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/lp/de-gruyter/the-diet-and-pellet-residue-taphonomy-of-barn-owls-tyto-alba-on-a-fzpTaoyF6Z
- Publisher
- de Gruyter
- Copyright
- © 2020 Anthony S. Cheke et al., published by Sciendo
- ISSN
- 0351-2851
- eISSN
- 2199-6067
- DOI
- 10.2478/acro-2020-0001
- Publisher site
- See Article on Publisher Site
Abstract
Acrocephal 41 us (184/185): 3–24, 2020 10.2478/acro-2020-0001 The diet, and pellet residue taphonomy, of Barn Owls Tyto alba on a Greek island reveals an exceptional diversity of avian prey Prehrana in tafonomija ostankov izbljuvkov pegaste sove Tyto alba na grškem otoku razkriva izjemno raznolikost ptičjega plena 1 2 Anthony S. Cheke , Julian P. Hume 139 Hurst Street, GB–OX4 1HE Oxford, UK, e–mail: anthony.cheke@dodobooks.com Bird Group, Department of Life Sciences, Natural History Museum, Akeman St, Tring, GB–HP23 6AP Herts, UK, e–mail: j.hume@nhm.ac.uk Barn OwT l yto alba pellets and loose bones on a cave floor from Amorgos (Cyclades, Greece) were examined and the birds found to have caught at least 39 species of bird, mostly identified from humeri, plus shrews Crocidura suaveolens, a few lizards and dung beetles, in addition to their principal diet of roden R ta s ( ttu rsat ras ttus, mice Apodemus spp. & Mus musculus). Amongst the birds, migrants appeared most vulnerable to owl predation, with some notable exceptions, while resident species were under-represented. The range of bird species found appears to be the largest recorded for any Barn Owl study of a single site. Considerable differences were found in species proportions of taxa in fresh pellets and in loose bones, probably due to differential rates of degradation. Photographs of all humeri are included to aid identification in other studies. 1. Introduction As part of wider studies on the fauna of Amorgos ( Cheke & Ashcroft 2017, Cheke et e d Th iet of owls is well studied due to the e al. 2 as0e2 0), during 2015–2019 Barn Owl roosts in of analysing the pellets they eject of un sd miaglel c stead v es and cavities notified to ASC by locals vertebrate bones and invertebrate exoskele w te or ne e s, a xn ad m ined by ASC and Ruth Ashcroft (REA) the food habits of the geographically wi fde or i spn rt ea ad c t pellets and loose bones on cave floors; owl Barn Owl Tyto alba are among the best know prn e y has not previously been studied on Amorgos. (e.g. Bunn et al. 1982, Taylor 1994, Romano et Thirty-nine whole pellets were collected from three al. 2020). In Barn Owls, small mammals alm lo os ct at ions, and hundreds of bones retrieved from the invariably make up the majority of food it floeo m r o s if o n ne cave, with a few from a fourth site (see both numbers and biomass, but reptiles and bib r e d ls oa w r)e . Th e sites are not occupied continuously by also taken, and in some circumstances the lt at ht e o er c wl as n , a nd there appears to have been a decline make a signic fi ant contribution. Here we rept oo n rt o ea nr e xtinction of these owls on the island in the the remarkable variety of avian species takel n b as y B t de ar c n a de, probably related to poisoning of rats – a Owls on Amorgos, a small island (1, 3 21 k 3 km m dead owl with no signs of injury was found by local long by 6 km at the widest point) in the Cyctlo au de r g s/uide Lonaïs Jallais in 2015. We found no Kiklades southeast of Naxos in the Aegean. recent (post 2015) pellets in 2016 or 2017, though 3 A. S. Cheke, J. P. Hume: The diet, and pellet residue taphonomy, of Barn O Tytow all bs a on a Greek island reveals an exceptional diversity of avian prey Figure 1: Location map of Amorgos (circled) in the Figure 2: Google Earth panoramic view of the Aegiale eastern Aegean sea. area, northeastern Amorgos, with location of Barn Owl sites indicated. Slika 1: Lokacija otoka Amorgos (obkroženo) v vzhodnem Egejskem morju. Slika 2: Google Earthov panoramski pogled območja Aegiale, severovzhodni Amorgos, z oznakami lokacij pegastih sov. e f Th our collection localities are all fairly close together around the village of Langada in the Aegiale area in the north-east of Amorgos (Figure 2). This part of the island consists of part-metamorphosed Triassic-Eocene limestone bedrock (‘massive marble’R , osenbaum et al. 2007), with numerous cavities and some caves. The area is also the wettest part of the island with the richest vegetation, a maquis that is almost forest in parts of the area between Langada to Theologos, and consequently has the highest bird diversity throughout the year (Cheke et al. 2020; see Table 5 for status of prey species). The principal site (D; Figure 3), about 1 Figure 3: Photo of entrance to Site D (Dhri cave), with km NE of the village at Dhri, is a small cave in the REA at the location of the main Barn Owl pellet deposit. wall of a wooded gorge, with a roost site on a dry- Barn Owl excreta is visible on the stone wall to the left of REA. The loose bones were found on the cave floor in stone wall near the entrance – there is no evidence the foreground. that the owls penetrate into the true interior. On a cliff above and just south of the village is the shrine Slika 3: Fotograjfi a vhoda na lokaliteto D (jama Dhri); of Aghia Triada (AT), were the owls used to nest REA označuje lokacijo glavnega nahajališča izbljuvkov (local informants, pers. comm. to ASC) but since c. pegaste sove. Na zidu levo od REA so vidni sovini iztrebki. Posamezne kosti so bile najdene na jamskih 2010 now only used sporadically for roosting. Just tleh (v ospredju fotograjfi e). NW, and as the Arakalos gorge opens out below the village, is a larger cave (A) in a cliff somewhat three, still wet, were collected in Apr -il 2018, al dic ffi ult of access where we only collected pellets though the owl was not seen. Only an old dry pellet once. Finally, south and up the talus slope from AT was found in 2019, and indeed most pelle-ts we re is a rock cleft (C) where ASC first found bones, but covered were dry with no indication of when they only a few, and not subsequently visited. had been produced. 4 Acrocephal 41 us (184/185): 3–24, 2020 2. Methods ones, were collected directly into ziplock bags, then sorted into major taxa back at ASC and REA’s Whole pellets were simply picked up from th pe flo ied-à o-t r erre in Langada, for later more spec-ific iden of a site, dried in the sun or oven, and prese t riv fice at d i io nn b y ASC in Oxford. ziplock bags for later analysis in Oxford, UKB . The ir ds (humeri, skull elements) were p-rovision bones were separated out in water and al al b lly i onde esn tified by ASC in Oxford using published and invertebrate remains removed, sortemd a atenrd ia l (principa Jáno lly ssy 1983, Brown et identie fi d to major taxa (birds, lizards) or s alp .e 1 c9i8 e7 a s nd online skullsite.com), but later re- (mammals) by ASC or Linda Losito (dung beet clh ee sc ).k ed together with JPH against ref -erence ma Rodents were identie fi d to genus using Lawre t n ec re & ial i n the skeleton collection at the bird section Brown (1973), its clear and extensive diagrao mf s t nh ote Natural History Museum, Tring, UK superceded by more recent guides; only one s (N pH ec M ieU s K). To aid the preliminary identifications each of Rattus and Mus are found on Amorgo a t s able of bone measurements of European birds in (Cheke & Ashcroft 2017), and the tw Ap o odemus the size range found was compiled from t - he litera species are separated by non-overlapping o tuvree ( ra alvla ilable from the first author on request). jaw sizei ( bid.), confirmed by the relatively larger tooth roots A.in mystacinus. Shrew bones were 3 . Results presumed to be from the only known species on the island, C rocidura suaveolens (Cheke & Ashcroft Both by number (72% of loose bones, 82% in pellets) 2017). Gecko Mediodactylus (Cyrtopodion) kotschyi and biomass the main prey of the Amorgos Barn dentaries were identie fi d from V et i al ll . (2 a 018), Owls were the four rodents present on the island – differing larger non-gekkonid lizard jaws aB ss lu am ck R ed at Rs attus rattus, some Rock Mic Ap e odemus to be from the abundant wall l Poidz aa rr cid s mystacinus, many remains assigned provisionally to erhardii, the only possibly candid Ch at ee ( ke & Wood Mice Apodemus sylvaticus, and a few House Ashcroft 2017). Bird names and sequence foll M oic w eMu s musculus. In addition to birds, other prey the standard English-language handbook o we f Gr re f e re ek q uent shre Crw oci s d ura suaveolens, a very few birds ( Handrinos & Akriotis 1997), with, in lizards (Reptilia – Squamata), beetles (Coleoptera) tables, more recently revised names in barn ac d koente s unidentie fi d dragony fl (Odonata) (Tables where relevant. Pellets and whole skulls o1 & 2 n th, e c Ch ae vk ee & Ashcroft 2017). floor were often host to larvae of tapestry mot Bh ro s wn Rat R. s norvegicus and voles Cricetidae, Trichophaga tapetzella and clothes mot Th ins ea common prey in mainland Europe, are absent from bisselliella, revealed when the adults emerged i An t mor hg e os (Masseti 201 Ch 2, eke & Ashcroft ziplock bags. 2017). The floor of cave D between the entrance and The proportions of the various prey species the roosting rock, semi-open to the ele am meonntgs, st loose bones and pellets are str -ikingly dif proved to be rich in bones from prey. The cav fe er is ent, and ver y significantly so o ten a χ st (Table 3); used by goats so the bones from decayed pell p e o ts s w sibe lr e r e easons for this are discussed below. scattered over several square metres of groun Bd o, b th uit n pellets (Table 1) and in the loose bones only in the top couple of centimetres after th oe l n t a h ye er cave floor (Table 2) there was a sma- ll but sig of goat droppings was removed. Digging n de ificea pn er t proportion of bird remains. Amongst the revealed no further bones. Collections, m floa ode b r boy nes, humeri were the most frequent and carefully surveying and raking through tg he e s ne ur ra fl al c y t e he best preserved avian bones recovered layer, were made on several occasions durin (T g a 2b 0l 1e 4 5–), and these are also the most diagnostic to 2019 until the site was more or less works ep d o eciu et s i . n the absence of complete skulls (Jánossy There was no new owl-generated input over 19 t8h3i)s . The bird bones proved on study to come period until four fresh pellets (birdless) we fr re f om a s ounu d rprisingly wide range of species, although in March 2018, and a partial fifth in April 2019, bu mt any were represented by only one or a fe -w individ old and including a bird, probably dating fu ra ol m t s. O hv e erall a total of 39 species (Table 5, Figure 4 same batch. All bones found, including ver& y s5m ) w ale l re recorded; in addition to loose bones, 5 A. S. Cheke, J. P. Hume: The diet, and pellet residue taphonomy, of Barn O Tytow all bs a on a Greek island reveals an exceptional diversity of avian prey there were eleven more or less complet - e a sist se o; i cin t at he other direction it is 7.8km over land + ed skeletons in pellets (Table 4), including a S 6.8k ism o kin ver sea to Liadi – in any case both are little Carduelis (Spinus) spinus (Fig. 8), the only specie m s ore than rocky outcrops and it is a lot further to not represented in the loose bones on the ca isl ve flo an o d r s w . ith suitable habitat. It is thus m - ost unlike The birds taken by the Barn Owls range fr lo y t m hat any but a tiny proportion of bones will have the very small (Chiffch Pa hyl ff loscopus collybita) to been brought in from outside Amorgos. quite bulky (Blackb Tu irrd dus merula), a weight e l Th ist of birds taken is by no means a random range of 6–10 to 80–125 g, though birds in t set o he f the island’s small and medium passerines. The lower part of this range (15–25 g) predom bi un lat k o e f the avian prey consists of migrants – summer (Table 4; weights from Snow & Perrins 1998)v . A islit l ors, passage migrants and winter visitors, as birds taken are passerines, except for W Jynr xy ne in cd k icated in Table 4. Resident species in the relevant torquilla, which is, apart from Common Qu sa izil e range are noticeably under-represented, the most Coturnix coturnix, the only non-passerine ( - oc ac ba undant (Sardinian WaS ryl bl ve ia r melanocephala, sional waders excepted) recorded on the iHo slau nsd i e Sn parrow Pas ser domesticus, and Crested Lark the same size range as the othCh er p er ke e y ( et al. Galerida cristata) barely making the list, and Blue 2020). Rock-thrus M hon ticola solitarius not showing up at all; some residents that are preyed on are species 4. Discussion supplemented by more abundant winter visitors (Stonecha St a xicola rubicola, Goldfinch Carduelis 4.1 Birds carduelis, LinnetC arduelis (Linaria) cannabina, Chaffinch Fringilla coelebs). This suggests the All but one of the bird species identified in rem mia gir na s nts are relatively naïve or vulnerable to Barn has been recorded live on the island by ASO C a wl p nd redation, whereas the residents are more REA (Cheke et al. 2020) in the years since 2007 a. ware of the threat. Migrants from mainland areas Some however occur infrequently, and in on w e c ill r asa e, rely have been subject to predation from Barn Siskin, bones were identified in a pellet be Ofw or le s, and in addition may on arrival be exhausted living birds had been seen, and Wryneck ho ur w mer ei ak and thus easy targets. We presume the owls were found (though not identified) before a catc sigh h t t he birds at night when roosting. There are record. The cave floor produced a single Crosa sb nio lm l alies however – in both spring and autumn Loxia curvirostra humerus – a species not yet sm eein gration Spotted Flycatc Muhse cr ics apa striata on the island, although known (but accidec na tn b al) o e t n he most abundant birds in the better nearby IraklGa ia ( valas 2014). vegetated parts of the island, yet they largely escape We believe owl prey was taken on Amoprrg ed oa s tion, whereas the less abundant, albeit still itself, not brought in from outside. In Greefcae m irloy c st ommon, Pied and Collared Flycatchers Barn Owls are resident, with a small num Fb ice er o dulaf hypoleuca/F.albicollis, indistinguishable immigrants from further n H oan rtdrino h ( s & osteologically, are rather frequent v ictims. Akriotis 1997). Home ranges in the Medite-rraO nt eher common migrants that the owls do not an area appear not to have been studied, but further north in Europe most birds range c.3km from their roosts, rarely to 16 T k am ( ylor 1994, 2002). The only confirmed cases of regular foraging across sea Please note: The following bone photo sets (Figs 4–7) gaps we have found are in the Balearic islands, where are composites made up of photos of pairs of bones owls cross up to 4.5km of sea to forage on adjacent photographed separately. Therefore, the scale islands G ( uerra et al. 2014) and on Skomer Island bar should be taken as a guide only; more exact measurements can be found in Table 5, column 2. (Wales, UKL ; oughran 2006) where the sea gap to the mainland is only 0.6km. On Amorgos the Opomba: Sledeče zbirke fotograj k fi osti (slike 4–7) over-sea distance to the nearest potentially useful so kolaži fotograj p fi arov kosti, ki so bili fotograrfi ani island is greater (6.6km across sea to Ano Antikeri), posamično. Merilo je zato zgolj vodilo, podrobne meritve and it is a further 24km (overland) from ts h o v t e ca a bv ele i 5. 6 Acrocephal 41 us (184/185): 3–24, 2020 A B C D E F G H I J K Figure 4: Humeri in caudal view of passerines (warblers & chats) discussed in the text. In this and subsequent figures (Figure 5, 6 and 7) the left-hand image(s) with numbered codes are reference material from the UK Natural History Museum (NHM), the right-hand bones, labelled Amorgos, are samples collected in this study. In the list the individual bone captions are separated by vertical line 'ǀ'. A – Blackcap Sylvia atricapilla NHMUK S/1968.6.36 u/s ǀ Garden Warbler Sylvia borin NHMUK S/1968.6.28 ♀ ǀ Amorgos, B – Great Reed Warbler Acrocephalus arundinaceus NHMUK S/1998.92.22 u/s ǀ Amorgos*, C – Nightingale Luscinia megarhynchus NHMUK S/1968.4.13 ♀ ǀ Amorgos, D – Orphean Warbler Sylvia hortensis** NHMUK S/1968.6.24 ♂ ǀ Amorgos, E – Common Whitethroat Sylvia communis NHMUK S/1968.4.25 ♂ ǀ Amorgos, F – Robin Erithacus rubecula NHMUK S/1968.1.22 ♂ ǀ Amorgos, G – Willow Warbler Phylloscopus trochilus NHMUK S/1968.1.11 ♂ ǀ Amorgos, H – Chiffchaff Phylloscopus collybita NHMUK S/1968.1.20 ♂ ǀ Amorgos, I – Sedge Warbler Acrocephalus schoenobaenus NHMUK 1930.3.24.457 u/s ǀ Amorgos, J – Sardinian Warbler Sylvia melanocephala NHMUK S/1998.29.2 u/s ǀ Amorgos, K – Subalpine Warbler Sylvia cantillans** NHMUK S/2002.40.1 u/s ǀ Amorgos. Scale bar = 10mm. * The Amorgos humerus in ‘B’ is not a Great Reed Warbler, but inferred to be from a Olive Tree WarbleH r ippolais olivetorum on the basis of size and morphology (our comparisons and J.Kessler pers. comm.); there being no specimens of H.olivetorum in the NHM skeleton collection we used the bone that came nearest in size and appearance. ** S.(h.) hortensis and the eastern form S.(h.) crassirostris are not separated in the NHM skeleton collections, and are almost certainly indistinguishable; the same applies to the forms/species of the S. cantillans complex. Slika 4: Nadlahtnice (kavdalno) pevk (trstnice in taščice), omenjenih v besedilu. V tej in sledečih tabelah (tabele 5, 6 in 7) so na levi strani referenčni primerki iz NHM, na desni, označeni Amorgos, pa primerki iz te raziskave. V opisu slik so posamezne vrste ločene s pokončno črto “ǀ”.A – Črnoglavka Sylvia atricapilla ǀ vrtna penica Sylvia borin ǀ Amorgos, B – rakar Acrocephalus arundinaceus ǀ Amorgos,* C – slavec Luscinia megarhynchus ǀ Amorgos, D – svetlooka penica Sylvia hortensis ǀ Amorgos,** E – rjava penica Sylvia communis ǀ Amorgos, F – taščica Erithacus rubecula ǀ Amorgos, G – severni kovaček Phylloscopus trochilus ǀ Amorgos, H – vrbji kovaček Phylloscopus collybita ǀ Amorgos, I –bičja trstnica Acrocephalus schoenobaenus ǀ Amorgos, J – žametna penica Sylvia melanocephala ǀ Amorgos, K – taščična penica Sylvia cantillans ǀ Amorgos,** Merilo = 10 mm. * Nadlahtnica B ne pripada rakarju, vendar na podlagi velikosti in morfologije najverjetneje oljčnemu vrtniku Hippolais olivetorum (lastna primerjava in osebna komunikacija z J. Kassler). V zbirki NHM ni oljčnega vrtnika, zato smo uporabili kost, ki mu je po velikosti in videzu najbolj podobna. ** svetlooka penica S. hortensis in vzhodna svetlooka penica S. crassirostris v zbirki okostji NHM nista ločeni in ju ni mogoče razlikovati. Enako velja za vrsto oz. obliko taščične penice S. cantillans. 7 A. S. Cheke, J. P. Hume: The diet, and pellet residue taphonomy, of Barn O Tytow all bs a on a Greek island reveals an exceptional diversity of avian prey A B C D E F G H I J Figure 5: Humeri in caudal view of passerines (n fi ches, buntings & sparrows) discussed in the text. A – Chafn fi ch Fringilla coelebs NHMUK S/1968.4.38 ♀ ǀ Brambling Fringilla montifringilla NHMUK S/1976.20.1 u/s ǀ Amorgos, B – Crossbill Loxia curvirostra NHMUK S/2008.6.1 u/s ǀ Amorgos, C – Ortolan Bunting Emberiza hortulana NHMUK S/1968.4.39 ♀ ǀ Cretzchmar’s Bunting Emberiza caesia NHMUK S/1968.4.41 ♂ ǀ Amorgos, D – Greenfinch Carduelis chloris NHMUK S/1982.22.2 u/s ǀ Amorgos, E – Hawn fi ch Coccothraustes coccothraustes NHMUK S/1984.82.1 ♂ ǀ Amorgos, F – House Sparrow Passer domesticus NHMUK S/1977.76.1 u/s ǀ Spanish Sparrow Passer hispaniolensis NHMUK S/1968.1.41 ♀ ǀ Amorgos*, G – Linnet Carduelis (Linaria) cannabina NHMUK S/1976.26.1 ♂ ǀ Amorgos, H – Goldn fi ch Carduelis carduelis NHMUK S/2017.8.2 ♀ ǀ Amorgos, I – Serin Serinus serinus NHMUK S/1961.13.30 ♂ ǀ Amorgos**, J – 31. Siskin Carduelis (Spinus) spinus NHMUK S/1982.40.1 ♂ ǀ Amorgos. Scale bar = 10mm. * Note the marked difference in morphology between House and Spanish Sparrow humeri, presumably ree fl cting the greater muscle mass required in the migratory Spanish Sparrow; Amorgos sparrow humeri are clearly from House Sparrows, although a mandible is from a Spanish Sparrow. ** The Serin from Amorgos is very much in the upper end of the size range for the species. Slika 5: Nadlahtnice (kavdalno) pevk (ščinkavci, strnadi in vrabci), omenjenih v besedilu. A – ščinkavec Fringilla coelebs ǀ pinoža Fringilla montifringilla ǀ Amorgos, B – krivokljun Loxia curvirostra ǀ Amorgos, C – vrtni strnad Emberiza hortulana ǀ balkanski strnad Emberiza caesia ǀ Amorgos, D – zelenec Chloris chloris ǀ Amorgos, E – dlesk Coccothraustes coccothraustes ǀ Amorgos, F – domači vrabec Passer domesticus ǀ travniški vrabec Passer hispaniolensis ǀ Amorgos,* G – repnik Carduelis (Linaria) cannabina ǀ Amorgos, H – lišček Carduelis carduelis ǀ Amorgos, I – grilček Serinus serinus ǀ Amorgos,** J – čižek Carduelis (Spinus) spinus ǀ Amorgos. Merilo = 10 mm. * Opazna morfološka razlika v nadlahtnici domačega in travniškega vrabca, ki je najverjetneje posledica večje mišične mase slednjega, ki je selivec. Nadlahtnice z otoka Amorgos nedvomno pripadajo domačemu vrabcu, najdene spodnje čeljusti pa travniškemu vrabcu. ** Grilček z otoka Amorgos dosega zgornjo mejo velikosti za to vrsto. 8 Acrocephal 41 us (184/185): 3–24, 2020 A B C D E F G H I J Figure 6: Humeri in caudal view of passerines discussed in the text. A – House Martin Delichon urbica NHMUK S/1973.29.3 u/s ǀ Barn Swallow Hirundo rustica NHMUK S/1985.7.1.237 ♂ ǀ Amorgos*, B – Stonechat Saxicola (torquata) rubicola ** NHMUK S/1968.1.28 ♂ ǀ Amorgos, C – Whinchat Saxicola rubetra NHMUK S/1983.113.2 ♂ ǀ Amorgos, D – Spotted Flycatcher Muscicapa striata NHMUK S/1968.6.82 ♂ ǀ Amorgos, E – Pied Flycatcher Ficedula hypoleuca NHMUK S1968.6.45 ♂ ǀ Collared Flycatcher Ficedula albicollis NHMUK S1968.6.79 ♂ ǀ Amorgos, F – Black Redstart Phoenicurus ochrurus NHMUK S/1968.4.12 ♂ ǀ Amorgos, G – Dunnock Prunella modularis NHMUK S/1996.50.6 ♂ ǀ Amorgos, H – Wryneck Jynx torquilla NHMUK S/1968.4.5 ♀ ǀ Amorgos, I – Woodchat Shrike Lanius senator NHMUK S/1983.41.1 u/s ǀ Amorgos, J – Red-backed Shrike Lanius collurio NHMUK S/1968.6.85 ♂ ǀ Amorgos. Scale bar = 10mm. * Although both species are common migrants, Amorgos hirundine humeri are clearly from House Martins not Swallows. ** The segregates in the Saxicola torquata complex are not separated in the NHM skeleton collections. Slika 6: Nadlahtnice (kavdalno) pevk, omenjenih v besedilu. A – mestna lastovka Delichon urbica ǀ kmečka lastovka Hirundo rustica ǀ Amorgos,* B – prosnik Saxicola (torquata) rubicola ǀ Amorgos,** C – repaljščica Saxicola rubetra ǀ Amorgos, D – sivi muhar Muscicapa striata ǀ Amorgos, E – črnoglavi muhar Ficedula hypoleuca ǀ belovrati muhar Ficedula albicollis ǀ Amorgos, F – šmarnica Phoenicurus ochrurus ǀ Amorgos, G – siva pevka Prunella modularis ǀ Amorgos, H – vijeglavka Jynx torquilla ǀ Amorgos, I – rjavoglavi srakoper Lanius senator ǀ Amorgos, J – rjavi srakoper Lanius collurio ǀ Amorgos. Merilo = 10 mm. * Čeprav sta obe vrsti pogosti selivki, pripadajo najdene nadlahtnice mestnim lastovkam. ** Oblike in podvrste Saxicola torquata v zbirki okostji NHM niso ločene. 9 A. S. Cheke, J. P. Hume: The diet, and pellet residue taphonomy, of Barn O Tytow all bs a on a Greek island reveals an exceptional diversity of avian prey A B C D E Figure 7: Humeri in caudal view of the largest passerines discussed in the text. A – Golden Oriole Oriolus oriolus NHMUK S/1968.6.10 ♂ ǀ Amorgos, B – Starling Sturnus vulgaris NHMUK S/1973.46.1 ♀ ǀ Amorgos, C – Blackbird Turdus merula NHMUK S/1982.134.1 ♂ ǀ Amorgos, D – Songthrush Turdus philomelos NHMUK S/1982.48.1 u/s ǀ Amorgos, E – Crested Lark Galerida cristata NHMUK S/1998.92.35 u/s ǀ Amorgos. Scale bar = 10mm. Slika 7: Nadlahtnice (kavdalno) večjih pevk, omenjenih v besedilu. A – kobilar Oriolus oriolus ǀ Amorgos, B – škorec Sturnus vulgaris ǀ Amorgos, C – kos Turdus merula ǀ Amorgos, D – cikovt Turdus philomelos ǀ Amorgos, E – čopasti škrjanec Galerida cristata ǀ Amorgos. Merilo = 10 mm. 10 Acrocephal 41 us (184/185): 3–24, 2020 Figure 8: An associated Siskin Carduelis (Spinus) spinus recovered from a single Barn Owl pellet. Scale bar = 10mm. Slika 8: Okostje čižka, sestavljeno iz enega samega izbljuvka pegaste sove. Merilo = 10 mm. 11 A. S. Cheke, J. P. Hume: The diet, and pellet residue taphonomy, of Barn O Tytow all bs a on a Greek island reveals an exceptional diversity of avian prey Table 1: Barn Owl prey in Amorgos analysed in 39 pellets. Codes: a – Pied/Collared Flycatcher Ficedula hypoleuca/albicollis, b – Woodchat Shrike Lanius senator, c – Blackcap Sylvia atricapilla, d – Chafn fi ch Fringilla coelebs, e – Blackbird Turdus merula, f – Siskin Carduelis (Spinus) spinus, g – Chiffchaff Phylloscopus collybita, h – gecko Mediodactylus kotschyi, j – wall lizard Podarcis erhardii, k – Copris hispanus, m – Thorectes cf. bruelli, n – Stonechat Saxicola (torquata) rubicola, x – not identie fi d. Localities & dates: D – Dhri cave (1–23: 21. 5. 2015, 24–27: 21. 4. 2018, 28: 5. 4. 2019), AT – Agia Triada Shrine (6. 3. 2015, none found subsequently), A – Araklos gorge cave (22. 5. 2015), UL – unlabelled sample (labels lost, but UL1 & 2 probably from D (30. 3. 2016, old pellets missed in 2015), 3 & 4 from A (22. 5. 2015). Sample / Vzorec D AT A UL Prey / Plen 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 Total:D 1 2 3 4 5 6 1 1 2 3 4 Grand Total / Skupaj Rattus rattus 1 1 1 1 1 1 1 2 2 1 1 13–13% (14) 1 1 15–12% Apodemus mystacinus 1 1 1 2 1 1 1 8–8% (8.5) 1 1 10–8% A. cf. sylvaticus 1 1 2 5 2 1 1 1 2 3 3 2 5 3 2 2 4 2 2 1 1 1 47–47% (50) 1 2 1 1 2 3 1 58–46% Mus musculus 2 1 1 2 2 1 1 1 2 1 14–14% (15) 1 15–12% Crocidura suaveolens 1 1 1 1 1 2 1 8–8% (8.5) 2 10–8% Bird / Ptica 1a 1a 1a 1n 4–4% (4) 1b 1c 2de 3dfg 11–9% Lizard /Kuščar 1h 1h 1j 3–3% - 3–2.5% Dung beetle / Govnač 1k 1k 1k 3–3% - 1x 1m 5–4% Dragonfly / Kačji pastir 1x 1–1% 1–0.8% TOTALS / SKUPAJ 101 (94) 125 (119) 12 Acrocephal 41 us (184/185): 3–24, 2020 Tabela 1: Plen pegastih sov na otoku Amorgos, analiziran iz 39 izbljuvkov. Oznake: a – črnoglavi/belovrati muhar Ficedula hypoleuca/albicollis, b – rjavoglavi srakoper Lanius senator, c – črnoglavka Sylvia atricapilla, d – ščinkavec Fringilla coelebs, e – kos Turdus merula, f – čižek Carduelis (Spinus) spinus, g – vrbji kovaček Phylloscopus collybita, h – gekon Mediodactylus kotschyi, j – Erhardova pozidna kuščarica Podarcis erhardii, k – Copris hispanus, m – Thorectes cf. bruelli, n – prosnik Saxicola (torquata) rubicola, x – ni določeno. Lokacije & datumi: D – jama Dhri (1–23: 21. 5. 2015, 24–27: 21. 4. 2018, 28: 5. 4. 2019), AT – svetišče Agia Triada (6. 3. 2015, brez novejših najdb), A – jama v soteski Araklos (22. 5. 2015), UL – neoznačeni vzorci (oznake izgubljene, UL1 in 2 najverjetneje z lokalitete D (30. 3. 2016, stari izbljuvki spregledani v letu 2015), 3 & 4 z lokalitete A (22. 5. 2015). Sample / Vzorec D AT A UL Prey / Plen 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 Total:D 1 2 3 4 5 6 1 1 2 3 4 Grand Total / Skupaj Rattus rattus 1 1 1 1 1 1 1 2 2 1 1 13–13% (14) 1 1 15–12% Apodemus mystacinus 1 1 1 2 1 1 1 8–8% (8.5) 1 1 10–8% A. cf. sylvaticus 1 1 2 5 2 1 1 1 2 3 3 2 5 3 2 2 4 2 2 1 1 1 47–47% (50) 1 2 1 1 2 3 1 58–46% Mus musculus 2 1 1 2 2 1 1 1 2 1 14–14% (15) 1 15–12% Crocidura suaveolens 1 1 1 1 1 2 1 8–8% (8.5) 2 10–8% Bird / Ptica 1a 1a 1a 1n 4–4% (4) 1b 1c 2de 3dfg 11–9% Lizard /Kuščar 1h 1h 1j 3–3% - 3–2.5% Dung beetle / Govnač 1k 1k 1k 3–3% - 1x 1m 5–4% Dragonfly / Kačji pastir 1x 1–1% 1–0.8% TOTALS / SKUPAJ 101 (94) 125 (119) Table 2: Numbers of rodent and shrew jaws, and minimum total of bird humeri (from Table 5) recovered loose from the floor of Dhri Cave (L – left, R – right jaws). Minimum number of mammal individuals indicated by the higher g fi ure for left or right in italics, numbers then used in Table 3. Minimum bird numbers as shown in Table 5.The lizard is Podarcis erhardii. Tabela 2: Število čeljusti glodalcev in rovk ter minimalno število nadlahtnic ptic (iz tabele 5), pridobljenih iz tal jame Dhri (L – leva, R – desna čeljust). Minimalno število osebkov sesalcev (višja vrednost za levo ali desno čeljust) je zapisano ležeče, vrednost pa je nato uporabljena v tabeli 3. Minimalno število ptic, kot jih prikazuje tabela 5. Kuščar je Podarcis erhardii. Species / Vrsta Date / Datum L R L R L R L R L R min. dentary May 2015 51 50 6 6 18 19 7 5 March 2016 63 60 10 13 30 29 2 12 4 October 2017 12 15 2 1 5 7 1 1 4 3 April 2018 42 49 11 9 38 30 4 3 16 21 1 April 2019 17 12 6 2 10 10 3 3 1 3 1 TOTALS (459) / 185 186 35 31 101 95 8 9 40 37 86 2 SKUPAJ (459) % (rounded) 40.5 7.5 22 2 9 19 Rattus rattus Apodemus mystacinus A.cf. sylvaticus Mus musculus Crocidura suaveloens Bird /Ptica Lizard / Kuščar A. S. Cheke, J. P. Hume: The diet, and pellet residue taphonomy, of Barn O Tytow all bs a on a Greek island reveals an exceptional diversity of avian prey or rarely catch are summer visitors Olsiv p. s acueo gu gse sts migrants were targeted, though the Warbler H ippolais (Iduna) pallida, and wheatearo s wls also took a young Chukar pa Ar let ctr or id isg e Oenanthe spp., passage migrants Common Redst ch a uk rt ar and even a Scops OO w tu l s scops. Milos, at Phoenicurus phoenicurus, Wood WarblP ehyl r loscopus 151 km , is a bit larger than Amorgos, and here also sibilatrix and Tree PipA it nthus trivialis, and wintert he owls at two sites took some larger species than visitors Meadow Pip Aint thus pratensis, Pied on Amorgos (e fl dgling kestrel, 2 species of pigeon; WagtaM il otacilla alba and Corn BuntiE nm g beriza Table 6), and also targetted resident species more calandra. The preponderance of Chaffinches may than in Amorgos, though migrants were captured be due to there being a large winter roost n in au ear c tu am ve n ( A livizatos & Andriopoulos 2016); D (ASC & REA unpublished data); likewise t bih re d numbers, at 6% of mammals, were lower than frequently predated Blac Syl kc via ap a s tricapilla are in Amorgos. The study was said to be ongoing, very common on migration, and frequent as w ali tn h t o eu rg h no further data have been published. visitors, sometimes also aggregating in the viciO nf m ity o ai fn land Greek sites, birds averaged 4% the owl rooCh stse ( ke & Ashcroft 2016). by number and less in biomass, but were 39.6% eTh re have been several other publish (wed in ter) to 43.4% (summer) of diet as biomass at studies of Barn Owl diet in Greece, onlM y t itw rio k ou, a wetland site in northeastern Greece of which involved a small island, Antik (Gyotuhtir na er & Alivizatos 2003). Although only (Alivizatos et al. 2005) and Milos ( Alivizatos 11–13% in number – the birds were not identie fi d & Andriopoulos 2016). Antikythira, off thet o species but the number/biomass ratio suggests southern Peloponnese, famous for quantiltayr a gind sh species, probably non-passerines (waders?) variety of migratory birD ds ( ime ak .gi . et al. 2006), were targetted at th O is s buch ite & . Benda (2009) is less than half the size of Amorgos and has o re n pl o y t rtw ed 1 o 4 bird species total from two sites in the species of roden Rt ( attus rattus & Mus musculus; Peloponnese (Greek mainland) and 15 from five ibid.), yet only about 9–10 species of birds w siteer s i e n Crete (Table 6), while A liet v i al zat . (2o 00 s 5) found in Barn Owl d A ie lt ( ivizatos et al. 2005, 2+ recorded only 2–7 species each in six mainland sites, species not identie fi d; Table 6), though the pre as nd en B c on e tzorlos et al. (2005) only two species in of a raiPl orzana sp., Barn SwalloH w irundo three sites, though up to 9% of items in winter at rustica, flycatcherF s icedula sp. and shrikLan es ius one of them. Table 3: Chi-square test comparing proportions of prey species in loose bones and pellets from the Dhri Cave, testing an expectation of equal proportions (calculated ‘expected’ figures in italics). χ = 79.6, df = 5, p << 0.0001. The individual χ in bold indicate major departures from the expected result that both loose bones and pellets would have the same proportions of prey. For loose bones the counts are for the most frequent bones by taxa: mammal jaws and bird humeri. Tabela 3: Hi-kvadrat primerjave razmerja vrst plena v izbljuvkih in njihovih ostankih (posamezne kosti) iz jame Dhri, test pričakovanih enakomernih razmerij (izračunane pričakovane vrednosti v ležeči pisavi). χ = 79.6, df = 5, p << 0.0001. Posamezen χ v krepki pisavi prikazuje velike odklone od pričakovanja, da so razmerja vrst plena v izbljuvkih in njihovih ostankih (posamezne kosti) enaka. Pri posameznih kosteh so podane vrednosti za najpogostejše kosti iz taksona: čeljusti sesalcev in nadlahtnice ptic. R attus Apodemus Apodemus cf. Mus Crocidura Birds / Totals / Taxa / Takson rattus mystacinus sylvaticus musculus suaveolens Ptice Skupaj 186 35 101 9 40 86 Loose bones / 165.1 35.7 122 .8 19.1 39.8 74.6 457 posamezne kosti (2.67) (0.01) (3.87) (5.34) (0.00) (1.74) 13 8 47 14 8 4 Pellets / 33.9 7.3 25.2 3.9 8.2 15.4 94 Izbljuvki (12.88) (0.07) (18.86) (26.16) (0.00) (8.43) Totals / Skupaj 199 43 148 23 48 90 551 14 Acrocephal 41 us (184/185): 3–24, 2020 The only report we have found that appe in ag s rs eabird), Common Quail, wa C de har rad s rius comparable to the Amorgos situation is fr ale om t xand h re inus, Calidris ferruginea andTr inga sp., a Balearic islands at the other end of t-he M maerd sih t ter er C n hlidonias sp. and Common Bee-eater ranean G . uerra et al. (2014) looked at the diet o Me f rops apiaster, reflecting in part the greater range Barn Owls on Formentera and islets adjao cf h ent abt io tats available – Amorgos has only tiny token neighbouring Eivissa (Ibiza), finding a ric - h c wo em tlands. Comparative results from an inland site bined haul of nearly 30 species, mostly mig in I ran bt izsa a : re added fr So om mmer et al. (2005) in individually, more than 21 on the small iT sl ae bt o le 6f . As in Greece, on these islands the bulk of s’Espalmador and more than 16 combining t thw e po rey was rodents. sites on Formentera (Table 6; numbers incluW de e were unable to study seasonal variation in some species identified only to genus). The specc ie atc s h, but as w A it liviz h atos & Andriopoulos range here was wider than on Amorgo-s, eo mn M braiclos B , osé & Guidali (2001) in north Italy ing Storm-petr H eyl drobates pelagicus (local bre -edfound an increase in birds taken in summer/ Table 4: Numbers of loose complete bird bones on Dhri cave floor, by collection date [excluding radiuses or scapulas as too few to be worth listing], compared with Denys et al.’s percentages (2017). Note that the collecting dates in 2015–2018 are sampling from the same initial bone ‘population’ as there was no new input in the intervals; the return of owls in 2018 may have added a few bones before the 2019 collection. Humerus heads, although identia fi ble, excluded to retain comparability with other bones. The low figures in 2019 ree fl ct diminishing returns as the site is worked out. Abbreviations: CMC – carpometacarpus, TbT – Tibiotarsus, TMT – Tarsometatarsus. Tabela 4: Število posameznih celih kosti ptic na tleh jame Dhri, glede na čas zbiranja [brez koželjnic in lopatic, ki jih je v vzorcu premalo] in v primerjavi z odstotki, navedenimi v Denys et al. (2017). Kosti, nabrane v obdobju 2015–2018, predstavljajo vzorec iz istega začetnega vzorca, saj v obdobju ni bilo najdenega novega, svežega materiala. Vrnitev sov v letu 2018 je prispevalo nekaj novih kosti pred vzorčenjem v letu 2019. Glave nadlahtnice, čeprav določene, so bile izločene zaradi zagotavljanja ustreznosti z drugimi kostmi. Majhne vrednosti v letu 2019 kažejo na zmanjševanje kosti v začetnem vzorcu. Okrajšave: CMC – karpometakarpus, TbT – golenično-nartna kost, TMT – Tarzusmetatarzus. Date / Datum May March October April April TOTALS/ % post-cranialD / enys et Bone / kost 2015 2016 2017 2018 2019 SKUPAJ % brez lobanje al (2017) coracoid / krokarnica 3 11 7 13 2 36 9.2 14.3 humerus / nadlahtnica 34 39 17 26 11 127 32.6 10.7 ulna / komolčnica 11 29 14 23 10 87 22.4 10.7 CMC 5 12 4 8 3 32 8.2 8.9 femur / stegnenica 11 17 10 11 1 50 12.9 19.6 TbT 6 7 4 2 - 19 4.9 17.9 TMT 13 6 7 8 4 38 9.8 17.9 Total post-cranial / 389 56 skupaj brez lobanje skull (cranium+bill) / 1 (partial 1 1 2 5 skupaj (lobanja+kljun) / delna) cranium (no bill) / 1 1 lobanja (brez kljuna) upper mandible / 1 (partial 4 9 2 20 zgornja čeljust / delna) lower mandible / 5 7 3 15 spodnja čeljust 15 A. S. Cheke, J. P. Hume: The diet, and pellet residue taphonomy, of Barn O Tytow all bs a on a Greek island reveals an exceptional diversity of avian prey Table 5: Avian prey of Barn Owls & their status in Amorgos, from loose bones on floor of Dhri cave (site D), listed by size of humerus. Conventions & Codes: n[x/y] – total [left/right] + h [head of humerus only]; under status: PM – passage migrant, R – resident, SV – summer visitor, WV – winter visitor, qualie fi d by a = abundant, c – common, f – frequent/fairly common, s – scarce, r – rare. > refers across to comment column. Weight data from Swon & Pnirre S (1998), except, due to an obvious printing error, Greenn fi ch from Pnirre S (1987); status data: ASC, REA & Apostolos Cristopoulos (pers. obs. 2007–19 in Cekeh et al. 2020). ‘+X’ in bold in column 7 indicates additional specimens in pellets, not included in total. Humerus Size of samples No. whole humeri [L/R] sample Ref./ (mm)/ Weight Min. No. of or heads only h[L/R] / Oznaka vzorc V a elikost vzorca range (g)/ birds / Min. Št. celih ali samo gl A ad v ditional comments [additional bones noted where identified - mostly only from larger species] / nadlahtnice (v mm) razpon teže (g)English name / Angleško imS ecientific name / Znanstveno ime Status št. ptic nadlahtnic Dodatni komentarji 1b 12 6–10 Chiffchaff Phylloscopus collybita PMc, W Vc 1+1> 1>[1/-] confirmed against pellet sample UL3 & NHM collections 1c 12.5 7–12 Willow Warbler P.trochilus PMc 2 2[-/2] consistent differences from Chiffchaff 25,25a 13 8–13 Subalpine Warbler Sylvia cantillans SVc 2 4[2/2] 33 13.5 9–15 Sedge Warbler Acrocephalus schoenobaenus PMs 1 1[-/1] 24 13.5 10–15 Sardinian Warbler Sylvia melanocephala Ra 1 1[1/-] 31 14–14.5 11–18 Siskin Carduelis (Spinus) spinus W V* 0+1 2 [1/1] [from pellet UL3] *irruptive; rare visitor, but common when arrives 32 14.5–15 13–18 Whitethroat Sylvia communis PMf, SVr 2 2 [1/1] + h[-/1] 2,2b 15 10–16 Pied / Collared Flycatcher Ficedula hypoleuca / albicollis both PMc 4+3> 5[3/2] + h[1/-] not separable; both common; close to St Fo .se nm ec ito hra qt u; ata also possible] [also pellets D3, D12, D16] 20,20a 15 13–17 Stonechat Saxicola (torquata) rubicola Rs, W Vf 3+1> 2[2/-] +h[1/-] [>also pellet D28] 13 15 16–22 House Martin Delichon urbicum PMa 1 2 [1/1] 4 16 14–19 Goldfinch Carduelis carduelis Rs,W Vf 1 2[1/1] 14 16 11–14 Serin Serinus serinus W Vs 1 1[-/1] 28 16 14–20 Spotted Flycatcher Muscicapa striata PMa 1 1[1/-] very close to Whinchat & similar also to R Phoe eni ds cu tr a u r s t phoenicurus 3abcefgh 16–17 14–19 Whinchat Saxicola rubetra PMc 8 15[8/7] close to Spotted Flycatcher 23 16.5 Robin Erithacus rubecula W Vc 1 1[1/-] 5,5abcde 16–18 16–25 Blackcap / Garden Warbler Sylvia atricapilla / borin PMa, W Vf / PMr 11+1> 15[5/10] +h[1/1] not separable, but Garden Warbler is very scarce [>also pellet sample AT4] 30 [15–17] 16–25 Dunnock Prunella modularis W Vr 1 h[1/-] head of bone only 4a 17 15–22 Linnet Carduelis (Linaria) cannabina Rc, W Vc 1 2[1/1] 7,7a 18 13–19 Black Redstart Phoenicurus ochruros W Vc 2 3[2/1] 8 18 24–38 House Sparrow Passer domesticus Ra 2 2[1/1] + h[-/1] 22 18 20–25 Ortolan / Cretzchmar’s BunE tm in bg eriza hortulana / caesia both PMs 2 2[2/-] not separable; both scarce 26,26a 18 17–24 Nightingale Luscinia megarhynchos SVf 1 2[1/1] 6,6a 18–19 25–34 Greenfinch Carduelis (Chloris) chloris W Vs 2 2[1/1] + h[-/1] 9,9abcdef 18–20.5 18–29 Chaffinch / Brambling Fringilla coelebs / montifringilla Rs,W Va / W Vs 12+3> 22[12/10] not separable, but Brambling is scarce; [>also pellet samples A1, UL3, D-partial 2017] 10 18.5 16–25 Orphean Warbler Sylvia hortensis (S. crassirostris) SVf 1 [1/-] not confirmed to species as no comparative material av H a.i ol lia vbl etoe r, b um u i t s the only member of 21 19 14–23 cf. Olive-tree Warbler Hippolais cf. olivetorum PMr 1 1[-/1] the genus in the size range 29 19+ 35–50 Crossbill Loxia curvirostra W Vr 1 1[-/1] more robust than Greenfinch/sparrow 11,11abc 20 25–35 Red-backed Shrike Lanius collurio PMc 5 9[5/4] also an ulna in floor samples 12,12a 22–22.5 30–40 Woodchat Shrike L. senator SVf 1+1> 2[1/1] also 2 ulnas in floor samples [>also pellet AT1] 27,27a 24 30–45 Wryneck Jynx torquilla PMs 2 2[1/1] + h [1/-] similar to shrike/lark 18 24 46–70 Hawfinch Coccothraustes coccothraustes W Vs 1> 2[1/1] – also 2 skulls & coracoids in floor samples 31 26 37–55 Crested Lark Galerida cristata Rc 1 1[1/-] also ulnas & upper mandible in floor samples 16 26 65–100 Song Thrush Turdus philomelos W Vc 2 3[1/2] also most other long-bones in floor samples 17 27–28 50–90 Starling Sturnus vulgaris W Vs 2 2[1/1] +h[1/-] 15,15abc 29–30 80–125 Blackbird Turdus merula W Vc 5+1> 6[5/1] + h[-/2] also most other long-bones in floor samples [>also pellet sample A1] 19 31 56–79 Golden Oriole Oriolus oriolus PMc 1 1[1/-] + h[-/1] also 2 ulnas, a coracoid & a carpometacarpus in floor samples Total humeri / Skupaj nadlhatnice 86+12 Additional species identified from other bones / Dodatne vrste, določene na podlagi drugih kosti: 2018 ulnas 26–50 Skylark Alauda arvensis W Vf 1 ulna TBT31 18–29 Tree Pipit Anthus trivialis PMc 1+ tibiotarsus Bill 6 22–36 Spanish Sparrow Passer hispaniolensis PMf 1 upper mandible 16 Acrocephal 41 us (184/185): 3–24, 2020 Tabela 5: Ptice kot plen pegaste sove in njihov status na otoku Amorgos iz vzorca posameznih kosti na tleh jame Dhri (lokaliteta D), navedene po velikosti nadlahtnice. Oznake: n[x/y]–skupaj [leva/desna] + h[le glava nadlahtnice], PM – selivka, R – stalnica, SV – polenta obiskovalka, WV – prezimovalka, a – zelo pogosta, c – pogosta, f – manj pogosta, s – redka, r – zelo redka. Podatki o teži iz Swon & Pnire S (1998), le zelenec zaradi napake v tisku iz Pirre S (1987). Podatki o status vrste: ASC, REA in Apostolos Cristopoulos (osebno opazovanje 2007–2019 in Cekeh in sod. v 2020). +X krepko označuje dodatne osebke v izbljuvkih, ki niso vključeni v seštevek. Humerus Size of samples No. whole humeri [L/R] sample Ref./ (mm)/ Weight Min. No. of or heads only h[L/R] / Oznaka vzorc V a elikost vzorca range (g)/ birds / Min. Št. celih ali samo gl A ad v ditional comments [additional bones noted where identified - mostly only from larger species] / nadlahtnice (v mm) razpon teže (g)English name / Angleško imS ecientific name / Znanstveno ime Status št. ptic nadlahtnic Dodatni komentarji 1b 12 6–10 Chiffchaff Phylloscopus collybita PMc, W Vc 1+1> 1>[1/-] confirmed against pellet sample UL3 & NHM collections 1c 12.5 7–12 Willow Warbler P.trochilus PMc 2 2[-/2] consistent differences from Chiffchaff 25,25a 13 8–13 Subalpine Warbler Sylvia cantillans SVc 2 4[2/2] 33 13.5 9–15 Sedge Warbler Acrocephalus schoenobaenus PMs 1 1[-/1] 24 13.5 10–15 Sardinian Warbler Sylvia melanocephala Ra 1 1[1/-] 31 14–14.5 11–18 Siskin Carduelis (Spinus) spinus W V* 0+1 2 [1/1] [from pellet UL3] *irruptive; rare visitor, but common when arrives 32 14.5–15 13–18 Whitethroat Sylvia communis PMf, SVr 2 2 [1/1] + h[-/1] 2,2b 15 10–16 Pied / Collared Flycatcher Ficedula hypoleuca / albicollis both PMc 4+3> 5[3/2] + h[1/-] not separable; both common; close to St Fo .se nm ec ito hra qt u; ata also possible] [also pellets D3, D12, D16] 20,20a 15 13–17 Stonechat Saxicola (torquata) rubicola Rs, W Vf 3+1> 2[2/-] +h[1/-] [>also pellet D28] 13 15 16–22 House Martin Delichon urbicum PMa 1 2 [1/1] 4 16 14–19 Goldfinch Carduelis carduelis Rs,W Vf 1 2[1/1] 14 16 11–14 Serin Serinus serinus W Vs 1 1[-/1] 28 16 14–20 Spotted Flycatcher Muscicapa striata PMa 1 1[1/-] very close to Whinchat & similar also to R Phoe eni ds cu tr a u r s t phoenicurus 3abcefgh 16–17 14–19 Whinchat Saxicola rubetra PMc 8 15[8/7] close to Spotted Flycatcher 23 16.5 Robin Erithacus rubecula W Vc 1 1[1/-] 5,5abcde 16–18 16–25 Blackcap / Garden Warbler Sylvia atricapilla / borin PMa, W Vf / PMr 11+1> 15[5/10] +h[1/1] not separable, but Garden Warbler is very scarce [>also pellet sample AT4] 30 [15–17] 16–25 Dunnock Prunella modularis W Vr 1 h[1/-] head of bone only 4a 17 15–22 Linnet Carduelis (Linaria) cannabina Rc, W Vc 1 2[1/1] 7,7a 18 13–19 Black Redstart Phoenicurus ochruros W Vc 2 3[2/1] 8 18 24–38 House Sparrow Passer domesticus Ra 2 2[1/1] + h[-/1] 22 18 20–25 Ortolan / Cretzchmar’s BunE tm in bg eriza hortulana / caesia both PMs 2 2[2/-] not separable; both scarce 26,26a 18 17–24 Nightingale Luscinia megarhynchos SVf 1 2[1/1] 6,6a 18–19 25–34 Greenfinch Carduelis (Chloris) chloris W Vs 2 2[1/1] + h[-/1] 9,9abcdef 18–20.5 18–29 Chaffinch / Brambling Fringilla coelebs / montifringilla Rs,W Va / W Vs 12+3> 22[12/10] not separable, but Brambling is scarce; [>also pellet samples A1, UL3, D-partial 2017] 10 18.5 16–25 Orphean Warbler Sylvia hortensis (S. crassirostris) SVf 1 [1/-] not confirmed to species as no comparative material av H a.i ol lia vbl etoe r, b um u i t s the only member of 21 19 14–23 cf. Olive-tree Warbler Hippolais cf. olivetorum PMr 1 1[-/1] the genus in the size range 29 19+ 35–50 Crossbill Loxia curvirostra W Vr 1 1[-/1] more robust than Greenfinch/sparrow 11,11abc 20 25–35 Red-backed Shrike Lanius collurio PMc 5 9[5/4] also an ulna in floor samples 12,12a 22–22.5 30–40 Woodchat Shrike L. senator SVf 1+1> 2[1/1] also 2 ulnas in floor samples [>also pellet AT1] 27,27a 24 30–45 Wryneck Jynx torquilla PMs 2 2[1/1] + h [1/-] similar to shrike/lark 18 24 46–70 Hawfinch Coccothraustes coccothraustes W Vs 1> 2[1/1] – also 2 skulls & coracoids in floor samples 31 26 37–55 Crested Lark Galerida cristata Rc 1 1[1/-] also ulnas & upper mandible in floor samples 16 26 65–100 Song Thrush Turdus philomelos W Vc 2 3[1/2] also most other long-bones in floor samples 17 27–28 50–90 Starling Sturnus vulgaris W Vs 2 2[1/1] +h[1/-] 15,15abc 29–30 80–125 Blackbird Turdus merula W Vc 5+1> 6[5/1] + h[-/2] also most other long-bones in floor samples [>also pellet sample A1] 19 31 56–79 Golden Oriole Oriolus oriolus PMc 1 1[1/-] + h[-/1] also 2 ulnas, a coracoid & a carpometacarpus in floor samples Total humeri / Skupaj nadlhatnice 86+12 Additional species identified from other bones / Dodatne vrste, določene na podlagi drugih kosti: 2018 ulnas 26–50 Skylark Alauda arvensis W Vf 1 ulna TBT31 18–29 Tree Pipit Anthus trivialis PMc 1+ tibiotarsus Bill 6 22–36 Spanish Sparrow Passer hispaniolensis PMf 1 upper mandible 17 A. S. Cheke, J. P. Hume: The diet, and pellet residue taphonomy, of Barn O Tytow all bs a on a Greek island reveals an exceptional diversity of avian prey Table 6. Avian prey on Mediterranean islands compared (see text for references). Crete is included to compare a large island on the same migration route as Antikythera and the Cyclades (Milos, Amorgos). Code: dno – does not occur (on specie fi d island). NB: 2 more species in the Ibiza islands were unidentie fi d – these could overlap with identie fi d species (different bones?) or could add a further 2+ species to the combined list. Equally in Amorgos there are probably unidentie fi d further species amongst harder-to-identify bones not checked in detail (ulna, femur etc). References: A – this paper, B – Obuch & Benda 2009, C – Alivizatos & Andriopoulos 2016, D – Guerra et al. (2014), E – Sommer et al. (2005). Notes: 1 – Lanius sp., 2 – Turdus/Sturnus, 3 – Sylvia sp., 4 – Ficedula sp. in Antikythera, as ‘Muscicapidae small size’ in Ibiza, 5 – Turdus sp. (large size), 6 – Emberiza sp. (uncertain), 7 – At San Carlos there were also numerous unidentie fi d bird remains, but they were not sorted by bone type, 8 – fledgling, 9 – Phylloscopus sp., 10 – 28 is combined total for s’Espalmador & Formentera. Tabela 6: Primerjava ptic kot plena na sredozemskih otokih. Kreta je vključena kot primerjava velikega otoka na isti selitveni poti kot otoki Antikitera in Kikladi (Milos, Amorgos). Oznake: dno – vrsta se ne pojavlja. Dve vrsti ptic na Ibizi nista bili določeni, kar se lahko prekriva z že določenimi vrstami (različne kosti) ali pa predstavljata dve dodatni vrsti. Podobno velja za Amorgos, kjer bi med težko določljivi kostmi verjetno našli še kakšno dodatno vrsto. Viri: A – ta raziskava, B – Obuch & Benda (2009), C – Alivizatos & Andriopoulos (2016), D – Guerra et al. (2014), E – Sommer et al. (2005). Opombe: 1 – Lanius sp., 2 – Turdus/Sturnus sp., 3 – Sylvia sp., 4 – Ficedula sp. na Antikiteri in majhna Muscicapidae na Ibizi, 5 – večja vrsta Turdus sp., 6 – Emberiza sp. (negotovo); 7 – v San Carlosu so bili številni nedoločeni ostanki ptic, ki pa niso bili razvrščeni po tipu kosti, 8 – mladič., 9 – Phylloscopus sp., 10 – 28 je skupno število vrst v Espalmador & Formentera. Species / Vrsta Lokaliteta Reference / Vir A B C B D D E Storm-petrel Hydrobates pelagicus dno dno dno dno * Common Kestrel Falco tinnununculus *8 Chukar Partridge Alectoris chukar, chick * Common Quail Coturnix coturnix * Rail sp. Porzana sp. dno * Kentish Plover Charadrius alexandrinus dno * Scops Owl Otus scops * Curlew sandpiper Calidris ferruginea dno * ‘shank ’ Tringa sp. * marsh tern Chlidonias sp. dno * Rock Dove Columba livia * Turtle-dove Streptopelia turtur * Collared-dove Streptopelia decaocto * Bee-eater Merops apiaster * Wryneck Jynx torquilla * * * Crested Lark Galerida cristata * * * Skylark Alauda arvensis * * Short-toed Lark Calandrella brachdacryla * * Barn Swallow Hirundo rustica * * House Martin Delichon urbica * Tree Pipit Anthus trivialis * Amorgos Antikythera Milos Crete Ibiza s’Espalmador Ibiza Formentera Ibiza San Carlos Acrocephal 41 us (184/185): 3–24, 2020 Continuation of Table 6 / Nadaljevanje tabele 6 Species / Vrsta Lokaliteta Reference / Vir A B C B D D E Meadow Pipit Anthus pratensis * Wren Troglodytes troglodytes * Woodchat Shrike Lanius senator * (*1) (*1) * Red-backed Shrike Lanius collurio * (*1) (*1) * Golden Oriole Oriolus oriolus * Starling Sturnus vulgaris * *2 Great Tit Parus major dno dno * Dunnock Prunella modularis * Blackcap / Garden Warbl Se yl rvia atricapilla / borin * *3 *3 * *3 *3 Whitethroat Sylvia communis * Subalpine Warbler Sylvia cantillans * Orphean Warbler Sylvia hortensis (S. crassirostris) * Sardinian Warbler Sylvia melanocephala * * Sedge Warbler Acrocephalus schoenobaenus * cf. Olive-tree Warbler Hippolais olivetorum * dno dno dno Chiffchaff Phylloscopus collybita * * (*9) Willow Warbler Phylloscopus trochilus * *9 (*9) Gold- / Firecrest Regulus sp. * Pied / Collared Flycatch F e ic redula hypoleuca / albicollis * *4 * * Spotted Flycatcher Muscicapa striata * * Whinchat Saxicola rubetra * Stonechat Saxicola torquata (S.rubicola) * Wheatear sp. Oenanthe sp. * Black Redstart Phoenicurus ochrurus * * * * Robin Erithacus rubecula * * * * * * Nightingale Luscinia megarhynchos * Blue Rock-thrush Monticola solitarius * Blackbird Turdus merula * * (*5) * Songthrush Turdus philomelos * * House Sparrow Passer domesticus * * * * * * Spanish Sparrow Passer hispaniolensis * Tree Sparrow Passer montanus dno dno * Chaffinch / Brambling Fringilla coelebs / montifringilla * * * * Amorgos Antikythera Milos Crete Ibiza s’Espalmador Ibiza Formentera Ibiza San Carlos A. S. Cheke, J. P. Hume: The diet, and pellet residue taphonomy, of Barn O Tytow all bs a on a Greek island reveals an exceptional diversity of avian prey Continuation of Table 6 / Nadaljevanje tabele 6 Species / Vrsta Lokaliteta Reference / Vir A B C B D D E Hawfinch Coccothraustes coccothraustes * Crossbill Loxia curvirostra * Serin Serinus serinus * * Greenfinch Carduelis (Chloris) chloris * * * * Goldfinch Carduelis carduelis * * * Linnet Carduelis (Linaria) cannabina * * * * Siskin Carduelis (Spinus) spinus * Corn Bunting Emberiza calandra * * Ortolan / Cretzschmar’s Emberiza hortulana / caesia * *6 Bunting No. of species / Št. vrst 39 7 14 15 21 16 7 (7) >28 <(10) autumn. However more striking were t -he d frieffeqr uent component of Barn Owl p Roul rey ( in ences between two pairs (roosts & nest sit& es) o Dn ube ly y 2012), notably in Greece on A - n a few kilometres apart in agricultural habtiitkat yt. O henre a and Milos islA an liziv ds ( atos et al. caught only 3.7% birds, the other 16.5%; in b 20o 05 t, h Alizivatos & Andriopoulos 2016), but case sparrow Pas ss er spp. (mostlP. y domesticus) were no lacertids were recorded for Gr Roul eecin e b y the major component (54–58%), but other sp& ecD ie ube s y (2012). However O buch & Benda re - were not fully identified so the diversity i pos n rteod u t nidentified lacertids from Crete, and known, though a total of 11 species are men L ta ice on rte ad t. rilineata was found in pellets on Milos Given that sparro P. w s do(mesticus, P. montanus, by Alizivatos & Andriopoulos (2016); our P. hispaniolensis) are the avian group that feat Pou dr ae rcs is appear to be the first confirmed wall most commonly in Barn Owl diet studie-s, it i liz s i ar n ds found in Greek Barn Owl pellets. teresting that the Amorgos owls make so littD lu e u ng b se eetles, which are nocturnal, feature of this abundant resource, presumably av qu oiit de r ing egularly ( D een .g. ys et al. 2017), though hunting in and around the village despi- te t th he p eir t rox otal contribution is small. imity of the caves to Langada. 4.3. Prey proportions and bone sur vival 4.2. Other taxa It is clear from the data that the cave floor held Rodents and shrews are normal compone fn at r m s o o fr e rat and bird bones, and fewer Wood Mice Barn Owl diet (e R .g o. mano et al. 2020) and will and House Mice, than would be expected from the not be discussed further here, although so rat me d ios i at n t a he complete pellets found in the same cave of taphonomic interest may be extracted f(r T o am t ble 3 he ). W hile changes in hunting behaviour over material at a later date (but see below). time cannot be ruled out, we think the explanation Lizards are rarely an important constil ta ur eg ne t o ly l f ies in the relative robustness of b - ones of dif the diet ib( id.), though geckos are a know -n if f eirnent species. Rat bones, even the freq-uent juve Amorgos Antikythera Milos Crete Ibiza s’Espalmador Ibiza Formentera Ibiza San Carlos Acrocephal 41 us (184/185): 3–24, 2020 niles, are bulky, so slower to degrade (thoua gn h m d t ah ne r y everse effect in rat and mouse bones – are eroded), while the bird bones found a - re n prootpo icretionately far fewer rat bones in the ably better preserved overall in this envisr eo dn im me en nt t than in pellets, and fa Mu r m s b or on e es than their rodent equivalents, hence bot (o h t f th weo s se pecies in both genera). The authors did taxa will have above-expected proportions n. o H t d ouisse cuss the compositional discrepancies, nor Mice bones seem particularly degradable (f d eiw j d ta h w ey p s ublish the proportions of different bone are intact) while Wood Mice are intermedit ay tp ee , b s w ut ithin totals of mammal and bird taxa. many more are found damaged than those of the larger Rock Mice, as previously n Cho et ke ed ( & 5. Conclusion Ashcroft 2017). Shrew jaws seem robust and are generally in good condition, though t - heO pu rr d opor ata set of 39 species appears to be the most tion in pellets and loose is the same, as it i dis w veris te r h ange of birds for a single site recorded for Rock Mice. In the circumstances of thi -s pB aa rt rin O cu wls anywhere, 51% of the total diversity lar cave, trampling by goats would seem to o b f s e t ma hle l passerines (and Wryneck) recorded for main agent of bone deterioration, followet d b he i y t slh ae nd (77 specCh iese ; ke et al. 2020). While action of goat urine and faeces, aided by ra eat inw in at g m er ainly rodents as elsewhere, Barn Owls on blown in during winter, which probably en xo plratih n-s eastern Amorgos also target an unusually the lack of bones below the surface. Surfaw ce b ide r one as nge of passerine migrants to the island, would rapidly dry out, those below woul - d p1 r 9e % o sum f i a ndividual vertebrate prey items in Dhri bly remain wet and decay. Cave D), although the percentage in fresh pellets In addition to chemical erosion post-eji en t ction he c , ave was only 4%; for all local pellets it was Denys et al (2017) found that small mamm9a% ( l Table 1). Apart from the case discussed in bones were more likely to be more dam Ia big ze ad / Formentera, this breadth of avian prey does by digestion prior to ejection than birdn b oon t se ese, m to occur elsewhere in Europe; indeed in another indication of their greater suscm ep ot si t o bil f m ity a inland Europe birds hardly feature. For to forces of degradation. However they d exia d n mpo le t, t he massive study in FraG nc ué e b rin y separate rodents and shrews in their anal(y 19 s2 is 8; o ) fu or u nd only a handful of birds (Chaffinch, data show shrew bones as being as resist spaan rt a row s Pa s sser sp.) in hundreds of rodent and birds, suggesting contrast with rodents iss s hr hea w i rpt er e ms, though one exceptional site, out of than Denys et al.’s results imply. They also1 9 studied, in the French Pyrenees has yielded a looked at which bones were most likely to s hiu gr h p vie vr ec entage of birds (7 Liboi 7.6%; s 1983), but ‘processing’ by Barn Owls (and other raptot rh s) a e b ni d r,d s were apparently not fully identified to albeit on a much smaller sample than ours sp , c ec aim es e, t hough 77% of them were spPa arsr se o r ws up with quite different proportions (Tablse 4 pp. A b ). ig survey in Britain only 2% by number However they were sampling at nest siteso , w f p hr ee ry w e as birds (Glue 1974); in central Italy Barn Owls dismember prey before feeding i Za t t gor oše k (2018) found no birds at all, though their young, whereas adults feeding theu mp t selv o 3 es% w , ere recorded in the Rome area by as in our Amorgos sites, swallow most prey Sa w lva ho ti l e e t al. (2002). A meta-analysis in Ireland (Bunn et al. 1982, Taylor 1989). In their studyf , ound birds ranged from 0.2% in areas with leg bones were more likely to survive tha bn otw h i inn gv asive shrC ew roci s dura russula and voles bones, in contrast to our data where hum My eroi a desn d (e x- Clethrionomys) glareolus, to 3.5% in ulnae were much more frequent than femu ar a rean s d y et to be reached by these a Smidd nimay l s ( tarsus elements amongst the loose bones. H et e al nc . 2 e i 01 n 8 ); 22 bird species were recorded from 29 abandoned sites or zooarchaeological co sl tl ue d cit eion s. Fu s rther east in mainland E Ou buch rope , th these different proportions may indicate wh ete al t.h ’s m er eta-analysis of 215 1 ce 6ntur y to modern the site was a roosting or nesting area. sites in Slovakia found 53 bird species overall, but In a similar pellet vs sediment study f wirto h l m ow percentages and diversity per site, and Ibiza ( Sommer et al. 2005), no such differencesi n Bulgari M a ilchev et al. (2006) recorded 40 were found in the proportion of bird b spon eceis e, s over 28 sites. 21 A. S. Cheke, J. P. Hume: The diet, and pellet residue taphonomy, of Barn O Tytow all bs a on a Greek island reveals an exceptional diversity of avian prey Overall the low proportion of birds pes r s imipt le i y lse ft avian remains as ‘birds’ with little or no true throughout the world Ro rm an ano ge ( et al. further identification. 2020), however in North Africa, the Middle Ea In a st nalysing bones found loose on the floor and Pakistan birds can form much of the die o tf a r , wito ho sting site of any pellet producing predator, 100% taken in one study in Tunisia (see discuit i ssio s i n mportant to take into account the relative in Bunn et al. 1982: 90). While in Pakistan bird de s terioration rate of bones from die ff rent species, as can make up 30% ( Nadeem et al. 2012), only four our data shows. In our case there is a very signic fi ant species were taken and one (Common Mybniaas i h n favour the avian bones and the largest Acridotheres tristis) predominated; in Algeria up t roo de nt, and this may well be the case elsewhere. The 36 species can be involved (over 19 s Rih it an ese , proportion of die ff rent skeletal elements recovered, 2003, 2004), though House Sparrows made u thp e ‘ taphonomic signat Pu ok re in ’ ( es & Peterhans 71% of the avian catch, and 100% in another s 199 t7 u)d , r y arely looked at, may also be signic fi ant for (Hadjoudj et al. 2012). Studies by Obuch & understanding the status of sites, abandoned or Benda (2009) yielded 64 avian prey species spr a en ac di ent, no longer in u Den se ( ys et al. 2017). over 27 sites in several countries in southern Eu Ac ro kp ne o ,w ledgements: ASC particularly thanks Israel and Eg ypt, the maximum numbers at oL no e s nia tïe s J allais for showing him sites A and D, and being 20 (perhaps up to 25) in Isr O a buch el; (2018) sharing his observations on local Barn Owls, Linda found site maxima of 16 and 19 in sites in JoL rd oa sint,o for identifying beetle remains, and Ruth though the total over 10 sites in that couAnsthrcy w roft f aso r helping to scour the cave floor for 43 species. We have not been able to revieb w t one h se ; h e is also indebted to the late Michael Rudd whole enormous literature on Barn Owl dwieht ( osse a ee ncient village house he and R -uth Ash Romano et al. 2020), but have attempted to finc dr oft had access to in Amorgos. We thank Joanne studies where birds have featured signic fi a C no to ly t per a o nd Judith White for access to the skeleton compare with ours. collection in the Natural History Museum, Tring, Clearly Barn Owls are adaptable a UKn , ad n d two anonymous referees whose comments opportunistic, and have been recorded els le ew d t he o u re seful untapped literature and a refinement targeting wintering thrushes in communao lf t ro h o e p sta s, p er. Also published after acceptan - ce, Rou and roosting or urban aggregations of sl pia nr 's ( ro 2w 02s0, ) new monograph, which, while c - ompar hirundines, starlings, finches, larks, bun intg B ing asr n Owl diets worldwide, does not drill down and mynas M ( ilchev et al. 2006, Rahine 2003, into the kind of detail we present here. 2004, Nadeem 2012). Chaffinches and possibly thrushes and Corn Buntings appear to hav Pe s ostu scc rh ip t roosts in Amorgos (ASC pers. obs.), but most of the species taken do not roost commS uinnacle lyt. his paper was submitted, reviewed and It would appear that in Amorgos the owa lc s fi cn ed pt ed, an important paper with direct bearing many migrants an easy target, though the v on o ar uir r ety e sults has been published. In a meta-analysis remains unexplained; however the accumu cl oa vt eion rin g Mediterranean islands from Sardinia down on the cave floor may represent a longer t to t imien y Espartar (0.2 ha, BaleJa an ri že cs k) ovič series, and thus more opportunity, than in o & K tl h en er ov šek (2020) found that, in addition to reported studies. a general trend of birds and reptiles being taken Our study, and likewise those of otherm s c orie t ted han on mainland Europe, “diet diversity with high bird catches, shows that Barn Ow w l p as g ellr ee tat s er on the larger Mediterranean islands. should be included when investigating b - irH d d ow ise t v r eir, a more diverse diet did not mean a higher bution and potentially also abundance, in a nd ud m ib te io r o n f taxa, but a wider range of abundant and to being a well-established method of sa em vepn li ln y r g epresented taxa. The smaller the islands, the mammals. Furthermore, and to facilitate s mo im ri e b lair rds and reptiles were consumed, compared studies in the future, it is hoped that the pt ho h oto ig s o he f r proportions of mammals on the larger humeri will aid others in identifying bir-ds iisl n E au nr do s. These findings support the idea of barn pean owl pellets – many previous researche or w s h lsa ’ f ve e eding flexibility and opportunistic predator 22 Acrocephal 41 us (184/185): 3–24, 2020 Brown R., Ferguson J., Lawrence M., Lees D. (1987): behaviour”. The results were largely independent Tracks signs of the birds of Britain and Europe. An of human urbanisation and island isolation. identification gui– d A e. C Black/Christopher Helm, Our results emphasize this pattern, and as Tina London. Klenovšek commented to ASC (email 21.09.2020) Bunn D. S., Warburton A. B., Wilson T. D. S. (1982): “What a coincidence. I am really sorry we could not The Barn Owl. – T&AD Poyser, London. use your results in our meta-analysis. They would fit Cheke A. S., Ashcroft R. E. (2016 [‘2015’]): Vocal ga- th so well into our findings”. erings of Blackc Sa ylpvs ia atricapilla in olive trees on a Greek island. - Rivista Italiana di Ornitologia – Research in Ornitholog y 85(2): 36–37. 6. Povzetek Cheke A. S., Ashcroft R. E. (2017): Mammals and b-ut terflies new to Amorgos (Kiklades), with notes on Raziskani so bili izbljuvki pegasT tyto e s a olv ba e in reptiles and amphibia –n P sa . rnassiana Archives 5: njihovi ostanki (posamezne kosti) na jamskih tleh 11–27. otoka Amorgos (Kikladi, Grčija). Sove so uplenile Cheke A. S., Tsiakiris R., Christopoloulos A., vsaj 39 vrst ptic, kot je bilo ugotovljeno na po Ad shcr lag oft i R. E. (2020): The breeding birds of a small Aegean island (Amorgos, Cyclades, Greece), najdenih nadlahtnic. 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Journal
Acrocephalus – de Gruyter
Published: Nov 1, 2020