Journal of Archaeological Science (1999) 26, 14491457

Article No. jasc.1999.0429, available online at http://www.idealibrary.com on

Tunel: A Case Study of Avian Zooarchaeology and Taphonomy

Jude Higgins
Natural History Museum and Biodiversity Research Center and the Department of Anthropology, The University
of Kansas, Lawrence, Kansas 660452454, U.S.A.
(Received 9 September 1998, revised manuscript accepted 22 February 1999)
Although not often considered, there are many osteological characters unique to the avian skeleton that influence the
taphonomy of bird bones. These characters are reviewed and their archaeological significance discussed herein. The
presence of marrow in many avian long bones is important to interpretation of avian remains from archaeological sites
because the presence of marrow aects bone density and, in turn, preservation. Other structural properties that aect
avian bone preservation include cortical wall thickness, length and pneumatic state. Based on an analysis of
approximately 10,000 bird bones from the archaeological site of Tunel, Tierra del Fuego, Argentina, I found that
specific breakage patterns resulted from natural taphonomic processes acting as a result of the unique avian bone
characteristics. This information may allow researchers to distinguish breakage patterns in avian bones resulting from
natural taphonomic processes from breakage patterns that are culturally induced.
 1999 Academic Press
Keywords: AVIAN OSTEOLOGY, ZOOARCHAEOLOGY, TAPHONOMY, PNEUMATICITY, AVIAN BONE
MARROW.

Introduction

vian remains found in archaeological sites are

frequently reported as simple lists of taxa
recovered from the faunal assemblage, and
some times only identified as birds. Although a few
recent studies have analysed bird remains in greater
detail (e.g. Rich, 1980; Ericson, 1987; Lefe`vre, 1989;
Livingston, 1989; Emslie, Speth & Wiseman, 1992),
it is unusual for in-depth analyses (i.e. identification
of taxa, analysis of fracture/breakage patterns, tool
marks, burning, specimen dimensions, preservation
and carnivore activity) to be performed on avian bones
(but see Cassoli & Tagliacozzo, 1997). Because any
archaeological bone assemblage is an estimate of the
relative importance of the animals represented
(Guilday, 1970), all remains, including avian, must be
included in detailed analyses of assemblages. If avian
remains are not analysed with the same rigor as other
remains, it will not be possible to determine accurately
prehistoric diet breadth.
Although several studies have determined subsistence strategies, time of site occupation and past
environmental conditions by identifying avian remains
(e.g. Howard, 1929; Yesner, 1977; Avery & Underhill
1986; Orquera, Piana & Haydee Tapia de Bradfors,
1984; Piana et al., 1992; International Journal of
Osteoarchaeology, 1997), too little attention has been
given to the cultural significance of bird remains. Avian
remains may have been ignored because some
researchers assume that birds contribute little energy to

the human diet (because of their low fat-to-lean ratio).

However, most bird bones are filled with marrow (a
fat-rich energy source), the quality of which may vary
among elements and/or taxa; thus, various birds may
contribute dierentially to the human diet. The best
method for determining if avian bones were exploited
for marrow is through detailed analyses of marrow
distribution in various avian taxa, coupled with comparable analyses of breakage and fracture patterns in
archaeological remains.
A second reason for the lack of attention to avian
remains may be the abundance of large-mammal
remains in many sites. Because analyses of dominant
taxa seemingly result in adequate depictions of general
subsistence strategies of prehistoric peoples, lower
abundance remains, such as bird bones, are ignored
(Dawson, 1969). However, the low abundance of avian
remains may not reflect their actual contribution to
prehistoric diets, but instead might be a result of the
vulnerability of bird bones to taphonomic eects.
Among the several processes that can aect bones in
archaeological assemblages are: fracture and breakage
patterns resulting from butchering; tool-marks left
from skinning and meat removal; burning and other
evidence of cooking; damage as a result of carnivore
activity; and the eects of natural degradation and
decay. Analysis of archaeological faunal assemblages
requires dierentiation between cultural and natural
processes (Schier, 1987). Cultural transforms (Ctransforms), as defined by Schier (1987), are human
behaviours that aect artefacts after their initial use,

1449
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 1999 Academic Press

1450 J. Higgins

and non-cultural transforms (N-transforms) are natural

events of the physical environment that modify artefacts. Patterned breakage may result from cultural, as
well as non-cultural, processes (Schier, 1987).
To distinguish between the eects of cultural and
non-cultural transforms, it is necessary to dierentiate
the patterns left by each process. As similar patterns
can be produced by dierent mechanisms (Binford,
1981), it is necessary to attempt to isolate the consistent
eects of N-transforms on avian remains. We can then
determine the patterning produced by C-transforms for
use in behavioural inference. Thus, to infer the significance of birds in prehistoric diets, we must first understand the natural taphonomic eects on avian bones.
Patterns of both cultural and non-cultural transforms
on bird bones are related directly to properties of the
avian skeleton such as marrow distribution, bone
density and cortical wall thickness. An understanding
of these properties is fundamental to accurate assessment of the eects of N- and C-transforms. The unique
characteristics of the avian skeleton and their relationship to cultural and non-cultural transformation
processes are described below. By applying these
properties to five representative bones, I predict
expected non-cultural breakage patterns and test these
predictions on the sample of avian remains from
the archaeological site of Tu nel, Tierra del Fuego,
Argentina.

Unique Characteristics of the Avian Skeleton

Ultrastructure
Among the unique microstructural properties of bird
bones are: (1) the presence of sandwich-bone construction; (2) fusion of bones, thin cortical walls; and (3)
cartilaginous epiphyses, and cones in developing long
bones. Sandwich-bone construction is characterized by
an inner and an outer layer of bone separated by small
perpendicular columns of bony spicules (Bu hler, 1992).
Birds typically have many flat, bar-like bones composing the braincase, palate, sternal plates and ends of
long bones composed of sandwich-bone (Figure 1).
Many elements of the avian skeleton are fused (e.g. the
skull, thorax, sacrum and pelvic girdle) and, in many
taxa, the thoracic and sacral vertebrae fuse at maturity
(via ossification of hyaline cartilage); this provides an
easy method of aging specimens. Fusion increases the
strength of individual elements and regions, and was
shown by Bu hler (1992) to arise independently of
pneumatization. Unlike in other tetrapods, the cortical
bone wall is thin in birds; moreover the structural
density may vary among elements and within a single
element. In larger birds, longer bones usually have
thinner cortical walls relative to their diameter. As
these bones also have larger diameters proportional to
body size, they are stronger (Bu hler, 1992).
The growth/ossification of avian long bones also is
unique because, in contrast to mammal bones, the

Figure 1. Saggital (right) and cross (left) sections of the humerus of

a herring gull showing sandwich bone construction. Note the bony
struts of sandwich bone within the pneumatic marrow cavity and the
relatively thin cortical wall. Adapted from Proctor & Lynch (1993)

epiphysis in the developing avian long bone is cartilaginous and, with the exception of the proximal
metatarsus and tibia, no secondary centre of ossification is present (Church & Johnson, 1964; Riddell,
1987). Ossification of the avian metaphysis (the area
between the bone end [epiphysis] and the shaft [diaphysis]) does not occur until more than 14 days after
hatching. Before this time, only cones of cartilage
(Figure 2) are present in the metaphysis. In the metaphysis of a mammal bone there is a disc (rather than
cone) of cartilage, known as the epiphyseal plate,
which separates the epiphysis and diaphysis.
Pneumatization and hollow bones
Pneumatization, the replacement of marrow by air, is
the most distinctive and best-known characteristic of
avian bones. The extent of pneumatization varies

Archaeological Implications of Avian Taphonomy 1451

Figure 3. Medial (right) and lateral (left) views of the proximal right
humerus of a chicken. Note the lateral view is sectioned to show the
interior of the bone. f.= foramen. Adapted from Marshall (1960).

Figure 2. Saggital section through the end of a typical long bone of

a bird showing regions of cartilage and bone deposition
(cart.=cartilage). Redrawn from Marshall (1960).

taxonomically and there are vast dierences among

closely related species. For example, only the humerus,
sternum and centra of a few cervical vertebrae of
cormorants (Phalacrocorax spp.) are pneumatic, but
almost all bones in the skeleton of the albatross
(Diomedeidae spp.) are pneumatized (Bellairs & Jenkin,
1960; Casler, 1973). Although the precise function of
pneumaticity in bird bones is unclear, it has been
suggested that lighter bones are an adaptation for
flight. It has also been postulated that the pneumatic
structure is associated with avian respiration (as it is a
connection to the atmospheric air) and/or to temperature regulation. As an example of the latter, it has been
suggested that a pneumatic skull roof helps keep the
brain warm in passerine finches (Bellairs & Jenkin,
1960; Warncke, 1987; J. Baumel, pers. comm., 1997; L.
D. Martin, pers. comm., 1998). Pneumatization occurs
in the cavities of long bones, between the layers of
sandwich bone, within vertebrae and in the cranium
and can be identified by the presence of pneumatic
pores, foramina or fossae (Figure 3). Pneumatic
foramina can be distinguished by trabeculae in the
fossae, whereas nutrient foramina lack trabeculae
inside the opening (L. D. Martin, pers. comm., 1998).
However, not all pneumatic foramina have trabeculae
and no simple set of characters can diagnose them
unequivocally (Bu hler, 1992).
There has been considerable confusion regarding the
marrow status of bird bones and the relationship of the

presence of marrow to pneumaticity. This is most likely

an artefact of misuse of terminology. Hollow bones,
characterized by a vacuous tube consisting of a
medullary cavity (which comprises the mid-section of
long bones) are present in all tetrapods. Therefore, all
long bones are hollow by definition. When applied
to bird bones, however, hollow often is misused to
designate pneumaticity (e.g. Trapani, 1998:481). This
misconception is further extended to the common idea
that all avian bones are hollow and, therefore,
pneumatic. However, many bird bones (e.g. distal limb
bones, femur, jugal bar, cervical vertebrae I and II and
the caudal vertebrae) are marrow-filled. In a study of
the pneumaticity of fowl bones, Hogg (1980) found
that only the humerus was pneumatic. Furthermore,
before females lay eggs, their long bones are filled with
medullary bone, which functions as a calcium deposit
for eggshell production (Kyes & Potter, 1934; Simkiss,
1967; Driver, 1982).
Bone marrow and medullary bone
All avian bone marrow is a cellularly rich (in terms of
cells/volume) and highly vascular, loose connective
tissue (Tavassoli & Yoey, 1983). Long bone and
vertebral marrow is the major site of haemopoiesis (red
blood cell production) in all post-hatching birds. However, dierent skeletal elements vary in their degree of
red blood cell production and fat content. McDonald
(1939) reported that the femoral bone marrow of
pigeons is hyperplastic (i.e. high in the production of
red blood cells), whereas the marrow of the radius and
ulna was hyper- to hypoplastic (i.e. low in the production of red blood cells); the marrow of subadult birds
often is more cellular than that of mature birds.
During haemopoesis, fat cells are present in large
quantities in the marrow (Tavassoli & Yoey,

1452 J. Higgins

1983:110). However, the concentration of fat in avian

marrow is a function of species, diet, age, sex and
health, along with numerous other factors (Grimiger,
1976:257). In adult birds, the peripheral portion of the
marrow cylinder is hyperplastic, whereas the axial
portion is hypoplastic and contains a large amount of
fat (Jordan, 1942). This resembles the condition found
in mammals (Tavassoli & Yoey, 1983). In birds, fat
replaces some haemopoietic tissue as the animals age;
however, the bone marrow of mature birds contains
a large amount of lymphatic tissue (unlike that of
mammals) (Sturkie, 1976).
It is important to distinguish between marrow and
medullary bone. In gravid females, calcium-rich
medullary bone forms in marrow cavities, coincident
with the distribution of marrow. The marrow cavity
becomes permeated with interlacing spicules and
loosely arranged fibres of medullary bone (Driver,
1982).

Archaeological Implications of Avian Skeletal

Properties
Because of the unique nature of the avian skeleton,
interpretations of bird remains may be confounded by
natural transformational processes. The structural
properties of avian bones may aect preservation.
Similarly, cultural transformation processes, such as
human butchering activities, may relate to the quality
and distribution of bone marrow and the articulation
of major muscle groups.
Ultrastructure
During the process of archaeodeposition, physical
forces such as strain, tension and uneven stress distribution aect each skeletal element. Depending on their
characters, bones will be preserved better or worse. For
example, the presence of any opening, such as a
foramen, in the wall of the bone will increase the
tendency toward uneven stress distribution, hence
bones are more likely to break in an area with a
foramen (Dullemeijer, 1974). Bone length is also an
important factor in the dierential preservation of
elements. Longer bones will break more easily than
shorter ones of the same diameter and bones with
larger diameters proportional to body size, such as
those of larger birds, are strongest (Bu hler, 1992).
Cortical wall thickness
The thickness of the cortical wall (Figure 1) varies
taxonomically and throughout the skeleton, as well as
within each individual element. The cortical wall of
pneumatized bones is generally thinner than that of
non-pneumatized bones. This may compromise the
suggestion that pneumaticity increases strength (Rich,
1980). In larger birds, longer bones generally have
thinner cortical walls relative to the diameter of the

bone. The cortical wall often is thinner at the ends of a

bone; however, this is not always the case. For
example, the distal end of the tibiotarsus is the thickest
part of the bone. Also, diving birds have relatively
thicker cortical walls (presumably to decrease buoyancy) (Bu hler, 1992); thus, distal tibiotarsi and the
bones of diving birds are more likely to survive
taphonomic processes.
Structural density
Structural density, measured as a ratio of mass to
volume (g/cm3), also aects natural taphonomic processes (Lyman, 1984, 1994a)viz., bones of higher
density will more likely survive in archaeological
assemblages. Because some bones are more porous
than others (Shipman, 1981), density can be measured
as either true density (in which the volume of pore
space is excluded) or bulk density (in which pore space
is included in volume) (Lyman, 1994a).
Pneumaticity
Pneumaticity is characterized by the presence of a
diverticulum, or air sac, within the marrow cavity. As
a long bone grows, the diverticulum displaces the
marrow and trabeculae (of sandwich bone) and the
trabeculae recede to the ends of the bone (Figure 1).
Taphonomically, this is important because the maximal stress and strain on a bending bone is found at the
ends (Dullemeijer, 1974), between the diverticulum and
the remaining sandwich bone. One eect of noncultural transforms on pneumatic long bones is the
breaking o, and consequent absence, of proximal
and/or distal ends.
Distribution of bone marrow
The location of marrow in the avian skeleton and, in
particular, the amounts of fatty marrow, is archaeologically significant, especially if prehistoric hunters
broke bones for marrow extraction. If marrow extraction was practised, one would expect to find evidence
of this in the breakage and fracture patterns of
archaeological remains (e.g. Jones, 1993:108). It follows that these patterns should be related to the
marrow content and quality in the various skeletal
elements.
Predicted dierential representation and preservation
All the factors discussed above must be considered
together as a basis for predictions of the natural eects
of taphonomy on avian bones. To elucidate the possible eects of skeletal characteristics on the survival
and cultural use of avian bones, I analysed the length,
cortical wall thickness, density, pneumatic nature
and marrow content of a collection of avian bones

Archaeological Implications of Avian Taphonomy 1453

Table 1. Eects of N-transforms on various elements of the avian skeleton
N-transform
eects

Distal end
missing

Proximal end
missing

Both ends
missing

Comments

X
X
X

X
X
X
X

Many break at nutrient foramen

Distal end most likely to survive
Third digit most likely to survive

Humerus
Coracoid
*Femur
Tibiotarsus
*Tarsometatarsus

*Bones that are least dense.

Distal end of coracoid is the humeral end.
Prominent pattern in penguins is for distal end of coracoid to be missing and proximal end to be present.

Table 2. Eects of C-transforms, specifically marrow extraction

C-transform
eects
Humerus
Coracoid
Femur
Tibiotarsus
Tarsometatarsus

Distal end
missing

Proximal end
missing

Both ends
missing

X
X
X

Distal end of coracoid is the humeral end.

comprising five elements (coracoid, femur, humerus,

tarsometatarsus and tibiotarsus) of the cormorant
(Phalacrocorax auritus).
Regarding the non-cultural breakage patterns of
bird long bones, the predicted eects attributable to
N-transforms are shown in Table 1. Because bird
humeri are pneumatic, have thin cortical walls and are
relatively long, they should be broken more often than
non-pneumatic bones. As relatively long bones, they
are most likely to break where the diverticulum ends
and sandwich bone begins (Figure 1); hence, they
should be missing both ends. The coracoid will
probably break at the sternal end, where it is least
dense; the humeral end also might break because of a
thinning cortical wall. The femur is likely to be missing
one or both ends because the cortical wall thins
proximally and distally. Because it is a relatively long
bone, the tibiotarsus is likely to suer damage to one
or both ends; however, the distal end is more likely to
be preserved because the cortical wall is thickest in that
area. The distal end of the tarsometatarsus is likely to
be missing, because of a thinning of the cortical wall.
The third digit is most likely to be preserved because its
cortical wall is relatively thicker than that of the other
digits.
The predicted patterns of breakage attributable to
C-transforms are slightly dierent from those for
N-transforms (Table 2). When dealing with smaller
animals, hunters usually break o the ends of bones to
form a cylinder, and then push or suck out the
marrow (Jones, 1993:108). If humans were targeting
and breaking bones for marrow extraction, we would

anticipate finding bones that have high-quality marrow

which were missing both ends. The femur and tibiotarsus contain thick marrow that fills the cavity
completely (Higgins, 1997; pers. obs.) and, therefore,
should be missing both ends. Marrow in the coracoid
and tarsometatarsus fills the cavity but is somewhat
thinner in consistency (McDonald, 1939; Hogg, 1984;
Higgins, 1997; pers. obs.). Because the marrow resides
nearer to the humeral end of the coracoid, one would
expect this portion of the bone to be missing more
often. These bones may be broken for marrow extraction, but because the marrow is of lesser quality they
may be less fragmented (sensu Lyman, 1994a, b) than
the femur and tibiotarsus. The marrow in the radius
and ulna is thin in consistency and low in fat content.
The humerus, a pneumatic bone lacking marrow,
should not be found to be broken.
Disarticulation and butchering of birds may introduce C-transformational patterns that dier from
those relating to marrow extraction. If patterns of
breakage and cut marks reflect meat removal and
disarticulation, these should be associated with the
origin and insertion of the major muscle groups on the
bones examined (Table 3). For example, about eight
major muscles originate/insert on the proximal end of
the tibiotarsus, with the muscle mass decreasing distally. It is likely that butchering would result in the
fragmentation/destruction of the proximal end of this
bone. Therefore, a large distribution of distal tibiotarsi
may indicate butchering activities.
It can be especially dicult to distinguish various
patterns associated with C-transforms and, depending
on the extent of carcass exploitation, these patterns
may be confounding. For example, an assemblage
containing a relatively high incidence of bones which
lack fatty marrow may be an artefact of rendering and
destroying the bones that contained fatty marrow
rather than a reflection of the bones themselves as a
food choice. Therefore, this process would leave a
faunal signature that produces a negative relationship between the abundance of the fatty-marrow bearing bones exploited and food value variables, a pattern
Brink (1977) identified in relation to the rendering of
bone grease.

1454 J. Higgins
Table 3. Location of cut marks if various muscles are removed
Element

Location of cut marks

Humerus

Glenoid fossa; proximal end; proximal

ventral surface; proximal dorsal surface;
deltoid tuberosity; medial epicondyle; lateral
epicondyle

Coracoid

Anterior medial surface; posterior (sternal)

end; anterior surface; anterior lateral surface
Caudal surface; distal medial condyle; distal
end; anterior surface; proximal head

Femur

Associated muscles

Tibiotarsus

Proximal tibial crest; caudal surface;

proximal medial condyle; proximal anterior
surface

Tarsometatarsus

Anterior surface; posterior surface; lateral

surface

Biceps brachii; humerotriceps; pectoralis

major; supracoracoideus; deltoideus; teres
major; latissimus dorsi; coracobrachialis;
triceps brachii; pronator longus and brevis;
flexor carpi radialis and ulnaris; anconeus;
ulnaris lateralis; extensor communis
Supracoracoideus; sternocoracoideus;
scapulo-humeralis anterior
Semitendinosus; adductor longus;
gastrocnemius; quadriceps femoris;
semimembranousus; iliotibialis;
caudofemoralis
Gluteus maximus; sartoris; femorotibialis
medius; ambiens; caudofemoralis; tibialis
anterior; gastrocnemius; peroneus longus;
flexor perforans and digitorum longus;
extensor digitorum longus
Tibialis anterior; gastrocnemius; peroneus
longus; flexor hallucis*; extensor hallucis*;
extensor digitorum longus; flexor digitorum
longus

* The only muscle tissue present along the entire length of the tarsometatarsus bone.

It is sometimes dicult to distinguish between Cand N-transforms because there are potential areas of
overlapping patterns (Tables 1 and 2). For example,
the absence of the humeral end of the coracoid, the
proximal and/or distal end of the tibiotarsus and the
proximal and/or distal end of the femur may result
either from the structural properties of these bones or
the extraction of marrow. Similarly, although a large
distribution of distal tibiotarsi may indicate butchering
activities, the cortical wall of this bone is thickest
distally. Therefore, the presence of distal tibiotarsi may
also may be an artefact of the structural properties of
this bone.
Importance to archaeological studies
In several recent studies (e.g. Emslie, Speth &
Wiseman, 1992; Godfredsen, 1997), bird remains have
been examined in an attempt to understand human
butchering activities. However, results of each of
these studies are confounded by natural transformational processes described above. Godfredsen
(1997) attributed the fragmentation of wing elements in
an assemblage of gull bones to cultural processes.
Acknowledging the poor meat content of the wing, she
argued that wing bones must have been valued for their
marrow. However, the marrow quality of the ulna is
poor and the humerus, a pneumatic bone, lacks
marrow. It is likely that non-cultural transformational
processes (Table 1) produced the breakage patterns
that Godfredsen (1997) attributed to human activity.
Livingston (1989) recognized that the dierential
representation of avian bones in archaeological

assemblages may relate to the physical structure of the

elements, which varies with the functional anatomy of
the bird. She (1989:545) suggested that the . . . relative
robusticity of appendicular elements is also related to
feeding and escape behavior: birds that feed in shallow
water and take flight easily (ducks of the genus Anas)
have more robust wind elements than birds that feed in
deep water and/or must build up momentum before
they can fly (grebes, ducks of the genus Aythya and
large rails). These results also suggest that a morphological factor such as bone density is aecting element
survivorship. Livingston was correct in assuming that
disparities exist in the preservation potential of various
avian skeletal elements. However, if her definition of
robusticity relates to bone size to density, she was
incorrect in assuming that the patterned survivorship
of various elements was because of density.
In a preliminary study, I examined bones representing five skeletal elements (coracoid, femur, humerus,
tarsometatarsus and tibiotarsus) of 13 species of ducks
in the genera Anas and Aythya and grebes in the family
Podicipedidae. These taxa were grouped as strong
flyers (Anas) and gracile-winged birds (Aythya and
Podicipedidae), as considered by Livingston (1989). I
calculated the bulk densities (Lyman, 1994a) by
measuring the volume (water displacement) and mass
of each bone. In both groups, the bones with the lowest
bulk density were the tarsometatarsi and femora. The
bulk density of the wing elements of the gracile-winged
birds was significantly higher than the densities of the
wing elements of the strong flyers; this was consistent
for all five elements (Table 4). However, many of the
bones of Anas (strong flyers) are larger than those of

Archaeological Implications of Avian Taphonomy 1455

Table 4. t-tests on dierences in bone density between Anas and
Aythya and Podicipedidae

1.8

Element

1.4

df

t-value

0680
0933

6
5

10559
19034

<00001
<00001

0717
1018

6
5

6133
8827

<00009
<00003

0416
0780

6
5

5635
6485

<00013
<00013

0476
0822

6
5

5159
5105

<00021
<00038

0760
0842

6
5

8989
4538

<00001
<00062

N=13.
Table 5. Average density (g/ml) for five skeletal elements grouped by
taxon
Taxon
Aythya and Podicipedidae

Anas

1.6

P-value

Element

Average bulk
density (g/ml)

Humerus
Coracoid
Femur
Tibiotarsus
Tarsometatarsus
Humerus
Coracoid
Femur
Tibiotarsus
Tarsometatarsus

093
084
082
10
078
068
075
048
072
043

Aythya and Podicipedidae. Therefore, robusticity

glossed as size is not an indicator of bone density.
However, if bone density is regressed against cortical
wall thickness (Figure 4), 47% (R2 =0468; P<00001)
of the variation in bulk density can be explained by
cortical wall thickness.
Emslie, Speth & Wiseman (1992), observing many
more Gruiform leg elements than wing elements in an
archaeological assemblage, concluded that these birds
were used for food. Similarly, Ericson (1987) compared
the distribution of avian wing elements versus leg
elements in archaeological and paleontological assemblages to uncover evidence of human activity. The
assumption underlying each of these studies is that if
humans were targeting the portion of the bird with the
most muscle mass, this should result in the dierential
selection of anterior and posterior elements. However,
analyses of bulk bone densities (above) indicate a
general trend for dierential survival of anterior and
posterior elements (Table 5). Dierential preservation
of elements may therefore relate merely to bone density
and may not reflect human activity.

Density (mg/ml)

Humerus
Anas
Aythya and Podicipedidae
Tibiotarsus
Anas
Aythya and Podicipedidae
Tarsometatarsus
Ana
Aythya and Podicipedidae
Femur
Anas
Aythya and Podicipedidae
Coracoid
Anas
Aythya and Podicipedidae

Mean

1.2
1.0
0.8
0.6
0.4
0.2
0
0.04

Density = 0.06 + 8.02

0.06

thickness; R = 0.468

0.08
0.10
0.12
0.14
Cortical wall thickness (mm)

0.16

Figure 4. Regression of bulk density to cortical wall thickness for 65

ducks and grebes (Aythya, Anas, and Podicipedidae).

Tu nelan archaeological case study

Examination of avian remains from the Tu nel archaeological site illustrates how detailed attention to the
natural transformational eects on bird bones can
clarify archaeological interpretations and increase our
resolution of prehistoric avian exploitation. Tu nel is
located along the shore of the Beagle Channel in Tierra
del Fuego, Argentina. It is a multi-component shell
midden site that dates from 6200100 years ago (ya)
to 45060 ya (Orquera, Piana & Haydee Tapia de
Bradfors, 1982). I identified approximately 10,000
bones (represented by the humerus, coracoid, femur,
tibiotarsus and tarsometatarsus), many of which
showed evidence of cultural modification. Preliminary
analysis of each of the five elements indicated numerous spiral fractures and patterned breakage. Although
spiral fractures do not necessarily denote human modification (Johnson, 1985), bones fractured to expose
marrow show curvilinear patterns that include spiral
or longitudinal fractures of the shaft and flakes
(Marshall, 1989:16). Consequently, if prehistoric
Fuegians pushed or sucked marrow out of the bird
bones, this behaviour should be reflected in the breakage and fracture patterns of the Tu nel assemblagei.e.
breakage patterns should duplicate the expected
frequencies of breakage (Table 2).
Comparisons of the relative frequency of breakage
(complete bones vs. bones missing a distal end) of each
of the five elements indicate a significant dierence
(
2 =36675; P>0001; df=4). The humerusalbeit
void of marrowwas the most frequently broken
bone. The tibiotarsus ranked next, followed by the
tarsometatarsus, coracoid, and femur. Because the
marrow-bearing bones, such as the femur, were
the least broken, I conclude that the Fuegians were not
specifically targeting marrow; hence, the breakage
patterns are the result of non-cultural transformation
processes. These patterns follow the predicted patterns
as a result of N-transformation processesthe longest
bones, the humerus and tibiotarsus, are more likely to

1456 J. Higgins

break and shorter elements, i.e. the coracoid and

femur, are most likely to remain complete.
There is also a statistical dierence (
2 =9935;
P>0001; df=4) in the skeletal representation of the
five elements. As predicted, the femur and tarsometatarsus, which are the least dense, were the least
represented.
Preliminary analyses of cut marks indicate that they
are numerous and their placement is consistent (sensu
Guilday et al., 1962) in reference to anatomical location and orientation (as suggested by Lyman, 1987;
Noe-Nygaard, 1989 and Gilbert & Jimenez, 1991. The
cut marks seem to be associated with major muscle
attachments (Table 3). Therefore, it seems logical
that the marks are the result of meat removal and
disarticulation. However, further study is necessary
to determine the exact butchering techniques and
disarticulation methods used by the Fuegians.
Natural taphonomic processes acting on avian bones
(and the unique properties associated with them) cause
patterns which may be mistaken for cultural modifications. Thus, as Schier (1987:23) noted, it is necessary for archaeologists to . . . demonstrate that the
traces to be used as evidence for inferences were not
caused by other processes, especially transformation
processes.

Acknowledgements
I thank Sandra Gray, Philip Humphrey, Anne Maglia,
Linda Trueb, Mark Robbins, R. Lee Lyman and Jack
Broughton for helpful comments on an earlier version
of this manuscript. Thanks to Larry Martin and Julian
Baumel for discussions and direction regarding avian
physiology. I would also like to acknowledge the entire
community at the Natural History & Biodiversity
Research Center at the University of Kansas for its
intellectual and financial support.

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