasian cuckoo. New Zealand Journal of Zoology 10: 371-382
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Hartert, E. 1903. On the birds of the Key and South-East Islands, and of Ceram-laut. Novitates Zoologicae 10: 232-254
Hartert, E. 1925. Review of the genus Cacomantis Müll. Novitates Zoologicae 32: 164-174
Hartert, E. & Stresemann, E. 1925. Ueber die indoaustralischen Glanzkuckucke (Chalcites). Novitates Zoologicae 32: 158-163
Junge, G.C.A. 1937. The birds of south New Guinea Part 1 Non Passeres. Nova Guinea ns 1: 125-188
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Mason, I.J. 1982. The identity of certain early Australian types referred to the Cuculidae. Bulletin of the British Ornithologists' Club 102: 99-106
Mathews, G.M. 1918. The Birds of Australia. London : Witherby & Co. Vol. 7 pts i-iv xii pp. 1-384 pls 325-362. [Date published 4 Mar. 1918: volume publication dated as 1918–1919; Publication of Pt 1 (4 Mar. 1918), Pt 2 (15 May 1918), Pt 3 (26 Aug.), Pt 4 (19 Dec 1918), Pt 5 (12 June 1919)]
Mathews, G.M. 1919. The Birds of Australia. London : Witherby & Co. Vol. 7 pt 5 pp. 385-499 + xii pls 363-370 Appendixes A & B. [Date published 12 June 1919: publication dated as from preface, 12 June 1919 given in Appendix B]
Mayr, E. 1932. Birds collected during the Whitney South Sea Expedition. XIX Notes on the Bronze Cuckoo Chalcites lucidus and its subspecies. American Museum Novitates 520: 1-9
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Mayr, E. 1944. The birds of Timor and Sumba. Bulletin of the American Museum of Natural History 83: 123-194
Mayr, E. 1964. On the taxonomy of Cuculus pallidus (Latham). The Emu 64: 41
Mayr, E. & Amadon, D. 1951. A classification of recent birds. American Museum Novitates 1496: 1-42
Parker, S.A. 1981. Prolegomenon to further studies in the Chrysococcyx "malayanus" group (Aves, Cuculidae). Zoologische Verhandelingen (Leiden) 187: 1-56
Rand, A.L. 1941. Results of the Archbold Expeditions. No. 32 New and interesting birds from New Guinea. American Museum Novitates 1102: 1-15
Sibley, C.G., Ahlquist, J.E. & Monroe, B.L., Jr 1988. A classification of living birds of the world based on DNA-DNA hybridization studies. Auk 105: 409-423
Stresemann, E. 1940–1941. Die Vögel von Celebes. Teil III. Systematik und Biologie. Journal of Ornithology 89: 1-102
Stresemann, V. & Stresemann, E. 1961. Die handschwingen-mauser der Kuckucke (Cuculidae). Journal of Ornithology 102: 317-353
Wetmore, A. 1960. A classification for the birds of the world. Smithsonian Miscellaneous Collections 139(11): 1-37