Fowling during the Gravettian: the avifauna of Pavlov I, the Czech Republic

This article appeared in a journal published by Elsevier. The attached

copy is furnished to the author for internal non-commercial research

and education use, including for instruction at the authors institution

and sharing with colleagues.

Other uses, including reproduction and distribution, or selling or

licensing copies, or posting to personal, institutional or third party

websites are prohibited.

In most cases authors are permitted to post their version of the

article (e.g. in Word or Tex form) to their personal website or

institutional repository. Authors requiring further information

regarding Elsevier’s archiving and manuscript policies are

encouraged to visit:

http://www.elsevier.com/copyright

http://www.elsevier.com/copyright

Author's personal copy

Fowling during the Gravettian: the avifauna of Pavlov I, the Czech Republic

Zbigniew M. Bochenski a,*, Teresa Tomek a, Jaros1aw Wilczynski a, Jiri Svoboda b, Krzysztof Wertz a,Piotr Wojtal a

a Institute of Systematics and Evolution of Animals, Polish Academy of Sciences. Sławkowska 17, 31-016 Krakow, PolandbDepartment of Palaeolithic and Palaeoethnology, Institute of Archaeology, ASCR, 69201 Dolnı Vestonice 25, Czech Republic

a r t i c l e i n f o

Article history:

Received 20 May 2009

Received in revised form

5 August 2009

Accepted 11 August 2009

Keywords:

Bird remains

Gravettian

Upper Palaeolithic

Hunting

Broad spectrum revolution

Corvus corax

Lagopus lagopus

Lagopus muta

Tetrao tetrix

a b s t r a c t

This is the first article that describes in detail the bird remains from the Gravettian mega-site Pavlov I in

the Czech Republic. More than 1000 bird bones represent at least 19 taxa, of which the most numerous

are tetraonids including black grouse (Tetrao tetrix), willow grouse (Lagopus lagopus) and ptarmigan

(Lagopus muta), and ravens (Corvus corax). The archaeological and biological contexts indicate that most

birds were hunted by people in the vicinity of the site, possibly with the help of knotted nets made from

plant fibres. We suggest that ravens were killed while feeding on carcasses and/or food remains that may

have been disposed of intentionally, and feathers may have been used for arrow fletching. Human-

modified bones indicate that birds were used not only for food but also as raw material for tools and

decoration. Although mammals were certainly more important in the subsistence of the Gravettian

people, it is clear that birds played a role in their culture. The location of Pavlov I enlarges the explanatory

scope of the Broad Spectrum Revolution hypothesis to include higher latitudes north of the

Mediterranean.

� 2009 Elsevier Ltd. All rights reserved.

1. Introduction

The open-air loess site of Pavlov I, situated in the Pavlovian Hills

in the region of Moravia, the Czech Republic, is one of the largest

and most important Upper Palaeolithic sites in Europe. It is part of

a large complex of Gravettian sites that extend from the village of

Dolnı Vestonice in the northwest to Pavlov in the southeast. The

site is located on a gentle slope between 190 and 200 m a.s.l.,

several dozen meters above the Dyje river. The 14C dates indicate

a relatively short time-span of occupation between 27 and 25 ka BP,

i.e. in the Evolved Pavlovian stage, an equivalent of the final Inter-

pleniglacial or OIS 3 (Svoboda, 2005). The great advantage of the

site is that it was excavated between 1952 and 1971 by one person,

Bohuslav Klıma, using a single research strategy of investigating the

area in squares measuring 1 or 4 m2.

An analysis of the environmental, archaeozoological, anthro-

pological, and archaeological records has been underway since the

1990s (Svoboda, 1994, 1997, 2005; Trinkaus et al., 2009). Both

pollen and charcoal analyses from the Gravettian cultural layers

show that the landscape was partly covered by woodland areas,

forming a parkland (arboreal pollen usually exceeds 50%), with

conifers such as pine, spruce and larch dominating, but accompa-

nied by deciduous trees such as willow and alder along the

riverbed. The sporadic occurrence of species such as oak, beach, and

yew is also indicated (Damblon, 1997; Mason et al., 1994; Opravil,

1994; Rybnıckova and Rybnıcek, 1991; Svobodova, 1991a,b). Large

meadow-steppe mixed with conifer patches, and subalpine grass

communities on the rocky summits were also present. Studies of

the vertebrate fauna confirm the environmental reconstructions

based on the plant remains (Musil, 1955, 1959, 2005; Nadachowski,

2005).

Pavlov I has been interpreted as a typical example of a large

settlement occupied for a long period, and used repeatedly by

hunters-gatherers, where the settlement levels accumulated as

a thick complex of layers, creating local palimpsests (Novak, 2005:

70). An abundance of artefacts indicate a very intensive occupation

of the site. For instance, the site yielded more than 200,000 stone

artefacts (Verpoorte, 2005), 1000 ceramic fragments (Soffer and

* Corresponding author.

E-mail addresses: [email protected] (Z.M. Bochenski), tomek@

isez.pan.krakow.pl (T. Tomek), [email protected] (J. Wilczynski),

[email protected] (J. Svoboda), [email protected] (K. Wertz), wojtal@

isez.pan.krakow.pl (P. Wojtal).

Contents lists available at ScienceDirect

Journal of Archaeological Science

journal homepage: ht tp: / /www.elsevier .com/locate/ jas

0305-4403/$ – see front matter � 2009 Elsevier Ltd. All rights reserved.

doi:10.1016/j.jas.2009.08.002

Journal of Archaeological Science 36 (2009) 2655–2665


Vandiver, 2005) and more than 600 bone, antler and ivory tools

(Bruhl, 2005). Current fieldwork focuses on the smaller ‘‘satellite’’

camps in the vicinity, which are characterised by having a more

restricted fauna (Pavlov VI; Svoboda et al., 2009).

The site of Pavlov I also yielded a large number of animal

remains (Musil, 1955, 1959, 1994, 1997). At present, more precise

data on faunal remains are available only for the material excavated

between 1952 and 1956 in the southeastern part of Pavlov I. This

area covers about a half the sitewhere some 46,000mammal bones

were found. The most numerous remains belonged to game

species, including hare (Lepus sp.), reindeer (Rangifer tarandus),

mammoth (Mammuthus primigenius), and horse (Equus sp.). A

characteristic feature of this archaeological site is the large number

of carnivores. These comprise nearly 40% of all mammal remains –

the most numerous being wolf (Canis lupus), red fox (Vulpes vulpes)

and Arctic fox (Alopex lagopus). Other less commonmammals found

at the site include the wolverine (Gulo gulo), cave lion (Panthera

spelaea), cave bear (Ursus spelaeus) and brown bear (Ursus arctos)

(Musil, 2005). The southeastern part of Pavlov I also yielded about

two dozen remains of small mammals including rodents (Nada-

chowski, 2005). The presence of all parts of their skeletons indi-

cated that most of the hunted herbivores and carnivores were

brought to the site and dismembered there. However, the over-

representation of foot bones of both cave lion and cave bear indi-

cates that their skins rather than whole carcasses were brought to

the site.

Mammal bones modified by people make up less that 2% of all

the remains. Of the 16,000 bones studied from the 1954 to 1956

excavations, 299 fragments had some traces of human activity such

as cut and burn marks. Human related marks were found on bones

representing all size categories: relatively small species (hare and

fox), medium size species (wolf and reindeer) and large mammals

(horse andmammoth). It is noteworthy that the marks observed on

the bones indicate that the Gravettian people hunted large and

medium-sizemammals for their skin andmeat (Wojtal et al., 2005).

Zooarchaeological studies in Pavlov I have so far focused on

mammals, with birds generally being treated superficially. Musil

(1958,1959) states that birds constitute some 8–9% of the total faunal

assemblage, and include raven (Corvus corax), black grouse (Lyrurus

tetrix), and partridge (Alectoris sp.) (see also Musil, 1994, 1997, 2003,

2005;Wojtal et al., 2005).Our aimhere is tofill the gap in information

about the birds and provide insights into Gravettian use of birds.

2. Material and methods

This study includes all avian remains found in the entire area of

Pavlov I during the excavations undertaken between 1952 and 1971.

Althoughwedonotknowwhether thematerialwas sieved, the large

number of very small objects recovered (e.g. isolated teeth of small

mammals, some fifteen hundred phalanges of hare and fox) indicate

that the excavations were carried out carefully and that the remains

available for study are representative. Due to the lack of precise data

we cannot exclude the possibility that the bird remains come from

two or more facies. Therefore they are interpreted here as a single

assemblage. The material belongs to the Moravian Museum (Brno,

Czech Republic) and is housed in Budisov castle.

The bird bones were identified by Teresa Tomek and Krzysztof

Wertz, based on comparisons with modern skeletons in the

collection of the Institute of Systematics and Evolution of Animals,

Polish Academy of Sciences, and with the help of available manuals

(Erbersdobler, 1968; Kraft, 1972; Tomek and Bochenski, 2000). For

practical reasons, in the case of body part representation we

combined the few taxa of the Tetraonidae family (black grouse

Tetrao tetrix, willow grouse Lagopus lagopus, ptarmigan Lagopus

muta, Lagopus sp. and Tetraonidae indet.) into one group

‘‘tetraonids’’, this being justified on morphological and ecological

grounds (Cramp and Simmons, 1980; Erbersdobler, 1968; Kraft,

1972). However, we analysed the remains of the raven (Corvus

corax) separately from other species of the Corvid family present at

Pavlov I, i.e. magpie (Pica pica), jay (Garrulus glandarius), chough

(Pyrrhocorax pyrrhocorax), and jackdaw (Corvus monedula) because

ravens were easily distinguished by their large size. We examined

the bone assemblage for element frequency, cut marks, traces of

burning, perforation of the distal humerus due to the overextension

of the elbow joint (Laroulandie, 2005b; Laroulandie et al., 2008),

predator activity marks including perforations made with beaks or

claws (Bochenski et al., 2009), root etching and weathering

(Bochenski and Tomek, 1997).

The results are presented in three different ways – as Number of

Identified Specimens (NISP), Minimum Number of Individuals

(MNI), and Minimum Number of Elements (MNE). The MNE was

calculated for each major type of bone only, i.e., omitting small

elements such as phalanges or vertebra. The MNE is the sum of

complete bones (left and right) and proximal (left and right) or

distal (left and right) parts – whichever is more numerous (Lyman,

1994, 2008).

The ratio of wing-to-leg elements was calculated as the number

of wing fragments (humerusþ ulnaþ carpometacarpus) divided by

the sum of wing and leg fragments (femurþ tibiotarsusþ tarsome-

tatarsus), and then expressed as a percentage (Ericson, 1987;

Livingston, 1989). The calculations were based on NISP and MNE.

A Chi-square test was used to evaluate the significance of differ-

ences in the proportion of wing and leg elements.

3. Results

The avian remains from Pavlov I include 1045 fragments rep-

resenting at least 19 taxa (Table 1, Fig. 1). Members of two families –

Tetraonidae and Corvidae – clearly predominate, comprising

respectively 48.8 and 45.8% of the total NISP, or 52.0 and 29.9% of the

Table 1

Avian species found at Pavlov I.

Taxon NISP MNI

N % N %

1 Cygnus cygnus Whooper swan 2 0.2 1 1.3

Cygnus cygnus/Cygnus olor Whooper/Mute swan 5 0.5 1 1.3

2 Cygnus columbianus Bewick’s swan 6 0.6 2 2.6

3 Anas querquedula Garganey 1 0.1 1 1.3

4 Asio flammeus Short-eared owl 1 0.1 1 1.3

5 Haliaetus albicilla White-tailed eagle 2 0.2 1 1.3

6 Gyps fulvus Griffon vulture 2 0.2 1 1.3

Accipitridae indet. 1 0.1 – 0.0

7 Falco cf. tinnunculus cf. Kestrel 1 0.1 1 1.3

8 Falco cf. peregrinus cf. Peregrine 2 0.2 1 1.3

9 Lagopus lagopus Willow grouse 312 29.9 31 40.3

10 Lagopus muta Ptarmigan 21 2.0 4 5.2

Lagopus sp. 58 5.6 – 0.0

11 Tetrao tetrix Black grouse 62 5.9 5 6.5

Tetraonidae indet. (Tetrao tetrix/Lagopus sp.) 56 5.4 – 0.0

12 Perdix perdix Partridge 1 0.1 1 1.3

13 Charadriiformes indet. (size Larus) 1 0.1 1 1.3

14 Turdus sp. Thrush indet. 1 0.1 1 1.3

15 Passeriformes indet. (size Emberiza) Perching birds 1 0.1 1 1.3

16 cf. Pica pica cf. Magpie 1 0.1 – 0.0

Pica pica/Garrulus glandarius Magpie/Jay 1 0.1 1 1.3

17 Pyrrhocorax pyrrhocorax Chough 7 0.7 1 1.3

Pyrrhocorax/Corvus monedula Chough/Jackdow 9 0.9 1 1.3

18 Corvus monedula Jackdow 12 1.1 2 2.6

small Corvidae 3 0.3 – 0.0

19 Corvus corax Raven 445 42.6 18 23.4

Total identifiable 1014 97.0 77 100

Aves indet. 31 3.0

Total 1045 77 100

Z.M. Bochenski et al. / Journal of Archaeological Science 36 (2009) 2655–26652656


MNI. Tetraonids are represented by three species of similar size – the

willowgrouse,ptarmigan, andblackgrouse,whereas corvid remains

are dominated by those of the raven with jackdaw, chough,

magpie, and jay being less numerous. Thirteen out of the 14

specimens of anseriforms recovered from Pavlov I belong to two

(or three?) species of swans (Cygnus sp.), whereas only one bone

represents a duck – the garganey (Anas querquedula). Diurnal birds

of prey are represented by at least four species (NISP¼ 8, MNI¼ 4),

including small hawks and a large eagle and a vulture. The

remaining species, represented by single specimens, include

a short-eared owl, a charadriiform, a thrush, and a small passerine.

With the exception of the head (skull and mandible) all

elements of the skeleton were found but their relative abundance

varies (Table 2, Fig. 2). Fragments of the trunk (sternum and pelvis)

are scarce, whereas those of long bones prevail both in tetraonids

and in the raven. Many small elements including vertebra and wing

and leg phalanges were also recovered.

When the results are expressed in NISP, the differences between

the frequencies of particular elements are larger than in MNE

(Tables 2 and 3). For raven, the highest values of the NISP were

obtained for the ulna (17.8%) followed by coracoid, carpometa-

carpus, tarsometatarsus, and leg phalanges (10.1% each). In tet-

raonids, two bones (ulna and coracoid) scored the highest NISP

(17.3% and 17.1%, respectively) followed by tarsometatarsus (13.0%),

humerus (11.4%) and carpometacarpus (11.4%).

With raven, values of the MNE for six out of nine long bones are

very similar and range between 12.1% and 14.6%; they are some-

what smaller only for the humerus and tibiotarsus (7.3% each) and

radius (4.9%) (Table 3). In tetraonids the most numerous element in

terms of the MNE was the coracoid (21.6%) followed by humerus,

ulna, carpometacarpus, and tarsometatarsus (ranging between 13.0

and 14.7%), with the remaining elements scoring below ten percent.

In tetraonids, wing elements represent 59.5% of the sum of the

wing and leg elements in terms of NISP, and 58% in terms ofMNE. In

both cases the deviation from the expected 50% (1:1 proportion)

was statistically significant (c2¼12.32, P< 0.01 and c2¼ 4.98,

P< 0.05, respectively). But such differences were not observed in

the case of ravens where the wing-to-leg ratio was approximately

1:1 both in terms of NISP (c2¼ 2.99, P> 0.05), and in terms of MNE

(c2¼ 0.03, P> 0.05).

Apart from fragmentation, the material is relatively well

preserved. No damage attributable toweathering, animal predation

or perforation of the distal humerus due to the overextension of the

elbowwas found, and the incidence of root etching on bone surfaces

is low. Yet thedegree of fragmentation is high–of the924 specimens

of long bones recovered, only 51 (5.5%)were complete (Table 4). The

percentages of proximal and distal ends (data pooled for all long

bones: Table 4, bottom row)were very similarwithin both groups of

birds (tetraonids and raven). Shafts (i.e., fragmentswithout articular

ends) were also relatively numerous. In both groups of birds

(tetraonids and ravens) the scapular part of the coracoid greatly

outnumbered its sternal end, and the distal end of the tibiotarsus

clearly predominated over the proximal end (Table 4). Similarly

large differences between the frequencies of articular ends were

observed in two other elements only in the case of raven, where the

distal parts of the humerus and tarsometatarsus were more than

twice as numerous as their proximal ends.

Direct traces of human activity were found on only a few bones.

Examples include one partly burned distal carpometacarpus of

Lagopus sp., a distal humerus of a black grouse with a cut mark

(Fig. 3), and a humerus of awhooper swan (Cygnus cygnus) with cut

marks on the anterior side of its proximal endwhichwe attribute to

carcass dismembering (Fig. 4).

Three other bones – all ulnae – were intentionally modified by

humans.Oneof them, identified as raven, is a 56.5-mmlong fragment

Fig. 1. Simplified spectrum of the birds of Pavlov I, based on Number of Identified

Specimens (NISP¼ 1045) and Minimum Number of Individuals (MNI¼ 77). Tetraonids

include Tetrao tetrix, Lagopus lagopus and Lagopus muta. For details see Table 1.

Table 2

Number of identified specimens found at Pavlov I. Category ‘‘other prey’’ includes

the remaining taxa listed in Table 1.

Element Corvus

corax

Tetraonids Other prey Total

NISP % NISP % NISP % NISP %

Quadratum 3 0.7 – – – – 3 0.3

Vertebra 10 2.2 17 3.3 6 6.5 32 3.1

Sternum – – 2 0.4 – – 2 0.2

Coracoid 45 10.1 87 17.1 4 4.3 136 13.0

Scapula 29 6.5 23 4.5 1 1.1 53 5.1

Furcula – – 3 0.6 – – 3 0.3

Humerus 31 7.0 58 11.4 8 8.7 97 9.3

Ulna 79 17.8 88 17.3 14 15.2 181 17.3

Radius 19 4.3 26 5.1 3 3.3 48 4.6

Carpometacarpus 45 10.1 58 11.4 5 5.4 108 10.3

Wing phalanges 7 1.6 2 0.4 1 1.1 10 1.0

Os carpi ulnare 3 0.7 – – – – 3 0.3

Pelvis 2 0.4 1 0.2 1 1.1 4 0.4

Femur 43 9.7 35 6.9 2 2.2 80 7.7

Tibiotarsus 38 8.5 38 7.5 10 10.9 86 8.2

Fibula 1 0.2 – – 1 1.1 2 0.2

Tarsometatarsus 45 10.1 66 13.0 13 14.1 124 11.9

Pedal phalanges 45 10.1 5 1.0 11 12.0 61 5.8

Element indet. – – – – 12 13.0 12 1.1

Total 445 100 509 100 92 100 1045 100

Z.M. Bochenski et al. / Journal of Archaeological Science 36 (2009) 2655–2665 2657


of an ulna shaftwith polished ends (Fig. 5). Another one is a complete

right ulna of Bewick’s swan (Cygnus columbianus), with five distinct

cuts forming a design on the ventral side of the proximal end (Fig. 6).

The third specimen is a 25-cm long fragment of a left ulna of griffon

vulture (Gyps fulvus), with two groups of cuts at its proximal section.

One group consists of eight clear cuts on the ventral side of the bone,

and theotherconsistsof18slight incisionson thedorsal side; allmade

at relatively regular intervals (Fig. 7a and b).

4. Discussion and comments

The archaeological background of Pavlov I, which is an open,

semi-permanent or perhaps even permanent ‘‘mega-site’’ settle-

ment, strongly implies that the accumulation of faunal remains is

the result of deliberate human selection. Caves and rockshelters,

Fig. 2. Frequency of particular elements expressed in %NISP. CMC¼ carpometacarpus; Os c. uln.¼ os carpi ulnare; TBT¼ tibiotarsus; TMT¼ tarsometatarsus. For details see Table 2.

Table 3

Minimum number of major elements (MNE) found at Pavlov I. Category ‘‘other prey’’

includes the remaining taxa listed in Table 1.

Element Corvus corax Tetraonids Other prey

MNE % MNE MNE % MNE MNE

Coracoid 26 12.6 63 21.6 3

Scapula 28 13.6 22 7.5 1

Humerus 15 7.3 31 10.6 4

Ulna 28 13.6 43 14.7 8

Radius 10 4.9 14 4.8 1

Carpometacarpus 29 14.1 38 13.0 4

Femur 25 12.1 17 5.8 2

Tibiotarsus 15 7.3 25 8.6 7

Tarsometatarsus 30 14.6 39 13.4 10

Total 206 100.0 292 91.5 40



Fig.3.Distal

humerusoftheblack

grouse

(Tetraotetrix)

with

acu

tmark

on

the

poste

riorsid

e.

Table 4

Fragmentation of long bones found at Pavlov I. Category ‘‘other prey’’ includes the remaining taxa listed in Table 1.

Element Corvus corax Tetraonids Other prey Total

N Whole

bone (%)

Proximal

part (%)

Distal part

(%)

Shaft

(%)

N Whole

bone (%)

Proximal

part (%)

Distal part

(%)

Shaft

(%)

N Whole

bone (%)

Proximal

part (%)

Distal part

(%)

Shaft

(%)

N Whole

bone (%)

Proximal

part (%)

Distal part

(%)

Shaft

(%)

Coracoid 45 13.3 51.1 22.2 13.3 87 5.7 66.7 12.6 14.9 4 50.0 25.0 25.0 – 136 9.6 60.3 16.2 14.0

Scapula 29 – 96.6 – 3.4 23 – 100 – – 1 – 100 – – 53 – 98.1 0.0 1.9

Humerus 31 – 19.4 48.4 32.3 58 – 37.9 53.4 8.6 8 – 50.0 37.5 12.5 97 – 33.0 50.5 16.5

Ulna 79 – 19.0 35.4 45.6 88 2.3 47.7 39.8 10.2 14 14.3 21.4 42.9 21.4 181 2.2 33.1 38.1 26.5

Radius 19 – 26.3 42.1 31.6 26 – 53.8 34.6 11.5 3 – – 33.3 66.7 48 – 39.6 37.5 22.9

Carpometacarpus 45 11.1 40.0 24.4 24.4 58 19.0 46.6 25.9 8.6 5 20.0 60.0 20.0 – 108 15.7 44.4 25.0 14.8

Femur 43 7.0 32.6 41.9 18.6 35 2.9 31.4 45.7 20.0 2 – 50.0 50.0 – 80 5.0 32.5 43.8 18.8

Tibiotarsus 38 – 10.5 39.5 50.0 38 – 15.8 65.8 18.4 10 – 10.0 70.0 20.0 86 – 12.8 54.7 32.6

Tarsometatarsus 45 8.9 20.0 57.8 13.3 66 13.6 28.8 45.5 12.1 13 – 23.1 76.9 – 124 10.5 25.0 53.2 11.3

Element indet. – – – – – – – – – – 11 – – – 100 11 – – 0.0 100

Total NISP 374 4.8 32.6 35.0 27.5 479 5.8 46.3 35.9 11.9 71 7.0 23.9 42.3 26.8 924 5.5 39.1 36.0 19.4

Fig.4.Proxim

alhumerusofthewhoopersw

an(Cygn

uscygn

us)

with

cutmark

sonthe

anterio

rsid

e.

Z.M

.Boch

enski

etal./JournalofArch

aeo

logica

lScien

ce36(2009)2655–2665

2659


especially those occupied seasonally by people, are more likely to

have mixed deposits of human debris, non-human prey remains,

and bones derived from animals dying naturally. We would expect

to find numerous remains of rodents and small birds in Pavlov I if

the site had been inhabited also by owls, but this is not the case.

Their absence cannot be attributed to poor recovery because many

very small phalanges of hare and fox are present. Theoretically, the

relatively large birds recovered from the deposits (tetraonids and

ravens) could be food remains of a large predator such as an eagle

or eagle owl, but it is very unlikely that a raptor would chose to live

within a large human settlement and even less likely that it would

not be quickly disturbed or killed by the human inhabitants.

Moreover, we did not notice characteristic perforations of certain

bones which large raptors often make with their beaks and claws

(Bochenski et al., 2009). Mammalian predators can also be ruled out

as accumulating agents because they would destroy most of the

fragile bird bones or at least leave gnaw marks (Andrews, 1990;

Lyman, 1994). Water transport is not an option either because the

site is situated high above the river. However, the species compo-

sition of the birds recovered from the site and the presence of the

aforementioned bones modified by humans, confirm the conclu-

sion reached on the basis of mammalian fauna that the remains

were accumulated by humans (Svoboda, 2005).

The relatively lownumberof identifiedbird species (19) certainly

does not reflect the real diversity of avifauna in the habitat. More-

over, the relative abundance of particular taxa in the assemblage

differs considerable from any known modern avifauna. The over-

representation of one ormore groups of birds (at Pavlov I, tetraonids

and ravens made up more than 90% of the total NISP) is a typical

feature of an assemblage deposited by a selective predator – in this

case humans. In Europe there are many examples of assemblages

attributable to human origin in which one or more bird taxa

predominate. These usually include galliforms (Alectoris, Lagopus,

Tetrao), ducks, bustards, or snowy owls, depending on the locally

available prey (Cassoli and Tagliacozzo, 1997; Koumouzelis et al.,

2001; Laroulandie, 1998, 2003, 2005a; Tagliacozzo and Gala, 2002;

Tyrberg, 1998). Nearby Late Pleistocene sites that contain large

accumulations of tetraonids include Mamutowa Cave, Deszczowa

Cave, Ob1azowa Cave, Krucza Ska1a and Komarowa Cave where

tetraonids account for 40–70%of all avian remains (Bochenski,1981;

Bochenski and Tomek, 2004; Cyrek et al., 2000; Tomek and

Bochenski, 2005; Tomek et al., 2003).

According to Mourer-Chauvire (1983), humeri and femora are

the most numerous elements in human-derived assemblages of

bird bones, whereas in avian derived assemblages tarsometatarsi

and carpometacarpi prevail in middle size prey (e.g. Lagopus), and

tarsometatarsi and coracoids in large prey (e.g. Tetrao). However, as

shown by Bochenski (2005), the problem is more complicated and

the predominance of the humerus is not specific only to materials

deposited by humans. Avian remains from Pavlov I are puzzling

because while on the one hand various elements are well repre-

sented which makes it difficult to apply the criterion of human

versus non-human origin of thematerial, on the other hand, femora

of ravens and tetraonids are somewhat less numerous than several

of the other elements which does not agree with the assumption

that the assemblage includes human food remains. Yet the

contextual data indicate that the birds – in any case tetraonids –

were probably eaten by people. A similar situation where in an

obviously human-derived assemblage femora were less numerous

Fig. 5. Ulna shaft of the raven (Corvus corax), with polished ends. The fragment is

65.5 mm long.

Fig. 6. Complete right ulna of the bewick’s swan (Cygnus columbianus), with five

distinct cuts forming a design on the ventral side of the proximal end.



than other elements was also observed in other archaeological sites

(Guminski, 2005; Steadman et al., 2002), which indicates that the

criterion must be used cautiously. A possible explanation for this

phenomenon has been suggested by Higgins (1999) who claims

that femora are most subjected to destruction in natural as well as

cultural depositions. This is on the one hand because femora

contain large quantities of marrow desirable for human

consumption (Higgins, 1999: 1453–4), and on the other hand, the

bones are more fragile due to their relatively thin cortical walls and

low compactness. However, we are unsure whether the latter

explanation is valid also for galliforms whose leg bones seem to be

particularly sturdy.

Another method often used to distinguish human food remains

from birds which have been subject only to natural decomposition

factors is the ratio of the wing-to-leg elements (Ericson, 1987).

According to Ericson (1987) in assemblages accumulated by people,

bones from the meat-rich leg bones prevail. Although the theory

has many supporters, it has also been partly criticized (Bochenski,

2005; Livingston, 1989; Serjeantson et al., 1993). Recent studies

have shown that the ratio may also be useful in distinguishing

pellet and uneaten food remains of owls and diurnal birds of prey.

Wing bones greatly predominate in uneaten food remains of many

diurnal birds of prey (Bochenski et al., 1999; Bochenski and Torn-

berg, 2003; Bramwell et al., 1987; Laroulandie, 2000, 2002; Mli-

kovsky, 1996) whereas their occurrence in owl and diurnal raptor

pellets is much lower or they are absent (Bochenski et al., 1993,

1998; Bochenski and Nekrasov, 2001; Bochenski and Tomek, 1994).

However, it may occasionally happen that leg elements prevail in

assemblages deposited by raptors as in the case of pellet material

derived from the imperial eagle (Bochenski et al., 1999; Bochenski,

2005) or that wing bones predominate in assemblages deposited by

humans (Guminski, 2005; Livingston, 1989; Serjeantson et al.,

1993). Our results, showing predominance of wing elements in

tetraonids and a nearly equal ratio of wing and leg bones in ravens,

confirm the conclusion reached by Serjeantson et al. (1993) that

anatomical distribution alone cannot provide an answer to the

question of who accumulated the bones – humans or animals.

Although it is an important factor in taphonomic analysis, other

data including the archaeological context may have priority. It is

also true that it is better to use a combination of many criteria to

successfully attribute analysed material to a particular predator or

a group of predators (Bochenski, 2005).

The overrepresentation of wing elements in archaeological sites

is a well-known phenomenon which cannot be explained by one

universal theory (Bovy, 2002; Guminski, 2005). Various factors

including cultural and behavioural as well as post-depositional

modifications play a role in producing this pattern, and it is difficult

to pinpoint the main reason in the case of tetraonid remains at

Pavlov I. While it is very likely that tetraonids and possibly also

ravens were eaten by the Gravettian people, the birds were used

also for other purposes. The few artefacts made from avian bone at

the site are direct evidence that birds were not just a source of food.

Wing bones, especially those of large taxa, were often selected for

tool-making in prehistory (d’Errico et al., 2003; Gal, 2005; Lar-

oulandie, 2003; Wijngaarden-Bakker, 1997), which was also the

case at Pavlov I where all three bones decorated by humans

belonged to large birds. It is also possible that wing feathers were

used as decoration or for utilitarian purposes. However, direct

evidence for the use of feathers in archaeological sites is hard to

obtain and therefore it often remains only a possible option (Bovy,

2002; Mulkeen and O’Connor, 1997; Reichstein and Pieper, 1986;

Reisch, 1976).

The small number of bones with human-induced modifications

is not unusual in materials attributable to human origin. Assem-

blages are known with high frequencies of cut marks and/or traces

of burning (Cassoli and Tagliacozzo, 1997; Laroulandie, 2005a;

Tagliacozzo and Gala, 2002; Reisch, 1976), but in other assemblages

such evidence is scarce even when the sites are unequivocally

cultural in origin (Alhaique et al., 2004; Steadman et al., 2002). In

comparison to large mammals, birds are relatively easy to

Fig. 7. A 25-cm long fragment of a left ulna of the griffon vulture (Gyps fulvus), with two groups of cuts at its proximal section. One group consists of 18 light cuts on the dorsal side

of the bone (a), and the other consists of eight clear cuts on the ventral side (b).



disarticulate and one can do it with one’s bare hands without using

any tools (Steadman et al., 2002). Such a procedure may result in

characteristic perforation of the distal humerus and damage to the

proximal ulna due to overextension of the elbow (Laroulandie,

2005b; Laroulandie et al., 2008) but no such damage was noted at

Pavlov I. One would expect to recover similar number of scapulae

and coracoids because the two bones are firmly articulated and

tend to stay together when bird carcasses are pulled apart by hand.

Yet, the assemblage shows a paucity of scapulae relative to cora-

coids whichmay be due to the fact that the thin blade of scapulae is

easily broken and the remaining articular part is small and less

likely to be recovered.

Skeletal representation for themain prey species, tetraonids and

ravens, indicates the presence of nearly all major and many minor

anatomical elements which suggests that whole carcasses were

brought to the site and subsequently processed there. The only

missing part of the body is the head (beak, brain case and

mandible). However, the presence of three quadrates of the raven

suggests that the birds were not decapitated outside the site. The

absence of heads in the material may be due to their fragility and

tendency to disintegrate in pre- and post-depositional processes

such as human trampling and diagenesis. The scarcity or lack of

elements of the head and neck has also been observed at other

human-derived sites (Steadman et al., 2002), indirectly supporting

our suggestions.

The high degree of fragmentation of the bones and over-

representation of shafts without articular ends may reflect the

consumption of the ends of the bones (Laroulandie, 2005b;

Steadman et al., 2002) or post-depositional damage resulting from

the unique osteological characters of avian bones (Higgins, 1999). It

is likely that somedamage at Pavlov I is due to tramplingbecause the

site was occupied intensively. The overrepresentation of some

portions of certain bonesmay be best explained by the properties of

thebones. For instance, the sternal endof the coracoid is thin and the

proximal end of the tibiotarsus also consists of some thin crests – in

both cases those parts of the bones may be easily damaged. As

a consequence, scapular parts of coracoids clearly predominate in

many cultural and non-cultural assemblages (Bochenski, 2005;

Bochenski et al., 1993, 1997, 1998; Steadman et al., 2002).

Members of the Tetraonidae family are ground-dwelling, weak

flyers, and relatively heavy with bodies weighing between 0.4 and

1.4 kg. This makes them vulnerable to predation. They have always

been a common source of food for various predators including

birds, mammals, and people (Boev, 1997; Laroulandie, 2005a;

Stewart, 2007; Tyrberg, 1995) and it seems that Pavlov I is no

exception. Although direct traces attributable to food processing

are very scarce on bones from the site, they include one partly

burned wing bone of Lagopus which may indicate cooking for

consumption.

The large number of ravens at Pavlov I is also interesting. Their

body weight is comparable to that of tetraonids (0.99–1.56 kg) but

unlike some galliforms they fly very well and their flight feathers

are stiff and strong (Cramp and Perrins, 1994). The habitat around

the site (rocks and trees not far from the dwellings) was certainly

suitable for them to nest in, but they tend to keep their territories to

themselves and, depending on the season of the year, one can see

solitary birds, couples or small family flocks only. Therefore the

Gravettian people must have either hunted ravens over larger

territories outside the immediate vicinity of their site or, which

seemsmore likely, they took advantage of the fact that ravens often

feed on carrion. It is tempting to speculate that ravenswere killed at

the nearby mammoth bone accumulation of Dolnı Vestonice

I. Large carcasses of mammoths and their bones left by people must

have attracted many carrion-eating animals. Of course, remains of

smaller game that are abundant at Pavlov I also could have

attracted ravens. It is impossible to determine whether the people

lured ravens, and if so, whether they did it intentionally by

exposing carcasses or merely took advantage of the fact that the

birds were already nearby. It is noteworthy that no other Upper

Palaeolithic site contains somany remains of ravens (Stewart, 2007;

Tyrberg, 1998). Pavlov I is unique in this respect. Unfortunately

there is no direct evidencewhy the Gravettian people specialized in

killing ravens. They may have hunted the birds because they were

easy prey close to their dwellings, because ravens stole their food,

or perhaps because they needed the birds for raw materials (bones

or feathers). The lack of cut marks and traces of burning on raven

remains may suggest that the birds were used for purposes other

than food. The fragment of a raven ulna with polished edges (Fig. 5)

supports this hypothesis. It may have been made into a tube, either

for use as a drinking ‘‘straw’’, blowing or sniffing, or with both ends

plugged, as a container for some substance such as ochre (Wijn-

gaarden-Bakker, 1997). Ravens have symbolic meaning in many

cultures throughout the world (Goodwin, 1976) and therefore it is

possible that they played a similar role at Pavlov I during the Upper

Palaeolithic. This would agree with the archaeological context that

suggests symbolic activities and social rituals performed at the

Moravian mega-sites of Pavlov I and Dolnı Vestonice I (Musil, 1994;

Svoboda, 2005).

The remaining smaller species of corvids make up about 3.2% of

all bird remains (NISP¼ 33, MNI¼ 5) and are representedmainly by

jackdaws that are currently considered synanthropes, living near

human settlements, but they also tend to nest among rocks and in

the hollows of older trees (Cramp and Perrins, 1994). It is likely that

this group of birds also nested in the vicinity of Pavlov I. Jackdaws,

like ravens, could have been hunted but because their remains are

scarce, it is difficult to draw further conclusions.

An interesting feature of the analysed material is the relatively

large number (NISP¼ 13, MNI¼ 4) of swan remains (Cygnus sp.),

especially in comparison to the number of the remaining anseri-

forms (only one bone of a duck was found). The presence of swans

indicates a largewater body in the vicinity of the site which implies

other waterfowl must have been present as well. Yet, it seems that

of all anseriforms only the swans were of particular interest to the

Gravettian people, which can be deduced not only from their

relative abundance but also from the fact that one of the three

decorated bones belonged to a swan (Fig. 6). It is difficult to know

why, but large size is a possible explanation. It has already been

noted in many archaeological sites that large species were

preferred to smaller ones (Guminski, 2005; Laroulandie, 2003).

Another interesting feature of the material is the relatively large

variety of birds of prey. We include in this category all meat-eating

species – accipitrids and owls. Although their remains are scarce, at

least five different species are represented. The griffon vulture ulna

with cut marks (Fig. 7a and b) – one of the only four artefacts made

from bird bones at the site – suggests that raptors may have had

some special significance to the Gravettian people. This would

agree with the result of other studies on the mammalian remains

that show the use of carnivores by the Gravettian people in Central

Europe (Fladerer, 2001; Lipecki and Wojtal, 1998; Munzel, 2004;

West, 1996; Wojtal, 2000; Wojtal et al., 2005). The regularity of the

cut marks on the vulture ulna suggests decoration. Ravens are in

some ways similar to birds of prey because they also eat meat and

they are capable of actively killing smaller prey (Cramp and Perrins,

1994). Therefore, the Gravettian huntersmay have treated ravens as

birds of prey and attributed special meaning to them too, which

could explain the large number of raven remains at the site.

Small animals made an increasingly important contribution to

human diet in the Mediterranean Basin from the early Middle

Palaeolithic onward, especially just before the Neolithic –

a hypothesis called ‘‘Broad Spectrum Revolution’’ (Davis, 2005;



Flannery, 1969; Munro, 2004; Stiner, 2001; Stiner et al., 2000).

While small-game assemblages of the Middle Palaeolithic were

dominated by slow-moving tortoises, more agile small vertebrates

including lagomorphs and birds were common prey beginning

from the Upper Palaeolithic. The integration of relatively fast-

moving (and therefore more difficult to catch) small animals into

the diet is interpreted as a necessary consequence of overhunting

slow-moving and slow-reproducing resources such as tortoises.

The bird-rich material from Pavlov I supports the hypothesis. A

unique feature of Pavlov I is the species composition: besides gal-

liforms which were typically hunted throughout Europe, ravens

played an important role. Most of the data in support of the

hypothesis of the Broad Spectrum Revolution derive from sites

around the Mediterranean (Davis, 2005; Stiner, 2001). The location

of Pavlov I enlarges the geographical zone of the hypothesis to

include higher latitudes.

Although we are unsure how the birds were procured at Pavlov

I, their high numbers indicate that the Gravettian people caught

them efficiently. Fully ossified bones indicate that the birds were

adult when killed, thus the people did not rely on collecting young

birds from their nests – a relatively easy task. Hunting so many

birds without a trap or a good weaponwould be a time-consuming

and ineffective procedure. One possible clue to themethods used to

procure birds comes in the form of negative clay impressions of

fibre-based constructions (Adovasio et al., 1996, 1997, 2005). Ado-

vasio and colleagues have proved that the sites of Pavlov I and Dolnı

Vestonice I and II contain the world’s oldest evidence for the use of

plant fibres in the manufacture of textiles, basketry, and netting.

Knotted nets may have served as highly efficient animal traps,

supported indirectly by our results. Another possibility is that bows

were used to hunt birds and medium-size mammals such as lep-

orids and foxes. It is tempting to speculate that the stiff and strong

flight feathers of ravens were a source of arrow fletching. Because

hard evidence of bows and arrows before the Mesolithic is lacking

(Cattelain, 2006; Rosendahl et al., 2006), only indirect pathways

remain to address the question in the Upper Palaeolithic. The

anatomical analysis of ligament attachments and enthesopathies

on the upper limb bones (Churchill, 1994; Villotte, 2008) cannot be

used as an unequivocal and convincing argument. What remains is

an analysis of the potential lithic projectiles, with emphasis on their

form, size, and the typical impact scars on their tips (Knecht, 1997;

Plisson and Geneste, 1989; Villa et al., 2009). Given their extremely

small size, the geometric microliths of Pavlov are better candidates

for arrowpoints than some of the other European specimens, which

are generally interpreted as spearpoints. Yaroshevich (personal

communication) who studied impact scars on the Pavlov microliths

concluded that some of them may have been used as projectiles,

most probably as arrows.

5. Conclusions

This is one of the very few studies of avian remains from the

Gravettian, and certainly the most abundant in bird bones. The

archaeological context with extremely numerous artefacts and tens

of thousands of mammal remains indicate a very intensive occu-

pation of Pavlov I. The skewed taxonomical profile of the avian

assemblage (i.e. more than 90% of the NISP being either grouse or

raven), and bird bones modified by humans are clear suggestions

that birds also played a role in the culture and subsistence of the

Gravettian people.

A unique feature of Pavlov I is the relative abundance of ravens

whose bones had some utilitarian purposes (an ulna made into

a tube), and whose feathers may have been used for decoration or

arrow fletching. It is likely that ravens were killed while feeding on

carcasses or food remains that may have been disposed

intentionally.

Anatomical distribution of particular bones and their fragments

in an assemblage should be used cautiously when studying the

taphonomy of a site. Relying exclusively on one taphonomic feature

such as the wing-to-leg ratio or the predominance of femora may

be misleading. It is advisable to use as wide a range of factors as

possible, including the archaeological and biological contexts.

The numerous bird remains (tetraonids and possible also other

taxa) from Pavlov I that contributed to the diet of its human

inhabitants can be interpreted as supporting the so-called Broad

Spectrum Revolution hypothesis, according to which people in

Europe and the Near East started to hunt swift, small animals from

the Upper Palaeolithic onwards. Until now, most of the data in

support of the theory derive from sites around the Mediterranean.

Pavlov I, which is situated far to the north, enlarges the

geographical zone of the hypothesis. The site also provides a clue as

to how the swift, fast-moving small-game animals were hunted.

Knotted nets made from plant fibres, whose world’s oldest

evidence was found at Pavlov I, may have been used as efficient

traps for catching birds and small mammals.

Acknowledgments

We thank Martin Oliva (Anthropos Institute, Moravian Museum,

Brno) and Moravske Zemske Museum for allowing this study of

bird bones from the Pavlov I site stored in Budisov castle. We are

very grateful to Erika Gal, Veronique Laroulandie, Cecile Mourer-

Chauvire and Tommy Tyrberg for fruitful discussions on various

aspects of this paper and supplying us with relevant literature, and

to Gary Haynes and Simon Davis for helpful comments and cor-

recting the English language of the manuscript. Five anonymous

reviewers provided useful suggestions how to improve the original

manuscript. Our research was partly supported by the Polish

Ministry of Science and Higher Education (grants No. 2 P04C 08130

for years 2006–2009 awarded to P. Wojtal and No. 0903/P01/2006/

31 for years 2007–2009 awarded to J. Wilczynski).

References

Adovasio, J.M., Soffer, O., Klıma, B., 1996. Upper palaeolithic fibre technology:interlaced woven finds from Pavlov I, Czech Republic, c. 26,000 years ago.Antiquity 70, 526–534.

Adovasio, J.M., Hyland, D.C., Soffer, O., 1997. Textiles and cordage: a preliminaryassessment. In: Svoboda, J. (Ed.), Pavlov I – Northwest. The Upper PaleolithicBurial and Settlement Context. Dolnı Vestonice Studies, vol. 4, pp. 432–443.

Adovasio, J.M., Soffer, O., Hyland, D.C., 2005. Textiles and cordage. In: Svoboda, J.(Ed.), Pavlov I Southeast. A Window into the Gravettian Lifestyles. Dolnı Ves-tonice Studies, vol. 14, pp. 432–443.

Alhaique, F., Bisconti, M., Castiglioni, E., Cilli, C., Fasani, L., Giacobini, G., Grifoni, R.,Guerreschi, A., Iacopini, A., Malerba, G., Peretto, C., Recchi, A., Rocci Ris, A.,Ronchitelli, A., Rottoli, M., Thun Hohenstein, U., Tozzi, C., Visentini, P.,Wilkens, B., 2004. Animal resources and subsistence strategies. CollegiumAntropologicum 28, 23–40.

Andrews, P., 1990. Owls, Caves and Fossils. Predation, Preservation and Accumula-tion of Small Mammal Bones in Caves, with an Analysis of the Pleistocene CaveFaunas from Westbury-Sub-Mendib, Somerset. UK. Natural History MuseumPublications, London.

Bochenski, Z., 1981. Fossil remains of birds from Mamutowa Cave. Folia Quaternaria54, 1–24 (in Polish with English summary).

Bochenski, Z.M., 2005. Owls, diurnal raptors and humans: signatures on avianbones. In: O’Connor, T. (Ed.), Biosphere to Lithosphere. New Studies in Verte-brate Taphonomy. Oxbow Books, Oxford, pp. 31–45.

Bochenski, Z.M., Boev, Z., Mitev, I., Tomek, T., 1993. Patterns of bird bone frag-mentation in pellets of the Tawny Owl (Strix aluco) and the Eagle Owl (Bubobubo) and their taphonomic implications. Acta Zoologica Cracoviensia 36 (2),313–328.

Bochenski, Z.M., Huhtala, K., Jussila, P., Pulliainen, E., Tornberg, R., Tunkkari, P.S.,1998. Damage to bird bones in pellets of Gyrfalcon Falco rusticolus. Journal ofArchaeological Science 25, 425–433.



Bochenski, Z.M., Huhtala, K., Sulkava, S., Tornberg, R., 1999. Fragmentation andpreservation of bird bones in food remains of the Golden Eagle Aquila chrys-aetos. Archaeofauna 8, 31–39.

Bochenski, Z.M., Korovin, V.A., Nekrasov, A.E., Tomek, T., 1997. Fragmentation of birdbones in food remains of Imperial Eagles Aquila heliaca. International Journal ofOsteoarchaeology 7 (2), 165–171.

Bochenski, Z.M., Nekrasov, A.E., 2001. The taphonomy of Sub-Atlantic bird remainsfrom Bazhukovo III, Ural mountains, Russia. Acta Zoologica Cracoviensia 44 (2),93–106.

Bochenski, Z.M., Tomek, T., 1994. Pattern of bird bone fragmentation in pellets of thelong-eared owl Asio otus and its taphonomic implications. Acta Zoologica Cra-coviensia 37 (1), 177–190.

Bochenski, Z.M., Tomek, T., 1997. Preservation of bird bones: erosion versus diges-tion by owls. International Journal of Osteoarchaeology 7 (4), 372–387.

Bochenski, Z., Tomek, T., 2004. Bird remains from rock-shelter in the Krucza Ska1a(Central Poland). Acta Zoologica Cracoviensia 47 (1–2), 27–47.

Bochenski, Z.M., Tomek, T., Tornberg, R., Wertz, K., 2009. Distinguishing nonhumanpredation on birds: pattern of damage done by the white-tailed eagle Haliaetusalbicilla, with comments on the punctures made by the golden eagle Aquilachrysaetos. Journal of Archaeological Science 36, 122–129. doi:10.1016/j.jas.2008.07.018.

Bochenski, Z.M., Tornberg, R., 2003. Fragmentation and preservation of bird bonesin uneaten food remains of the Gyrfalcon Falco rusticolus. Journal of Archaeo-logical Science 30, 1665–1671.

Boev, Z., 1997. Wild galliform and gruiform birds (Avis, Galliformes and Gruiformes)in the archaeological record of Bulgaria. International Journal of Osteo-archaeology 7, 430–439.

Bovy, K.M., 2002. Differential avian skeletal part distribution: explaining theabundance of wings. Journal of Archaeological Science 29, 965–978.

Bramwell, D., Yalden, W., Yalden, P.E., 1987. Black grouse as the prey of the goldeneagle at an archaeological site. Journal of Archaeological Science 14, 195–200.

Bruhl, E., 2005. Bone, antler, and ivory tools. In: Svoboda, J. (Ed.), Pavlov I Southeast.A Window into the Gravettian Lifestyles. Dolnı Vestonice Studies, 14, pp.252–293.

Cassoli, P.F., Tagliacozzo, A., 1997. Butchering and cooking of birds in the Palaeolithicsite of Grotta Romanelli (Italy). International Journal of Osteoarchaeology 7,303–320.

Cattelain, P., 2006. Apparition et evolution de larc et des pointes de fleches dans laprehistoire europeene (Paleo-, Meso-, Neolithique). In: Bellintani, P. (Ed.), Cat-ene operative dellarco preistorico. Giunta della Provincia Autonoma di Trento,Trento, pp. 45–66.

Cramp, S., Perrins, C.M., 1994. The Birds of the Western Palearctic, vol. VIII. OxfordUniversity Press, Oxford, New York.

Cramp, S., Simmons, K.E.L., 1980. The Birds of the Western Palearctic, vol. II. OxfordUniversity Press, Oxford, London, New York.

Churchill, S.E., 1994. Human Upper Body Evolution in the Eurasian Later Pleistocene.PhD thesis, University of New Mexico, Albuquerque.

Cyrek, K., Nadachowski, A., Madeyska, T., Bochenski, Z., Tomek, T., Wojtal, P.,Miekina, B., Lipecki, G., Garapich, A., Rzebik-Kowalska, B., Stworzewicz, E.,Wolsan, M., Godawa, J., Ko�sciow, R., Fostowicz-Frelik, q., Szyndlar, Z., 2000.Excavation in the Deszczowa Cave (Kroczyckie Rocks, Czestochowa Upland,Central Poland). Folia Quaternaria 71, 5–84.

Damblon, F., 1997. Anthracology and past vegetation reconstruction. In: Svoboda, J.(Ed.), Pavlov I – Northwest. Dolnı Vestonice Studies, 4, pp. 437–442.

Davis, S.J.M., 2005. Why domestic food animals? Some zoo-archaeological evidencefrom the Levant. Journal of Archaeological Science 32, 1408–1416.

d’Errico, F., Henshilwood, C., Lawson, G., Vanhaeren, M., Tillier, A.-M., Soressi, M.,Bresson, F., Maureille, B., Nowell, A., Lakarra, J., Backwell, L., Julien, M., 2003.Archaeological evidence for the emergence of language, symbolism, and music– an alternative multidisciplinary perspective. Journal of World Prehistory 17(1), 1–70.

Erbersdobler, K., 1968. Vergleichend morphologische Untersuchungen an Einzel-knochen des postcranialen Skeletts in Mitteleuropa vorkommender mittel-grober Huhnervogel. Ludwig-Maximilians-Universitat Munchen.

Ericson, P.G.P., 1987. Interpretations of archaeological bird remains: a taphonomicapproach. Journal of Archaeological Science 14, 65–75.

Fladerer, F.A., 2001. Die Faunareste vom jungpalaolithischen Lagerplatz Krems-Wachtberg, Ausgrabung 1930. Jagdwild und Tierkorpernutzung an der Donauvor 27.000 Jahren, 39. Mitteilungen der Prahistorischen Kommision, Osterrei-chische Akademie der Wissenschaften, Wien. Philosophish-historiche Klasse.

Flannery, K.V., 1969. Origins and ecological effects of early domestication in Iran andthe near East. In: Ucko, P.J., Dimbleby, G.W. (Eds.), The Domestication andExploitation of Plants and Animals. Duckworth, London, pp. 73–100.

Gal, E., 2005. New data to the bird bone artefacts from Hungary and Romania. In:Luik, H., Choyke, A.M., Batey, C.E., L}ougas, L. (Eds.), From Hooves to Horns, fromMollusc to Mammoth. Manufacture and Use of Bone Artefacts from PrehistoricTimes to the Present. Proceedings of the 4th Meeting of the ICAZ Worked BoneResearch Group at Tallinn, 26th–31st of August 2003, 15. Muinasaja teadus,Tallinn, pp. 325–338.

Goodwin, D., 1976. Crows of the World. Cornell University Press, Ithaca, New York.Guminski, W., 2005. Stone age hunters of Dudka and Szczepanki, Masurian Lake-

land, NE Poland. Acta Archaeologica 76, 111–143.Higgins, J., 1999. Tunel: a case study of avian zooarchaeology and taphonomy.

Journal of Archaeological Science 26, 1449–1457.Knecht, H., 1997. Projectile Technology. Plenum, New York.

Koumouzelis, M., Ginter, B., Koz1owski, J.K., Pawlikowski, M., Bar-Yosef, O.,Albert, R.M., Litynska-Zajac, M., Stworzewicz, E., Wojtal, P., Lipecki, G., Tomek, T.,Bochenski, Z.M., Pazdur, A., 2001. The early upper Palaeolithic in Greece: theexcavations in Klisoura cave. Journal of Archaeological Science 28, 515–539.

Kraft, E., 1972. Vergleichend morphologische Untersuchungen an Einzelknochennord- und mitteleuropaischer kleinerer Huhnervogel. Ludwig-Maximilians-Universitat Munchen.

Laroulandie, V., 1998. Etudes archeozoologique et taphonomique des Lagopedes dessaules de la grotte magdalenienne des Eglises (Ariege). Anthropozoologica 28,45–54.

Laroulandie, V., 2000. Taphonomie et archeozoologie des oiseaux en grotte:applications aux sites Paleolithiques du Bois-Ragot (Vienne), de Combe Sau-niere (Dordogne) et de la Vache (Ariege). These d’Universite, Universite deBordeaux I.

Laroulandie, V., 2002. Damage to pigeon long bones in pellets of the eagle owl Bubobubo and food remains of peregrine falcon Falco peregrinus: zooarchaeologicalimplications. In: Bochenski, Z.M., Bochenski, Z., Stewart, J.R. (Eds.), Proceedingsof the 4th Meeting of the ICAZ Bird Working Group, Krakow, Poland, 11–15September, 2001. Acta Zoologica Cracoviensia 45 (special issue), 331–339.

Laroulandie, V., 2003. Exploitation des Oiseaux au Magdalenien en France: Etat deslieux. In: Costamagno, S., Laroulandie, V. (Eds.), Mode de vie au Magdalenien:Apports de l’archeozoologie/Zooarchaeological insights into Magdalenian life-ways. Actes du colloque 6.4 du XIVe Congres du l’UISPP, Liege, Belgique, 2–8septembre 2001. BAR n� 1144. Royaume-Uni, Oxford.

Laroulandie, V., 2005a. Bird exploitation pattern: the case of Ptarmigan Lagopus sp.in the upper Magdalenian site of La Vache (Ariege, France). DocumentaArchaeobiologiae. In: Grupe, G., Peters, J. (Eds.), Feathers, Grit and Symbolism.Birds and Humans in the Ancient Old and New Worlds, Proceedings of the 5thMeeting of the ICAZ Bird Working Group, Munich, 26–28 July 2004, 3, pp.165–178.

Laroulandie, V., 2005b. Anthropogenic versus non-anthropogenic bird boneassemblages: new criteria for their distinction. In: O’Connor, T. (Ed.), Biosphereto Lithosphere. New Studies in Vertebrate Taphonomy. Oxbow Books, Oxford,pp. 25–30.

Laroulandie, V., Costamagno, S., Cochard, D., Mallye, J.-B., Beauval, C., Castel, J.-C.,Ferrie, J.-G., Gourichon, L., Rendu, W., 2008. Quand desarticuler laisse destraces: le cas de l’hyperextension du coude. Annales de Paleontologie 94,287–302.

Lipecki, G., Wojtal, P., 1998. Mammal remains. In: Kozlowski, J.K. (Ed.), Complex ofUpper Palaeolihic Sites near Moravany, Western Slovakia, vol. 2. Institute ofArchaeology, Jagellonian University, Cracow/Archaeological Institute, SlovakAcademy of Sciences, Nitra, pp. 103–126. Moravany-Lopata (Excavations1993–1996).

Livingston, S.D., 1989. The taphonomic interpretation of avian skeletal partfrequencies. Journal of Archaeological Science 16, 537–547.

Lyman, R.L., 1994. Vertebrate Taphonomy. Cambridge Manuals in Archaeology.Cambridge University Press, Cambridge.

Lyman, R.L., 2008. Quantitative Paleozoology. Cambridge Manuals in Archaeology.Cambridge University Press, Cambridge.

Mason, S., Hather, J., Hillman, G., 1994. Preliminary investigation of the plant macro-remains from Dolnı Vestonice II and its implications for the role of plant foodsin Palaeolithic and Mesolithic Europe. Antiquity 68, 48–57.

Mlikovsky, J., 1996. New data on the food of the White-tailed Sea Eagle (Haliaetusalbicilla) in the Svjatoj Nos wetlands, Lake Baikal. Buteo 8, 115–118.

Mourer-Chauvire, C., 1983. Les oiseaux dans les habitats paleolithiques: gibier deshommes ou proies des rapaces? In: Grigson, C., Clutton-Brock, J. (Eds.), Animaland Archaeology: 2. Shell Middens, Fishes and Birds. British ArchaeologicalReports, International Series, 183, pp. 111–124.

Mulkeen, S., O’Connor, T.P.O., 1997. Raptors in towns: towards an ecological model.International Journal of Osteoarchaeology 7, 440–449.

Munro, N.D., 2004. Zooarchaeological measures of hunting pressure and occupationintensity in the Natufian. Current Anthropology 45 (s4), s5–s34.

Munzel, S.C., 2004. Subsistence patterns in the Gravettian of the Ach valley,a former tributary of the Danube in the Swabian Jura. In: Svoboda, J., Sedlack-ova, L. (Eds.), The Gravettian Along the Danube. Dolnı Vestonice Studies, 11, pp.71–85.

Musil, R., 1955. Osteologick�y material z paleolitickeho sidliste v Pavlove. (Dasosteologische Material aus der palaolithischen Siedlungsstatte in Pollau). ActaAcademiae Scientarum �Cechoslovenicae, Basis Brunensis 27/318 (6), 279–320.

Musil, R., 1958. Funde von Wirbeltieren in Mahren (Tschechoslowakei) aus demletzten Wurm-Stadial. Eiszeitalter und Gegenwart 9, 61–68.

Musil, R., 1959. Osteologick�y material z paleolitickeho sidliste v Pavlove. �Cast II. (Dasosteologische Material aus der palaolithischen Siedlungsstatte in Pavlov. II Teil).Anthropozoikum 8, 83–106.

Musil, R., 1994. The fauna. Hunting game of the culture layer of Pavlov. In:Svoboda, J. (Ed.), Pavlov I. Excavations 1952–53. Eraul 66/Dolnı VestoniceStudies, 2, pp. 181–209.

Musil, R., 1997. Hunting analysis. In: Svoboda, J. (Ed.), Pavlov IdNorthwest. DolnıVestonice Studies, 4, pp. 443–468.

Musil, R., 2003. The middle and upper palaeolithic game suit in central andsoutheastern Europe. In: Andel, T.H.van, Davies, W. (Eds.), Neanderthals andModern Humans in the European Landscape During the Last Glaciation. Cam-bridge, pp. 167–190.

Musil, R., 2005. Animal prey. In: Svoboda, J. (Ed.), Pavlov I Southeast. AWindow intothe Gravettian Lifestyles. Dolnı Vestonice Studies, 14, pp. 190–228.



Nadachowski, A., 2005. Small vertebrates and environmental reconstruction. In:Svoboda, J. (Ed.), Pavlov I Southeast. A Window into the Gravettian Lifestyles.Dolnı Vestonice Studies, 14, pp. 187–189.

Novak, M., 2005. Pavlov I – Southeast. Review of spatial distributions. In: Svoboda, J.(Ed.), Pavlov I Southeast. A Window into the Gravettian Lifestyles. Dolnı Ves-tonice Studies, 14, pp. 53–71.

Opravil, E., 1994. The vegetation. In: Svoboda, J. (Ed.), Pavlov I. Excavations 1952–53.Eraul 66/Dolnı Vestonice Studies, 2, pp. 175–180.

Plisson, H., Geneste, J.M., 1989. Analyse technologique des pointes a cran solutreensdu Placard (Charente), du Fourneau du Diable, du Pech de la Boissiere et deCombe Sauniere (Dordogne). Paleo 1, 65–106.

Reichstein, H., Pieper, H., 1986. Untersuchungen an Skelettresten von Vogeln ausHaithabu (Ausgrabung 1966-1969). In: Berichte uber die Ausgrabungen inHaithabu Bericht, 22. Karl Wachholz Verlag, Neumunster.

Reisch, L., 1976. Beobachtungen an Vogelknochen aus dem Spatpleistozan der Hohlevon Kephalari (Argolis, Griechenland). Archaologisches Korrespondenzblatt 6,261–265.

Rosendahl, G., Beinhauer, K.-W., Loscher, M., Kreipl, K., Walter, R., Rosendahl, W.,2006. Le plus vieil arc du monde? Une piece interessante en provenance deMannheim, Allemagne. L’Anthropologie 110, 371–382.

Rybnıckova, E., Rybnıcek, K., 1991. The environment of the Pavlovian – Palae-oecological results from Bulhary, South Moravia. In: Edler-Kovacs, J. (Ed.),Palaeovegetational Development in Europe, Pan-European PalaeobotanicalConference Vienna, pp. 73–79.

Serjeantson, D., Irving, B., Hamilton-Dyer, S., 1993. Bird taphonomy from the insideout: the evidence of gull predation on the manx shearwater Puffinus puffinus.Archaeofauna 2, 191–204.

Soffer, O., Vandiver, P., 2005. Ceramic fragments. In: Svoboda, J. (Ed.), Pavlov ISoutheast. AWindow into the Gravettian Lifestyles. Dolnı Vestonice Studies, 14,pp. 415–431.

Steadman, D.W., Plourde, A., Burley, D., 2002. Prehistoric butchery and consumptionof birds in the Kingdom of Tonga, South Pacific. Journal of ArchaeologicalScience 29, 571–584.

Stewart, J.R., 2007. An Evolutionary Study of Some Archaeologically SignificantAvian Taxa in the Quaternary of the Western Palaearctic. In: BAR InternationalSeries, 1653.

Stiner, M.C., 2001. Thirty years on the ‘‘Broad Spectrum Revolution’’ and paleolithicdemography. Proceedings of the National Academy of Sciences 98 (13),6993–6996.

Stiner, M.C., Munro, N.D., Surovell, T.A., 2000. The tortoise and the hare: small-gameuse, the broad spectrum revolution, and Paleolithic demography. CurrentAnthropology 41, 39–73.

Svoboda, J. (Ed.), 1994, Pavlov I. Excavations 1952–53. Eraul 66/Dolnı VestoniceStudies, 2 Liege.

Svoboda, J. (Ed.), 1997. Pavlov I – Northwest. Dolnı Vestonice Studies, 4 Brno.Svoboda, J., 2005. Pavlov I – Southeast. Location, Stratigraphy, Microstratigraphies,

and Features. In: Svoboda, J. (Ed.), Pavlov I Southeast. A Window into theGravettian Lifestyles. Dolnı Vestonice Studies, 14, pp. 25–52.

Svoboda, J., Kralık, M., �Culıkova, V., Hladilova, S., Novak, M., N�yvltova Fisakova, M.,N�yvlt, D., Zelinkova, M., 2009. Pavlov VI: an upper paleolithic living unit.Antiquity 83, 282–295.

Svobodova, H., 1991a. The pollen analysis of Dolnı Vestonice II, section No 1. In:Svoboda, J. (Ed.), Dolnı Vestonice II – Western Slope, 54, pp. 75–88. Eraul.

Svobodova, H., 1991b. Pollen analysis of the upper paleolithic triple burial at DolnıVestonice. Archeologicke rozhledy 43, 505–510.

Tagliacozzo, A., Gala, M., 2002. Exploitation of Anseriformes at two Upper Palae-olithic sites in Southern Italy: Grotta Romanelli (Lecce, Apulia) and Grotta delSantuario della Madonna a Praia a Mare (Cosenza, Calabria). In: Bochenski, Z.M.,Bochenski, Z., Stewart, J.R. (Eds.), Proceedings of the 4th Meeting of the ICAZBird Working Group, Krakow, Poland, 11–15 September, 2001. Acta ZoologicaCracoviensia 45 (special issue), 117–131.

Tomek, T., Bochenski, Z.M., 2000. The Comparative Osteology of European Corvids(Aves: Corvidae), with a Key to the Identification of their Skeletal Remains.Publications of the Institute of Systematics and Evolution of Animals, Krakow.

Tomek, T., Bochenski, Z., 2005. Weichselian and Holocene bird remains fromKomarowa Cave, Central Poland. Acta Zoologica Cracoviensia 48A (1–2),43–65.

Tomek, T., Bochenski, Z., Bochenski, Z.M., 2003. Birds (Aves). In: Valde-Novak, P.,Nadachowski, A., Madeyska, T. (Eds.), Oblazowa Cave: Human Activity, Stra-tigraphy and Palaeoenvironment. Institute of Archaeology and Ethnology PolishAcademy of Sciences, Krakow, pp. 102–113.

Trinkaus, A.E., Svoboda, J.A., Wojtal, P., N�yvltova Fisakova, M., Wilczynski, J., 2009.Human remains from the Moravian Gravettian: morphology and taphonomy ofadditional elements from dolnı Vestonice II and Pavlov I. International Journalof Osteoarchaeology. doi:10.1002/oa.1088.

Tyrberg, T., 1995. Palaeobiogeography of the genus Lagopus in the West Palearctic.Courier Forschungsinstitut Senckenberg 181, 275–291.

Tyrberg, T., 1998. Pleistocene Birds of the Palearctic: a Catalogue. In: Publication ofthe Nuttall Ornithological Club, 27 Cambridge, Massatchusetts, and updates athttp://web.telia.com/wu11502098/pleistocene.pdf (2008-02-24).

Verpoorte, A., 2005. Lithic assemblage of Pavlov I South-Central (1954, 1956, 1963,1964). In: Svoboda, J. (Ed.), Pavlov I Southeast. A Window into the GravettianLifestyles. Dolnı Vestonice Studies, 14, pp. 75–111.

Villa, P., Boscato, P., Ranaldo, F., Ronchitelli, A., 2009. Stone tools for the hunt: pointswith impact scars from a Middle Paleolithic site in southern Italy. Journal ofArchaeological Science 36, 850–859.

Villotte, S., 2008. Enthesopathies et Activites des hommes prehistoriques. Thesepresente a l’Universite de Bordeaux 1 pour obtenir le grade de Docteur.Bordeaux.

West, D.L., 1996. Krakow-Spadzista, unit E and unit F. Faunal remains. In:Koz1owski, J.K. (Ed.), The Upper Palaeolithic site Krakow-Spadzista. Units D, Eand F (Excavations1986–1989). Folia Quaternaria 67, 21–34.

Wijngaarden-Bakker, L.H., 1997. The selection of bird bones for artefact productionat Dutch Neolithic sites. International Journal of Osteoarchaeology 7, 339–345.

Wojtal, P., 2000. Taphonomy. In: Cyrek, K., Nadachowski, A., Madeyska, T.,Bochenski, Z., Tomek, T., Wojtal, P., Miekina, B., Lipecki, G., Garapich, A., Rzebik-Kowalska, B., Stworzewicz, E., Wolsan, M., Godawa, J., Ko�sciow, R., Fostowicz-Frelik, q., Szyndlar, S. (Eds.), Excavation in the Deszczowa Cave (KroczyckieRocks, Czestochowa Upland, central Poland). Folia Quaternaria 71, 5–84.

Wojtal, P., Sedlackova, L., Wilczynski, J., 2005. Human activities on the faunalmaterial. In: Svoboda, J. (Ed.), Pavlov I Southeast. AWindow into the GravettianLifestyles. Dolnı Vestonice Studies, 14, pp. 229–231.


Fowling during the Gravettian: the avifauna of Pavlov I, the Czech Republic

Documents