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Diets of Breeding Peregrine and Lanner Falcons in South Africa

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Diets of Breeding Peregrine and Lanner Falcons in South Africa j RaptorRes. 33(3):190-206 ¸ 1999 The Raptor ResearchFoundation, Inc. DIETS OF BREEDING PEREGRINE AND LANNER FALCONS IN SOUTH AFRICA ANDREWR. JENKINS PercyFitzPatrick Institute of African Ornithology, University of CapeTown, Rondebosch 7701, SouthAfrica GRAHAM M. AVERY SouthAfrican Museum, P.O. Box 61, CapeTown 8000, SouthAfrica ABSTRACT.-•Thediets of breedingPeregrine (Falcoperegrinus) and Lanner (F. biarmicus)Falcons in South Africa were determined from the analysisof prey remains collected at nest sites and through direct observationsto determinethe regionalvariation in PeregrineFalcon prey and to measurediet overlap, and the potential for competition between Peregrine Falconsand sympatriccongeners. Direct obser- vationssuggested that remainsunder-sampled small prey by about 10% and over-sampledlarge prey by about 8%. Peregrineand Lanner Falconspreyed mostlyon birds. Pigeonsand Streptopeliadoves com- prised the bulk (38-66% by frequency;68-85% by mass)of the PeregrineFalcon prey in each of three study areas. Columbidswere supplementedby starlings(mostly European Starling [Sturnusvulgaris]) on the Cape Peninsula,sandgrouse (Pterocles spp.) and swifts(Apus spp.) on the Orange River, and mousebirds(Colius spp.) in the Soutpansberg.Cape PeninsulaPeregrine Falcons had the leastdiverse diet, the narrowestfeeding niche and they took the largestproportion of juvenile birds. Peregrine Falconson the Orange River had the broadestfeeding niche and preyedmainly on 'commuter'species rather than sedentaryresidents. Lanner Falconsin the Soutpansbergtook mainlyterrestrial or cursorial species,particularly young chickens (Gallus gallus, 40%; 37%) and charadriids,but columbidswere also important.The dietsof sympatricPeregrine and Lanner Falconsoverlapped by about 35%. Peregrines Falconsconcentrated their foraging on woodland and cliff-dwellingprey, while Lanner Falconstook mainly open-countryspecies. Close-neighboring pairs of congenersdid not obviouslyaffect the food- niche parametersof either speciessuggesting that they were not activelycompeting for food. KEYWORDS: PeregrineFalcon; Falco peregrinus; LannerFalcon; Falco biarmicus; diet;,niche dimensions; competition. Dieta de Falcoperegrinus y Falco biarmicus en reproduccitn en Surifrica RESUMEN.--Lasdietas de Falcoperegrinus y Falcobiarmicus en reproduccitn en Surfifricafueron deter- minadasa partir del anilisis de restosde presasrecolectados en los sitiosde anidacitn y a travts de observacionesdirectas para determinar la variacitn regional en las presasde halctn peregrino,medir su coincidenciay la competenciapotencial entre halconesperegrinos y suscongtneres simpitricos. Las observacionesdirectas sugirieron que las presaspequefias fueron subvaloradasen un 10% y que las presasgrandes fueron sobrevaloradasen un 8%. Loshalcones peregrinos y lanariosse alimentaron mas que todo de aves.Palomas y otrasaves del gtnero Streptopeliarepresentaron la mayoria(38-66% de la frecuencia; 68-85% de la biomasa) de las presasde los halconesperegrinos en cada una de las tres ireas de estudio.Las Columbiformesfueron suplementadaspor Sturnusvulga•is en la Peninsuladel Cabo, Pteroclesspp. y Apusspp. en el Rio Orange y Coliusspp. en el Soutpansberg.Los halconespere- grinos de la Peninsuladel Cabo tuvieron la dieta menosdiversa y el nicho alimenticiomis restringido cazando avesjuveniles. Los halconesperegrinos del R/o Orange tuvieron el nicho alimenticio mils amplio y depredaron en "especiespasajeras" contrario alas sedentarias.Los halconeslanarios en el Soutpansbergse alimentaron de especiesterrestres particularmente de pollos. (Gallusgallus, 40%; 37%) y Charadriformes,las Columbiformes fueron tambitn importantes.Las dietas de loshalcones peregrinos y lanariossimpitricos coincidi6 en un 35%. Los halconesperegrinos concetraron su forrajeo en presas de bosquesy riscosmientras que los halconeslanarios simpitricos coincidi6 en un 35%. Loshalcones peregrinosconcentraron su forrajeo en presasde bosquesy riscos,mientras que los halconeslanarios en presasde espaciosabiertos. La cercania de parejas de congtneres obviamente no afect6 los pari- metrosdel nicho alimenticiode ninguna de las dos especieslo que sugiereque estasdos especiesno estan compitiendo por cornida. 190 SEPTEMBER 1999 PEREGRINE AND LANNER DIETS IN SOUTH AFRICA 191 [Traducci6n de Cfsar M/trquez] Food habits of the Peregrine Falcon (Falcopere- northwest to southeast, and temperatures are mild to warm. grinus)have been describedfor arctic and temper- Prey remains were collected from falcon nest ledges ate regions (Mearns 1983, Hunter et al. 1988, Rat- and from below roost sitesand feeding perches.Uneaten cliffe 1993, Rosenfield et al. 1995), and for remains, regurgitated pellets and plucked feathers were southwesternAustralia (Pruett-Joneset al. 1981, used in combination to minimize bias in diet analyses Marchant and Higgins 1993). However,in the trop- (Simmons et al. 1991, Oro and Tella 1995). Collections were made from just after egg laying to soon after fledg- ics, the diet is poorly known and data for African ing. The frequency of collectionsvaried between sites, populationsare limited (Mendelsohn1988). The years,and areas (Table 1). Care was taken to remove all only studiesof Lanner Falcon (E biarmicus)diets material on each collection to prevent duplication in sub- have been in the northern periphery of its range sequent samples. Pellets were broken up with tweezers and individual bone remnants were separated from the in southern Europe, eastern Sahara, and Israel feather matrix. (Massa et al. 1991, Yosef 1991, Goodman and All avian osteological material was identified using Haynes 1992). Although Peregrine Falcons share comparativeskeletons in the South African Museum and many parts of their distribution with other large additional material loaned from the Transvaal Museum, falconshaving similar resourceneeds (Cade 1982), Pretoria and the National Museum, Bloemfontein. Indi- vidual body parts were separatedaccording to taxon and few studieshave comparedPeregrine Falcon diets recorded as the number of identified specimens (NISP). with thoseof sympatriccongeners (Cade 1960, Por- The minimum number of individuals (MNI) was calcu- ter and White 1973). This paperdescribes the diets lated from the most common body part among the iden- of breeding Peregrine and Lanner Falcons in tified specimens,after accounting for paired elements South Africa, the type and diversityof prey taken (Klein and Cruz-Uribe 1984). Incompletely ossified boneswere consideredto be those of juveniles. Mammal by Peregrine Falcons in different environments, remains were identified according to cranio-dental char- defines the feeding niches of the two speciesin an acteristics. area of sympatry and assessesthe potential for Bird and mammal remains were identified to the low- competitionbetween these two congenersin terms est possibletaxonomic level. All prey were assignedto a of diet overlap. size classbased on bone size for unidentified prey and on body massdata from the literature (Brownet al. 1982, METHODS Smithers1983, Maclean 1993) when prey were identified to at least the family level. Size classeswere small (up to We conductedour studyat sitesin three areasof South sparrowsize, averageabout 20 g), small to medium (star- Africa. Sixteen pairs of Peregrine Falconswere studied ling size, averageabout 60 g), medium (dove size, aver- on the Cape Peninsula,Western Cape Province(34ø10'S, age about 130 g), medium-large (large dove size,average 18ø25'E) from 1989-95. This area had both inland and about 220 g), large (pigeon sizeand larger, averageabout coastalnest sitesdispersed in a mosaicof urban, forest, 350 g), and very large (francolin size and larger, average and heathland habitats. Altitude varied from 0-1100 m about 600 g). In biomasscalculations, unidentified prey and the climatewas temperate with locallyvariable winter were given these averagemass values. Mean body mass rainfall (400-2000 mm per year). Seven pairs of Pere- valuesfrom the literature were used for prey identified grine Falconswere also monitored annually from 1989- to species.Prey identified only to higher taxonomiclevels 95 on the lower Orange River, Northern Cape Province were assignedmass estimates based on published weights (28ø30'S,17ø00'-20ø40'E). This is an arid, hilly to moun- of similar or related forms. tainous area, sparselyvegetated except for narrow strips Arthropod, amphibian and reptile remains were iden- of riparian bush or irrigated croplandsalong the river tified at a grosslevel only. A small number of nonavian banks. Altitude varied from 30-700 m and the climate remains were consideredto be unlikely prey of falcons, washot and dry (mean annualrainfall 60-130 mm). Prey particularlyPeregrine Falcons, on the basisof sizeor hab- remains were collected at seven Peregrine Falcon nest it. For example, observationssuggest that African Pere- sites in the Soutpansberg range, Northern Province grine Falconshunt only flying prey (Hustler 1983, Tar- (23ø00'S, 29ø40'E) from 1988-95, and the diet of these boton 1984) and, therefore, are not likely to take birds was compared with that of a sympatricpopulation terrestrial mammals and reptiles. These taxa (Appendix of nine pairs of Lanner Falconsover a 3-yTperiod from 1) were excludedfrom the analyses.Some relatively com- 1991-93. Evergreen forest and moist woodland occurred plete, easilyidentifiable prey were identified in the field along the eastern foot of the Soutpansbergescarpment, and discarded. These were not included in the data and, with dry thornveldin
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