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Taphonomic signature of the Eurasian eagle-owl(Bubo bubo) on the avian accumulation of Cau delDuc (Lleida, Spain)

Goizane Alonso , Anna Rufà , Maite Arilla & Ruth Blasco

To cite this article: Goizane Alonso , Anna Rufà , Maite Arilla & Ruth Blasco (2020) Taphonomicsignature of the Eurasian eagle-owl (Bubo�bubo) on the avian accumulation of Cau del Duc (Lleida,Spain), Historical Biology, 32:10, 1320-1333, DOI: 10.1080/08912963.2019.1587614

To link to this article: https://doi.org/10.1080/08912963.2019.1587614

Published online: 08 Mar 2019.

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ARTICLE

Taphonomic signature of the Eurasian eagle-owl (Bubo bubo) on the avianaccumulation of Cau del Duc (Lleida, Spain)Goizane Alonsoa, Anna Rufà b, Maite Arillac,d and Ruth Blasco e

aUniversidad de Burgos (UBU), Burgos, Spain; bPACEA-UMR 5199, Université de Bordeaux, Pessac Cedex, France; cIPHES, Institut Català dePaleoecologia Humana i Evolució Social, Tarragona, Spain; dÀrea de Prehistòria, Universitat Rovira i Vigili (URV), Tarragona, Spain; eCentro Nacionalde Investigación sobre la Evolución Humana (CENIEH), Burgos, Spain

ABSTRACTThe study of small prey has been the focus of interest during the past few decades, especially due to itsimplications for the subsistence and cultural behaviours of human populations. In this regard, a growingbody of evidence has shed light on the human exploitation of small prey, including birds. Nevertheless,small animal accumulations are not always a result of anthropogenic inputs, as they are important prey formany other predators (e.g. mammalian carnivores and birds of prey). As a consequence, establishing thetaphonomic pattern of each predator is a basic step towards determining the origin of faunal accumula-tions, and actualistic studies provide essential information in that respect. The present work aims tocharacterise the avian accumulations generated by the Eurasian eagle-owl for its subsequent applicationto the archaeological record. To this end, a modern avian accumulation from the small cave of Cau del Duc(Lleida, Spain) was analysed following a taphonomic approach. Our results show specific characteristicsfitting with the previous studies carried out on Eurasian-eagle-owl-made bone accumulations, as well assome inputs of mammalian carnivores indicating secondary actions in the faunal assemblage.

ARTICLE HISTORYReceived 18 October 2018Accepted 23 February 2019

KEYWORDSActualistic research; birds;Eurasian eagle-owl (Bubobubo); Cau del Duc; IberianPeninsula

Introduction

The study of faunal specimens at archaeological sites has provento be a key issue for prehistoric research, as it allows for dealingwith different facets of past human groups. For this reason, it isa field of great interest for prehistoric research. The faunalaccumulation at archaeological sites can be the consequence ofa wide variety of natural, exogenous or intrusive processes.Therefore, identifying the agents or processes that generatedeach assemblage could be a difficult task. Small animals – suchas leporids or birds – are subject to similar taphonomic histories.Their assemblages can usually be attributed to predation byraptors and/or mammalian carnivores (Núñez-Lahuerta et al.2016, 2017; Rufà et al. 2016, 2017; Arriaza et al. 2017), but also tohuman activities (Díez Fernández-Lomana et al. 1995; Fioreet al. 2004; Laroulandie 2005; Peresani et al. 2011; Cochardet al. 2012; Finlayson et al. 2012; Blasco and Fernández Peris2012a, 2012b; Blasco et al. 2013, 2014, 2016; Gabucio et al. 2014;Laroulandie et al. 2016; Rufà et al. 2018) or natural deaths(Oliver and Graham 1994; Laroulandie 2000; Serjeantson 2009;Bovy et al. 2016). In that sense, it is common to find assemblageswith a combination of anthropic and non-anthropic inputs(Binford 1981; Hockett and Haws 2002; Sanchis Serra andFernández Peris 2008; Lloveras et al. 2010, 2011; Rufà et al.2014; Laroulandie et al. 2016). For that reason, it is essential tocharacterise the taphonomic pattern in each case.

Regarding avian specimens, both archaeological and actualisticstudies have focused their attention on how to distinguish anthro-pic accumulations (Laroulandie 2000; Laroulandie et al. 2008;Blasco and Fernández Peris 2009, 2012b; Blasco et al. 2013;

Laroulandie and Lefèvre 2014; Radovčić et al. 2015; Romandiniet al. 2016) from those produced by different birds of prey ormammalian carnivores (Bochenski et al. 1993, 1997; Bochenskiet al. 1999, 2009; Bochenski and Tomek 1994, 1997; Bochenski1997; Laroulandie 2000, 2002, 2010; Bochenski and Nekrasov2001; Bochenski and Tornberg 2003; Mallye et al. 2008;Monchot and Gendron 2011; Lloveras et al. 2014a, 2014b;Laroulandie et al. 2016; Rodríguez-Hidalgo et al. 2016). Most ofthose studies have assessed the basic features of each taphonomicpattern, such as anatomical profiles, fragmentation degree andmodifications over the bone surface. In the particular case of theEurasian eagle-owl (Bubo bubo), the knowledge about the maintaphonomic traits that this raptor produces through avian remainsis limited (Bochenski et al. 1993; Laroulandie 2002; De Cupereet al. 2009). The Eurasian eagle-owl is a bird widely distributedover Europe, Asia and the north of Africa and it is the largestnocturnal bird of prey existing inEurope. It inhabits a great varietyof ecosystems: boreal conifer forests, mixed deciduous forests,Mediterranean scrubs, steppes and deserts (Mikkola 1994;Penteriani et al. 2002). Although young individuals can makedispersive movements of about 200 kilometres, it is a territorialand eminently sedentary animal. It tends to nest in fissures, holesor rocky ridges of themountains, but also in small caves protectedby scrubs and in tree trunks (Mikkola 1994). Its diet is based on theconsumption of small rodents and birds, together with largeranimals such as rabbits, hares and small carnivores. However, itis also able to prey upon reptiles, fishes, frogs, insects and evensmall ungulates. One of the main types of prey of the Eurasianeagle-owl in areas where it is abundant – such as the IberianPeninsula and southern France – is the European rabbit

CONTACT Goizane Alonso goizaneac@gmail.com Universidad de Burgos (UBU), Calle Don Juan de Austria 1, 09001 Burgos, Spain

HISTORICAL BIOLOGY2020, VOL. 32, NO. 10, 1320–1333https://doi.org/10.1080/08912963.2019.1587614

© 2019 Informa UK Limited, trading as Taylor & Francis Group

Published online 08 Mar 2019

(Oryctolagus cuniculus) (Pérez Mellado 1977; Donázar 1989;Serrano 1998; Zarco et al. 2016). The Eurasian eagle-owl swallowsthe prey whole if its size is small enough. In the case of largeranimals– like adult rabbits or large birds – it separates the skeletoninto smaller pieces and it does not consume the entirety of theremains (Yravedra 2006; Lloveras et al. 2009). Afterwards, theingested remains are discarded through the regurgitation of pel-lets, which are mixes of fur, feathers and the bones of the ingestedprey. Those elements are discarded in the nests or in their vicinity,leading to considerable osseous accumulations over time (Hockett1995; Lloveras et al. 2009) and mixing with other discarded non-ingested remains accumulated close to the nest. Thus, the assem-blages generated by this raptor are usually a combination ofingested and non-ingested remains (Hockett 1995; Cochard2008; Lloveras et al. 2009). The study of taphonomic bone patternsgenerated by the Eurasian eagle-owl is of interest for the inter-pretation of the fossil faunal assemblages, because it wasa common inhabitant of European Pleistocene archaeologicalsites. This research attempts to contribute to the characterisationof the Eurasian eagle-owl’s avian accumulations using a modernbird assemblage generated by this raptor in the Cau del Duc cave(Lleida, Spain) and discern, whenever possible, its taphonomicsignature from those generated by mammalian carnivores andother birds of prey.

Materials and methods

Study area: Cau del Duc

Cau del Duc is a small cave located in the northeast part ofthe Iberian Peninsula, in the province of Lleida (Catalonia,Spain), at 819 metres above sea level (Figure 1). This

region is characterised by a temperate climate (Rivas-Martínez et al. 2002). However, due to the proximity to thePyrenees, the average annual temperature oscillatesbetween 10ºC and 12.5ºC, and the average annual precipi-tation is 600–800 mm (Ninyerola et al. 2005). The entranceto the cave is located on the upper part of a limestone cliff(Figure 2), ~115 metres above the river Flamisell. The cavehas a tubular shape and is nine metres in length from theentrance to the innermost zone. Coinciding with this innerarea is a small chamber to the west (called a ‘covacha’),giving the cave an L-shaped morphology. The presence ofthe Eurasian eagle-owl in proximity to the cave has beendocumented for years by the forestry people in the area,and it is considered to be the main accumulator of bones.The faunal sample presented here was recovered inJuly 2015. In order to accurately control the density ofthe material, bones were taken according to their locationin the cave. Samples were recovered at one-metre intervalsfrom the entrance to the innermost area (the back of thecave) in separate plastic bags. Each sample contained seedsand bones of micro- and macro-fauna. The majority of themacro-faunal accumulation was composed of leporids,including rabbits and hares, followed by avian bones.Other than that, a small portion of the macro-faunal sam-ple comprised bones from various taxa, consisting ofa hemi-mandible from an immature wild boar (Sus scrofa),a tooth from a modern human (Homo sapiens) and somelong bones from an unidentified ovicaprid species. Themicro-faunal bones included rodents and reptiles, amongothers. Once brought to the laboratory, a sample thatinvolved 1912 avian bones was selected for the presentstudy.

Figure 1. Location of Cau del Duc (red dot) in the Iberian Peninsula.

Figure 2. Limestone cliff (a) and entrance to the cave (b).

HISTORICAL BIOLOGY 1321

Anatomical and taxonomical analysis

The avian sample was taxonomically and anatomically classifiedusing reference collections and anatomy-specialised bird atlases(e.g. Woelfle 1967; Kraft 1972; Janossy 1983; Moreno 1985;Cohen and Serjeantson 1996; Tomek and Bocheński 2000,2009; Bochenski and Tomek 2009). In cases where specificassignment to a species category was not possible, the specimenswere classified according to the genus (e.g.Alectoris), family (e.g.Phasianidae) or order (e.g. Galliformes). Bones that could not beclassified were categorised as ‘indeterminate’. Several estimationindexes were calculated to assess the completeness of the sample:number of specimens (NSP), minimum number of elements(MNE) and minimum number of individuals (MNI) (Lyman1994). The NSP refers to all recovered specimens, includinganatomically and taxonomically identifiable bone fragments.In order to compare the number of recovered bones in relationto the expected number of bones to be recovered, the relativeabundance (RA) index was determined, which is shown asa percentage (Dodson and Wexlar 1979; Lyman 1994). In addi-tion to these indexes, some ratios were calculated to evaluate theproportion of wing elements with respect to leg elements(Ericson 1987), the proportion of core elements with respect tolimb elements and the proportion of proximal elements withrespect to distal elements (Bramwell et al. 1987; Bochenski andNekrasov 2001). Those ratios were used on studies alreadycarried out by other authors (e.g. Bochenski 2005; Laroulandie2010; Lloveras et al. 2014a, 2014b). Although core-to-limb andproximal-to-distal ratios do not show an equal proportion ofelements, we have applied the same equation as the aforemen-tioned authors in order to be able to make accurate compar-isons. All of the indexes were calculated based on the MNE andshown as a percentage.

● Wing-to-leg ratio. The total number of wing elements(humeri, ulnae, carpometacarpi) was divided by the sumof wing and leg elements (femora, tibiotarsi,tarsometatarsi).

● Core-to-limb ratio. The total number of core elements(sternums, coracoids, scapulae, pelvises) was divided bythe sum of core and limb elements (humeri, femora,radii, ulnae, tibiotarsi, carpometacarpi, tarsometatarsi).

● Proximal-to-distal ratio. The total number of proximalelements (scapulae, coracoids, humeri, femora, tibio-tarsi) was divided by the sum of proximal and distalelements (ulnae, radii, carpometacarpi, tarsometatarsi).

Taphonomic analysis

FragmentationIn order to determine the fragmentation degree, differentskeletal categories were established (Bochenski et al. 1993);for long bones, phalanges, vertebrae and ribs, the classifica-tion proposed by Lloveras et al. (2014b) was applied. Thefragmentation degree was calculated based on the NSP andshown as a percentage.

● Long bones (humeri, radii, ulnae, carpometacarpi,femora, tibiotarsi, tarsometatarsi) were classified as

complete, proximal joint, proximal joint and shaft,shaft, shaft and distal joint or distal joint.

● Crania were classified as whole skull with beak, craniawith beak and brain case without back part, brain casewithout back part, brain case, complete beak or frag-mented beak.

● Mandibles were classified as whole, one branch, articularpart, distal part or medium portion of the branch.

● Scapulae were classified as complete or acromial process.● Coracoids were classified as complete, proximal frag-

ment or distal fragment.● Pelvises were classified as synsacrum with one or both

ilio-isquio-pubic bones, ilio-isquio-pubic bone, synsa-crum (partial or complete) or acetabulum.

● Sternums were classified as more than one-half withrostrum or less than one-half with rostrum.

● Ribs were classified as complete or fragmented.● Vertebrae were classified as complete, vertebral body,

vertebral epiphysis or spinous process.● Phalanges were classified as complete, proximal frag-

ment or distal fragment.

Likewise, the bone completeness was assessed according tothree degrees (Bochenski 2005): high (>60% complete), mod-erate (30–60% complete) and low (<30% complete). Forquantifying the proportion of complete bones, the followingequation modified from Laroulandie (2000) was applied:number of complete bones × 100/total NSP.

Digestion and beak/talon marksThe avian sample was analysed using an Olympus SZX7 stereo-microscope. Mechanical marks produced by the beak or talonswere defined as grooves with a flat or rounded bottom (Cáceres2002); even though the predominant marks are punctures(Bochenski and Tornberg 2003; Bochenski et al. 2009). In orderto characterise the digestion damage, the range of five corrosiondegrees established by Andrews (1990) was considered: absent(0), light (1), moderate (2), strong (3) and extreme (4).

Post-depositional modificationsPost-depositional modifications, including weathering, calcitecoating, manganese oxide deposits and chemical corrosion,were also documented in the avian sample. Following thecriteria proposed by Behrensmeyer (1978) andBehrensmeyer et al. (2003), weathering damage was classifiedinto six stages: stage 0) no signs of alteration; stage 1) crack-ing; stage 2) flaking at the thinner layers; stage 3) the bonesurface is homogeneously weathered; stage 4) the bone sur-face is homogeneously weathered and bears small splinters;and stage 5) the bone is completely broken, fragile and withlarge splinters (Behrensmeyer 1978; Behrensmeyer et al. 2003;Cruz 2008). Calcite coating and manganese oxide depositslinked to humidity were also recorded. The manganeseappears in the sample as black stains on the bone surface inisolated, clustered or dispersed patterns. Additionally, thepresence or absence of chemical corrosion was documented,which implies a loss of soluble material and punctual colour-ing changes due to leaching processes in the cave. Eventually,root etching was documented in the form of branched

1322 G. ALONSO ET AL.

grooves varying in length, width and depth (Behrensmeyer1978; Fernández-Jalvo 1992; Blasco 2011; Fernández-Jalvoand Andrews 2016).

Results

Taxonomical and anatomical representation

The analysed sample involved a total of 1912 avian specimenscorresponding to an MNI of 177. The highest density of boneswas found in the middle area of the cave (at 3, 4, 5 and 6metres),whereas both the entrance and the innermost areas showeda lower quantity. Phasianidae and Columbidae were the mostabundant groups (NSP = 190 and NSP = 222, respectively). ThePhasianidae groupmainly consisted ofAlectoris rufa (NSP = 31).The third-largest group, Corvidae (NSP = 117), was composedof a wide variety of species. The families with the lowest numberof bones were Phalacrocoracidae and Turdidae, represented byjust one specimen each. Aside from that, the majority of speci-mens (NSP = 1115) could not be attributed to any taxonomicalcategory, so they were classified as indeterminate (Table 1).Adult individuals were predominant, representing 100% of thesamples taken from most sections of the cave. Only 84 bones ofimmature individuals were recovered, which corresponded toan MNI of 9. Although these specimens were mainly foundwithin the indeterminate group, some of them belonged to thePhasianidae and Columbidae families and to the Anatinae sub-family (Table 1). A predominance of post-cranial elements overcranial elements was observed. Nevertheless, all skeletal ele-ments were represented to a greater or lesser degree.Generally, posterior phalanges and humeri dominated theassemblage, although coracoids, vertebrae, ulnae, carpometa-carpi and elements of lower limbs (femora, tibiotarsi and tar-sometatarsi) were also well represented. Contrarily, cranial andcore elements showed the lowest values. There were no differ-ences when comparing the anatomical elements found at differ-ent depths within the cave (Table 2). Considering the RApercentage, the best-preserved specimens were coracoids andposterior phalanges, followed by humeri, carpometacarpi andtarsometatarsi. Mandibles, furculae, vertebrae and ribs showedthe lowest representation, indicating a significant loss of theseskeletal elements (Figure 3).

Regarding the skeletal profile, a higher proportion of upperlimb bones was observed in the wing-to-leg ratio (56.1%), as theresulting percentage surpassed 50%. The core-to-limb ratioshowed a predominance of limb bones over core elements inthe whole assemblage. This can be observed in the percentageobtained (18%), which showed that core elements only repre-sented a small proportion of elements in comparison to limbelements. In addition, the percentage for the proximal-to-distalratio was 55.7%, since a higher number of proximal elementswas observed throughout most of the cave (Table 3).

Taphonomic modifications

FragmentationCompleteness was moderate, as complete bones were recov-ered in 40.2% of the avian sample, especially regarding ver-tebrae and phalanges. The most fragmented anatomical

elements were the cranial and core bones. In some cases, itwas possible to find complete scapulae, coracoids or ribs,whereas the cranium, pelvis and sternum always appearedfragmented (Table 4). Limb bones were highly fragmentedwith only 9.5% completeness (both proximal and distal endswere the most preserved portions, comprising 94.6% of thelimb bones). Among complete limbs, tarsometatarsi and car-pometacarpi were the least fragmented bones (29.7% and31.3% of complete elements, respectively). In the case ofcarpometacarpi, about half of the elements (51.6%) lackedthe most fragile part (minor metacarpal). Finally, the percen-tage of complete radii and ulnae was between 8% and 20%(radii = 8.8%; ulnae = 16.5%), whereas in the case of thehumeri, femora and tibiotarsi, the proportion narrowlyexceeded 3% (7.1%, 3.8% and 2.7%, respectively).

Mechanical modifications and digestion marksNo triangular punctures typically associated with beak or talonmarks were identified in the avian sample. However, several pitswere found, occasionally related to crenulated edges, whichcould be linked to secondary accesses of small mammaliancarnivores (Figure 4). These tooth marks were detected in0.14% of the whole assemblage, and in no case reached morethan 5 mm in size (width and length). In most cases (62.5%),they were found on the proximal ends of humeri.

Table 1. NSP (%), MNE and MNI values of the avian specimens. Note thatimmature individuals are marked with ‘(i)’.

NSP NSP % MNE MNI

PhasianidaeAlectoris rufa 31 1.62 31 8 + 1(i)Perdix perdix 7 0.31 7 2Tetrao urogallus 3 0.15 3 1Alectoris rufa/Perdix perdix 140 7.32 105 14Phasianidae indet. 9 0.47 9 3AnatidaeAnas plathyrynchos 1 0.05 1 1Anas querquedula 4 0.21 4 2Anatinae 71 3.71 60 13 + 1(i)Anatidae indet. 3 0.15 3 2ColumbidaeColumba livia f. domestica 1 0.05 1 1Columba palumbus 2 0.10 2 1Columba livia/Columba oenas 1 0.05 1 1Columbidae indet. 218 11.40 160 25 + 1(i)CorvidaeCorvus monedula 7 0.36 7 2Garrulus glandarius 2 0.10 2 1Pica pica 2 0.10 2 1Pyrrhocorax pyrrhocorax 3 0.15 3 1Corvus corone/Corvus frugilegus 2 0.10 2 1Corvus monedula/Pyrrhocorax graculus 2 0.10 2 1Corvus monedula/Pyrrhocorax pyrrhocorax 1 0.05 1 1Garrulus glandarius/Pica pica 5 0.26 5 2Corvidae indet. 93 4.86 77 12Falconidae 5 0.26 5 2Phalacrocoracidae 1 0.05 1 1Rallidae 28 1.46 26 4Scolopacidae 78 4.08 63 14Strigidae 5 0.26 5 2Turdidae 1 0.05 1 1Falconidae/Accipitridae 1 0.05 1 1Falconiformes 5 0.26 5 2Passeriformes 60 3.13 50 11Small passeriformes 5 0.26 5 2Indeterminate 1115 58.31 345 32 + 6(i)Total 1912 100 992 177

HISTORICAL BIOLOGY 1323

Table2.

NSP

(%)andMNEby

leng

thof

thecave

(metres)(Cmc=carpom

etacarpu

s,Wingph

g=wingph

alange,Tib

=tib

iotarsus,Tmt=tarsom

etatarsus,Posteriorph

g=po

steriorph

alange,Ind

et.=

indeterm

inate).

1m2m

3m4m

5m6m

7m8m

9mCo

vacha

Total

NSP

(%)

MNE

NSP

(%)

MNE

NSP

(%)

MNE

NSP

(%)

MNE

NSP

(%)

MNE

NSP

(%)

MNE

NSP

(%)

MNE

NSP

(%)

MNE

NSP

(%)

MNE

NSP

(%)

MNE

NPS

(%)

MNE

Skeletal

elem

ent

Beak

--

--

1(0.46)

14(0.68)

43(0.49)

34(1.10)

4-

-2(5)

2-

-1(2.27)

115

(0.78)

14Mandible

--

--

1(0.46)

15(0.86)

5-

--

--

--

--

--

-6(0.31)

6Furcula

--

--

1(0.46)

1-

-2(0.32)

21(0.27)

1-

-1(2.5)

1-

--

-5(0.26)

5Co

racoid

--

3(8.33)

319

(8.79)

1644

(7.58)

3826

(4.26)

2413

(3.59)

12-

-3(7.5)

3-

-4(9.09)

4112(5.87)

100

Scapula

--

1(2.77)

15(2.31)

59(1.55)

93(0.49)

31(0.27)

1-

--

-2(10.52)

23(6.81)

324

(1.25)

24Vertebra

--

--

26(12.03)

2629

(5)

2934

(5.57)

3418

(4.97)

18-

--

--

-2(4.54)

2109(5.70)

109

Rib

--

--

--

--

--

--

--

1(2.5)

1-

-1(2.27)

12(0.10)

2Sternu

m-

-1(2.77)

12(0.92)

22(0.34)

26(0.98)

65(1.38)

5-

--

-1(5.26)

12(4.54)

219

(0.99)

19Hum

erus

--

2(5.55)

123

(10.64)

1961

(10.51)

4056

(9.18)

3782

(22.65)

611(25)

14(10)

24(21.05)

44(9.09)

4237(12.39)

169

Radius

--

3(8.33)

38(3.70)

613

(2.24)

1019

(3.11)

159(2.48)

5-

--

-3(15.78)

35(11.36)

460

(3.13)

46Ulna

1(100)

11(2.77)

116

(7.40)

1348

(8.27)

3039

(6.39)

2651

(14.08)

39-

-5(12.5)

5-

-5(11.36)

5166(8.68)

120

Cmc

--

3(8.33)

310

(4.63)

936

(6.20)

3242

(6.88)

3623

(6.35)

212(50)

22(5)

23(15.78)

35(11.36)

5126(6.58)

113

Wingph

g-

--

-4(1.85)

415

(2.58)

1412

(1.96)

129(2.48)

7-

--

-2(10.52)

21(2.27)

143

(2.24)

40Pelvis

--

2(5.55)

25(2.31)

54(0.68)

43(0.49)

37(1.93)

7-

-1(2.5)

1-

--

-22

(1.17)

22Femur

--

2(5.55)

110

(4.63)

950

(8.62)

3050

(8.19)

3632

(8.84)

23-

-5(12.5)

41(5.26)

13(6.81)

3153(8)

107

Tib

--

3(8.33)

213

(6.01)

1051

(8.79)

3539

(6.39)

2630

(8.28)

19-

-3(7.5)

2-

-2(4.54)

1141(7.37)

95Tm

t-

-2(5.55)

212

(5.55)

1136

(6.20)

2748

(7.86)

3731

(8.56)

301(25)

12(5)

2-

-3(6.81)

3135(7.06)

113

Posteriorph

g-

-13

(36.11)

1343

(19.90)

43161(27.75)

158

214(35.08)

209

37(10.22)

37-

-11

(27.5)

11-

--

-479(25.05)

471

Indet.

--

--

17(7.87)

1712

(2.06)

1214

(2.29)

149(2.48)

9-

--

-3(15.78)

33(6.81)

358

(3.03)

58Total

1(100)

136

(100)

33216(100)

198

580(100)

498

610(100)

554

362(100)

299

4(100)

440

(100)

3619

(100)

1944

(100)

421912

(100)

1633

1324 G. ALONSO ET AL.

Digestive damage (Figure 4) was documented on 790bones (41.3%) with a predominance of light and moderatedegrees of damage (79.6% and 16.7% respectively) (Table 5).The majority of digested bones came from the middle area ofthe cave, but it should also be noted that the highest numberof bones came precisely from this part of the cave. There wasno difference regarding the digestion percentage when com-paring the distinct sampled sections of the cave, as more than15.8% of the bones were affected throughout. It is worthmentioning that none of the specimens showed an extremedegree of digestion, and only 3.7% of the affected bones (1.5%of the whole sample) presented strong alteration (Table 5).The most affected bones were the humeri, followed by bothupper (ulnae, radii and carpometacarpi) and lower (femora,tibiotarsi and tarsometararsi) limbs. Regarding core and cra-nial elements, the digestion damage did not exceed 10.3%(Table 6). Proximal and distal ends of the bones were espe-cially damaged.

Post-depositional modificationsPost-depositional modifications were documented in samplesfrom all sections of the cave. The most common modificationwas calcite coating, which was registered on more than half(58.3%) of damaged bones. In most cases, the deposits cov-ered nearly the whole surface of the bones (81.7%). Fissuresand longitudinal cracks affected 31.6% of specimens.Manganese oxide deposits, chemical corrosions and rootetching (Figure 5) were observed on less than 5% of theaffected bones (Table 7). Manganese oxides were more fre-quent at the innermost part of the cave, as it is the mosthumid area. In fact, 73.5% of bones bearing this alterationwere recovered from the five-metre point to the ‘covacha’chamber.

Discussion

Based on the obtained results, it can be confirmed that thesample of avian bones recovered at Cau del Duc fits well witha typical nest accumulation of the Eurasian eagle-owl(Bochenski et al. 1993; Bochenski and Tomek 1994;Laroulandie 2000, 2002; Bochenski and Nekrasov 2001).

Nevertheless, the intervention of different predators cannotbe ruled out, since some tooth marks have also been recog-nised. In fact, based on the pit sizes and their morphology(Laroulandie 2000; Mallye et al. 2008; Monchot and Gendron2011; Rodríguez-Hidalgo et al. 2016), we could suggest theoccasional presence of small carnivores in the cave. Thus, it isimportant to clarify some issues mentioned by other authorsto make accurate inferences about the Cau del Duc birdaccumulation.

At the taxonomical level, in the analysed sample,a predominance of Phasianidae was noticed, and Alectorisrufa particularly stood out. However, as the most abundanttype of prey in a specific region can condition the diet of eachpredator (Donázar 1987; Donázar et al. 1989; Iezekiel et al.2004; Ontiveros et al. 2005; Lozano et al. 2006; Moleón et al.2009; Lloveras et al. 2014b), it is common to find bones ofAlectoris rufa in places where there is a high proportion ofphasianids. In addition, it has to be considered that most of thetaphonomic studies done on accumulations of avian bonesproduced by the Eurasian eagle-owl do not describe the taxarepresented. Accordingly, the data for the taxonomical profileon accumulations generated by this predator are still scarce.Therefore, the taxonomical representation might not always bea characteristic feature of the accumulation generated by eachpredator. For example, the Phasianidae family is abundant inaccumulations generated by Bonelli’s eagle (Aquila fasciata),and the main species represented is Alectoris rufa (Lloveraset al. 2014b). In some studies on avian assemblages producedby the golden eagle (Aquila chrysaetos), the Phasianidae familyis also abundant; nevertheless, Tetrao and Lagopus are themain genera represented (Bochenski et al. 1999). This can beexplained by the fact that the study done by these researcherswas performed on golden eagle accumulations in Finland (thecircumpolar area), a region where both genera (Tetrao andLagopus) are well adapted (BirdLife International 2018).Furthermore, Lagopus represents the majority of non-ingested bones left by the gyrfalcon (Falco rusticolus)(Bochenski and Tornberg 2003). On the other hand, avianaccumulations produced by the eastern imperial eagle(Aquila heliaca) are characterised by Corvidae with represen-tation percentages of ~70% (Bochenski et al. 1997); while theVerreaux’s eagle’s (Aquila verreauxii) accumulations show

Figure 3. The RA percentage of the different skeletal portions for avian specimens (Cmc = carpometacarpus, Wing phg = wing phalange, Tib = tibiotarsus,Tmt = tarsometatarsus, Posterior phg = posterior phalange).

HISTORICAL BIOLOGY 1325

Table3.

MNEvalues

forwing,

leg,

core,limb,

proximalanddistalelem

ents

andtheirratio

s,expressedas

apercentage,aton

e-metre

intervalsof

thecave.

1m2m

3m4m

5m6m

7m8m

9mCo

vacha

Total

TotalN

SP1

36216

580

610

362

440

1944

1912

Wing/leg

wing>

leg

wing=leg

wing>

leg

wing>

leg

wing=leg

wing>

leg

wing>

leg

wing>

leg

wing>

leg

wing>

leg

wing>

leg

Wing

15

41102

99121

39

714

402

Leg

05

3092

9972

18

17

315

Ratio

100

5057.74

52.57

5062.69

7552.94

87.5

66.66

56.06

Core/limb

core<lim

bcore<lim

bcore<lim

bcore<lim

bcore<lim

bcore<lim

bcore<lim

bcore<lim

bcore<lim

bcore<lim

bcore<lim

bCo

re0

728

5336

280

43

7166

Limb

013

77204

213

198

117

1125

759

Ratio

035

26.66

20.62

14.45

12.38

019.04

21.42

21.87

17.94

Proximal/distal

proximal<distal

proximal<distal

proximal>distal

proximal>distal

proximal>distal

proximal>distal

proximal<distal

proximal>distal

proximal>distal

proximal<distal

proximal>distal

Proximal

08

59152

126

116

111

715

495

Distal

09

4299

114

953

96

17394

Ratio

047.05

58.41

60.55

52.5

54.20

2555

53.84

46.87

55.68

1326 G. ALONSO ET AL.

Table 4. Number and percentage (%) of complete bones at the different metre intervals (length) of the cave. These percentages were calculated considering thetotal number of specimens recovered within each metre.

1m 2m 3m 4m 5m 6m 7m 8m 9m Covacha Total

Total NSP 1 36 216 580 610 362 4 40 19 44 1912Complete (%) 1 (100) 17 (47.22) 90 (41.66) 224 (38.62) 297 (48.68) 108 (29.83) 3 (75) 15 (37.5) 3 (15.78) 10 (22.72) 768 (40.16)Limb bones 1 (100) 2 (11.76) 20 (22.22) 35 (15.62) 54 (18.18) 55 (50.92) 3 (100) 3 (20) 2 (66.66) 9 (90) 182 (23.69)Crania - - - - - - - - - - -Coracoids - 2 (11.76) 3 (3.33) 7 (3.12) 3 (1.01) 1 (0.92) - - - - 15 (1.95)Scapulae - - - 1 (0.44) - 1 (0.92) - - - - 2 (0.26)Sternum - - - - - - - - - - -Ribs - - - - - - - 1 (6.66)Pelvises - - - - - - - - - - -Vertebrae - - 24 (26.66) 20 (8.92) 27 (9.09) 10 (9.25) - - - - 81 (10.54)Phalanges - 13 (76.47) 43 (47.77) 161 (71.87) 213 (71.71) 41 (37.96) - 11 (73.33) 1 (33.33) 1 (10) 484 (63.02)

Figure 4. Different degrees of digestive damage to avian bones (a1: light degree to a distal left femur of a Phasianidae; a2: moderate degree to a distal right femurof an indeterminate species; a3: strong degree to a distal right femur of an indeterminate species) and modifications (pits) produced by small mammalian carnivores(b).

Table 5. Number and percentage (%) of specimens showing digestive damage by metre intervals of the cave.

1m 2m 3m 4m 5m 6m 7m 8m 9m Covacha Total

Total NSP 1 36 216 580 610 362 4 40 19 44 1912Digestive damage (%) - 19 (52.77) 79 (36.57) 285 (49.13) 254 (41.63) 121 (33.42) 3 (75) 17 (42.5) 3 (15.78) 9 (20.45) 790 (41.31)Light (%) - 13 (68.42) 59 (74.68) 228 (80) 198 (77.95) 102 (84.29) 3 (100) 15 (88.23) 3 (100) 8 (88.88) 629 (79.62)Moderate (%) - 3 (15.78) 17 (21.51) 48 (16.84) 45 (17.71) 16 (13.22) - 2 (11.76) - 1 (11.11) 132 (16.70)Strong (%) - 3 (15.78) 3 (3.79) 9 (3.15) 11 (4.33) 3 (2.47) - - - - 29 (3.67)

Table 6. Number and percentage (%) of bones showing digestive damage by metre intervals. These percentages were calculated considering the total number ofdigested specimens recovered within each metre.

1m 2m 3m 4m 5m 6m 7m 8m 9m Covacha Total

Total NSP 1 36 216 580 610 362 4 40 19 44 1912Digestive damage (%) - 19 (52.77) 79 (36.57) 285 (49.13) 254 (41.63) 121 (33.42) 3 (75) 17 (42.5) 3 (15.78) 9 (20.45) 790 (41.31)Beak - - - - - - - - - - -Mandible - - - - - - - - - - -Furcula - - - - - - - - - - -Coracoid - 2 (10.52) 4 (5.06) 23 (8.07) 20 (7.87) 6 (4.95) - 2 (11.76) - - 57 (7.21)Scapula - 1 (5.26) 1 (1.26) 3 (1.05) 1 (0.39) - - - - - 6 (0.75)Vertebrae - - 1 (1.26) 2 (0.70) - - - - - - 3 (0.37)Ribs - - - - - - - - - - -Sternum - 1 (5.26) 2 (2.53) 2 (0.70) 2 (0.78) - - - - - 7 (0.88)Humerus - 2 (10.52) 19 (24.05) 56 (19.64) 41 (16.14) 43 (35.53) 1 (33.33) 2 (11.76) 1 (33.33) 2 (22.22) 167 (21.13)Radius - 1 (5.26) 3 (3.79) 11 (3.85) 10 (3.93) 2 (1.65) - - - 1 (11.11) 28 (3.54)Ulna - 1 (5.26) 8 (10.12) 37 (12.98) 30 (11.81) 19 (15.70) - 4 (23.52) 1 (33.33) 1 (11.11) 101 (12.78)Carpometacarpus - 3 (15.78) 3 (3.79) 22 (7.71) 28 (11.02) 9 (7.43) 2 (66.66) - - 2 (22.22) 69 (8.73)Wing phx - - - 1 (0.35) 3 (1.18) - - - - - 4 (0.50)Pelvis - 2 (10.52) 4 (5.06) 3 (1.05) - - - - - - 9 (1.13)Femur - 2 (10.52) 8 (10.12) 48 (16.84) 45 (17.71) 17 (14.04) - 4 (23.52) 1 (33.33) 2 (22.22) 127 (16.07)Tibiotarsus - 3 (15.78) 9 (11.39) 48 (16.84) 31 (12.20) 17 (14.04) - 3 (17.64) - - 111 (14.05)Tarsometatarsus - 1 (5.26) 5 (6.32) 26 (9.12) 37 (14.56) 8 (6.61) - 2 (11.76) - 1 (11.11) 80 (10.12)Posterior phx - - 4 (5.06) 3 (1.05) 2 (0.78) - - - - - 9 (1.13)Indeterminate bones - - 8 (10.12) - 4 (1.57) - - - - - 12 (1.51)

HISTORICAL BIOLOGY 1327

high proportions of pigeons (genusColumba) and birds of prey(genera Aquila and Bubo africanus) (Armstrong and Avery2014). Mammalian carnivores, such as the red fox (Vulpesvulpes) and European badger (Meles meles), can also generateavian accumulations where Corvidae, Phasianidae (Mallyeet al. 2008), Anatidae and Alcidae (Monchot and Gendron2011) families stand out. However, it should be taken intoaccount that the study done by Monchot and Gendron(2011) was carried out in an aquatic environment; thus, water-dwelling species were common. Therefore, it is important tohighlight that the dietary habits of each predator depend lar-gely on the prey availability or abundance.

Anatomical profiles constitute an important tool to deter-mine the main predator agent. In our sample, the low numberof cranial elements can be explained through the Eurasianeagle-owl’s behaviour. One of the habits of this raptor is toseparate the head of its prey before eating it (Bochenski et al.1993). Consequently, in its avian accumulations, it is com-mon to find scarce cranial elements compared to postcranialelements. This is a distinct characteristic since other owlspecies – such as the tawny owl (Strix aluco) and the snowyowl (Nyctea scandiaca) – also decapitate their prey, but theyconsume the head as well (Bochenski et al. 1993; Bochenski1997). Nevertheless, it has to be taken into account thata lower representation of cranial elements could also bea consequence of bone fragility. The taphonomic patternobserved at Cau del Duc also differs from the results obtainedby Lloveras et al. (2014a) in their study about the habits of theEgyptian vulture (Neophron percnopterus). This predatortransports the whole body of its prey to the nest when theprey animal is small, whereas in the case of larger birds it haspreyed upon, it only carries elements of the cranium or of theextremities. Concerning postcranial elements and following

the work of Bochenski et al. (1993), tarsometatarsi are themost abundant elements in the bird accumulations generatedby the Eurasian eagle-owl.

In the case of wing-to-leg proportions, the values observedat Cau del Duc fit with the anatomical profile described forthe Eurasian eagle-owl (Bochenski and Nekrasov 2001). Ina natural accumulation, a similar percentage of both upperand lower limb bones should be expected, since there wouldnot be the intervention of any predator (Ericson 1987).However, this is not the case at Cau del Duc, sincea preferential consumption of upper limb bones is inferred.These results agree with those obtained in studies on accu-mulations of pellets produced by nocturnal raptors and non-ingested bones of diurnal birds of prey (Bochenski et al. 1993,1997; Bochenski et al. 1999, 2009; Bochenski and Tomek1997; Laroulandie 2000, 2002; Bochenski and Nekrasov2001; Monchot and Gendron 2011; Lloveras et al. 2014a,2014b), with the exception of avian accumulations generatedby the snowy owl (Nyctea scandiaca), where lower limb bonespredominate (Bochenski 1997; Royer et al. 2019). In the caseof ingested bones of diurnal birds of prey and small mamma-lian carnivore accumulations, lower limb bones predominateover upper limb bones (Bochenski et al. 1997; Mallye et al.2008; Lloveras et al. 2014b; Rodríguez-Hidalgo et al. 2016).Nevertheless, studies on avian assemblages produced by thered fox (Vulpes vulpes) have identified a higher number ofwing elements (Monchot and Gendron 2011) (Table 8).However, some authors have pointed to the differential pre-servation, which depends on the bone density, to explainanatomical profiles (Hanson 1991; Higgins 1999; Dirrigl2001; Bovy 2002; Cruz 2005; Broughton et al. 2007). Basedon this hypothesis, the densest bones – and, therefore, theones with better preservation – would be directly related to

Figure 5. Root etching on a humerus shaft of an indeterminate species (a); manganese oxide deposits on a proximal right carpometacarpus of a Columbidae (b).

Table 7. Number and percentage (%) of post-depositional damage at the different metre intervals in the cave (length).

1m 2m 3m 4m 5m 6m 7m 8m 9m Covacha Total

Total NSP 1 36 216 580 610 362 4 40 19 44 1912Damaged bones 1 (100) 15 (41.66) 113 (52.31) 115 (19.82) 221 (36.22) 304 (83.97) 4 (100) 12 (30) 11 (57.89) 38 (86.36) 834 (43.62)Fissures - 8 (53.33) 37 (32.74) 57 (49.56) 72 (32.58) 80 (26.31) 3 (75) 6 (50) - 1 (2.63) 264 (31.65)Calcite-coating 1 (100) 4 (26.66) 65 (60.17) 46 (40) 133 (60.18) 202 (66.44) 1 (25) 3 (25) 3 (27.27) 28 (73.68) 486 (58.27)Manganese - - 1 (0.88) 8 (6.95) 5 (2.26) 6 (1.97) - - 7 (63.63) 7 (18.42) 34 (4.07)Chemical corrosion - 3 (20) 6 (5.30) 2 (1.74) 9 (4.07) 3 (0.98) - 1 (8.33) - 2 (5.26) 26 (3.11)Root-etching 1 (100) - 4 (3.53) 2 (1.74) 2 (0.90) 13 (4.27) - 2 (16.66) 1 (11.09) - 26 (3.11)

1328 G. ALONSO ET AL.

Table8.

Accumulationpatternof

differentbirdsof

prey

andmam

maliancarnivores

inavianassemblages.

Predator

Accumulation

Wing/leg

Core/limb

Proximal/distal

Completeness

(%)

Digestive

damage

References

Nocturnal

birdsof

prey

Eagleow

lIngested

Wing>

leg

Core<lim

bProximal>distal

Mod

erate

Ligh

t-mod

erate

Thisstud

y

Eagleow

lIngested

Wing>

leg

Core<lim

bProximal>distal

Mod

erate

Ligh

t-mod

erate

Bochenskie

tal.(1993);Bo

chenskiand

Nekrasov(2001);B

ochenskiandTomek

(1997);

Laroulandie(2000,

2002)

Long

-eared

owl

Ingested

Wing=leg

Core<lim

bProximal>distal

Mod

erate

Ligh

tBo

chenskiand

Tomek

(1994,

1997)

Snow

yow

lIngested

Wing<

leg

--

High

Ligh

tBo

chenski(1997);Royeret

al.(2019)

Tawny

owl

Ingested

Wing>

leg

Core<lim

bProximal>distal

Mod

erate

Ligh

t-mod

erate

Bochenskie

tal.(1993);Bo

chenskiand

Tomek

(1997)

Diurnal

birdsof

prey

Goldeneagle

Non

-ingested

Wing>

leg

Core>lim

bProximal>distal

High

-Bo

chenskie

tal.(1999)

Gyrfalcon

Non

-ingested

Wing=leg

Core<lim

bProximal>distal

High

-Bo

chenskiand

Tornberg

(2003)

Imperialeagle

Non

-ingested

Wing>

leg

Core<lim

bProximal>distal

High

Strong

Bochenskie

tal.(1997)

Ingested

Wing<

leg

Core<lim

bProximal=distal

Low

Strong

Bochenskie

tal.(1997)

White-tailed

eagle

Non

-ingested

Wing>

leg

Core<lim

bProximal>distal

High

-Bo

chenskie

tal.(2009)

Peregrine

falcon

Non

-ingested

Wing>

leg

Core<lim

bProximal>distal

High

-Laroulandie(2000,

2002)

Bonelli´s

eagle

Ingested

Wing<

leg

Core<lim

bProximal<distal

Mod

erate

Strong

-extrem

eLloveras

etal.(2014a)

Non

-ingested

Wing>

leg

Core>lim

bProximal>distal

Mod

erate

-Lloveras

etal.(2014a)

Egyptian

vultu

reIngested/Non

-ingested

Wing>

leg

Core>lim

bProximal>distal

High

Ligh

t-mod

erate

Lloveras

etal.(2014b)

Mam

malian

carnivores

Redfox

Non

-ingested

Wing>

leg

Core<lim

bProximal>distal

Mod

erate

-Mon

chot

andGendron

(2011)

Fox/Badg

erNon

-ingested

Wing<

leg

--

--

Mallyeet

al.(2008)

Lynx

Non

-ingested

Wing<

leg

Core<lim

bProximal>distal

High

-Rodríguez-Hidalgo

etal.(2016)

Genet

Ingested

--

-Low

Strong

Laroulandie(2000)

HISTORICAL BIOLOGY 1329

the method of locomotion. That is, in accumulations withflying bird bones it would be expected to find a better pre-servation of wing elements, as they are denser than otherskeletal elements and, consequently, less likely to be fragmen-ted or destroyed by post-depositional processes. Nevertheless,it has to be pointed out that bone density data from mostavian species has not hitherto been obtained that deeplyexamines this question. If we analyse the core-to-limb ratioin our sample, no significant information can be highlighted,as the proportion of limbs is always higher than the coreelements, a common characteristic in many predator accu-mulations. The proximal-to-distal index also does not providedistinct information about the taphonomic pattern of theEurasian eagle-owl, as in accumulations of birds of prey –both diurnal and nocturnal – and mammalian carnivores, theelements of the proximal part of the skeleton predominateover the distal, excluding the case of Bonelli’s eagle (Aquilafasciata) (Bochenski et al. 1993, 1997; Bochenski et al. 1999,2009; Bochenski and Tomek 1994, 1997; Bochenski 1997;Laroulandie 2000, 2002; Bochenski and Tornberg 2003;Mallye et al. 2008; Monchot and Gendron 2011; Lloveraset al. 2014a, 2014b; Rodríguez-Hidalgo et al. 2016) (Table 8).

According to Laroulandie (2000) and Bochenski (2005),the fragmentation degree observed in the avian assemblage ofCau del Duc does not correspond with those found iningested bones of diurnal birds of prey or mammalian carni-vores, where the number of complete bones is lower(Bochenski et al. 1997; Laroulandie 2000) or higher(Lloveras et al. 2014a). Our results also differ from thoseobserved in bird assemblages originated from non-ingestedbones of diurnal birds of prey and mammalian carnivores,which show a higher number of complete bones (Bochenskiet al. 1997, 1999, 2009; Laroulandie 2000, 2002; Bochenskiand Tornberg 2003; Lloveras et al. 2014a; Rodríguez-Hidalgoet al. 2016). Nevertheless, in the case of the red fox (Vulpesvulpes), a moderate degree of complete bones can be found, asdocumented by Monchot and Gendron (2011). Furthermore,the percentage of complete skeletal elements in the presentstudy is within the range of nocturnal birds of prey, includingthe Eurasian eagle-owl (Bochenski et al. 1993; Bochenski andTomek 1994, 1997; Laroulandie 2000, 2002; Bochenski andNekrasov 2001) (Table 8). The fragmentation of the pectoralgirdle could also be a consequence of the Eurasian eagle-owl’shabit of separating the wing joint from the core of their prey.Afterwards it ingests the whole wing (Bochenski 1960;Bochenski et al. 1993). Complete tarsometatarsi and carpo-metacarpi were found in relatively high percentages (29.7%and 31.3% respectively) owing to their lower flesh quantity.Thus, it is not necessary to break them before consumption(Laroulandie 2002). The radius is also considered a fragilebone, and is expected to be subject to higher fragmentation;however, its good preservation in some assemblages could bedue to it being protected by the more robust ulna (Bochenskiet al. 1993). The bone fragmentation in avian accumulationscan be also explained through the relative size differencesbetween the Eurasian eagle-owl and its prey. In most cases,the prey is not small enough to ingest entirely (Bochenskiet al. 1993). Additionally, the differential preservation of eachanatomical part depends on its robustness and density

(Bochenski et al. 1993). Consequently, the most compactareas have higher probabilities of being completely preserved.In the present study, a better preservation of the scapular partof coracoids has been observed compared to the sternal part.Regarding vertebrae and phalanges, as they are small andcompact bones, it is understandable that they do not showhigh fragmentation since they do not suffer direct impact(Bochenski et al. 1997).

Both pits and punctures seem to be a consequence of thechewing activities of small mammalian carnivores,a circumstance that suggests a sporadic presence of theseanimals in the cave. Nevertheless, the scarce number ofaffected bones makes it difficult to identify the carnivoreinvolved in the avian accumulation. With regards to thedigestion degree, the percentages documented in the pre-sent study coincide with the results of nocturnal raptoraccumulations, including the Eurasian eagle-owl(Bochenski et al. 1993; Bochenski and Tomek 1994, 1997;Bochenski 1997; Laroulandie 2000, 2002; Bochenski andNekrasov 2001; Royer et al. 2019). In bird assemblagesproduced by diurnal birds of prey or mammalian carni-vores, the digestive damage tends to be more intense, reach-ing strong and extreme damage (Bochenski et al. 1997;Laroulandie 2000; Lloveras et al. 2014b), except in thecase of the Egyptian vulture (Lloveras et al. 2014a) (Table8). With all variables together, we can propose that theEurasian eagle-owl is the main accumulator agent in theCau del Duc cave.

Finally, the good preservation of bones in general can beexplained due to the low affectation by post-depositionalprocesses, as shown by the fact that the bones barely presentany superficial modifications. The presence of manganeseoxide deposits at the innermost area of the cave indicatesthat it is a more humid zone compared to the entrance.

Conclusions

The results obtained from the avian assemblage of Cau delDuc fit well with the accumulation pattern originated bythe Eurasian eagle-owl. Among raptors, the Eurasian eagle-owl preferentially consumes limb bones, especially upperlimbs. As indicated in previous works, the absence ofcranial fragments and the possible loss of bones due todifferential preservation have been noticed. This raptorusually consumes preys larger in comparison to its bodysize, such as partridges or corvids (although its prey spec-trum is considerably broad). As a result, it is not able toingest them entirely, and therefore, tends to split the wingsfrom the core in order to make the consumption easier.Both wings and legs are the preferentially selected parts,probably owing to their higher alimentary content. Finally,the digestive damage differs from the damage obtained indiurnal birds of prey and mammalian carnivore accumula-tions, which produce a higher alteration of bones. Theoccasional presence of small mammalian carnivores wasalso documented at Cau del Duc, although it was notdetermined what type of predator was producing theobserved modifications because of the low number oftooth-marked bones. Conclusively, it is confirmed that it

1330 G. ALONSO ET AL.

is possible to characterise the pattern of accumulationsproduced by the Eurasian eagle-owl through specific fea-tures, and to distinguish it from the pattern generated byother birds of prey – both diurnal and nocturnal – andsmall mammalian carnivores.

Acknowledgments

We thank Jordi Fàbregas for his very useful help during fieldwork anddata recovery. This work was supported by the Spanish MINECO/FEDER projects CGL2015-68604-P and HAR2016-76760-C3-1-P, andthe Generalitat de Catalunya-AGAUR projects CLT009/18/00055 and2017 SGR 836. M. Arilla is the beneficiary of a research fellowship (FI)from AGAUR (2017FI-B-00096). A. Rufà is a beneficiary ofa postdoctoral research grant funded by the IdEx University ofBordeaux Investments for the Future program.

Disclosure statement

No potential conflict of interest was reported by the authors.

Funding

This work was supported by the Generalitat de Catalunya-AGAUR [2017SGR 836, 2017FI-B-00096]; MINECO/FEDER [CGL2015-68604-P andHAR2016-76760-C3-1-P]; Generalitat de Catalunya-AGAUR [CLT009/18/00055] and the IdEx University of Bordeaux Investments for theFuture program.

ORCID

Anna Rufà http://orcid.org/0000-0003-1278-4220Ruth Blasco http://orcid.org/0000-0001-9804-739X

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