Family ALCEDINIDAE

Todiramphus (Todiramphus) sanctus

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Family ALCEDINIDAE

Compiler and date details

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

Introduction

Alcedinidae (kingfishers) comprise about 90–95 species in 12–27 genera; ten species in five genera occur in Australia and its territories. The Australian fossil record is of unpublished material from the Miocene (W.E. Boles, pers. comm.) and Pleistocene-Holocene material from SE Queensland, eastern South Australia, and southern Victoria; it includes all Australian-occurring genera except Syma Lesson, 1827 and Tanysiptera Vigors, 1827. Kingfishers are cosmopolitan and widespread in warmer parts of the world, but are most diverse in Australo-Papuasia where some 31 species in eight or nine genera are recognised.

Kingfishers are perching birds that are commonly solitary and feed mainly on arthropods, fishes, amphibians and reptiles captured in plunging sallies from a vantage perch on to ground or into water. Nests are unlined burrows or holes excavated in ground or trees; eggs are glossy, plain-white, spherical; and young are altricial, nidicolous, and becoming spiny due to the retention of horny sheaths on their developing feathers; nidificational duties are shared by both sexes.

Family-group Systematics

There are three principal groups of kingfishers usually combined in one family, Alcedinidae (Sharpe 1892; Beddard 1898; Peters 1945; Mayr & Amadon 1951; Wetmore 1960; Wolters 1975–1982; Fry 1980a, 1980b; Burton 1984). Within Alcedinidae, these groups are treated by convention as subfamilies: Halcyoninae Vigors, 1825 (often as Daceloninae Bonaparte, 1837) (wood kingfishers), Alcedininae Rafinesque, 1815 (river kingfishers), and Cerylinae Reichenbach, 1851 (belted kingfishers), see Miller (1912), Peters (loc. cit.), Stresemann & Stresemann (1961), Fry (loc. cit.), Cramp (1985), cf. Maurer & Raikow (1981).

Sibley et al. (1988), Sibley & Ahlquist (1990) and Sibley & Monroe (1990) elevated the three groups to familial status based on their genomic distance according to DNA-DNA hybridization. Miller (loc. cit.), Maurer & Raikow (loc. cit.) and Christidis (1990) also recorded consistent structural differences between them in remigial sequence and moult, appendicular myology and karyotype respectively. Even so, the first two revisers still kept the three groups as subfamilies within Alcedinidae; and the significance of karyotypic differences between halcyonine and alcedinine kingfishers remains unclear, particularly because the condition in ceryline kingfishers is not established. Moreover, the three groups are monophyletic with respect to nearest relatives among coraciiform birds (Sibley et al. loc. cit.; Sibley & Ahlquist loc. cit.). Thus familial groupings in kingfishers, as in the owls, hang in the balance. So, pending further evidence, convention is followed here in treating the three groups of kingfishers as subfamilies, just as it is in separating strigid and tytonid owls into two families, cf. Christidis & Boles (1994).

Two of the subfamilies occur in Australia and its territories: Halcyoninae with four genera and Alcedininae with one, Alcedo Linnaeus, 1758. Australian halcyonine kingfishers capture prey mainly on the ground, are medium to large in size and have graduated tails that are longer than their thick bills; their wings are diastatasic or eutaxic and their primaries moult by simple descending mode beginning with the first (innermost). Australian alcedinine kingfishers capture prey mainly in water, are small to tiny in size, and have square-cut stumpy tails much shorter than their narrowly compressed bills; their wings are strictly eutaxic and their primaries moult by descending mode coincidentally from two foci, at the first and seventh. For differences in musculature and karyotype, see family diagnosis.

Alcyone bella Diggles, 1878 (= Ceyx lepidus Temminck, 1836) is excluded from the Australian fauna, see Mathews (1915).

The genera are arranged here in their subfamilies, commencing with Halcyoninae, after Fry (1980a, 1980b).

Genus-group Systematics

Dacelo Leach, 1815—A broad definition of Dacelo Leach, 1815 is followed, after Peters (1945) and Fry (1980a). It includes the two small, round-winged species of New Guinean kookaburras with only 14 cervical vertebrae, a group nevertheless accepted subgenerically here under Sauromarptis Cabanis and Heine, 1860 (cf. Mayr 1941a; Wolters 1975–1982). Generic limits in relation to Sauromarptis and the shovel-billed New Guinean Clytoceyx Sharpe, 1880, with its kookaburra-like form and sexual dimorphism, need evaluation.

Syma Lesson, 1827—Although often included in Halcyon Swainson, 1821, Syma Lesson, 1827 is separated here, following Mayr (1941a), Keast (1957), Condon (1975) and Wolters (1975–1982). Judged by their colour patterns, occipital sexual dimorphism, 'toothed' bills and aspects of osteology and myology (W.E. Boles, pers. comm.), its two species are paraphyletic with Halcyon and its other generic segregates (see below), pace Peters (1945) and Fry (1980a).

Tanysiptera Vigors, 1825—This well-defined genus comprises three groups of species (Fry 1980a) which are treated here as subgenera: white-breasted, blue-backed and spatulate-tailed Tanysiptera Vigors, 1825, the red-breasted and -rumped and spatulate-tailed T. nympha G.R. Gray, 1841 group, and buff-breasted, black-and-white backed and sagittate-tailed Uralcyon Heine, 1859. Only the last occurs in Australia.

Todiramphus Lesson, 1827—The remaining species of Australian halcyonine kingfishers have hitherto been placed by convention in a broadly circumscribed Afro-Australasian genus, Halcyon Swainson, 1821 (Peters 1945; Keast 1957; Condon 1975; Fry 1980a). Nevertheless, patterns of wing, eutaxy, territorial display, feather proteins and DNAxDNA hybridization indicate that Halcyon in its broad sense is polyphyletic and comprises at least two groups of species: Afro-Asian Halcyon and Australasian-southwest Pacific Todiramphus Lesson, 1827 (Mitchell 1901; Burton 1978; Fry 1980a; Knox 1980; Maurer & Raikow 1981; Sibley & Ahlquist 1990). The finer generic divisions circumscribed among Australo-Papuasian species of Todiramphus by Mathews (1918–1919) and implied by Mukherjee & Dasgupta (1973) are adopted here only at subgeneric level; Mayr (1941b) and Fry (1980a) have stressed links between these species and the midget, flat-billed Polynesian kingfishers (nominotypical Todiramphus) through the Sacred Kingfisher group (Sauropatis Cabanis & Heine, 1860).

Alcedo Linnaeus, 1758—Historically, Australian species have been referred to Alcyone Swainson, 1837, distinguished from Alcedo Linnaeus, 1758 by their two, not three forward toes and concolorous dark blue dorsum (Sharpe 1892; Keast 1957; Wolters 1975–1982; Devillers 1976–1980), cf. Peters (1945) and Condon (1975) who referred these species to red-billed, brown-plumaged Ceyx Lacépède, 1799. There is, however, no pattern of vicariance to suggest that the Australasian Alcyone kingfishers are paraphyletic with Alcedo. Rather it seems likely that the azurea Latham, 1802 superspecies and pusilla Temminck, 1836 are the respective vicariants of South-East Asian Alcedo meninting Horsfield, 1821 and Indonesian Alcedo coerulescens Vieillot, 1818, as pointed out by Fry (1980a). Such relationships imply that toe number and back colour have been acquired independently in the two Australian species-groups from different Alcedo-like ancestors (cf. also Delacour 1951). Accordingly, I follow Delacour (loc. cit.) and Fry (loc. cit.) in referring them to Alcedo and in separating the two species groups subgenerically.

Species-group Systematics

Dacelo leachii Vigors & Horsfield, 1827—Geographic variation appears to be clinal through the continuously distributed populations between central east Queensland and the Kimberley Division, Western Australia across the nexus at the head of the Gulf of Carpentaria (cf. Keast 1957; Mees 1961). These populations become progressively darker and smaller northwards to become convergently similar in northern Arnhem Land and northern Cape York Peninsula. The isolate in the Pilbara, Western Australia, is distinctively large, pale-headed and cinnamon-buff-breasted; its abrupt relictual intergradation with nominotypical leachii in Dampier Land and along the lower Fitzroy River, Western Australia, indicates past connections around the Eighty-mile Beach, Western Australia (Keast 1957).

Syma torotoro Lesson, 1827—The Australian isolate on Cape York Peninsula differs from all New Guinean populations in the duller, more drab brown cast to its dorsum and duskier toning of its bill in adulthood, rather than in size and ventral tone (cf. Keast 1957; Mees 1982).

Todiramphus chloris (Boddaert, 1783)—Apart from Johnstone's (1983) discovery of a small, pallid population in the Pilbara, Western Australia, it has been customary to recognise only one other subspecies in Australia: T. c. sordidus (Gould, 1842), see Keast (1957), Condon (1975), and Ford (1983). Even so, both Keast (loc. cit.) and Ford (loc. cit.) stressed the distinctiveness of a further large, bluer-toned form in central and south-eastern Queensland, a form which Mathews (1918–1919, 1946) had singled out and treated as a separate species; it is recognised subspecifically here. Populations on Cape York Peninsula and in the Kimberley and Arnhem Land, which appear to vary in dorsal tone and proportions as well, also need further evaluation.

Todiramphus sanctus (Vigors & Horsfield, 1827)—Mainland Australian populations are evidently monotypic (Keast 1957), their ventral toning varying largely according to wear. The status of resident forms on Norfolk and Lord Howe islands, however, needs resolution. Here the population on Lord Howe Island is treated as consubspecific with T. s. vagans (Lesson, 1828) of New Zealand (also Wolters 1975–1982), from which it is distinguished with difficulty as it may be self-introduced from New Zealand; but the bluer-backed, finer-billed form on Norfolk Island is recognised as distinct, following correction in Schodde et al. (1983). Relationships with Lesser Sundan T. australasia (Vieillot, 1818) are also in need of clarification, see Mayr (1944), Fry (1980a) and White & Bruce (1986).

Todiramphus macleayii (Jardine & Selby, 1830)—Eastern Australian populations, which differ from nominotypical macleayii Jardine and Selby, 1830 of Arnhem Land in their turquoise back and smaller white remigial patch (Keast 1957), change in their breeding range from large in the south to small on Cape York Peninsula (cf. Mayr 1937). Whether there are any steps indicative of secondary intergradation is not known.

Alcedo azurea Latham, 1802—Although Ford (1987) disputed differentiation and intergradation between nominotypical azurea Latham, 1802 and A. a. ruficollaris (Bankier, 1841) along the eastern foot of Cape York Peninsula (Schodde & Mason 1976), no supporting data was provided. Accordingly, Schodde & Mason's arrangement is kept, pending more detailed analysis of populations from north-eastern Queensland with additional material.

Alcedo pusilla (Temminck, 1836)—While the different forms in Arnhem Land-Gulf of Carpentaria and coastal eastern Queensland south of the Endeavour River are evidently monomorphic (Mayr 1941a; Keast 1957; Ford 1983), populations on northern and eastern Cape York Peninsula are variable. They change from smaller and more royal blue on the north-west coast to larger and darker around to the east and south-east. Whether this is ecophenotypic or due to gene flow from A. p. ramsayi (North, 1912) has yet to be clarified. So, too, does Mayr's (loc. cit) inference that nominotypical pusilla of southern New Guinea merges with Australian populations across Torres Strait; there is no material in AMNH to confirm it.

Excluded Taxa

Vagrant Species

CAVS:0753
ALCEDINIDAE: Alcedo (Alcedo) atthis Linnaeus, 1758 [Common Kingfisher; vagrant to Christmas Island] — Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp. [169]

CAVS:9934
ALCEDINIDAE: Halcyon pileata (Boddaert, 1783) [Black-capped Kingfisher] — Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp. [170]

CAVS:8081
ALCEDINIDAE: Tanysiptera hydrocharis Gray, 1858 [Little Paradise Kingfisher]

Diagnosis

Stout dagger-billed birds with broad heads and thick-set bodies with much blue in plumage; body feathering rather coarse and often glossed in defined tracts; aftershafts vestigial or absent; uropygial gland large, tufted (naked in Tanysiptera). Feet weak and anisodactylous; short tarsi finely reticulately scaled to scaleless, and flat-soled toes extensively syndactylous towards terminal joints. Sexes alike or dimorphic. Wings rounded: 10 primaries plus remicle without covert, moulting in descending sequence from different foci, and 11–14 eutaxic or diastataxic secondaries; tail short and square: 12 rectrices (10 in Tanysiptera) moulting centrifugally. Nares holorhinal and impervious, nasal septum impervious; desmognathous palate, bifid vomer fused with and beneath palatine shelf, palatines notched with attenuate caudolateral processes, lachrymals much expanded out from frontals and in to enclose reduced ectethmoids; basipterygoid processes absent; cervical vertebrae 14–15; sternum moderately two-notched on each side, only long single spina externa present (truncated in Ceryle), furcula without hypocleideum. Musculus expansor secundariorum present (Halcyoninae) or apparently absent (Alcedininae), biceps slip absent; pelvic muscle formula AX, no M. ambiens; deep plantar tendons Type V, but characteristically variable and fusing above or below slip to hallux. Carotid arteries paired. Syrinx tracheo-bronchial. Tongue a small rounded flap; caeca rudimentary to absent. Diploid karyotype of 76+ chromosomes, with one pair of macrochromosomes (Halcyoninae) or of 120+ chromosomes without obvious macrochromosomes (Alcedininae).

General References

Beddard, F.E. 1896. Contributions to the anatomy of picarian birds-Part III. On some points on the anatomy of the kingfishers. Proceedings of the Zoological Society of London 1896: 603-606

Beddard, F.E. 1898. The Structure and Classification of Birds. London : Longmans, Green xx 548 pp.

Burton, P.J.K. 1978. The basisphenoid notch of kingfishers. Bulletin of the British Ornithologists' Club 98: 68-74

Burton, P.J.K. 1984. Anatomy and evolution of the feeding apparatus in the avian orders Coraciiformes and Piciformes. Bulletin of the British Museum (Natural History) Zoology 47: 331-443

Christidis, L. 1990. Chordata 3B. Aves. Animal Cytogenetics 4. Berlin : Gebrüder Borntraeger 116 pp.

Christidis, L. & Boles, W.E. 1994. The Taxonomy and Species of Birds of Australia and its Territories. Monograph 2. Melbourne : Royal Australasian Ornithologists Union iv 112 pp.

Condon, H.T. 1975. Checklist of the Birds of Australia. Part 1 Non-Passerines. Melbourne : Royal Australasian Ornithologists Union xx 311 pp.

Cramp, S. (ed.) 1985. Handbook of the Birds of Europe, the Middle East and North Africa. The Birds of the Western Palaearctic. Oxford : Oxford University Press Vol. 4 960 pp. 98 pls.

Delacour, J. 1951. The significance of the number of toes in some woodpeckers and kingfishers. Auk 68: 49-51

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

Ford, J. 1983. Taxonomic notes on some mangrove-inhabiting birds in Australia. Records of the Western Australian Museum 10: 381-451

Ford, J. 1987. Hybrid zones in Australian birds. The Emu 87: 158-178

Fry, C.H. 1980. The evolutionary biology of kingfishers (Alcedinidae). Living Bird 18: 113-160

Fry, C.H. 1980. The origin of Afrotropical kingfishers. Ibis 122: 57-72

Johnstone, R.E. 1983. A review of the Mangrove Kingfisher, Halcyon chloris (Boddaert) in Australia, with a description of a new subspecies from Western Australia. Records of the Western Australian Museum 11: 25-31

Keast, A. 1957. Variation in the Australian kingfishers (Aves: Alcedinidae). Records of the Australian Museum 24: 61-72

Knox, A.G. 1980. Feather protein evidence for the relationships of African Halcyon species. Ibis 122: 72-74

Mathews, G.M. 1915. Diggles' Ornithology of Australia and other works. Austral Avian Records 2: 137-153

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]

Mathews, G.M. 1946. A Working List of Australian Birds including the Australian Quadrant and New Zealand. Sydney : G.M. Mathews 184 pp.

Maurer, D.R. & Raikow, R.J. 1981. Appendicular myology, phylogeny, and classification of the avian order Coraciiformes (including Trogoniformes). Annals of the Carnegie Museum 50: 417-434

Mayr, E. 1937. Birds collected during the Whitney South Sea Expedition. XXXV Notes on New Guinea Birds. II. American Museum Novitates 939: 1-14

Mayr, E. 1941. Birds collected during the Whitney South Sea Expedition. XLVII Notes on the genera Halcyon, Turdus and Eurostopodus. American Museum Novitates 1152: 1-7

Mayr, E. 1941. List of New Guinea Birds. A Systematic and Faunal List of the Birds of New Guinea and Adjacent Islands. New York : American Museum of Natural History xi 260 pp.

Mayr, E. 1944. Timor and the colonization of Australia by birds. The Emu 44: 113-130

Mayr, E. & Amadon, D. 1951. A classification of recent birds. American Museum Novitates 1496: 1-42

Mees, G.F. 1961. An annotated catalogue of a collection of bird-skins from West Pilbara, Western Australia. Journal of the Royal Society of Western Australia 44: 97-143

Mees, G.F. 1982. Birds from the lowlands of southern New Guinea (Merauke and Koembe). Zoologische Verhandelingen (Leiden) 191: 1-188 4 pls

Miller, W. de W. 1912. A revision of the classification of the kingfishers. Bulletin of the American Museum of Natural History 31: 239-311

Mitchell, P.C. 1901. On the anatomy of the kingfishers, with special reference to the conditions in the wing known as eutaxy and diastataxy. Ibis 8 1: 97-123