Family APODIDAE

Apus (Apus) pacificus

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Family APODIDAE

Compiler and date details

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

Introduction

Apodidae (swifts) comprise about 75–99 species in 14–20 genera; five species in four genera are accepted here as occurring regularly in Australia and its territories. The Australian fossil record is represented in mid Tertiary deposits in northwest Queensland and in the Holocene-Pleistocene of southeast Victoria and Kangaroo Island, including extant species. Swifts are cosmopolitan and widespread in warmer parts of the world, with centres of diversity in South America, Africa, South-East Asia and adjacent archipelagos.

Swifts are aerial, often colonial insectivores that feed on the wing at speed, for which their long wings and stream-lined bodies are adapted, allowing many species (e.g. of Apus) to rest or sleep on the wing. Nests are not built or are of hanging half-cups often bound with saliva and situated in crevices or holes in caves, cliffs, trees or even buildings; eggs are ellipsoid and dull plain-white; young are altricial, nidicolous, and long in fledging; nidificational duties are shared by both sexes.

Family-group Systematics

There are two or three subfamilies; representatives of only one—cosmopolitan Apodinae Olphe-Galliard, 1887—are present in Australia (after Brooke 1970). Apodinae are distinguished by minimal sexual dimorphism, eutaxic secondaries, single left carotid artery, well-developed trans-palatine processes, complex Musculus splenius capitis, cementing saliva in nests and usually naked nestlings. Three tribes are represented among Australian apodinine swifts: Collocaliini Bonaparte, 1853 (swiftlets), Chaeturini Bonaparte, 1857 (needle-tailed swifts), and Apodini Olphe-Galliard, 1887 (swifts of the genus Apus Scopoli, 1777 and allies, with pamprodactyl feet in which all four toes are directed forwards)—see Brooke (loc. cit.).

The genera are arranged in their tribes, commencing with Collocaliini, in the sequence according to Brooke (1970).

Genus-group Systematics

Aerodramus Oberholser, 1906—Until Brooke (1972) and Medway & Pye (1977) redefined the species groups of Oberholser (1906), Stresemann (1925, 1931) and Brooke (1970) generically, swiftlets of the tribe Collocaliini were combined by convention in a single genus Collocalia G.R. Gray, 1840. Brooke (loc. cit.) and Medway & Pye (loc. cit.) recognized three genera:

(1) Collocalia for the four glossy, white-bellied, non-echo-locating species;

(2) giant monospecific Hydrochous Brooke, 1970 which is dull-plumaged but does not echo-locate; and

(3) Aerodromus Oberholser, 1906 for the plain dull grey, echo-locating species, a number of which build nests prevailingly of solidified saliva. Aerodramus thus comprises 13–16 species (27 according to Sibley & Monroe 1990) centred in the Indonesian archipelagos and New Guinea (Medway 1966); it may be diphyletic within itself and comprise several groups of species.

This definition of Aerodramus is followed here, first because it has been explained comprehensively, secondly because of its biological basis, and thirdly because those recent revisers rejecting it (Salomonsen 1983; Beehler & Finch 1985; Sibley & Monroe 1990) provide no rigorously argued alternative. Even so, Medway & Pye's (1966: 235–236) pro–Aerodramus argument is somewhat confused, implying on one side that the Aerodramus group is paraphyletic because of nest structure and on the other that it is monophyletic judged by capacity to echo-locate. Given the consistent, shared traits of dull, grey-bellied plumage and echo-location in the group, and the adaptability of nest structure, the monophyly of Aerodramus, sensu lato, seems favoured over paraphyly on available information; but the issue remains open and needs investigation, cf. Christidis & Boles (1994).

Hirundapus Hodgson, 1837—Since Collins & Brooke (1976), the separation of Hirundapus Hodgson, 1837 from similarly needle-tailed Chaetura Stephens, 1826 has become widely accepted. Hirundapus differs by its great size, long-pointed and broad palatine processes, white flank stripe, and nest site in tree-hollows. Three of its four species occur exclusively between India, South-East Asia, the Philippines and Sulawesi; the fourth is more widespread, reaching Australia on migration.

Apus Scopoli, 1777—Opinion is divided as to whether Tachymarptis Roberts, 1922, comprising two Afro-Asian species, should be recognised generically (Brooke 1972; Wolters 1975–1982; Sibley & Monroe 1990) or combined as a subgenus within Apus Scopoli, 1777 (Brooke 1970). The latter view is followed here with reservation, as both cases have been argued well by Brooke (loc. cit.); on present evidence, the division between Tachymarptis and Apus appears minor in comparison with limits between other genera in Apodini. Although Brooke's (1972) argument implies that Apus in its customary sense is polyphyletic with respect to Cypsiurus Lesson, 1843, more information is needed. Whatever the outcome, the generic and specific nomenclature of the one species reaching Australia is not affected. Subgenus Apus, as accepted here, comprises about 17 species and is centred in Africa with species breeding eastwards through the Eurasian tropics and temperate regions to the Greater Sundas.

Species-group Systematics

Collocalia esculenta (Linnaeus, 1758)—The analyses of Somadikarta (1982, 1986) and Salomonsen (1983), integrated in Sibley & Monroe (1990), show that Collocalia esculenta (Linnaeus, 1758) in its earlier broad sense comprises at least three species: west Malesian C. linchi Horsfield & Moore, 1854, Philippines pallid-rumped C. marginata Salvadori, 1882 and pan Malesian and Papuasian C. esculenta. From the first two, C. esculenta itself is distinguished by a geographically varying combination of bluish rather than greenish glossed plumage, dark rump, white spotting on the inner rectrices in eastern forms, and a feather tuft on the hind toe in western forms (Somadikarta 1986). Among the 20–22 subspecies of C. esculenta (after Salomonsen 1983), C. e. natalis Lister, 1889 of Christmas Is. is in turn distinguished by dull green-grey dorsum, white-flecked rump, ill-defined elongate white spots in the rectrices, and naked hind toe; its position with respect to linchi is thus uncertain (Somadikarta 1986). It may prove specifically distinct; here, however, it is kept as a form of esculenta following Stresemann (1940–1941). Richly blue metallic New Guinean C. e. nitens Ogilvie-Grant, 1914, which has been recorded several times from north-eastern Queensland, is treated in the Appendices as a rare, erratic vagrant.

Aerodramus spodiopygius (Peale, 1848)—Mayr (1937), Medway (1966) and Salomonsen (1983) treated the resident pale-rumped Australian swiftlets as allopatric members of south-west Pacific A. spodiopygius (Peale, 1848), whereas Condon (1975), Bruce in White & Bruce (1986), and Sibley & Monroe (1990) separated them specifically. Only Bruce (loc. cit.) provided reasons; but his adoption of Salomonsen's (loc. cit.) comment that Australian terraereginae Ramsay, 1875 and other principal allotaxa could form a superspecies took little heed of similarities in dorsal pattern and nest structure recorded between terraereginae and spodiopygius by Medway (loc. cit.) and Salomonsen (loc. cit.). Accordingly, Australian populations are kept as subspecies of A. spodiopygius. Storr (1984) and Ingram (1991) treated the two endemic Australian forms as separate species, but their roosting and breeding ranges appear not to overlap, being separated on eastern and western watersheds of the Great Dividing Range. Specific relationships in the complex are still not resolved satisfactorily; in earlier Australian literature, Mascarene A. francicus (Gmelin, 1789) was implicated.

Aerodromus vanikorensis (Quoy & Gaimard, 1830)—Notwithstanding its few records (Mathews 1917; Bravery 1971; Blakers et al. 1984), this species is included here as a member of the Australian avifauna because of the likelihood that it is a much more regular visitor from New Guinea than its records suggest. Unlike Collocalia esculenta (Linnaeus, 1758), it is difficult to identify from Aerodromus spodiopygius (Peale, 1848) in the field, and it forages more widely at higher levels above forest and so is more prone to dispersal, particularly during cyclonic weather. A. vanikorensis (Quoy and Gaimard, 1830) is a plain-plumaged, medium-sized swiftlet differing from other similar species in the region by its dark rump and feathered tarsus. Comprising about 12 subspecies, it occurs at low altitudes on islands in the south-west Pacific from Sulawesi to all Papuasia, Santa Cruz and New Caledonia (Salomonsen 1983). In earlier Australian literature it has been confused with south-east Asian A. fuciphagus (Thunberg, 1812). Note, as of December 2015, AFD is treating this species as a vagrant.

Hirundapus caudacutus (Latham, 1802)—This is the most northerly breeding and largest of the four species of Hirundapus and is otherwise distinguished by its clearly defined white throat and more finely spined tail. Extensive loral mark and paler, less contrasting dorsal pattern indicate that the form reaching Australia on migration is nominotypical caudacutus which breeds in north-east Asia.

Apus pacificus (Latham, 1802)—Among Apus Scopoli, 1777, this species is distinguished by its combination of a white rump and deeply forked tail with crescentic white edging to its contour plumage. There are four subspecies (Deignan 1956; Vaurie 1959), their breeding ranges replacing one another north to south from central and north-eastern to south-eastern Asia; the more northerly breeding subspecies winter successively further south than the southerly ones. Only nominotypical pacificus, distinguished by its broad white rump band, brownish dusky body, deeply furcate tail and large size, reaches Australia and its territories, as implicit in Mees (1973).

Excluded Taxa

Vagrant Species

CAVS:0881
APODIDAE: Aerodramus vanikorensis (Quoy & Gaimard, 1830) [Uniform Swiftlet; vagrant to Qld and Torres Straight Islands. Also found in Indonesia — Irian Jaya, Sulawesi (Celebes); New Caledonia; Papua New Guinea — Bismarck Archipelago; Vanuatu; Philippines] — Mathews, G.M. 1917. Some new Australian birds. The Emu 16: 180-185; Salomonsen, F. 1983. Revision of the Melanesian swiftlets (Apodes, Aves) and their conspecific forms in the Indo-Australian and Polynesian region. Noona Dan Papers no. 141. Biologiske Skrifter Kongelige Danske Videnskabernes Selskab 23(5): 1-112; Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp. [17, 80]

CAVS:8964
APODIDAE: Aerodramus vanikorensis yorki (Mathews, 1916) [New Guinean Uniform Swiftlet; vagrant to Qld and Torres Straight Islands; Melanesia to Micronesia; E Cape York Peninsula south to Atherton Tableland—also S and E New Guinea and adjacent islands (conventionally as A. v. granti (Mayr, 1937)] — Bravery, J.A. & Schodde, R. 1973. Sight-records of swiftlets. The Emu 73: 29-30 (occurrence)

CAVS:0828
APODIDAE: Apus nipalensis (Hodgson, 1937) [House Swift; Specimen record from Point Stuart, Arnhem Land in March 1979, see Robertson (1980), accepted by RAOU Records Appraisal Committee (1988: case no. 91, accepted later). Also sight record from Caboolture, QLD, in January 1994, accepted by RAOU Records Appraisal Committee (case no. 177). Other tentative sight records have not been investigated, see, for example, Robertson (loc. cit.). Robertson, followed by Christidis & Boles (1994), referred his specimen to the western Greater Sundan subspecies subfurcatus Blyth, 1849, but without specifying its critical character states. Much more likely on geographical grounds is the Javan form, furcatus Brooke (1971), see, for example, Brooke (1971). Here the group is separated specifically under nipalensis Hodgson (1836), following Brooke in Snow (1978: 287) and Sibley & Monroe (1990)] — Brooke, R.K. 1971. Geographical variation in the Little Swift Apus affinis (Aves: Apodidae). Durban Museum Novitates 9: 93-103; Snow, D.W. (ed.) 1978. An Atlas of Speciation in African Non-Passerine Birds. London : British Museum (Natural History) vii 390 pp.; Robertson, D.G. 1980. First record of the House Swift Apus affinis (Apodidae) in Australia. Australian Bird Watcher 8: 239-242; RAOU Records Appraisal Committee 1988. Second Report of the Records Appraisal Committee. The Emu 88: 54-57 (case no. 91, accepted later); Sibley, C.G. & Monroe, B.L., Jr 1990. Distribution and Taxonomy of Birds of the World. New Haven : Yale University Press xxiv 1111 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.; Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp. [80]

CAVS:8091
APODIDAE: Apus nipalensis subfurcatus (Blyth, 1849) [Indonesian House Swift] — Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp. [80]

APODIDAE: Collocalia esculenta nitens Ogilvie-Grant, 1914 [New Guinean Glossy Swiftlet; vagrant to NT and N QLD (5 records accepted by Birds Australia Rarities Committee)]

CAVS:see 0881
APODIDAE: Collocalia vanikorensis (Quoy & Gaimard, 1830)

CAVS:9941
APODIDAE: Mearnsia novaeguineae (Albertis & Salvadori, 1879) [Papuan Spine-tailed Swift; vagrant to Torres Strait Islands] — Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp. [81]

Other Excluded

CAVS:0828
APODIDAE: Apus (Colletoptera) affinis (Hodgson, 1937) [Following Christidis and Boles (2008: 80), nipalensis is assigned as a subspecies of A. affinis, but see alternative arrangement of Schodde and Mason (1997: 402)] — Schodde, R. & Mason, I.J. 1997. Aves (Columbidae to Coraciidae). In, Houston, W.W.K. & Wells, A. (eds). Zoological Catalogue of Australia. Melbourne : CSIRO Publishing, Australia Vol. 37.2 xiii 440 pp.; Christidis, L. & Boles, W.E. 2008. Systematics and Taxonomy of Australian Birds. Melbourne : CSIRO Publishing 288 pp.

Diagnosis

Slender, small, dull-plumaged birds with long-tapered, bow-shaped wings and very short and wide bristle-less bills; body feathering sleek in defined tracts; dense underdown; long aftershafts present; uropygial gland large, naked. Feet small, pamprodactylous or aniso-dactylous; very short tarsi often feathered, with or without hind-pointed hallux; hypotarsus with single deep furrow. Sexes alike. Wings much attenuated with prolonged manus and a very short, thick humerus: 10 long primaries, the outermost usually longest, moulting in descending sequence, and about 8–11 short eutaxic or diastataxic secondaries; short tail square to forked: 10 rectrices (Australian species) moulting centripetally. Nares holorhinal and impervious, nasal septum extensively perforate; aegithognathous palate, with vomer bifid at level of departure of palatine processes from maxillaries, premaxilla unossified, maxillary processes undeveloped, palatine shelf unexpanded with narrowed processes, lachrymals vestigial, and nasals and maxillaries fused in a bar anteriorly to inarticulate naso-frontal hinge; basipterygoid processes absent; cervical vertebrae 13–14; sternum entire, both spina externa and interna fused in short spina communis, furcula stout with well-developed hypocleideum. Musculus tensor patagium brevis strong and extensively fleshed, but M. expansor secundariorum, M. sternocoracoideus and biceps slip absent; pelvic muscle formula A, no M. ambiens; deep plantar tendons Type V, usually fully fused. Carotid arteries paired or only left present. Syrinx tracheo-bronchial, with single pair of extrinsic and intrinsic muscles. Tongue short, bifid at tip, often with well-developed salivary glands that enlarge and become active during breeding; no crop; no caeca. Diploid karyotype of 64–76 chromosomes, with five pairs of macrochromosomes.