Breeding Biology and Food Habits of the Madagascar Kestrel (Falco Newtoni) in Northeastern Madagascar
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j. RaptorRes. 39(2):149-155 ¸ 2005 The Raptor ResearchFoundation, Inc. BREEDING BIOLOGY AND FOOD HABITS OF THE MADAGASCAR KESTREL (FALCO NEWTONI) IN NORTHEASTERN MADAGASCAR LILY-ARISONRENE DE ROLAND,JEANNENEY RABEARIVONY, HARILALAINAROBENARIMANGASON, AND GILBERTRAZAFIMANJATO ThePeregrine Fund's Project in Madagascar,B.P 4113, Antananarivo(101), Madagascar RUSSELL THORSTROM 1 ThePeregrine Fund, 5668 WestFlying Hawk Lane, Boise,ID 83709 U.S.A. ABSTRACT.--Westudied MadagascarKestrels (Falconewtoni) on Masoala Peninsula, northeastern Mada- gascarduring the 1997 and 1998breeding seasons. We locatedfive nestsites and observedeight nesting attemptsduring the two breeding seasons.All nestswere in tree cavitiesand averaged13.8 + 2.0 m (SE) above the ground in trees averaging22.8 +- 0.8 m (SE) in height (N = 5 nests). Egg laying took place from mid-Septemberto the first week of October.The modal clutch sizewas 4 + 0.9 (N = 6 nests, range 3-5 eggs).The incubationperiod averaged28 d, varying from 27-29 d (N = 5 nests).Hatching occurred from the middle of October to the first week of November with young fledging in late No- vember.Of 24 eggslaid in six nests,13 (54%) hatched,and seven(54%) of thosehatchlings fledged; thus, a total of 1.2 young fledged per breeding attempt were produced and overall nest successwas 50%. The MadagascarKestrel diet of 338 identifiedprey wascomposed of 93.8% lizards(N = 317), 2.6% insects(N = 9), 2.4% amphibians(N = 8), and 1.2% birds (N = 4). KEYWORDS: MadagascarKestrel; Falco newtoni; nestingbiology; food habits;nests; productivity. BIOLOG• REPRODUCTIVA Y I-IJd3ITOS ALIMENTICIOS DE FALCO NEWTONI EN EL NORESTE DE MADAGASCAR RESUMEN.-•Estudiamosindividuos de la especieP•tlco newtoni en la peninsulaMasoala en el norestede Madagascardurante las estacionesreproductivas de 1997y 1998.Localizamos cinco nidos y observamos ocho intentos de nidificaci0n durante los dos periodos reproductivos.Todos los nidos se encontraron en cavidadesen •rboles a una altura promedio de 13.8 + 2.0 m (EE) sobre el sueloyen •rboles con una altura promedio de 22.8 + 0.8 m (EE) (N = 5 nidos). La puestade huevosocurri6 desdemediados de septiembre hasta la primera semanade octubre. E1 tamafio modal de la nidada fue de 4 -+ 0.9 huevos (N = 6 nidos, rango 3-5 huevos).E1 periodo promedio de incubacitn fue de 28 dias, variando entre 27-29 dias (N = 5 nidos). La eclositn ocurri6 desdemediados de octubre hastala primera semanade noviembrey el abandonodel nido por parte de los polluelosocurri6 a fines de noviembre.De los 24 huevospuestos en seisnidos, 13 (54%) eclosionaron,y siete (54%) de los polluelosque eclosionaron abandonaron el nido. Por lo tanto, se produjeron un total de 1.2 juveniles que abandonaron el nido por intento de nidificaci0n,y el fixito de nidificaci0ngeneral fue del 50%. La dieta de F. newtoni,basada en 338 presasidentificadas, estuvo compuesta en un 93.8% por lagartijas(N = 317), en un 2.6% por insectos(N = 9), en un 2.4% por antibios(N = 8) yen un 1.2% por aves(N = 4). [Traduccitn del equipo editorial] Madagascarhas three residentspecies of falcons: most common raptor in Madagascar(Siegfried and MadagascarKestrel (Falconewtoni newtoni), Banded Frost 1970), detailed information on its biology Kestrel (F. zoniventris),and Peregrine Falcon (F. pe- and natural history are lacking (Langrand and regrinusradama), and two wintering species,Eleo- Meyburg 1984, Langrand 1990). This speciesis nora's Falcon (F. eleonorae)and Sooty Falcon (F. widely distributed throughout Madagascar and is concolor).Although the MadagascarKestrel is the found in open grasslands,savannah habitat, de- graded forests,and in the vicinity of villagesand towns. This falcon has two distinctive color • Email address:[email protected] morphs:pale and rufous (Siegfriedand Frost1970, 149 150 RENE DE ROLAND ET AL. VOL. 39, NO. 2 Cade 1982, Langrand 1990). In this paper, we re- sions,nest depth, and internest distance) were measured port basic information on the breeding biology after the young had fledged. and food habits of the MadagascarKestrel. RESULTS STUDY AREA AND METHODS One pair of MadagascarKestrels was trapped This studywas conducted within the secondaryhabitat and marked during this study. The adult female and cultivated terrain surrounding Ambanizana Village measurements were wing chord, 200 mm; tail (49ø57'E, 15%7'S), situated at the western boundary of length, 129 mm; tarsuslength, 38.2 mm; and body the Masoala National Park (MNP; Robenarimangason 1999). mass, 122 g. The adult male measurementswere Most of the peripheral zones of the MNP, are com- wing chord, 195 mm; tail length, 110.5 mm; tarsus posed of degraded forest, secondaryforest, and fallow length, 37.0 mm; and body mass,110 g. land intermixed with cultivated areas (crops include Five nesting attemptswere observedduring the cloves, coffee, vanilla, and bananas). The MNP itself con- first season, 1997-98, and three were documented sistsof 230 000 ha of primary forest with a typicalcanopy height of 25 m and with some emergent trees exceeding during the second field session,1998-99. Of the 30 m (Guillaumet 1984). The forested terrain is moun- five nesting pairs observed during the two breed- tainous, and lacks roads and a trail system.The altitude ing seasons,three pairs were composedof a pale- in the park ranges from sea level to 1230 m (Nicoll and morph male and a rufous-morphfemale, one pair Langrand 1989). The current study was undertaken in the lower altitude and cultivated areas ranging from sea was of a rufous-morphmale paired with a pale- level to 200 m above sea level. The annual mean rainfall morph female, and one pair included both rufous- at AndranobeField Station,7 km southof the studysite, morph individuals. is 6049 mm (Thorstrom et al. 1997). The dry seasonis Nest Characteristics. During this study, all nests from October to mid-December (Rene de Roland 2000). observed for MadagascarKestrels were placed in The climate is mild and the annual mean temperature varies from 18-31øC. natural-tree cavities,with a decayedwood substrate, The studyperiod wasfrom September1997-January in secondary and human-modified habitat. The 1998 and September1998-January 1999, which coincides trees identified for nestingwere mandrorofo (Tra- with the kestrel's breeding season. During September chylobiumrerrucosum), hintsia (Afzeliabijuga), lalona and October, we searchedfor nesting pairs by following vocalizing and flying adults. Observationswere made at (Weinmania sp.), and dead unidentified snags. least 50 m from nests, and five and three nests were mon- MadagascarKestrel nestsaveraged 13.8 -+ 4.5 m (N itored during the 1997 and 1998 breeding seasons,re- = 5) above the ground in trees averaging22.8 -+ spectively. 1.9 m (N = 5) in height. Nest cavitieswere oval- Nestswere observedfrom courtship up to post-fledg- shaped and measured 61 _+ 55.1 cm (N = 2) by ling period, and 521 hr 22 min of observationwere com- pleted. We recorded copulationfrequency (per hr and 33.5 _+ 16.2 cm (N = 2) with an interior depth of d) and duration (sec), clutch size (number), incubation 22.5 + 3.5 cm (N = 2). The distance between two period (d), nest attendance (percent of observation neighboringnests averaged 675 _+386.2 m (N = ume), dispersalof young (d), and productivity.Egg di- 4, range 300-1200 m). mensions (length and breadth), and egg and nestling Nesting Biology. Courtship activity involved vo- masswere measuredusing vernier calipers (with 0.01 mm of precision)and Pesolaspring-balance scales (0.1 g pre- calizations, nest site visits with food deliveries to cision), respectively.Daily nest observationswere made the female, and copulations.The courtshipperiod from 0600-1800 H with 10x binocularsand a zoom spot- wasmarked by flights (e.g., in open areasand over ting scope.While making observationsat nestsprey items trees) and accompaniedby moderate fluttering of were identified during prey deliveries. Kestrelswere trapped with a bal-chatri,a noosecarpet wings. During periods of inactivity,the kestrel pair fixed over the nest entrance, or a "wire hoop trap" (Ber- perched together in the top of dead branchesof ger and Mueller 1959, Thorstrom 1996). Morphometric trees. Courtship behavior,either flights or periods measurementstaken were: wing chord (mm), tail and of inactivity,was associatedwith loud calls "itsi, kit- tarsilength (mm), and body mass(g). One breeding pair si, kitsi, kitsi." During this period, the male's pri- of adult kestrelswas radiotaggedto estimate their rang- lng area. The kestrelswere radio-trackedfrom October- mary role seemed to be showingthe female poten- December 1998 by homing on foot, and locational fixes tial nesting sites. were recorded using an Eagle Explorer Global Position Copulationsusually occurred after prey deliver- System(GPS; Eagle Electronics,Catoosa, OK U.S.A.) with ies from the male to the female. Copulationsoc- 30-m of accuracy.The home range was estimatedby the minimum convex polygon (MCP) of the locationsusing curred on the top of dead branches.During cop- Ranges1V software (Kenward 1990). Severalnest site pa- ulations, the male emitted a "iiitsi, kitsi, kitsi, rameters (i.e., nest height, nest tree height, nest dimen- kitsi..." that continued to the end of this activity. JUNE 2005 BREEDINGBIOLOGY OF THE M•'d)^G^SC•V.KESTREL 151 Table 1. Number of prey items deliveredby male and female MadagascarKestrels at MasoalaPeninsula, Madagascar, during the breedingseasons of 1997 and 1998. SEX COURTSHIP INCUBATION NESTLING POST-FLEDGING TOTAL Male 13 110 130 18 271 Female 0 0 28 39 67 Total 13 110 158 57 338 The highest frequency of copulationsduring a 1 (Table 1). During this period, the adult female was hr period occurredbetween 0600-0700 H (N = 13 very aggressivetoward other cavity-nestingbirds of 91 observedcopulations). Copulationswere also (e.g., Madagascar Starlings [Hartlaubius aurata] frequent during the 0800-1000 H (N = 28) and and Broad-billed Rollers [Eurystomusglaucurus]), 1300-1500 H (N = 28). Ten daysprior to the onset and occasionallyattacked them. of incubation, the frequency of copulationsaver- Nestlings were observed wing exercising at the aged 9.1 - 0.4 (SE) times per day (N = 91 copu- cavity entrance at 3 wk of age.