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Australian terraereginae

Description: The only known to breed on mainland . The white to pale grey rump distinguishes this from vagrant Uniform and Glossy that occasionally reach mainland Australia. The wingspan being more than twice the length of the , and the practice of never folding the wings in against the body in flight, separates this bird from and martins, whose wingspan is less than twice the body length.. Swiftlets do not perch on top of objects other than their nests, but hang below or at the side of their roost. When roosting their wings project beyond the end of the tail but the tails of roosting swallows and martins, project well beyond their folded wings. In flight the swiftlets wings may be held at right angles to the body in which instance they appear quite broad, but when held back in the typical sickle-shape of swifts they are narrower and reduce drag. The tail displays a shallow fork when folded but can look square-ended when half open, or rounded when fully spread – particularly when manouevreing to catch prey or make sharp turns in .

Males and females are similar but juveniles have a fine pale edging to their primary feathers that distinguishes them from adults, either of which might be brooding an in a breeding colony.

Aerodramus t. terraereginae Aerodramus t. chillagoensis Mean ± SD (mm) Range (mm) N Mean ± SD (mm) Range (mm) N Wing:flat & straight 110.6 ± 3.2 102 - 117 75 107.0 ± 2.7 99 - 116 516 Outer Rectrix 51.0 ± 2.0 46 - 54 42 47.6 ± 2.1 41 - 56 119 Central Rectrix 41.9 ± 2.4 38 - 49 43 44.0 ± 5.3 40 - 53 6 Tail 52.7 ± 1.6 49 - 55 31 46.9 ± 2.0 43 - 51 13 Length 109.9 ± 4.8 101 - 118 29 103.3 ± 4.6 94 - 112 173 Wing Span 246.3 ± 8.8 232 - 260 15 249.9 ± 5.1 238 - 262 122 Tarsus 10.1 ± 0.8 8.2 – 12.1 45 9.6 ± 0.5 9 - 11 56 Mid-Toe 5.5 ± 0.6 4.2 – 7.1 43 5.5 ± 0.4 5 – 6.2 6 Claw 4.1 ± 0.3 3.5 – 5.2 43 4.1 ± 0.3 3.7 – 4.4 6 Culmen 7.4 ± 0.7 6.2 – 9.2 37 6.9 ± 0.5 6.4 – 7.5 6 Exposed Culmen 4.4 ± 0.5 2.4 – 5.3 37 4.3 ± 0.4 3.7 – 4.8 14 Weight (g) 10.9 ± 0.9 8.5 - 13 47 9.4 ± 0.7 7.7 – 12.1 598

The majority of these measurements were taken by M.K. Tarburton from live and all the skins in Australian museums. 49 A. t. chillagoensis measurements were provided by the Australian Bird and Banding scheme.

Distribution: A.t. terraereginae is usually seen within the cream-shaded area of the accompanying figure, while A.t. chillagoensis is moderately common in the light green area (Tarburton 2013).

Observations have also been made on these Continental Islands: Fitzroy (Warham 1962), Dunk (Banfield 1911), Timana (Thorpe), Bedarra (Richards), Goold (Campbell & Barnard 1917), & Hinchinbrook (Storr 1984a). On Magnetic Island they are occasionally observed during summer (Wieneke 1988).

The vagrants seen at Weipa and Cape York are from the New Atlas of Australian Birds (Barret et al 2003). Vagrants have also been recorded from near Brisbane and in northern N.S.W.

Habitat: Feeding habitat is air-space over rainforest, sclerophyll forest, savanna, & plantations. Breeding & sleeping habitat as under breeding behaviour.

Breeding Behaviour: Clutch: 1 invariably (Storr 1984) 2? (Mathews 1918). 20 nests with single egg (Banfield 1911).

The coastal A.t. terraereginae builds its nest in moderately dark situations to the underside of suspended granite boulders, in creek beds or above coastal shores, one to three metres from the ground. Sometimes it uses abandoned mine sites (Cassels 1961, Smyth et al 1980). Nests number between one and 600 in each colony (Griffin 1968, Smyth et al 1980). Nests may be built of moss (Griffin 1968), leaves, Kangaroo Grass, vines, feathers, egg shells, Eucalyptus leaves, vines, twigs, or mixtures of these components held together and to the rock face by the glue in the birds saliva (Smyth et al 1980, Crouther 1983).

The inland form A.t. chillagoensis builds its nest in totally dark situations well inside limestone caves, where nest numbers may also range from one to 600 in a colony. The nests are built of Kangraoo Grass, Black Spear Grass and saliva, and are attached to smooth overhanging surfaces from two to 30m above the floor. This location makes it impossible for Childrens Pythons and Brown Tree to reach most of the nests. I have watched both species fall from the rock in an effort to reach swiftlet nests.

This rock houses the Lower Dooloomai colony of Australian swiftlets. Photos: Copyright M. K. Tarburton.

Aerodramus t. chillagoensis builds its nests in genuine totally dark limestone caves not just overhangs. This colony is in Bottle Cave 2011, Photo Copyright M. K. Tarburton

Nesting in caves has its theoretical advantages for swiftlets just as nesting in tree hollows has for birds in general. These include lower predation rates (e.g. Tarburton 2009) and lower losses due to high winds. But do these advantages always convert into reality? It is possible that the clustered nature of their nesting activities may make at least some populations of swifts vulnerable to local weather events that could lead to local extinction. The , like all swiftlets studied to date, has a slow reproductive rate (e.g. clutch of two once a year compared to clutch of 4 twice a year for Welcome Swallows Hirundo neoxena), long incubation period (27 days compared to 16 for Welcome Swallows), and long fledgling period for its size (46-51 days compared to 19 days for Welcome Swallows) (Tarburton 1988b, 1993b). This means replacement of losses due to catastrophic events is slow compared with most . The two they lay each year will only be increased to three if one of the first two eggs or young nestlings is lost (Tarburton 1988b).

The advantage of slow development in the egg and nestling stage is that the young can successfully go without food for 2-3 days during a cyclone or some other disruption. This is well beyond the starvation endurance of birds of similar size. The disadvantage of slow development in the short breeding season presented by the Australian climate is that they cannot raise two broods in the one year. I first learnt of this problem after I had determined that their close relative, the White-rumped Swiftlet in Fiji laid two eggs, but only a single egg was ever found in Australian nests. This phenomenon is the opposite to what we would expect, because the harsher dry season of Australia should require the birds to lay more eggs not less than their island counterparts. Research was started in Queensland to determine if the published clutch size of one was correct.

To everyones surprise it was found that while Qld birds (both sub-species) initially lay only one egg, on average of 32 days later the parents present the first nestling with a second egg, which it accomodatingly incubates for its parents while they spend all their daylight hours catching food for the first nestling. The second egg was always found to hatch after the elder sibling fledged and the parents just kept on feeding the younger sibling. This trick saves the parents three weeks and allows them to raise two progeny in a “normal” or even a dry season. This sibling incubation was the first new method of incubation to be discovered in 150 years (Tarburton & Minot 1987). Although researchers studying other birds breeding under climate stress, thought they might find this phenomenon, the Australian Swiftlet is still the only bird in the world known to employ sibling incubation. Experiments demonstrated that Australian Swiftlets cannot raise two nestings simultaneously. There are not enough flying invertebrates for them to do so (Tarburton 1993b). When colonies are small compared to the area of suitable & available rock surface; nests are spaced out. When space is at an optimum, nests are built against or even onto neighbouring nests.

Breeding Colonies:

This is what the Lower Dooloomai Colony looked like before it was decimated by a land-based predator. Photo copyright M.K. Tarburton

Number of known Breeding Name of Colonies # of Active Census Date # of Birds at Census Date Colonies in Breeding Nests at Last Count Last Count Districts. (Tarburton 2013). _____

A.t. terraereginae: Broken River (4). Dip Creek Cave 1 2.9.1988. 2 Geologists Find 40 ?.4.1988 80 Millenium Cave large # 2000-2001. Trippers Delight 30 2.7.2010. 60 Cairns – Cooktown (7). Russell River Cave – Cairns district 12 ?.1.1993. 24 Red Peak – Cairns distr. 50 ?.?.1978. 100 Thornton Peak – Cairns district ? Jampee Cave, Mt Bartle Frere 20 ?.3.1997. 40 Cave B, - Cooktown district P ?

Family Islands (5). Bedarra (Richards) Isl. 167 ?.?.1980. “ “ 0 ?.1.2007. 0 Dunk Isl. 173 9.9.1989. 246 Timana Isl. (Thorpe) small #. 1948. Wheeler (Toolgbar) Isl. 500 27.12.1977. 1,000 Innisfail (4). Downey Creek Falls 250 1990. 500 N. Johnston River P ?.10.1995. Golden Crown Mine Shaft 1 5.3.1960. 2 The Twins Mine, 3 ?.12.1977. 6

Mackay (2). Above Dooloomai Falls 168 24.11.2011 50 9.2.2015 Below Dooloomai Falls 160 24.11.2011 48 9.2.2015 Tully (4). Tully Gorge Cave 1. 204 1981 500 1981. Tully Gorge Cave 2. 236 ?.8.1981 400 ?.8.1981. Tully Gorge Cave 3. 1981 30 1981. Tully Gorge Cave 4. 400+ 1982 1,000 Douglas Creek, Kirrama S.F. 1 1997 2

A.t. chillagoensis: Chillagoe - CH 5, Tower of London Cave 23 12.1.1987 0 14.12.2011. Mungana (42). CH 9, Swiftlet Cavern 37 3.7.2012. CH 24, Keef’s Cavern 85 6.7.2012. CH 26, Clam Cavern 63 4.7.2012. CH 30, Stop Press 29 9.7.2009. CH 46, Snakey Cavern 3old ?.12.1969 0 31.12.1986. CH 52, Swiftlet Cave 250 2.7.2012. CH 69, Rescue 8 17.4.2011. CH 81, New Southlander 36 23.8.2012. CH 124, Flow Cavern 17 29.6.2012. CH 133, But Good 0 26.6.1992. CH 144, Christmas Main 144 19.6.2012. CH 146, Pot 48 11.7.2012. CH 147, Crocodile Pot 253 22.6.2012. CH 149, Christmas Pot Lower 40 20.6.2012. CH 167, Crack Pot 481 20.6.2012. CH 169, Squeeze Pot 236 19.6.2012. CH 176, Capricorn Cave 0 12.1.1986. CH 187, Gordale Scar Pot 287 24.6.2012. CH 192, Wallabadori 2 2006. CH 229, September Cave 108 5.7.2012. CH 252, Good Black Friday Cave 248 26.6.2012. CH 306, Mudlark Cave 51 14.1.2007. 0 9.12.2011. CH 309, Barramundi Eyes P 4.10.1996. CH 312, Project 31 Cave 160 27.12.1989. CH 322, Swiftlet Scallops #1. Marks of old nests 13.1.1987. 0 CH 362, Hercules Cave Front colony 14 15.1.2007. 0 20.12.2011. CH 362, Hercules Cave Rear Colony 41 17.7.2012. CH 374, Swiftlet Scallops #2 Cave 4 13.7.2012. CH 397, Tarby’s Pot 230 24.6.2012. CH 380, Golgotha 211 18.6.2012. CH 381, Swiftrimlet 66 4.7.2012. CH 397, Shirl’s Triple Twirl 36 29.6.2012. CH 398, Swiftlet Swallet 380 18.7.2012. CH 459, Swiftlet Surprise 447 19.6.2012. CH 496, Satu Lagi 25 2.1.1992. CH 499, Cactus Cleft 43 12.7.2012. CH 501, Bottle Cave 141 24.6.2012. CH 532, Kerrin’s Dilemma 193 28.6.2012. CH 533, Krystelle’s Delight 97 27.6.2012. CH 535, Dirk’s Backyard 22 26.6.2012. CH 536, Lisette’s Art Shoppe 53 25.6.2012. CH 537, Vertical Mums 25 5.7.2012. CH 542, Ella Bella Cave 21 19.7.2012. CH 543, Beef-eater Cave 42 24.8.2012 97 16.7.2012. CH 544, Gruntagrike Cave SE 226 10.7.2012. CH 545, Vertical Terraces SE 70 6.7.2012. CH 547, Mikael’s Drop In 15 20.7.2012. CH 550, Cam’s Shaft 52 23.7.2012. CH 552, Telicis’s Twin 5 22.7.2012. CH 553, First Prussik Cave #2 23 1.7.2012. CH 554, Joseph’s Pit 38 28.7.2012. CH 556, Keith’s Offer 549+ 24.6.2014. CH 557, Shirl’s Swirl 26 24.6.2014. CH 558, Creation’s Speedsters 261 23.8.2014. CHMH1, Granite Rock Cavity Upper 14 26.7.2012. CHMH2, Granite Rock Cavity Middle 111 26.7.2012. CHMH3, Granite Rock Cavity Lower 227 26.7.2012. Mitchell – Palmer (15). MP 1, Guano Chute 169 1.10.1981. MP 8, Laundry Chute 32 2.10.1981. 64 MP 86,87, Gargoyle 4 ?.10.1981. 8 MP 88, Swiss Cheese 14 ?.10.1981. 28 MP 92?, Leopard Cave 2 15.8.1981. MP 92?, Leuberger Cave 126 1.10.1981. 252 MP 94, Here It Is 38 15.8.1981. MP 185, Fifty-fifty Cave P 17.4.1987. MP 188, Porthole Cave P 19.4.1987. MP 189, Delegation Cave P 19.4.1987. MP 194, David’s Delight Cave 50 2?.7.1987 MP 216, Whistle Stop P at 2 sites ? <2001. MP 223, BBC 50 ?.6.1990. 100 MP 225, FIB Swiftlet Extension. 80 ?.10.1990. 160 MP 228, 20 ?.4.1990. 40 MP 244, Split Level 8 old 26.8.1995. 16 MP 247, P 30.8.1995. P MP 254, Flutter Cave P old 23.6.1994. P MP 271, Retreat Cave 100 old ?.8.1997. 200 MP 277, Big Brother Cave P old 18.8.1999. P MP 291, Pure Joy Cave 50 7.7.2000.

WC 12 Swiftlet Cave 50+ 30.12.2013.

Nestling incubating egg Tarby’s Pot Dec 2011. Photo copyright M.K. Tarburton. Feeding Behaviour: Feeds over all types of country but prefers natural and man-made clearings (Storr 1973).

To reduce travel time back to the nests swifts carry pellets (boluses) of dead and dying areial invertebrates in their throat. If birds are intercepted between the feeding grounds and their occupied nests they usually disgorge this pellet of food. Smyth (1980) examined ten such pellets from the colony on Bedarra Island. The pellets contained an average of 94 invertebrates ranging from 1-8 mm. 50.5% of the prey were Dipterans () which we noticed in the introduction are considered by entomologist to be the most manoureavble of aerial . The other proportions were Hemiptera 26.9%, Coleoptera 12.4%, 7.0%, Araneida 2.7% and Odonata 0.5%.

I collected 45 food boluses from swiftlets at Chillagoe over two years and found that the average number of invertebrates per bolus was 146 ranging from 1-10 mm (Tarburton 1993a). Again the most manouvreable group, the flies, were dominant in 11 of the boluses. Homopterans (Plant Hoppers) were dominant in 18 boluses, Hymenopterans (Social Insects) dominant in 13 and Isoptera (Termites) dominant in three. The 1985/6 season was a normal rainfall year and my samples of available food in the sky averaged 9.4 invertebrates of the size swiftlets take but the 1986/7 season was a dry La Niña and the average number dropped to 5.4 insects per sample. The average size of available prey for the two years was 2.43 mm which is significantly smaller than captured prey that averaged 3.89 mm.

Conservation Status: Very common in central part of range around Tully (Storr 1984) and in the south on Paluma Range, & Mt Elliot, as well as open areas of the Atherton Tableland (Wieneke 1992).

From 1985 to 2011 I censused cave populations by counting shiny (ie occupied) nests present during the peak of the breeding season, that is between December and February. Since then my wife and I have switched to a less invasive method. We now count the birds into or out of a cave entrance at sunrise or sunset during the non-breeding season (ie (June or July) Tarburton & T 2013? By using the data in the cave populations given above we can say that the latest population estimate of A.t. terraereginae in 2011 is 4748 and for A.t. chillagoensis is 7004 birds. This gives us a total for the species of 11,752. Because these birds breed and sleep in the same part of the same caves every night of the year, humans are capable of and sometimes cats have destoyed a major part of a local population very easilly. However, because the 90 or so known colonies are stretched along 1,100 km of the NE coast of Australia often away from roads in rugged country I do not consider them as vulnerable as they may at first seem to be.

History: The first known specimen to be collected was taken by MacGillivray on Dunk Island, but it was ignored by Gould for 25 years (Campbell 1909, Mathews 1918).

The first known description from a specimen was made by Ramsay (1874), from several birds caught near Cardwell, Qld. between October 1873 & April 1874.

The first known breeding colony to be discovered by a european was in 1908 when beach comber Banfield found the first of four colonies on Dunk Island in 1908, about ten years after he took up residence on the island (Campbell 1909, Banfield 1911). The first mainland breeding colony to be reported was found by Mrs G. Skinner 24/8/1953, when she located 20 nests under a granite boulder, from where some adults flew, as a National Parks excursion climbed up Finch Hatton Gorge, west of Mackay. It turns out that the local parks ranger Charles Fry, had seen “” nests there two years earlier (Wheeler 1959).

The first colony in a disused mine was located by Cassels (1961) in March 1960, when a nest with nestling was found in the Golden Crown Mine at Mt Peter.

Cypselus terrae-reginae (Ramsay, 1874). terraereginae (Ramsay 1878). Collocalia francica (Hartert 1892). Collocalia francica terraereginae (Oberholser 1906). Salangana francica (Mathews 1908). Collocalia francica terraereginae Grey-rumped Swiftlet. (Mathews, 1913). Collocalia francica zoonava (Mathews 1916). Zoonava francica Grey-rumped Swiftlet. (Mathews 1918). Collocalia (Zoonava) francica Grey Swiftlet. (RAOU, 1926). Collocalia francica Grey Swiftlet, Mothbird. (Cayley 1959). Aerodramus spodiopygius (Brooke 1970). This is the paper that separated swiftlets that could echolocate into Aerodramus. Collocalia spodiopygia terraereginae (Ramsay). Grey Swiftlet. (Storr, 1973). Aerodramus spodiopygia terraereginae Grey Swiftlet. (Pecotich 1982). Aerodramus spodiopygius chillagoensis (Pecotich 1982). Separated this ssp on basis of being paler, smaller and having a broader & whiter rump than coastal terraereginaebirds. Collocalia spodiopygia terraereginae (Ramsay). White-rumped Swiftlet. (Storr, 1984). Collocalia chillagoensis (Pecotich). Chillagoe Swiftlet. (Storr, 1984). Aerodramus spodiopygius White-rumped Swiftlet. (Tarburton & Minot 1987). Aerodramus terraereginae (Lee et al 1996). They believe the DNA they looked at is sufficient to separate the Australian Swiftlet from the South Pacific’s A. spodiopygius. Aerodramus terraereginae (Price et al 2004, 2005). Further DNA evidence that Aerodramus is distinct from Collocalia and that terrareginae is distinct from A. spodiopygius, of the South Pacific. Aerodramus terraereginae Australian Swiftlet. (Christidus & Boles 2008).

Australian Swiftlet braking by exposing maximum surface area upon entering a cave at Chillagoe. Note full throat pouch. Photo copyright G. Anderson.

References:

Banfield, E.J. 1911. My Tropic Isle. Chapter 20. “Swifts & Eagles”” pp 164-167. London. Fisher Unwin. Barrett, G., Silcocks, A., Barry, Simon., Cunningam, Ross., Poulter, Rory. 2003. The New Atlas of Australian Birds. R.A.O.U. Melbourne. Blakers, M., Davies, S.J.J.F., Reilly, P.N. 1984. The Atlas of Australian Birds. Royal Australian Ornitologists Union. Melbourne Unvesity Press. Brooke, R.K. 1970. Taxonomic and evolutionary notes on the subfamilies, tribes, genera and subgenera of the swifts (Aves, Apodidae). Durban Mus. Nov. 9: 13-24. Busst, J. 1956. Nesting of Grey Swiftlet on Bedarra Island. N. Qld. Nat. 116: 1-3. Campbell, A.J. 1909. Annotations. Emu 8: 142-148. Campbell, A.J. & Barnard, H.G. 1917. Birds of Rockingham Bay district, N. Queensland. Emu 17: 2-38. Cassels, M.L. 1961. Second mainland nesting of the Grey Swiftlets. N. Qld. Nat. 29 (128): 3. Cayley, N.W. 1959. What Bird is That?. Halstead Press. Sydney. Christidus, L. & Boles. W.E. 2008. 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Witherby. xii 499 pp. swifts = 7: 246-281. Oberholser, H.C. 1906. A monograph of the Collocalia. Proc. Acad. Nat. Sci. Philadelphia 53: 177-212. Pecotich, L. 1974. Grey Swiftlets in the Tully River Gorge and Chillagoe Caves. Sunbird 5(1), 16-21. Pecotch, L. 1982. Speciation of the Grey Swiftlet Aerodramus spodiopygius in Australia. Tower Karst 4, 53-57. Price, J.J; Johnson, K.P; Clayton, D.H.2004. The evolution of echolocation in swiftlets. Jour. Avian Biol. 35, 135-143. Price, J.J., K.P. Johnson, S.E. Bush and D.H. Clayton. 2005. Phylogenetic relationships of the Papuan Swiftlet Aerodramus papuensis and implications for the evolution of avian echolocation. Ibis 147: 790-796. Ramsay, E.P. 1874. Descriptions of five new species of birds from Queensland, and of the egg of Chlamydodera maculata. Proc. Zool. Soc. Lond. 39: 601-605. Ramsay, E.P. 1878. Proc. Linn. Soc. N.S.W., vol II, p. 178. RAOU 1926. Official Checklist of the Birds of Australia 2n Ed. 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