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The diet of Eurasian Griffons (Gyps fulvus) in Crete

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JUNE2005 SHORTCOMMUNICATIONS 179

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Received2 November 2003; accepted 4 October 2004

j. RaptorRes.39(2):179-183
¸ 2005 The Raptor ResearchFoundation, Inc.

THE DIET OF EUP•S• GRIFFONS(GYPSFULVUS)

IN CRETE
STAVROS M. XIROUCHAKIS 1
Natural HistoryMuseumof Crete,University
of Crete,PO. Box2208, Heraklion71409, Crete,Greece

KEYWORDS:Eurasian Griffon;Gyps fulvus;pelletanalysis; Europe (Arroyo 1994). The speciesis distributedmainly

diet;,Crete. in countries bordering the Mediterranean basin and
feeds primarily on livestock carrion (Cramp and Sim-
Between a quarter and half of the global range of the mons 1980, Don•tzar 1993). In Crete, the griffon's feed-
EurasianGriffon (Gypsfulvus) populationoccurswithin ing ecologymay be more closelyrelated to pastoralism
than anywhere else in Europe. Paleontologicalfindings
i Email address:sxirouch@nhmc.uoc.gr indicate that all autochthonous-mammalspecies(apart
180 SHORT COMMUNICATIONS VOL. 39, NO. 2

from some rodents) disappearedfrom the island by the Table 1. Items found in Eurasian Griffon pellets in
end of the Pleistocene (Sondaar et al. 1996, Willemsen Crete (Greece, 2000-01).
1996), while the presence of griffons dates back to the
upper Pleistoceneand Holocene (Rich 1983). Moreover, ITEM N (PERCENT)
the introduction of sheepand goatsby farmers occurred
7000 yearsago (Cavalli-Sforza 1996,Jarman 1996). Domesticsheephair (Ovisaries) 127 (89.4)
Since the last century the main potential food sources Domesticgoat hair (Capra hircus) 22 (15.5)
for this specieson Crete consistedof domesticlivestock Rabbit hair (Oryct0lagus
cuniculus) 4 (2.8)
Beech marten hair (Martesfoina) 2 (1.4)
and one wild species(i.e., the Cretan wild goat [Capra
Bones (Caprines) 9 (6.3)
aegagn•scretica]).Given that (1) studiesof the Eurasian Gramineae spp. 61 (42.9)
Griffon's diet are completelylackingin Greece, (2) that Holm-oak leavesQuercuscoccifera 24 (16.9)
the most important griffon population in the country is Dirt 14 (9.86)
found on Crete (Handrinos 1985, Handrinos and Akri- Coleoptera (Family Dermestidae) 8 (5.6)
otis 1997), and (3) that the number of speciesthat are Olive core 1 (0.7)
availableas carrion on the island is low, this studyon the
gnffons' diet should lead to more effectivemanagement
of the species.Pellet analysisis the most common tech- EurasianGriffons are exclusivelycarrion eaters,feed-
nique for the studyof the diet of carrion-eatingspecies ing mainlyon softtissues,thus only the frequencyof the
(Hiraldo 1976, Coleman and Fraser 1987, Ceballos and presenceof prey items was estimated (namely the num-
Donfizar 1990, Thibault et al. 1993, Larraz, 1999). How- ber of pellets containing each prey speciesX 100/total
ever, an analyticalproblem arisesdue to the low occur- number of pellets;Ceballosand Donfizar 1990). The null
rence or even absenceof bonesand a high concentration hypothesisthat sheep and goat remains were found in
of hair or feathersin pellets(Houston1976). The latter the pellets in exact ratio to their proportion in livestock
numbers was tested by a chi-squaretestwith a Yates'cor-
provide information concerning the qualitative compo- rection (Zar 1996). When the remains of one or more
sition of the diet, but they do not reveal the number of animalswere found in the same pellet it was assumed
animalson which the bird hasfed upon. Thus, no quan- that they provided information on the frequencythat
utative data or biomassquantity can be estimated from griffonsfed on them (Hiraldo 1976).
suchpellets(Marti 1987). Here, I report qualitativedata
RESULTS
on the diet of the Eurasian Griffin on the island of Crete,
basedon the analysisof regurgitatedpellets. My field assistantand I collected 811 different items
METHODS among nest ruins. However, only 142 items were consid-
ered asregurgitatedmaterialand only93 (65.5%) had the
Fieldworkwasundertaken in 11 griffon coloniesat the typicalovalshapeof the pelletsproducedby raptors(Fitz-
end of the 2000 and 2001 breeding seasons.This period
was considered to be most suitable, because it offers the
her et al. 1977).The restof the itemsweretuftsof sheep
opportunity to collectpelletswhile causingminimum dis- and goat hair, which probably originatedfrom dissolved
turbanceto the birds.As nestsare progressively destroyed pellets,and were not includedin this analysis.
Pellet size
by the chicks,a relativelylarge sampleof pellets may be was rather small, with a mean diameter of 19.3 mm (SD
found among the nest material at the bottom of the = 8 mm, range = 5-50 mm), mean length of 57.7 mm
breeding cliffs. Unfortunately,collectionof pelletsbelow (SD = 24.5 mm, range = 18-138 mm), and mean width
roost sitesthat would accountfor the diet of the species of 33.5 mm (SD = 14.6 mm, range = 10-105 mm). Their
throughout the year wasnot feasible.The basesof most massranged from 0.6--15.3 g (i = 4.05 + 2.86 g) and
cliffs where roosting siteswere located were searchedin- contained 10 different constituents (Table 1).
tensivelybut did not provideany pellet material. The dominant speciesin the griffons' diet wasdomes-
Pellets were found below 14 nests in four colonies (two
in western and two in eastern Crete, which hosted 90 andtic sheep,whichwaspresentin almost90% of the pellets.
79 individualsin 2000and 2001,respectively). Theywere Domesticgoat wasthe secondmost important prey item
sun-dried in order to remove water (Coleman and Fraser (Table 1). Sheepwere representedin the pelletsmore
1987), their massmeasuredand separatedfrom bones than goats(X2 = 16.5,P < 0.001). The rabbit(Oryctolagus
and feathers. They were dissectedinto small piecesfor cuniculus)and the beech marten were the only additional
subsequent identification. The hair was identified speciesdetected in the analysis,but in a substantially
through a photonic stereoscopeLeica MZ8 (Leica Micro- smaller number of pellets. Piecesof long bones or whole
systemsLtd., Heerbrugg, Switzerland),with the aid of an vertebrawere found in 6.3% (N = 9) of the pellets,all
identification-key (Papageorgiou and Sfougaris 1989)
originatingfrom adult caprines.
and by comparisonto reference samplesfrom specimens
of local mammal species,which constitutepotentialprey Plant materialoccurredin 42.9% (N = 61) of the pel-
for griffons (i.e., domesticsheep [Ovisaries]and domes- lets and was mainly grass(i.e., Gramineae), while holm
tic goat [Capra hircus],beech marten [Martesfoina],bad- oak leaves(Quercuscocdfera) were found in 16.9% (N =
ger [Melesme/es],brown hare [Lepuseuropaeus], and the 24) of them. Fourteen pellets (9.86%) consistedexclu-
Cretan wild goat). sivelyof dirt with a few stones,one contained an olive
JUNE2005 SHORTCOMMUNICATIONS ] 81

core (0.7%) and eight (5.6%) contained cocoonsof co- The rabbit remains recorded were suspectedto have
leoptera of the family Dermestidae. been taken from a Bearded Vulture (Gypaetusbarbatus)
nest.A breeding pair had been suppliedwith dead rabbits
DISCUSSION
during the chick-rearingperiod (Xirouchakiset al. 2003)
In Europe, the griffon usuallyfeeds on cattle (Bostau- 3 km from the colonywhere the pelletscontaining rabbit
rus), equines (Equus caballus,Equus asinus), caprines hair were collected. After the Bearded Vultures' breeding
( Ovis aries, Capra hircus,Rupicaprarupicapra),red deer attempt, griffons were observedto visit the nest several
( Cervuselaphus),pigs ( Susscrofavar. domesticus), canids times and consumebones and prey remains. In other cas-
(Canisfamiliaris, Vulpesvulpes),lagomorphs(Lepuscapen- es, griffons were observed to take bone fragments from
the base of a cliff 4 m below another nest while a Bearded
s•s,Oryctolagus cuniculus),birds and insects(FernSndez
1975, Beven 1979, Cramp and Simmons1980, Gonzfilez Vulture wasincubating.This habit has alsobeen reported
et al. 1984), but it prefers large ungulateswhen theseare in Spain, where griffonsoccasionallyexplore the ossuaries
available (DonJzar 1993). Fernfindez (1975) reports that of Bearded Vultures, where the latter drop the bones to
the species'diet in southern Spain consistsmainly of do- break them (Bertran and Margalida 1997). Basedon the
mestic animals such as goats (34.9%), sheep (20.9%), present results, bone consumption seemed frequent in
donkeys (11.6%), and cows (7%); while in northern Crete, suggestingthat griffons do not suffer from calcium
Spain, sheep were found in more than 75% of pellets deficiencyassomeAfrican vulturesdo in areaswhere large
carnivores are extinct. This substance is not available in
analyzed (Marco and Garcia 1981). The resultsof the
present studydo not differ from relevantones in other large herbivores' carrion if no broken bones are found
Mediterranean regionsin the sensethat griffonsfeed op- resulting from the action of large mammalian predators
portunisticallyon the carrion of the most common spe- (Houston 1978, Richardsonet al. 1986). In contrast,grlf-
fons in Crete have been seen to break the bones of small
cies in their foraging range.
The findings of my pellet analysisprobablyreflect the ungulates easily,and thus are not dependent on mam-
availabilityof the different prey speciesand the way that malian carnivoresor other vulture specieswith powerful
grlffonsfeed on different typesof carrion. Medium sized beaks,which are absentfrom the island (e.g., Cinereous
domestic animals, such as sheep and goats,constitute the Vulture [Aegypius monachus] ).
majorityof livestock(i.e., 95.3%; National StatisticalSer- Vegetation in the griffon's diet wasprobablyconsumed
vice of Greece 1991) and the totality of the free grazing accidentally as well as the dirt, or was contained in the
livestockof Crete. They are present in the uplandsfrom ungulates' digestive tract. However, griffons have been
late April until late October, and suffer greater losses observed to eat green plants after feeding bouts, a pro-
than those that are homebred due to accidents, malnu- cessthat probably facilitates pellet formation (Hiraldo
trition, and sickness.Moreover, they are well represented 1976). Finally,the coleopteraprey that wasrecordedwas
in the pellets becausegriffons start feeding on their car- likely consumedincidentally along with hair as dormes-
cassesfrom any part of the body, thus consuminglots of tids (Dermestidae) feed on the wool of dead animals.
hair (pers. obs.). By contrast,large ungulateswere com- Overall, irrespectiveof the source of food used by grif-
pletely absentfrom the pellets. In Crete the number of fons (i.e., carrion from waste dumps or from transhu-
such animals has decreasedby 86% (National Statistical mance herds), pellet analysisshowedthat their diet was
Service of Greece, pers. comm.) in recent decades (i.e., dominated by small domesticungulates.Considering the
since 1961) and their corpsesare buried currently. Typ- low use of poisoned baitsfor vermin (due to the absence
icallyduring the 1960sand early1970sold equineswere of large predators) and the dramatic increasein livestock
left to die in remote upland areasor their carcasseswere numbers (•70%; National Statistical Service of Greece
thrown in a few wastedumps near the villages(primarily 1991) promoted by subsidypolicieswhen Greecejoined
in gorges).However,griffonsare incapableof tearing the the European Union in 1981, the future of the species
skin of these animals. Instead, griffons accessnutrition seems rather favorable. However, increased food abun-
from carcassesmainly from the natural orifices (e.g., dance does not necessarilyimply higher food availability.
mouth), which producesminimal intake of hair in their The latter is also determined by other factors such as
food (Houston 1976). stock-rearingmethods, sanitary conditions, closingdown
I attributedthe high frequencyof sheephair in the pel- of rubbish dumps, or outdoor abattoirs. In the mean-
lets to the spatialdistributionof domesticanimalsup to time, the strong dependence of griffons on pastoralism
the time that pellet collectiontook place.Pelletswere col- meansthat socio-economic factorsmay place them at risk
lected from nest material and accountfor the period that of a human-induced food shortage.
nestswere occupiedby incubatingindividualsor by parent
birds attending their young,specificallyfrom earlyJanuary LA DIETA DE GYPSFULVUS EN CRETA
to late May/early June. The majority of livestockin the
vicinityof the coloniesduringthisperiodwassheep.Small RESUMEN.--Recolectamos egagrtpilas de 14 nidos de
waste dumps that operate illegallynear these stockyards Gypsfulvus en Creta al final de las estacionesreproduc-
also provided dead biomassto the griffons. tivas de 2000 y 2001. E1 anfilisismostr6 que G. fulvus se
182 SHORT COMMUNICATIONS VOL. 39, NO. 2

ahmenta casi exclusivamentede carrofia proveniente de GONZALEZ,L.M., J.L. GONZALEZ,AND C. LLANDRES. 1984
ovejasy cabrasdomCsticas.Los equinosy bovinosestuvi- Tree-nesting colony of griffon vultures in Spain. Vul-
eron ausentesde las egagrCpilas,probablemente debido ture News 11:12-13.
al modo en que los buitres consumenlos cadgveresy a HANDRINOS, g. 1985. The status of vultures in Greece.
la escasez de estos animales como carrofia. La deficiencia Pages 103-115 in I. Newton and R.D. Chancellor
de calcio en los volantones no fue considerada como un [EDs.], Conservationstudiesin raptors. World Work-
problema porque se encontraron huesosde pequefios ing Group on Birds of Prey, Salonica, Greece.
unguladosen el 6.3% de las egagrCpilas.Se considera --AND T. AtomOTiS. 1997. The birds of Greece.
que el incremento en la abundancia de ganado de las Helm-A. & C. Black Ltd., London, U.K.
filtimas dos dCcadasha sido beneficiosopara la especie; H•RALDO,F. 1976. Diet of Black Vulture (Aegypius mona-
sin embargo, los mCtodos de cria del ganado tambiCn chus) in the Iberian Peninsula. Do•. Acta Verteb.3:19-
pueden afectar su estadode conservaciCnal largo plazo. 31.

[TraducciCndel equipo editorial] HOUSTON,D.C. 1976. Breeding of the White-backedand

ACKNOWLEDGMENTS
I thank C. Grivas for assistance in the fieldwork, Dr. I.
Robert (Natural History Museum of Paris) for assistance
w•th bone identification,A. Copland for improving the
English text; A. Margalida,V. Penteriani, and F. Sergio
JAmUAN,M.R. 1996. Human influence in the develop-
for their commentson an earlier draft of thismanuscript.
This studywas supportedby the Universityof Crete and
the Ministry of Education through the PYTHAGORAS II
research grants.
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