CSIRO PUBLISHING www.publish.csiro.au/journals/emu Emu, 2004, 104, 1–6 The breeding ecology and behaviour of a colour-marked population of Brown Falcons (Falco berigora) Paul G. McDonald School of Botany and Zoology, Australian National University, Canberra, ACT 0200, Australia. Email: [email protected] Abstract. This study took advantage of a large, closely monitored colour-banded population of Brown Falcons to describe aspects of this species’ breeding ecology and behaviour over three consecutive years. Both pair members aggressively defended territories throughout the year from conspecifics and other species alike. Males performed territorial displays more frequently than females, which rarely displayed unprompted. Strong differences in the types of parental care provided by each sex were evident, with females contributing most to incubation, brooding and feeding of nestlings and fledglings. Males, on the other hand, provided most of the food to both females and broods, from well before the first eggs were laid until nestlings were 2–3 weeks old. Prey deliveries were more frequent early in the morning and late in the evening; however, remains of larger prey were cached, presumably to provision the offspring more regularly throughout the day. Introduction Details of the study site (Baker-Gabb 1982, 1984a) and population, which included 44–49 breeding pairs, have been described elsewhere Despite being one of Australia’s most commonly encoun- (McDonald 2003a, 2003b; McDonald et al. 2003). tered raptors (Blakers et al. 1984), the behaviour and breed- Falcons were captured with bal-chatri and modified goshawk traps ing biology of the Brown Falcon (Falco berigora) has (see Bloom 1987 for details), fitted with a unique combination of colour received little scientific attention. In the most comprehensive bands approved by the Australian Bird and Bat Banding Scheme and released at the point of capture. During frequent visits to the study site studies to date, Baker-Gabb (1982) monitored up to 25 pairs banded birds were actively sought, identified and their behaviour noted. in southern Victoria over three seasons and Mooney (1976) When found, nest trees were climbed daily until clutches were described several aspects of the behaviour of captive indi- complete. If not observed directly, laying dates of first eggs were viduals. However, this work is largely confined to unpub- estimated using either a formula based on egg weight (P. Olsen, lished theses. Bollen (1993) published descriptions of the unpublished data) or, if eggs hatched, by backdating using an average incubation period of 36 days (this study) and estimating chick age from behaviour from one pair, while Debus (1991) described two formulae based on wing length (McDonald 2003b). Nests were visited displays of a single male and Olsen and Olsen (1980) have weekly after hatching to determine nestling periods and the number of described nest-defence behaviour in response to human birds successfully fledged from each nest. At ~4 weeks of age nestlings intruders. Additional information on the Tasmanian popu- were fitted with colour bands. Following fledging, the length of post- lation has been provided by Mooney in Cade (1982) and fledging parental care was determined by visiting the nest area bi-weekly. If fledglings could not be located in three successive visits Marchant and Higgins (1993). The latter also contains a they were deemed to have successfully reached independence. During review of additional anecdotal evidence. the breeding seasons of 2000 and 2001 intensive assessments of This study, as part of a larger investigation into the breed- parental roles at the nest were conducted using surveillance cameras ing biology and life-history strategies of Brown Falcons, throughout the nestling stage. Small cameras with infrared lights (Model 43150674; Radio Parts Group, Melbourne) were placed at nests took advantage of a large, colour-banded population to for 48-h periods throughout the nestling phase. Cameras were powered investigate and describe additional behaviours of birds of by deep-cycle batteries (Besco N70T; Battery World, Canberra) and known sex and background. In addition, footage from nest- connected to time-lapse video-recorders (Hitachi VT1200E; Radio surveillance cameras over two seasons allowed a detailed Parts Group, Melbourne) run at one-eighth normal speed; this examination of the degree of sex-role partitioning during minimised visits to the nest tree to 24-h intervals to change batteries and tapes. The behaviour of parent birds and offspring was then noted on parental care. subsequent viewing of video footage. Materials and Methods Statistical analyses The study was conducted between July 1999 and June 2002 ~35 km Goodness-of-fit tests were used to assess the significance of differences south-west of Melbourne, at the Western Treatment Plant, Werribee in prey-delivery rates throughout the day as well as sexual differences (38°0′S, 144°34′E), adjacent to Avalon Airport (38°2′S, 144°28′E) and in the recorded frequency of each display type. Where appropriate, small areas of surrounding private land, a total area of ~150 km2. Yates’ corrections for continuity were used (Zar 1996). One-way © Royal Australasian Ornithologists Union 2004 10.1071/MU02042 0158-4197/04/010001 2 Emu P. G. McDonald ANOVAs and linear regressions were used to assess yearly differences torial disputes, usually given with much calling. Equal in the duration of various phases of parental care. The methods used in numbers of males and females were observed performing this project was approved by the Australian National University Animal this display. Another apparently territorial display was Experimentation Ethics Committee (F.BTZ.02.99). observed in 11.1% (total n = 126) of observations, with Results falcons flying in a series of large, undulating ‘U’ shapes, Agonistic interactions giving quick wingbeats during upsurges, again with much calling and usually while rolling from side to side with the Both members of Brown Falcon pairs aggressively defended wings held in a half-closed position, particularly during territories from intraspecific intruders of either sex through- descents. More rarely, this display was performed while out the year, although agonistic interactions were more flying at a constant elevation in a figure-of-eight pattern. common and were of a longer duration between July and U-displays were given significantly more often by males November (the breeding season, see below). Interactions χ2 (85.7%, total n = 14, c = 5.21, P = 0.02); females were often escalated into physical contact between birds, with observed giving this display on just two occasions (both perched combatants observed jumping to the ground to avoid times in the company of a displaying male). U-displays were being struck by conspecifics on seven occasions. Talon- usually performed over a male’s own territory during/follow- presentation displays were also common and, in one extreme ing agonistic interactions, with neighbouring males observed case, two females locked talons and spiralled downwards giving this display while soaring 200 m apart. Typically, 20 m in a ‘cartwheeling’ flight, completing four full revolu- U-displays were given at great height, often culminating in tions before releasing their grip close to the ground. spectacular, steep and rapid descents to nest trees, with marked side-slipping and frequent calling throughout. After Territorial and courtship displays birds returned to perches following territorial disputes, occa- Displays were also frequently encountered throughout the sionally both members of a pair would continue to call, year, becoming more frequent from July to November. Of a raising their wings above their back, fully outstretched, while total of 126 displays given by banded birds, ‘side-slipping’ facing the direction from which the intrusion came. At dusk was the most frequently observed (62.7%). During this males would fly around their territory with fast wing beats display birds would rotate rapidly from side to side while in giving single, sharp ‘ack’ calls, with the ‘arrrrrk-ark-ark’ a gliding descent, often from great height, with wings held call often heard during this and all other displays. stiffly behind their back, alternatively displaying their dorsal and ventral plumage. Males were usually more exaggerated Courtship feeds and copulation behaviour than females in their movements, often using strong winds to The first courtship feeds of the season were recorded in perform the display in a stationary position above their mid-July each year. Males would bring prey, house mice eventual perch, rotating through an axis of up to 270° with (Mus musculus) in all cases identified (n = 13), towards each turn. This display was usually given by both sexes upon females with fast, exaggerated wing beats, frequently giving landing near the nest tree or returning to perches following ‘arrrrrk-ark-ark’ calls and performing the side-slipping agonistic interactions. Females (40.5%) were observed display before landing. Females responded with trilling calls giving this display less frequently than males (59.5%, n = 79, or occasionally a sharp ‘kark’, taking the prey from males’ χ2 c = 2.86, P = 0.09). In all cases, females were prompted in beak to beak or, less often, with her talons. Copulation was some manner before giving the display, whereas males often observed following a courtship feed on only one occasion performed displays alone. For example, following territorial (6.7%, total n = 15). Females initiated copulation on 27.8% disputes, females would usually follow males back to perches, of all occasions that it was witnessed (n = 18), by giving usually 3–5 m behind. Upon landing, males routinely gave the short, sharp monosyllable calls, bowing their heads forwards side-slipping display, with females following suit with fewer, and lifting up their tail and rump. After this behaviour males smaller rotations. The only other occasions when females would approach the female and land upon her back before were recorded side-slipping was after they were flushed from copulation (lasting 5–7 s) ensued.