Wilson Bulletin, 110(l), 1998, pp. 71-76

BREEDING OF THE WHITE-RUMPED PYGMY

ALAN C. KEMP ’ AND ADISAK VIDHIDHARM*

ABSTRACT.-A nest of the White-rumped , insi@s, was observed in Thailand for five days during the incubation period. The nest site in a woodpecker hole and the white eggs are described for the first time. Observations on diet, hunting and breeding behavior are also described. The systematic position of the is discussed. Received8 May 1997, accepted4 Oct. 1997.

The breeding biology of only one species distinctive male song. At the end of the study, the nest of pygmy falcon, the African Pygmy Falcon tree was climbed and the cavity inspected. Observations were made at the peak of the hot dry (Polihierax semitorquatus), has been studied season, with daily temperatures exceeding 40” C and in any detail (Maclean 1970). We report on only brief drizzle falling from the afternoon build-up five days of observation at a nest of its con- of clouds. The pair were quite approachable and ob- gener, the White-rumped Pygmy Falcon (P. servations were made either on foot or from a ground insignis). The six other very small , hide of saplings and foliage erected 20 m south of the falconets in the genera Spiziapteryx and Mi- nest tree. A pair of 10 X 40 binoculars and a 15-60X crohierax, are also relatively little known telescope facilitated observation. Vocalizations were recorded with an ECM-KlOO microphone to a Sony (Bierregaard 1994, Clark 1994, Kemp and WM-D3 cassette recorder and were analyzed with AV- Crowe 1994). ISOFT-SONOGRAPH PRO Windows version 2.5 soft- We watched White-rumped Pygmy Falcons ware. near the headquarters of the Huay Kha Khaeng Wildlife Sanctuary, western Thailand, RESULTS during 28 March to 1 April 1996. A nest of Nest site.-The nest tree was an emergent P. insignis had been photographed nearby sev- 24 m-tall Shorea obtusa tree with a dbh of en years earlier, at about the same time of year about 90 cm, one of several larger trees along (Pilai Poonswad, pers. comm.). Nakhasathien the crest of a low ridge in dry deciduous dip- and Stewart-Cox (1990) describe the sanctu- terocarp forest. Trees in the forest were about ary in detail. 15-20 m high, with some larger emergents, METHODS and the overall impression of the vegetation was a dense 2 m-high scrub layer, with nu- We discovered an active nest about 1 km south of the headquarters, close to the start of the Khao Hin merous bare tree trunks up to about 10 m, Dhaeng Nature Trail, at 1.5” 36 ’ 10.9” N, 99” 19 ’ 1.5” topped by an uneven and partly open canopy. E and an altitude of 210 m a.s.1. The nest was discov- The ridge with the nest tree was about 1 km ered at 1250 on 28 March after watching for a pair of east of the foothills of the Thanon Thongchai pygmy falcons that had been seen copulating in the mountains, which run north-south down the area several weeks before (Thannyalax Suntaramat, Thailand-Myanmar border. The foothills were pers. comm.). The nest area was watched from 12:00 to 19:00 on covered in denser, taller bamboo and ever- 28 March (except when erecting a blind at 14:Ot-15:00X green forest. dawn (06:00, local sunrise 06:25) to dusk (19:00, local The nest was in an old woodpecker hole on sunset 18:25) on 29 March, 06:00-12:00 and 14:00- the north-east side of the trunk, the entrance 19:00 on 30 March (for photography) and on 31 being 12 m above ground and 2 m below March, and 06:00-11:00 on 1 April. Three man-days where the first branches emerged. Below and also were spent searching the surrounding area for neighboring pairs, including the use of playback of the on the same side as the falcon nest, at between 7-10 m, were two old overgrown and four new holes in which three (two males and a I Dept. of , Transvaal Museum, PO. Box 413, female) Greater Flameback (Chrysocolaptes Pretoria, 0001 Republic of South ; E-mail: [email protected] lucidus) woodpeckers returned to roost each 2 Hornbill Project Thailand, Mahidol Univ., Rama evening. The tree, which appeared healthy up VI Rd., Bangkok, 10400 Thailand. to the lowest hole, had growths on the trunk

71 72 THE WILSON BULLETIN * Vol. 110, No. I, March 1998

TABLE 1. Hourly rates in Polihierux insignis of prey delivery to the female by the male and of hunting from the nest by the female, during three days of observation within the incubation period.

Delivery or hunting rates per hour, starting at: Number of Sex 06 07 08 09 10 II 12 13 14 I5 16 17 I8 We”@

Male 0.7 0.7 1.3 0.7 0.0 0.0 1.0 1.0 0.5 1.3 0.3 0.7 0.0 22 Female 0.0 0.0 0.0 0.0 0.7 0.0 0.0 0.0 3.5 1.3 1.7 0.3 0.0 19 Davs observed 3 3 3 3 3 12 2 2 3 3 3 3

above, sparse foliage and a large dead limb fied items. Most of the lizards appeared to be above the canopy. Druco flying lizards (or at least slender aga- The oval nest entrance was 9 cm high by 8 mids with long tails, large hind legs and, in cm wide and the nest floor was 9 cm below two individuals, expandable throat flaps). Un- the lower lip of the entrance. The nest cavity, identified items were small and were probably which had a diameter of about 32 cm, was insects. The male of a neighboring territory lined with fine wood chips. Overall, the nest was seen to catch a Draco lizard at 13:20 on hole was similar to the woodpecker roost 1 April, darting 25 m from a perch 3 m up to holes below. On the afternoon of 1 April, the snatch the lizard from 2 m above the ground nest contained two plain white, rounded-oval off a 6 cm diameter branch of a small tree. eggs that measured 35.4 X 25.6 mm and 32.5 The female confined her hunting to within X 24.7 mm. 50 m of the nest tree, striking either from the Incubation behavior.-The female incubat- nest entrance or from a nearby perch after a ed during the day, although the male some- previous attempt. Most prey (N = 17) were times entered the cavity briefly during or after not identified because they were consumed delivering food. The male roosted in the nest where captured, usually behind foliage, or with the female, entering at 18:45 on 28 were carried rapidly to the nest. A cicada and March, leaving again at 06:09 on 29 March a grasshopper were identified and probably and returning to sleep at 18:47 that evening. most prey was of this size, given that con- Both entered the nest at 18:55 on 1 April as sumption time was usually less than a minute soon as the nest inspection was completed. and no large prey, such as lizards, was seen The female spent most of the time in the being carried to the nest. nest cavity, only coming to the entrance when We saw the female attempt to capture prey hearing the male or other birds nearby (see 13 times. Six attempts were to the ground, below). During the heat of the day she also four into foliage, and three to a branch or perched at the nest entrance, from which she twig. Strike distances estimated to the nearest sometimes pursued nearby prey. Rates of prey 10 m were 40 m (l), 30 m (l), 20 m (2), 10 delivery and capture (Table 1) indicate that, m (6) and 0 m (vertical, 1). Four of these except for a late morning break, the male pro- strikes, one to the ground among dry leaves visioned the nest throughout the day and that and three into foliage (3 l%), were successful. the female hunted from the nest in the heat of Two of the successful strikes were upwards the afternoon. from the perch, at the underside of the foliage, Hunting behavior and diet.-We never saw and one downwards. the male hunt, despite numerous attempts to Calls and displays.-The male repeated a follow him as he left the nest area. His small shrill flight call chwirr chur chur (Fig. la), size made it difficult to detect him as he singly or in series, when flying to the nest seemed to hunt from low perches or from with food. The female responded with a mo- deep within the foliage. Judging from where notonously repeated and less harsh cheep call he disappeared and then re-appeared with (Fig. lb). This call was also used when the prey, the male appeared to hunt within about male was absent. It appeared to be a food so- 400-600 m of the nest. Prey delivered to the licitation call since it was often given for long nest by the male included eight lizards, a ci- periods when the female perched at the nest cada, an unidentified insect and 12 unidenti- entrance and hunted for herself in the after- Kemp and Vi&i&arm - WHITE-RUMPED PYGMY FALCON 73

kHz IO- la 8- ’ ’ z

kH; kH; IO lb 10 IC

8 8

6 6

4 4

2 2

0.5 S S

kH; 10 Id 8

6

4

2

u.s 1 1.5

FIG. 1. Sonograms of the main calls of Polihierax insignis. Male flight call (la), female food solicitation call (lb) and male song at low (lc) and high (Id) intensity. See text for further details. 74 THE WILSON BULLETIN l Vol. IZO, No. I, March 1998

noon. The intensity of the call was a soft purr Both birds performed exaggerated head when the female was alone (Fig. lb), accel- bobbing when examining potential prey. Apart erating to a harsh screech when the female from mantling prey, no other display postures was perched next to the male carrying food. were recorded under natural conditions. How- The female sometimes flew calling up to 200 ever, playback of selected calls were per- m from the nest to meet the male and solicit formed once each on the male when he was prey. about 200 m away from the nest. Playback of After the flight call, and especially after the male flight call and female solicitation call presenting prey, the male often uttered a short produced no visible reaction. Playback of the musical song. This was delivered either in ab- male song caused the male to fly towards the breviated form as a simple whistle-like phrase nest, stopping several times en route to sing wee-you (Fig. lc) or as a more developed se- in reply and once, during an intense period of ries wee-you-pity-you (Fig. Id). The latter 6-8 songs, to pump his tail up and down five somewhat resembles the twee twee twip song times through about 45”; the effect, with the of the African P. semitorquatus (Maclean wings drooped, was to flash the white rump 1970; sonagram in Maclean 1993). The song and display the white-spotted tail. was given either in the nest, if prey was pre- One evening, at 17:41, the female emerged sented there, or after returning to a nearby from the nest to perch with her plumage perch. The song was repeated, sometimes sev- fluffed. She then preened vigorously before eral times, especially if the female was out of flying east towards a sandy track. After 15 the nest and perched nearby. The male also min she returned to the nest without food and uttered a harsh insistent sheer, either as a sin- when the track was examined later, signs on gle note in alarm or rapidly repeated if ner- the sand suggested recent dust bathing. A pair vous. This was often given as he hung at the of pygmy falcons had been sketched one eve- nest entrance and once when he was dived at ning a few months earlier while dust bathing by an Indian Cuckoo (Cuculus micropterus) together on a sandy track just north of the away from the nest area. headquarters (Thannyalax Suntaramat, pers. The male carried prey in his feet until near comm.). the nest and then, at a perch or in flight, trans- Presence of and reaction to neighbors.- ferred it to his bill. This enabled him to land No neighboring pairs of pygmy falcons were at the nest entrance or on a perch with the found by searching. However, on 1 April at food ready to pass to the female. Several times 13:20 we were called to see a male in the the male refused initially to pass prey to the grounds of the headquarters. We watched it female. Sometimes he ate some himself and catch a lizard, consume the forequarters and at least twice he eventually consumed the then carry the hindquarters off in a north-east- whole lizard without feeding the female. Once erly direction. Although no nest was located, the male appeared to threaten the female, the behavior suggested a second breeding pair, mantling with wings drooped and three-quar- probably neighbors, with a nest at least 1.5 km ters open, after which he flew off to eat the north of the study nest. prey. Such encounters were noisy, accompa- The falcons at the nest were especially ag- nied by loud soliciting (female) and singing gressive to a pair of Hill Mynahs (Gracula or alarm calls (male). religiosa) that visited the nest tree as many as The pair clung at the nest almost like a five times a day. The female always came to woodpecker or parrot, braced on their tails, the nest entrance when she heard the mynahs and they were also able to run up the trunk call upon landing in the nest tree and once, without opening their wings. Twice the male when she had hunted away from the nest, she landed lower down on the nest trunk without dived at the mynahs on her way back to the food and scuttled between the woodpecker nest entrance. The female also came to the roost holes, entering one or other for a brief entrance on hearing a Shikra (Accipiter bad- period and even calling from within. Their an- ius) calling overhead, or when Hair-crested tics on the nest trunk or when trying to snatch Drongos (Dicrurus hottentottus), themselves prey from branches indicated unusual flexi- building a nest 50 m away, uttered alarm calls. bility of their leg and toe joints. At least three pairs of Shikra had nests Kemp and Vi&i&arm * WHITE-RUMPED PYGMY FALCON 75 within earshot of the falcon nest, the nearest 1994, and Asia, Clark 1994). Both have about 400 m away, and the area was also vis- marked sexual dimorphism, the male with ited several times by Collared Falconets (Mi- more gray and the female with more rufous crohierux caerulescens) and Black Baza (Avi- markings. Both pump the tail in display, the cedu Zeuphotes). At least five other species of African species more frequently (Maclean large woodpeckers also visited the falcon nest 1970), but so do some true falcons in the area and could have provided alternate nest Americas and on some Indian Ocean islands sites: Common Flameback (D. juvanense), (Kemp and Crowe 1994), including the Mad- Greater Yellownape (PicusJlavinucha), Lesser agascar Kestrel (F. newtoni, Siegfried and Yellownape (P. chlorolophus), White-bellied Frost 1970). Males of both Polihierax species Woodpecker (Dryocopus javensis) and Black- also utter a distinct song of similar structure. headed Woodpecker (P. erythropygius). Polihierax insignis has been separated previ- ously, on tail and primary dimensions, into the DISCUSSION Neohierux (Swann 1922). This separa- Our observations corroborate most aspects tion is supported by a phylogenetic study of of a recent summary of the biology of the syringeal morphology (Griffiths 1994). In a White-rumped Pygmy Falcon (Clark 1994). further distinction, the African P. semitorqua- They emphasize the importance of lizards in tus nests in cavities within the nest structure the diet, especially as large prey items deliv- of other birds, in the same way as the South ered to the nest when breeding. Specialization American Spot-winged Falconet (Spiziupteryx for hunting on trunks and foliage may explain circumcinctus; Bierregaard 1994), while the the limited overall range of this species in dry Asian P. insignis is now known to nest in tree deciduous forests. Such arboreal and terrestri- holes excavated by other birds, like the five al hunting differs notably from the more aerial species of Asian Microhierux falconets (Clark hunting above the canopy of sympatric Mi- 1994). crohierux falconets (Kemp and Crowe 1994). However, previous reports of the White- ACKNOWLEDGMENTS rumped Pygmy Falcon nesting in old stick We thank especially Dr Pilai Poonswad for arrang- nests and laying off-white eggs (Swann 1933, ing all logistics and funding for the study. We much Clark 1994) appear erroneous; the white eggs appreciated Preecha and Thannyalax Suntaramat for placed in a cavity nest, as described here, are sharing their knowledge and images of falconets, and typical of all other Old World falconets. Drs. Jaboury Ghazoul and Gnome Liston for teaching us to rope-climb safely. Phylogenies of the 60 species of suggests three phylogenetic and ecological LITERATURE CITED clades: (1) caracaras (polybori, 8 species), (2) forest and laughing falcons (micrasturi, 6 spe- AMADON, D AND J. BULL. 1988. Hawks and owls of the world: a distributional and taxonomic list, with cies) and (3) true falcons (falconi, 46 species) the genus Otus by J. T Marshall and B. E King. [Amadon and Bull 1988, Kemp and Crowe Proc. West. Found. Vert. Zool. 3:295-357. 1990, Griffiths 1994 (who includes the laugh- BIERREGAARD,R. 0. 1994. Neotropical raptor species. ing falcon with the true falcons)]. The first Pp. 254-255 in Handbook of birds of the world. two clades are Neotropical in distribution, but Vol. 2 (J. de1 Hoyo, A. Elliott, and J. Sargatal, the true falcons are cosmopolitan. The true Eds.). Lynx Editions, Barcelona. falcons include the three genera and eight spe- CLARK, W. S. 1994. Indomalayan raptor species. Pp. 255-256 in Handbook of birds of the world. Vol. cies of pygmy falcons and falconets, which 2 (J. de1 Hoyo, A. Elliott, and J. Sargatal, Eds.). are scattered across the Neotropical, Afrotrop- Lynx Editions, Barcelona. ical and Indomalayan biogeographic realms GRIFFITHS, C. S. 1994. Syringeal morphology and the (Amadon and Bull 1988). Phylogenetically, phylogeny of the Falconidae. Condor 96: 127-140. these dimunutive species represent basal KEMP, A. C. 1994. Afrotropical raptor species. l? 255 clades to the 38 species of larger falcons and in Handbook of birds of the world. Vol. 2 (J. de1 Hoyo, A. Elliott, and J. Sargatal, Eds.). Lynx Ed- kestrels in the genus FuZco (Kemp and Crowe icions, Barcelona. 1990, Griffiths 1994). KEMP, A. C. AND T. M. CROWE. 1990. A preliminary The two species in the genus Polihierax oc- phylogenetic and biogeographic analysis of the cur on different continents (Africa, Kemp genera of diurnal birds of prey. Pp. 161-175 in 76 THE WILSON BULLETIN . Vol. 110, No. I, March 1998

Vertebrates in the tropics (G. Peters and R. Hut- NAKHASATHIEN, S. AND B. STEWART-C• X. 1990. terer, Ed.%). Museum Alexander Koenig, Bonn. Thung Yai-Huay Kha Khaeng World Heritage KEMP, A. C. AND T M. CROWE. 1994. Morphometrics Nomination. Royal Forest Dept., Bangkok. of falconets and hunting behaviour of the Black- SIEGFRIED,W. R. AND l? FROST. 1970. Notes on the thighed Falconet Microhierax fringillaris. Ibis Madagascar Kestrel F&o newtoni. Ibis 112:400- 136:44-49. 402. MACLEAN, G. L. 1970. The Pygmy Falcon Polihierax SWANN, H. K. 1922. A synopsis of the Accipitres. semitorquatus. Koedoe 13: l-2 1. Wheldon and Wesley, London. MACLEAN,G. L. 1993. Roberts ’ birds of southern Africa, SWANN, H. K. 1933. A monograph of the birds of 6th ed. John Voelcker Book Fund, Cape Town. prey. Vol. 2, Wheldon and Wesley, London.