j. RaptorRes. 31 (1):40-43 ¸ 1997 The Raptor ResearchFoundation, Inc.

FOOD HABITS OF THE LANNER (FALCO BIARMICUS FEI,DEGGII) IN CENTRAL ITALY

FEDERICO MORIMANDO, FRANCESCOPEZZO Dipartimentodi BiologiaEvolutiva, Gruppo di Etologiae EcologiaComportamentale, Universitddi Siena,via PA. Mattioli 4, 53100 Siena,Italy

ALESSANDRO DRAGHI ProecoGestione Fauna E AmbienteStudio Associato, via Uopini19, 53100 Siena,Italy

ABSTR•CT.--Thediet of the Lanner Falcon (Falcobiarmicus feldeggii) was studied in an area of central Italy for one yr. Diet compositiondiffered accordingto the methodologyused for data collectionwith mammals,small passerines and insectsfound more frequentlyor exclusivelyin pellets.Conversely, anal- ysisbased on pluckingsand observationsof prey taken to the nest indicatedthe diet consistedof only nonpasserineand large passerinebirds. Overall, the birdsincluding European Starling (Sturnus vulgaris), Green Woodpecker(Picus viridis), and Jay (Garrulusglandarius) were the most commonprey taken by Lanner Falcon.Mammals taken includedwood mouse (Apodemussp.), commondormouse (Musca'rdinus avellanarius)and Savi'spine vole (MicrotussaviO. I•v WOP,DS: Lanner Falcon;Falco bia,rmicu• diet; feeding ecology;Italy.

Habitos alimenticiosdel Falcobiarmicusfeldeggii en la regi6n central de Italia. RESUMEN.--Ladieta del Falcobiarmicus feldeggii fue estudiadadurante un afio en un itrea de la regitn central de Italia. Se observ6que la composicitnde la dieta variabade acuerdocon la metodologia utilzada para la recoleccitn de datos,de tal manera que se encontraronpequefias aves passerinas, mamiferose insectosmits frecuentementeo exclusivamenteen las egagr6pilas.Por el contrario, al analizarla presaen el nido y al realizar desplumamiento,se hall6 que la diem consistiafinicamente de avesno passerinasy grandespasserinas. Sobre todo avescomo Sturnusvulgaris, Picus viridis, Garrulus glandarius,fueron la presamits comfin de E biarmicusfeldeggii.Los mamiferosincluidos en la diem eran Apodemussp., Muscardinusavellanarius y Microtussavii. [Traduccitn de AgustinaLanusse]

The European subspeciesof the Lanner Falcon find perches and roostswhere food remains may (Falco biarmicusfeldeggii) is limited in its distribu- be collected for dietary studies (Massaet al. 1991, tion to southern Italy and the Balkanswith a pop- Chiavetta 1992). Apart from studiesconducted in ulation estimated to be just a few hundred pairs Sicily (Mebs 1959, Massa et al. 1991), no detailed (Gensbo11992). There are <60 pairs in the Italian research has been done so far on the food habits peninsula but, in Sicily,the population consistsof of Lanner . Herein, we present results of >80 pairs (Massa et al. 1991). In Tuscany,central an analysisof the diet of a pair of Lanner Falcons Italy, five breeding pairs are known in a 8300 km2 in central Italy based on remains collected at area, with a densityof 1 pair/1660 km2. plucking areas and the nest site and observations The biology of the Lanner Falcon in Europe is of prey delivered to the nest. poorly studied (Cramp and Simmons 1980). Most work has been done on the African STUDY AREA (Cade 1965, Maclean 1984, Goodmanand Haynes The studywas conducted in a rural area named "Crete 1989, 1992) and very few studies have been con- Senesi"in Tuscany,central Italy, at 200-300 m elevation. The climate of the area is temperate, locallysubarid, with ducted in the Mediterranean area (Mebs 1959, an averageannual temperature of 18øC.It is very hilly, Bonora and Chiavetta 1975, Massa et al. 1991). open and eroded at the basesof hills which has created Lanner falcons nest on cliffs, making it difficult to an abundanceof smallclay/sand cliffs ranging from 10-

40 MARCH 1997 L•NER FAI•CO• FOOO HABITS 41

n= 15 n= 78 n= 71 100%

80%

60% ß Passeriformes

[] Non Passeriformes

ß Mammalia 40%

:.

Observations Pellets Pluckings

Figure 1. Compositionof the diet of Lanner Falconsduring the breeding season(December-June) using three different methods:direct observationsof prey taken to the nest, pellets,and plucked remains.Numbers above the bars refer to sample size for each method.

40 m in height. These small cliffs are usedfor nestingby collected 59 pellets under perches.Altogether, we Lanner Falcons (Morimando et al. 1994). The vegetation of the area is dominated by pasturelandwith small oak- identified 15 prey items delivered to the nest, 78 woods (Quercuspubescens) interspersed with cereal crops prey items from pellets, and 71 prey items from and small patchesof olive trees (Olea europea). plucked remains (Table 1). The mean number of

METHODS prey per collecting event was 0.94 prey for obser- We observeda pair of Lanner Falconsthrough two re- vations,1.32 for pelletsand 2.73 for pluckings. productive seasonsfrom spring 1994-surnmer 1995. We Direct observationsat the nest showedthat pas- made direct observationson prey brought to the nest sefine were delivered to the nest more fre- and we collected prey remains at the basesof nest cliffs. quently than nonpasserines(Table 1). Although By watching activity patterns, we also found two main most of these birds were not identified, the main percheswhere the falconsplucked and ate their prey. From June 1994, we collected prey remains at these prey speciestaken to the nest appeared to be the perchestwice a month by searchingthe ground carefully European Starling (Sturnusvulgaris) and Blackbird under each perch. Prey were often identified in the field (Turdusmerula). The MPW of prey brought to the based on feathers and fresh remains. Remains that were not readily recognizedas well as pellets were examined nest was 121 g (SE = 38.27). During nest obser- using a dissectingmicroscope and remains were identi- vations,we twice saw the male storing small prey fied using museum specimensand identification guides in a cliff hole that was used as a cache. (Brown et al. 1987, Svensson1992). We also used taxo- Nearly identical percentagesof passerinesand nomic keys(Chaline et al. 1974) and a sampleof speci- mens collectedlocally to compare bones and skeletalre- nonpasserineswere identified from plucked re- mains. mains (Table 1). The MPW of prey based on this We calculated the % occurrence and % biomass of of analysiswas 182.01 g (SE = 216.94), con- eachprey categoryin the diet basedon eachof the three siderablylarger than that of prey taken to the nest methods. Prey biomasswas determined based on the av- erage weight of prey taxa reported in the literature (t = 2.47, df = 20; P < 0.02). Basedon this type (Cramp 1977-94, for birds;Macdonald and Barret 1993, of analysis,the main prey were Green Woodpecker for mammals). We calculated the mean prey weight (Picusviridis), Jay (Garrulus glandarius)and Euro- (MPW) using the formula: Total Biomass/TotalNumbers pean Starling. Most of the biomass in the Lanner of each prey species. diet was from nonpasserinesof which the Green P•St;I•TS Woodpeckerwas the largest and most frequently We observed the nest 16 times for a total of 96 taken species. hr. We alsocollected plucked remains26 timesand Bone fragments in pellets were small and diffi- 42 MORIMANDOET AL. VOL. 31, NO. 1

Table 1. Numbers(N), percentageof biomass(%B) and percentageof occurrence(%0) of preyitems in Lanner Falcon diet in Tuscany,Italy, accordingto collectingmethods.

PELLETS PLUCKINGS OBSERVATIONS

PREYITEM N %B %0 N %B %0 N %B %0

MAMMALS Muscardinus avellanarius I 0.38 1.28 0 0 0 0 0 0 Apodemusspp. 3 0.96 3.85 0 0 0 0 0 0 Microtus savii I 0.38 1.28 0 0 0 0 0 0 Rodentia ind. 4 0.9 5.13 0 0 0 0 0 0 Total mammals 9 2.62 11.54 0 0 0 0 0 0

BIRDS Falco tinnunculus 0 0 0 4 6.19 5.63 0 0 0 Phasianus colchicus 0 0 0 2 10.83 2.82 I 38.57 6.67 Columba livia 3 11.49 3.85 7 16.25 9.86 I 16.53 6.67 Columbapalumbus I 5.74 1.28 1 3.48 1.31 0 0 0 Streptopeliaturtur 1 19.1 1.28 3 3.48 4.23 0 0 0 Athene noctua 0 0 0 1 1.08 1.41 0 0 0 Apus apus 0 0 0 1 0.31 1.41 0 0 0 Picus viridus 11 35.10 14.10 16 30.95 22.54 0 0 0 Total nonpasserines 16 54.24 20.51 35 72.58 49.30 2 55.10 13.33 Alauda arvensis 0 0 0 2 0.57 2.82 0 0 0 Turdus merula 6 6.13 7.69 I 0.62 1.41 3 13.22 20 Turdusphilomelos 0 0 0 3 1.58 4.23 0 0 0 Garrulusglandarius 0 0 0 11 14.47 15.49 0 0 0 Pica pica 3 6.89 3.85 3 4.18 4.23 0 0 0 Crrrvus monedula 0 0 0 0 0 0 1 10.74 6.67 Sturnusvulgaris 37 28.33 47.44 11 5.11 15.49 5 16.53 33.33 Emberiza calandra 0 0 0 I 0.29 1.41 0 0 0 Anthsupratensis 0 0 0 I 0.14 1.41 0 0 0 Passerifarmesind. 7 1.79 8.97 3 0.46 4.23 4 4.41 26.67 Total passerines 53 43.14 67.95 36 27.42 50.70 13 44.90 86.67 Total birds 69 97.38 88.46 71 100 100 15 100 100

cult to recognize. Nevertheless, this form of anal- 163.01) which was smaller than the MPW found in ysisshowed that mammals also occur in the diet of pluckings (t = 2.80, df = 20; P < 0.01). the Lanner Falcon (Table 1). Mammals identified Medium-sizedpasserines (• = 98.41 g) and large were wood mouse (Apodemusspp.), common dor- nonpasserines(• = 268.70g) were the primaryspe- mouse (Muscardinusavellanarius) and Savi'spine ciesfound in pluckedremains. In pellets,mammals vole (Microtussavii). Also, some pellets were made (• = 22.71g) and mainlypasserines of smallsize (• solely of insects remains, mainly Formicidae and = 63.70g) werepresent. Small passerines (• = 62.69 small Coleopterans.A careful inspectionof pellets g) and few nonpasserinesof large size (• = 265.62 showed that most insectswere inside the gizzard g) were hakento the nest. The differencesamong remains of Green Woodpeckersand Starlings.Pas- plucked remains,pellets and direct observationsas to the three main categoriesof prey found (passer- serineswere the most frequent prey found in pel- ines, nonpasserinesand mammals),were highly sig- lets, but nonpasserinespecies accounted for slight- nificant (X2 = 13.46, df = 4, P < 0.009). ly more biomass using this method of analysis. Main prey speciesrepresented in the pelletswere DISCUSSION the European Starling, Green Woodpeckerand We concluded that use of a single method to Blackbird.The MPW in pelletswas 100.44 g (SE = determine the diet of Lanner Falcon biases the re- MARCH 1997 L•NER FALCON Foor• HABITS 43

suits. Data from plucking sites provided more in- Siena and to P. Galeotti, G. Bogliani and F. Spina for formation about diet compositionper searchingef- critical comments on the manuscript. S. Lovari made use- ful suggestion on the first draft of the paper and im- fort than other collecting methods. Pellets are also proved the English text. J. B. Buchanan and two anony- important becausethey provided more complete mous referees provided useful suggestionsand improved information on the compositionof prey,especially data presentation. that of mammals and insects. Direct observations LITERATURE CITED of prey taken to the nest appear to be unnecessary since they add little to quantitative information on BONORA, M. AND M. CHIAVETFA. 1975. Contribution l'•tude du Faucon lanier Falcobiarmicus feldeggi en It- the Lanner Falcon'sdiet. The use of plucking data alie. Nos Oiseaux 33:153-168. alone may result in overestimatesof large- and me- BROWN,R., J. FERGUSON,M. LAWRENCEAND D. LEES. dium-size birds, as generally longer and heavier 1987. Tracks and signs of the birds of Britain and feathers fall directly beneath the perches,while the Europe. Croom Helm, Bromley, London, UK• light feathers of small birds tend to disperse(e.g., tEADE,T.J. 1965. Relations between raptors and colum- in the wind). Analysisof pellets alone tends to over- biforms at a desert water hole. Wilson Bull. 77:340- estimate the amount of small passerinesand mam- 345. mals. Lanner Falconsgenerally swallowsmall birds CHALINE,J., H. BAUDVIN,O. JAMMONAND M.C. SAINTGI- almost entirely, whereas they tear flesh and feath- RONS. 1974. Les proies des rapaces. Doin, Paris, ers from larger birds. This probably accountsfor France. the high frequency of small passerine remains CHIAVETTA,M. 1992. I1 Lanario. Pages674-678 in P. Bri- chetti, P. De Franceschi and N. Baccetti [Eds.], Fauna found in pellets. We feel that it would be best to d'Italia-AVESI. Vol. I. Edizioni Calderini, Bologna,It- evaluate information obtained from all the collect- aly. ing methods to most accurately assessthe diet of CRAMP,S. [ED.]. 1977-94. The birds of the Western Pa- Lanner Falcons. learcftc, Vol. I-IX. Oxford Univ. Press, Oxford, We found the diet of Lanner Falcon in central CRAMP, S. AND K.E.L. SIMMONS [EDS.]. 1980. The birds Italy to be qualitatively different from that of Lan- of the Western Palearctic, Vol. II. Oxford Univ. Press, her Falcon in other areas. Birds comprised 88- Oxford, UK. 100% of the diet in our studyarea dependingon GENSBOL,B. 1992. Collins guide to the birds of prey of the method of diet analysisused. These results Britain and Europe with North and the Middle East. Harper Collins, London, were consistentwith those of Cramp and Simmons GOODMAN, S.M. AND C.V. HAYNES. 1989. The distribu- (1980), except we found for the first time that tion, breeding seasonand food habits of the Lanner Green Woodpecker is taken by Lanners. We also from the Eastern Sahara.Nat. Geog•Res. 5:126-131. confirmed that they eat small mammals but that 1992. The diet of the Lanner Falco biarmicus in they do not appear to eat reptiles and insectsas a hyper arid region of the Eastern Sahara.J. Arid En- reported by Massa et al. (1991). In Sicily, birds viron. 22:93-98. (90.4%), reptiles (4.1%) and mammals(2.7%) are MACDONALD, D.W. AND P. BARRET. 1993. Mammals of in the diet of Lanners (Mebs 1959), with Jackdaw Britain and Europe. Harper Collins, London, UK. (Corvus monedula),Lesser Kestrel (Falconaumanni) MACLEAN, G. L. 1984. Robert's birds of Southern Africa. and feral Pigeon (C01umbalivia) preyed upon the New Holland Publishers, London, MASSA, B., F. LO VALVO, M. SIRACUSAAND A. CIACCIO. most. Magpie (Pica pica) and Spanish Sparrow 1991. I1 lanario (Falcobiarmicus feldeggi Schlegel) in (Passerhispaniolensis) are alsopreyed upon in Sicily Italia: status,biologia e tassonomia.Naturalista siciliano (Massa et al. 1991) and small mammals and rep- 15:27-63. tiles account for only a small percentage of the diet MEBS,T. 1959. Beitrag zur Biologie des Feldeggsfalken (4% and 2.3%, respectively). Falcobiarmicus feldeggi. Vogelwelt 80:142-149. MORIMANDO, F., E PEZZO, A. DRAGHI AND G. FRATALOC- ACKNOWLEDGMENTS CHI. 1994. Prima nidificazione di Lanario Falco biar- We are grateful to G. Fratalocchi, R. Nardi and G. Cap- micusin provincia di Siena e note sulla locale distri- pelli for their help with fieldwork. We are indebted also buzione storica. Avocetta 18:157-159. to G. Manganelli and N. Baccetti for their invaluable as- sistancein identifying feathers, bones and skeleton re- SVENSSON,L. 1992. Identificationguide to Europeanpas- mains of birds. Thanks are also due to F. Cancelli for serines. Stockholm, Sweden. allowing us to get free accessto the material of the Zoo- logical Museum of the "Accademia dei Fisiocritici" in Received2 April 1996; accepted1 December 1996