Peter Andrews Surface modifications of the Sima de

Department of Palaeontology, Natural los Huesos fossil humans

History Museum, London SW7 5BD,
U.K.
Yolanda Fernandez
Jalvo
Museo Nacional de Ciencias Naturales,
José Gutiérrez Abascal, 28006
Madrid, Spain
Received 4 March 1996
Revision received 22 October
1996 and accepted 1 December
1996
Keywords: Atapuerca, human
remains, taphonomy, carnivore
chewing, skeletal elements, bone
breakage, scavenging, Canidae,
Felidae, Ursidae, trampling,
human behaviour.
The sample of fossil human bones from the Sima de los Huesos, Atapuerca,
has been analysed to trace parts of its taphonomic history. The work reported
here is restricted to analysis of the skeletal elements preserved and their surface
modifications. Preliminary plans of specimen distribution published 6 years
ago indicate that the skeletal elements are dispersed within the cave, but more
recent data are not yet available. Most of the fossils are broken, with some
breakage when the bone was fresh and some when already partly mineralized,
both types showing some rounding. There are few longitudinal breaks on
shafts of long bones and so very few bone splinters. All skeletal elements are
preserved but in unequal proportions, with elements like femora, humeri and
mandibles and teeth with greater structural density being best represented.
There is no evidence of weathering or of human damage such as cut marks on
any of the human assemblage, but trampling damage is present on most bones.
Carnivore damage is also common, with some present on more than half the
sample, but it is mostly superficial, either on the surfaces of shafts and articular
ends or on the edges of spiral breaks. The sizes and distribution of the
carnivore pits indicate extensive canid activity, and this is interpreted as
scavenging of the bones in place in the cave. Indications of tooth marks from
a larger carnivore indicate the activity possibly of a large felid: the marks are
too large to be produced by small canids, with the larger marks concentrated
on spiral breaks on the more robust bones, and there is no evidence of bone
crushing and splintering in the manner of hyaenas. The nature of the SH
human assemblage is also consistent with accumulation by humans, the
evidence for this being the lack of other animals, especially the lack of
herbivorous animals, associated with the humans, and the high number of
individuals preserved.
? 1997 Academic Press Limited

Journal of Human Evolution (1997) 33, 191–217

Introduction
The human fossils described here were collected during the past 10 years from Middle
Pleistocene deposits in the chamber known as the Sima de los Huesos in Cueva Mayor cave.
This is part of the Atapuerca cave system near Ibeas, Burgos, Spain. The Sima de los Huesos
(SH) deposits consist of silty clays containing the human and some carnivore remains overlain
by a cave breccia containing very abundant remains of the Middle Pleistocene cave bear Ursus
deningeri (according to Torres, 1984). The first human fossils were found in 1976 (Aguirre et al.,
1976), and at this stage the deposits had been much disturbed by cavers hunting for cave bear
teeth. Some of the human fossils came from this disturbed material, but the great majority of
them have been excavated from undisturbed sediments. The deposits have been dated to the
Middle Pleistocene on the basis of the identification of the cave bear, and in addition there are
absolute dates on some of the fossils and a calcite layer indicating an age of between 200 and
300 ka.
The fauna associated with the human (MNI maximum of 32) and cave bear (MNI
minimum of 158) remains consists entirely of carnivores. There are both large and small
carnivore species, identified as follows with MNI (according to Cervera, 1992 and more
particularly Garcia et al., 1997) in parentheses: Ursus deningeri (158), Panthera leo (5), Lynx pardina
(2), Felis silvestris (1), two canids including Vulpes vulpes (23) and Canis lupus (2) formerly assigned
to Cuoninae, and five mustelids (García et al., 1997). All of these taxa are extremely

0047–2484/97/080191+27 $25.00/0/hu970137 ? 1997 Academic Press Limited

192      

fragmentary. The total absence of any herbivores may be taphonomically significant, since it
indicates a high degree of selection, and it excludes, for instance, such taphonomic processes
as water transport or natural death accumulations. The lower part of the SH fossiliferous
infilling has abundant human remains with only a few cave bear fossils, and the upper part has
abundant bears and only a few human fossils.
The SH chamber is a small blind chamber, the bottom part of which has been infilled with
sediment. The chamber is about 17 m2 in size and is reached today by a 13 m vertical shaft.
Access to the chamber in the past is difficult to reconstruct, but it may not have been restricted
to the single vertical drop existing today. There are at least two more possible passages leading
down to it, now blocked by boulders, and at the access end near the bottom of the present
vertical shaft there is an indication of another passage at approximately the same level as the
present floor level, but again this is blocked by boulders. These are described by Arsuaga et al.
(1997), although discarded by these authors as likely access points for the Sima because of the
present blockage. Further investigation is needed to identify the probable source of both
sediments and fossils in the SH chamber.
Several models have been proposed as to the means of entry of the human fossils into the
Sima de los Huesos. Most of these have been based on the early samples which contained
many specimens from the sediments disturbed by earlier caving activity. The apparent lack of
surface modifications, the presence of abrasion, and the predominance of transverse breaks on
the long bones may have been the result of recent damage, but the analysis of the present
sample is mainly based on excavated specimens from undisturbed deposits. The taphonomic
models include natural trap, with individuals falling down the vertical access shaft (Garcia
et al., 1997), catastrophic death in or near the cave (Aguirre, 1995), with or without subsequent
transport of the bones into the chamber, hibernation deaths (of the cave bears only, Díez,
1990), carnivore accumulation, or, as suggested earlier by one of us (Y.F.-J.), anthropogenic
accumulation (Arsuaga et al., 1990). Natural traps and transport are not processes that select
by species, so that they do not explain the lack of herbivorous animals in the SH fauna. In
addition, animals trapped in a natural trap to which there was no access should have all
skeletal elements from the trapped animals more or less equally abundant. Anthropogenic
selection is consistent with the lack of herbivores, with the carnivores entering the chamber
independently (Garcia et al., 1997). Accumulation by carnivore presupposes a remarkable
degree of prey selection on the part of whatever species of carnivore was the accumulating
agent, and while this is possible it is not likely.

Methods
Tooth mark terminology follows that of Binford (1981) for the most part. Pits consist of
superficial marks lacking a long axis, and punctures are deep pits that penetrate deep through
cortical bone or along the edges of intact or broken bone. Scores are surface marks with a long
axis more than four times the perpendicular axis. Measured dimensions of pits, punctures and
scores are in all cases the minimum diameter of the feature. Multiple marks made by
multi-cusped teeth were measured in two dimensions and the total area covered by the mark
given by multiplying the length by the breadth. Dimensions of carnivore tooth marks are
provided as means, modes and ranges, and they are divided into eight categories, defined as
follows: (a) pits anywhere on the surface of shafts but not on articular or broken ends [Figure
5(a)]; (b) scores anywhere on the surface of shafts but not on articular or broken ends [Figure
5(b)]; (c) puncture marks on articular ends of bones up to and including epiphyseal lines
           193

(Figure 6); (d) puncture marks on the edges of spiral breaks exposed in profile and probably
associated with the break (figure 3 bottom); (e) puncture marks on the edges of transverse
breaks exposed in profile and probably associated with the break (Figure 3); (f) puncture marks
along the edges of split bones exposed in profile and probably associated with the break; (g)
molar punctures made by multi-cusped teeth usually on flat areas of bone [Figure 5(a)]; (h)
puncture marks on the angular edges of some elements [e.g., on ribs and on scapular spines,
Figure 5(d), Figure 7 top].

The human sample

We analysed 1646 human fossils from the Sima de los Huesos. Most of these come from
excavation of undisturbed sediments, and although detailed field records of the distribution of
the fossils have been kept, these results have not yet been analysed. Some distribution data are
available from the early stages of the excavation (Arsuaga et al., 1990: Figure 1), and these
indicate a lack of direct association between elements. One example is given of the remains of
several parietal and occipital skull fragments, a mandible, an atlas, cervical and two thoracic
vertebrae and two clavicles and scapulae scattered within an area of about 40 cm2, although
it is not clear if these belonged to a single individual (Arsuaga et al., 1990: Figure 1). Arsuaga
et al. (1997) describe five instances of bones from single individuals being found in different
parts of the cave; for example fragments of Cranium 8 were recovered from areas extending
over at least 11 m. In the other four cases, fragments from single individuals were separated by
2–4 m. More detailed distribution data will soon be available, but the evidence available at
present suggests disturbance of the skeletal elements either during accumulation or post-
depositionally. This could have come about either by movement of the sediments into or
within the SH chamber or by the action of some biogenic factor such as scavenging by
carnivores or hibernation of cave bears.
All parts of the human skeleton are represented in the collection (Arsuaga et al., 1990),
although not in equal numbers [Figure 1(a)]. Phalanges and teeth are the most common
elements, with tarsals and carpals, femora and tibiae being the next most common (Table 1:
the number of teeth shown here is an underestimate because part of the sample had been
removed for study and we were not able to study them). It is evident from this distribution that
all sizes of bones are being recovered, with no bias against any particular size class, but the
absolute numbers can be misleading unless they are related to the expected numbers of
elements based on the numbers of individuals.
The number of human individuals (MNI) is presently 32 (Bermúdez de Castro & Díez,
1995; Bermudez de Castro & Nicolas, 1997) based on detailed age analysis of teeth. Age classes
ranged from 4–35 years, with young individuals (less than 11 years) and old (more than 27
years) being under-represented. Well over half the sample (19 individuals, 59·4%) are
estimated at less than 19 years in age, and over 90% estimated as less than 27 years old at time
of death (Bermúdez de Castro & Díez, 1995). This detailed an age breakdown is possible for
the teeth but more difficult for postcranial elements, and as an alternative we have calculated
MNIs for each element taking into account divisions into proximal and distal ends and
numbers of shafts (not all of which had been given registration numbers), but not size and
robusticity differences or differences in age. For example, we counted the numbers of proximal
and distal ends of each bone and took which ever was the greater, and then we added shafts
(or the opposite end plus shaft) if the level at which the shaft was broken impinged on the end
for which we were basing the count (Table 2).
194      

300
(a)

Numbers of specimens
250

200

150

100

50

0
Humerus
Ulna
Radius
Clavicle
Femur
Fibula
Tibia
Ribs
Metatarsal
Metacarpal
Vertebrae
Sacrum
Patella
Pelvis
Scapula
Calcaneum
Astragalus
Tarsal & carpals
Phalanges
Mandible
Skull
Left lower M1
Teeth
100.00%
(b)
90.00%
Relative abundances

80.00%
70.00%
60.00%
50.00%
40.00%
30.00%
20.00%
10.00%
0.00%
Humerus
Ulna
Radius
Clavicle
Femur
Fibula
Tibia
Ribs
Metatarsal
Metacarpal
Vertebrae
Sacrum
Patella
Pelvis
Scapula
Calcaneum
Astragalus
Tarsal & carpals
Phalanges
Mandible
Skull
Left lower M1
Teeth

90
(c)
80
70
60
MNE

50
40
30
20
10
0
1 1.2 1.4 1.6 1.8 2 2.2
Structural density
Figure 1. (a) Numbers of specimens of skeletal elements for the SH human assemblage. (b) Relative
abundance of skeletal elements based on MNIs of 20 (.), based on the most abundant postcranial element
(femur), and 32 (/), based on age distribution of the teeth. (c) Numbers of elements compared with structural
density of the elements.
           195

Table 1 Relative abundances of all skeletal elements based on MNI=20

(femur) and MNI=32 (teeth and jaws, Bermudez de Castro and
Diez, 1995)

Relative Relative
Individuals abundance abundance
Total no. represented MNI=20 MNI=32

Humerus 33 12 60% 37·50%

Ulna 24 6 30% 18·80%
Radius 26 6 30% 18·80%
Clavicle 17 8 40% 25·00%
Femur 52 20 100% 62·50%
Fibula 30 8 40% 25·00%
Tibia 61 17 85% 53·10%
Ribs 25 1 5% 3·10%
Metatarsal 23 3 15% 9·40%
Metacarpal 25 3 15% 9·40%
Vertebrae 32 2 10% 6·30%
Sacrum 12 6 30% 18·80%
Patella 14 7 35% 29·90%
Pelvis 35 8 40% 25·00%
Scapula 15 6 30% 18·80%
Calcaneum 8 4 20% 12·50%
Astragalus 10 5 25% 15·60%
Tarsal and carpals 65 4 20% 12·50%
Phalanges 278 5 25% 15·60%
Mandible 18 13 65% 40·60%
Skull 14 14* 70% 43·80%
Left lower M1 13 13 65% 40·60%
Teeth 83 32† 100% 100%

*Based on fragments of frontal bone with supra-orbital torus.

†Based on Bermudez de Castro & Diez (1995) taking ages of individuals into
account.

The greatest MNI value is 20 for the femur. The tibia is also well represented, but the MNI
for most elements is one to eight (Table 1). MNI figures used in Figure 1(b) are 32 based on
teeth and 20 based on femora. The distributions (relative abundances) of skeletal elements are
similar for the two MNI values, but the elements appear to have greater relative abundances
with the lower MNI value. In both, there is clear differentiation between the robust elements
such as femur, tibia, mandible and humerus, and the weaker, more easily destroyed, elements
such as radius, clavicle and fibula in the limb bones and vertebrae and ribs in the axial skeleton
(Boaz & Behrensmeyer, 1976). The skull is relatively well represented, as are also the sacrum,
scapula and pelvis, and the distribution of elements compared with the structural density
figures from Boaz & Behrensmeyer (1976) is shown in Figure 1(c) (r=0·7929, y=55·8751
x"57·4967).
This analysis indicates that the human fossil assemblage is derived from a number of
relatively complete skeletons, since all skeletal elements are preserved. To what extent they
were articulated is unknown because the bones appear to have become dispersed within the
SH chamber and no information is presently available as to associations between elements.
The size and density differences between different elements, complete skulls and pelves, for
example, compared with the large numbers of phalanges and teeth, are such that would not
be produced by transport of disarticulated bones, although mass transport with the sediments,
196      

Table 2 Numbers of specimens by bone portion

Prox. end Distal end Split

Complete with shaft with shaft Shaft shaft Blade MNI

Humerus 3 8 15 7 — — 12
Ulna 0 14 1 9 — — 6
Radius 0 7 6 13 — — 6
Clavicle 0 7 0 10 — — 8
Femur 2 26 14 10 7 — 20
Fibula 1 9 7 13 — — 8
Tibia 3 13 20 25 — — 17
Ribs — 4 1 20 — — 2
Metatarsal 23 — — — 2 — 3
Sacrum 2 2 8 — — — 6
Patella 14 — — — — — 7
Pelvis 4 — — — — — —
Pelvis acetab — 13 — — — — —
Pelvis ilium — — — — — 18 8

for example in a mud flow, is possible. Another possibility for the loss of the weaker elements
is that it was produced by some form of post-depositional or diagenetic destruction. This
possibility will be considered below when surface modifications are described, but in general
the bones in SH are well preserved with little sign of diagenetic change.

Breakage
Most of the fossil human bones from SH are broken. We investigated this breakage at two
levels, one making a simple distinction between transverse, spiral and longitudinal breaks, and
the other assessing completeness in terms of numbers of shafts and articular ends.
On the major long bones (comprising humerus, ulna, radius, clavicle, femur, tibia, fibula
and ribs) there were 487 broken ends on 283 specimens. Of these, 54% were classified as
transverse breaks and 46% as spiral breaks. Some of these breaks could have been due to
recent damage caused by the disturbance of the deposits by cavers, and so it was necessary to
investigate the types of breakage on fossils excavated from undisturbed deposits. Using data
kindly provided by J. L. Arsuaga, we were able to identify 280 breaks on bones excavated from
undisturbed sediment, and 52% of these were classified as transverse and 48% as spiral, similar
to the total sample. We conclude that both types of breakage are almost equally represented
in the SH assemblage, indicating both green-bone breakage, when the bone was fresh, and
dry-stick breakage some time after deposition but not recent. In making these counts we
included both isolated fragments of limb bone and the breaks on fragments that have since
been discovered to be associated when there was evidence that the fragments were not in direct
association, i.e., when they were found separately, often in different years, and given different
registration numbers.
Most of the breaks described above were the broken ends of bone cylinders. In other words
the shafts were not split longitudinally to produce shaft splinters. These are commonly found
in both carnivore-ravaged assemblages and are also the product of human processing (Lyman,
1994). Only 38 limb bone specimens in the SH sample show evidence of splitting (n=268),
which at 13·4% is a very low figure. It is possible that additional specimens will be found that
           197

100%

Cumulative percent of fragments

90%
80%
70%
60%
50%
40%
30%
20%
10%
0%
Humerus

Ulna

Radius

Clavicle

Femur

Fibula

Tibia

Ribs
Figure 2. Fragmentation of seven long bone elements and ribs. ( ) shaft; ( ) distal end with shaft; ( )
proximal end with shaft; ( ) complete.

could not be attributed to skeletal element, i.e., unidentified bone splinters, but we did not see
any, and their apparent absence excludes the possibility that the SH human assemblage has
been ravaged by bone crushing carnivores like hyaenas.
The same set of 283 major limb bones was also examined for degree of completeness. Only
nine bones could be classified as complete, three each of humeri and tibiae, two femora and
one fibula. This is only 3% of the specimens, and 5% based on MNI. The breakdown into
proximal, distal ends and shafts is shown in Figure 2, with the percentages of complete bones
shown at the bottoms of the bars and proportions of broken fragments shown above. The
pattern is the classic one for ravaged assemblages, with distal humerus, proximal femur and
ulna, and shafts of radius, clavicle, fibula and tibia being the most abundant body parts. This
result is consistent with the breakage data given above and also with the element analysis in the
previous section, where it was shown that the weaker elements are under-represented in the
assemblage. This confirms the action of destructive forces on the SH human bone assemblage,
that the weaker bones are under-represented because of post-mortem loss.

Surface modifications
The SH sample of human fossils was examined for the following modifications: weathering,
both surface and subsurface; rounding or scarring of bone surfaces from abrasion by water
transport or trampling; cut marks or cut mark mimics made by human action or trampling
again; insect damage; and evidence of carnivore action. Some of these proved not to be
relevant to the SH assemblage, for example weathering and cut marks, both of which are
totally absent (also observed by Díez, 1990). This is a clear indication of the absence of surface
exposure of the bones before burial in the cave and absence of human action in the
disarticulation of the bodies. Evidence of trampling was ubiquitous, with most of the fossils
showing some degree of surface damage, with flaking, scarring of the surface, cut mark mimics,
grooving and rounding of broken ends (Figure 3). No pattern was observed in the distribution
of trampling marks except in the degree of rounding of broken ends. It was also noted that
198      

Figure 3. Abrasion and rounding of broken ends, transverse breaks above and spiral breaks below. Note also
the colour change in the two conjoined fragments, below.

conjoining bone fragments may have differential colour and degree of post-depositional
damage even when both fragments came from the undisturbed excavation levels (Figure 3,
bottom). Without knowledge of the relative positions of associated fragments, it is not possible
to say why these differences exist.
All broken ends of long bones were scored for rounding or absence of rounding. It was
found that slightly more of the transverse breaks were rounded than were the spiral breaks.
Fossils from the undisturbed deposits showing evidence of rounding had 24·8% of transverse
breaks rounded and 20·7% of spiral breaks. Corresponding figures for the total sample,
including material from the disturbed sediments on the surface in addition to that from the
excavations, were 22·5 and 18·6%, respectively, showing that the rounding was not the
product of recent cave activity. Both spiral and transverse breaks with some degree of rounding
are shown in Figure 3.
It is assumed that transverse breaks occur some time after death, when bones are partially
mineralized, and spiral breaks occur at or soon after death. The greater degree of rounding on
the transverse breaks indicates that the process producing the rounding must have operated at
a time considerably after burial and it suggests possible reworking. This could have been the
result of movement of the bones in the sediment or of sediment transport, although there is no
           199

Figure 4. Surface modifications of one of the crania. Left, probably insect action; right, a surface scrape
where the skull rubbed against the cave wall.

clear sedimentological evidence for the latter. It has been demonstrated that abrasion by silty
clay matrix may produce rounding of fossilized bones more rapidly than fresh bone, which
again suggests modification some time after deposition (Fernandez-Jalvo, 1988).
One of the skulls, Cranium number 5 (Arsuaga et al., 1993), has two surface modifications
on the vault (Figure 4). One of these is a series of grooves with rounded U-shaped
cross-sections apparently resulting from chemical solution (Figure 4, left). These grooves are
too long and regular to have been caused by plant roots, and although we lack comparative
data we are interpreting this as the result of insect action. This was the only instance of this
modification that we observed. On the same skull, at the top of the vault, is another series of
grooves and gouges cut into the surface of the bone (Figure 4, right). These have striations
along the bottom of the marks, and since this specimen was found with this part of the skull
in contact with and pushed up against the wall of the cave, the gauges and the striations are
interpreted as the result of movement of the skull against the cave wall.

Evidence for carnivore activity

Apart from post-depositional trampling marks, the most abundant surface modifications on the
SH fossil accumulation resulted from carnivore damage. Carnivore damage was found on
more than half the sample of major limb bones and ribs, with the highest number on femora
(50 out of 52 specimens had carnivore damage) and the lowest number on the pelvis (13 out
of 36)—see Table 3 and Figures 5–7. There is no apparent relation between the robusticity of
the skeletal element and the incidence of carnivore damage.
The distribution of carnivore damage is shown on Table 3 divided into seven categories as
defined above. In each case the average size of the modification is shown on the left and the
number of specimens from which the mean value was calculated shown on the right. Gnawing
of articular ends is also present, for example on the glenoid area of the scapula and the
mandibular condyle [Figure 5(c)], and rounding due to carnivore action is also occasionally
present [Figure 5(f)]. Carnivore damage occurs most frequently on the shafts of limb bones, on
their articular ends, and on spiral breaks. For example, there are 235 pits or puncture marks
on the surface of shafts (category a in Table 3), with a distribution on the skeletal elements
similar to the general distribution mentioned above, although there are differences: the
greatest number of marks is on the scapula, which of course has a greater surface area, and
there are also many marks on the femur, humerus and fibula. There are just under half this
number of marks on the articular ends (category c in Table 3), but again with a similar
distribution.
 200

Table 3 Average size of puncture marks and grooves (mm)

Nc N % a n b n c n d n e n f n h n

Humerus 17 33 51·5 1·8 34 1·0 21 2·5 22 4·7 9 2·1 7 4·0 1

Ulna 10 24 41·7 2·8 17 1·9 16 2·2 8 2·1 19 2·4 5
Radius 6 13 46·2 1·3 5 0·7 2 1·9 1 6·4 1 1·8 1
Clavicle 12 17 70·6 1·1 15 2·7 8 2·2 12 2·4 7
Femur 50 52 96·2 1·8 36 0·8 14 2·3 28 4·2 18 3·5 8 3·4 1
Fibula 15 26 57·7 1·0 40 0·4 9 1·1 5 2·1 15 2·4 7 2·5 3
Tibia 19 46 41·3 2·5 16 1·1 29 2·8 14 2·5 11
Ribs 13 25 52·0 1·7 17 0·6 18 3·5 8 2·2 8 5·8 3
Sacrum 8 12 66·7 5·0 5 4·4 5 2·4 10 3·8 2
Pelvis 13 36 36·1 3·6 7 3·1 3 3·6 1 4·2 5
Scapula 10 15 66·7 2·9 43 3·2 20 3·8 2

Nc=Catalogued specimen with carnivore damage.

N=Total number of catalogued specimens.
n=Total number of marks.
a=Carnivore punctures on surface of bone (minimum dimension).
b=Carnivore gnawing on surface of bone (transverse measurement of grooves).
c=Carnivore punctures on articular surfaces.
     

d=Carnivore punctures on spiral breaks.

e=Carnivore punctures on transverse breaks.
f=Carnivore punctures on split shafts.
h=Carnivore punctures on intact bone edges.
           201

Figure 5. Carnivore action. (a) surface pits, with both isolated pit upper right, and a molar pit left; (b) surface
groove; (c) removal of the mandibular condyle by chewing; (d) edge puncture on the side of a rib; (e) surface
pits on the shaft of a humerus; (f) extensive chewing of a phalanx.

Figure 6. Carnivore punctures in the acetabulum of a pelvis made after disarticulation of the pelvis and
femurs.

The distribution of carnivore damage is shown on Figure 8. This shows categories (a)–(h)
from the bottom to the top in the cumulative bar chart. All of the long bones follow the same
general pattern, but the sacrum, pelvis and scapula have a different pattern because of their
different morphology. On all the major limb elements, with the exception of the clavicle, 50%
202      

Figure 7. Chewing of the scapular blade.

100%

80%
Total features

60%

40%

20%

0%
Humerus

Ulna

Radius

Clavicle

Femur

Fibula

Tibia

Ribs

Sacrum

Pelvis

Scapula

Figure 8. Numbers of carnivore features, expressed as cumulative percentages and divided into seven
chewing categories for the major postcranial elements: see text for definitions of the categories. ( ) Punctures
on intact edges; ( ) punctures on split shafts; ( ) punctures on transverse breaks; ( ) punctures on spiral
breaks; ( ) punctures on articular surfaces; ( ) scoring on surface of bone; ( ) pits on surface of bone.

or more of the carnivore damage is confined to the surfaces of the shafts (categories a and b).
Damage on broken ends of bones varies between 30–50%, and the total of 135 marks (Table
3) were found on 77 of the 487 broken ends. This distribution pattern of carnivore damage
shows that the SH assemblage has been extensively modified by carnivores that had limited
capacity to break human bones, and this is consistent with the lack of breakage described
earlier and particularly the low frequency of bone splinters.
           203

7.0

6.0

5.0
Size (mm)

4.0

3.0

2.0

1.0

0.0
Humerus Ulna Radius Clavicle Femur Fibula Tibia
Figure 9. Mean sizes of carnivore features on the seven major long bone elements, divided into the seven
categories of carnivore damage: see text for definitions. ( ) Pits on surface of bone; ( ) scoring on surface
of bone; ( ) punctures on articular surfaces; ( ) punctures on spiral breaks; ( ) punctures on transverse
breaks; ( ) punctures on split shaft; ( ) punctures on intact edges.

Also shown on Table 3 are the mean sizes of the different categories of carnivore damage.
The mean sizes of punctures on articular ends are generally larger than those on the surfaces
of limb bone shafts (Figure 9) as would be expected in the softer bone of the articular surfaces.
Also the punctures exposed in the broken ends of bone shafts are generally larger than the
surface pits, and those on spiral breaks are larger than the ones on transverse breaks (Figure
9). Finally, punctures on soft bone, such as sacrum, pelvis and scapula, are consistently larger
and deeper (penetrating the cortical bone) than those on more compact bone (Tables 3 and 4).
The size distribution of carnivore punctures is shown in composite form in Table 4 for each
of the main skeletal elements. The modes of the size distributions are as follows:

0·6–1·0 mm—humerus, tibia, ribs

1·1–1·5 mm—femur, fibula, radius
1·6–2·0 mm—clavicle
2·1–2·5 mm—ulna, sacrum
2·6–3·0 mm—scapula
3·6–4·0 mm—pelvis

Although they have the size distributions skewed towards small diameter pits, the major limb
elements like humerus, femur and tibia also have the widest range of pit sizes, all of them
having marks ranging above 7·6 mm (Table 4). This is an indication that more than one
carnivore might be responsible for the marks, although none of the elements has a bimodal
distribution of pit sizes. Some of the skulls also have evidence of carnivore tooth marks: Skull
6 has a puncture 6·4#7·2 mm on the right side behind the broken zygomatic arch and
another slightly smaller one (5·0#5·2 mm) on the left, and there is scoring on Skull 3 with
cross-sectional diameter 2·4 mm.
 204

Table 4 Size distribution of carnivore chewing features by skeletal element

Humerus Ulna Radius Clavicle Scapula Femur Fibula Tibia Sacrum Pelvis Ribs

0·0–0·5 4 — — 1 — 13 13 2 — — 10
0·6–1·0 26 5 2 9 1 53 22 22 — — 10
1·1–1·5 13 10 4 6 4 58 23 16 — — 7
1·6–2·0 6 12 3 12 5 26 3 15 3 1 8
2·1–2·5 8 20 — 3 10 17 8 2 7 2 6
2·6–3·0 4 11 — 5 13 12 3 9 5 2 2
3·1–3·5 6 4 — 1 5 10 — — 1 3 1
3·6–4·0 1 3 — — 6 13 1 2 1 4 1
4·1–4·5 1 2 — 2 2 7 1 1 — 1 1
4·6–5·0 — — — 1 3 6 — 1 1 1 1
5·1–5·5 2 — — 1 2 2 — 3 1 — 2
5·6–6·0 1 — — — — 2 — — 1 1 —
6·1–6·5 1 — 1 — — — — — — — 1
6·6–7·0 1 — — — — 1 — — — — 1
7·5–7·5 — — — — — 2 — 1 — 1 —
>7·6 2 — — — — 3 — 3 1 — —
     
           205

As an alternative to the composite analysis in Table 4, we have analysed two types of

carnivore damage separately for all elements (Table 5). The size distribution of the 235 pits on
the surfaces of limb bone shafts is shown on Figure 10(a). They make up more than one third
of all the carnivore modifications, and they have a distribution heavily skewed towards the
small end of the size range. They are also the marks most abundant on robust elements such
as femora and humeri [shown by the arrow on Figure 10(a)] so that they are influencing the
distributions shown in Table 3. By contrast, there is a much wider size distribution of carnivore
punctures on spiral breaks [Figure 10(b)], which make up 15% of the carnivore damage, and
the sizes of these punctures on robust elements are much larger than are the surface pits on the
same bones, i.e., humerus [shown by the black areas at the bottoms of the bars of Figure 10(b)]
and femur [the shading indicated by the arrows on Figure 10(b)]. This difference in size
distribution is highly significant (t-test difference in means P=0·001), and the differences
between the superficial marks on femora and humeri and the punctures on spiral breaks on the
same bones is an indication that more than one size of carnivore was producing the damage,
with a larger carnivore producing the spiral breaks and a smaller species producing the surface
damage but failing to break the bones.
One further type of carnivore damage, labelled as category g but not shown on any of the
preceding analyses, was investigated. This is damage we have interpreted as caused by
multi-cusped teeth or molar punctures. They are few in number (Table 6), and they vary
greatly in size. The average size of the tooth puncture is calculated from the length
plus breadth divided by two, and the average area is the two dimensions multiplied together
(Table 6). There is much variation in size, with one very large feature 78 mm2 in area on a
sacrum.

Comparison with canid scavengers

The size distribution of the carnivore tooth marks is extremely close to that of a recent
assemblage of sheep bones scavenged by foxes Vulpes vulpes (Andrews, 1995). This assemblage
is part of an ongoing taphonomic project started in 1978 based on the accumulation of bones
from natural deaths in an undisturbed part of central Wales. The bones from both wild and
domestic animals are being observed in a long-term experiment, and the assemblage analysed
here consists of the lag deposit of sheep bones recovered from an area of approximately 16 km2
in the late 1970s (Andrews & Armour-Chelu, 1997). The bones have been dispersed both by
scavengers and by water transport, and many have been modified by trampling and
weathering. The scavengers are mainly foxes, but some sheep dog activity is also present, the
Welsh border collie being a dog only slightly larger than male foxes. Badgers and other
mustelids are also abundant in the area and probably also contribute to the scavenging impact,
although there is no direct evidence of this.
The great majority of sheep bones from the Welsh fox-scavenged assemblage remain intact.
A sample of 73 bones have been analysed for five limb elements that are comparable between
sheep and humans (humerus, radius, femur, tibia and ribs). Of these 48 are still complete, or
66% compared with 3% complete in the SH human sample.
The overall distributions of carnivore marks on the Welsh sheep bones are extremely similar
to the SH human assemblage both in proportional numbers of marks and in their size
distribution, although there are fewer surface pits on most of the recent bones. The pattern on
the latter seems to be that some bones have a large number of surface pits whereas most bones
have few or none. There are many puncture marks on the articular ends of limb bones
(category c), and the size of these punctures is consistently larger than the surface marks on the
 206

Table 5 Size of punctures on the surface (a) and on spiral breaks (d) of major postcranial elements

Humerus Ulna Radius Clavicle Femur Fibula Tibia Ribs

a d a d a d a d a d a d a d a d

0·0–0·5 1 — — — — — 1 — — — 4 — — — — —
0·6–1·0 4 — — — 4 — 10 — 22 — 19 — 1 2 3 —
1·1–1·5 9 — 2 3 1 — 4 — 45 — 14 7 4 — 5 1
1·6–2·0 1 — 1 8 — — 1 6 12 4 1 1 6 2 4 1
2·1–2·5 — 1 4 7 — — — 2 5 2 — 3 — 2 3 2
2·6–3·0 — — 4 — — — — 4 5 1 — 2 2 3 — —
3·1–3·5 — 2 3 — — — — — 2 2 — — — — — —
3·6–4·0 — — 2 — — — — — 1 8 — — 1 1 — 1
4·1–4·5 — 1 — 1 — — — — 1 4 — 1 — — — —
4·6–5·0 — — 1 — — — — — 1 2 — — — — — 1
5·1–5·5 — 2 — — — — — — — 2 — — — 1 — 2
5·6–6·0 — 1 — — — — — — — 1 — — — — — —
6·1–6·5 — 1 — — — 1 — — — — — — — — — —
6·6–7·0 — 1 — — — — — — — — — — 1 — — —
7·5–7·5 — — — — — — — — — 2 — — 1 — — —
     
           207

90
(a)
80
Punctures on surface of bone
70
60
50
40
30
20
10
0
0.0–0.5

0.6–1.0

1.1–1.5

1.6–2.0

2.1–2.5

2.6–3.0

3.1–3.5

3.6–4.0

4.1–4.5

4.6–5.0

5.1–5.5

5.6–6.0

6.1–6.5

6.6–7.0

7.1–7.5
Size classes (mm)
25
(b)
Punctures on spiral breaks

20

15

10

0
0.0–0.5

0.6–1.0

1.1–1.5

1.6–2.0

2.1–2.5

2.6–3.0

3.1–3.5

3.6–4.0

4.1–4.5

4.6–5.0

5.1–5.5

5.6–6.0

6.1–6.5

6.6–7.0

7.1–7.5

Size classes (mm)

Figure 10. (a) Size distribution of surface pits on the seven long bone elements plus ribs. (b) size distribution
of punctures on spiral breaks on seven long bone elements plus ribs. Cumulative percentages for the eight
postcranial elements. ( ) Ribs; ( ) tibia; ( ) fibula; ( ) femur; ( ) clavicle; ( ) radius; ( ) ulna; ( )
humerus.

Table 6 Number and dimensions of molar punctures (mm)

Average Average
Number size area Minimum Maximum

Humerus 2 6·3 34·1 26·6 41·5

Ulna 2 3·1 9·3 6·5 12·0
Femur 8 4·9 26·6 4·7 31·4
Ribs 1 1·5 2·4
Sacrum 2 7·5 57·4 36·0 78·8
Pelvis 1 4·5 19·1
Scapula 1 6·0 35·6
208      

same bones. There are also puncture marks on spiral breaks (category d), as in the SH sample,
with a size range from 1·1–4·7 mm and mean values from 2·3–3·1 mm compared with values
for the SH sample in Table 3 of 2·1–6·4 mm on the comparable bones, with maximum values
reaching 7·5 mm (Table 5). The size distribution for surface pits on the fox-scavenged sheep
assemblage (Figure 11) is not significantly different from the SH sample despite the fact that
there are some larger pits on the latter, but the distribution of punctures on spiral breaks on
the SH sample is significantly different (t-test on distribution of means P>0·05) from that of the
fox-scavenged sheep assemblage (Figure 11) because there are many more large punctures,
although the lower parts of the size distributions are again rather similar. This is a strong
indication that the SH sample has been scavenged by a fox/small dog type of carnivore and
that a larger carnivore also produced a limited amount of damage. The relative impact of the
two sizes of carnivore is difficult to assess because it is likely that some of the smaller marks
would also have been made by the large carnivore. It can be emphasized again that there is
an upper size limit on the size of the tooth marks made by small carnivores, so that they could
not have made the marks larger than this size, but the larger carnivores could have made both
the large and some of the smaller marks.
There are also few longitudinal breaks producing bone splinters in the Welsh fox-scavenged
assemblage. Only 16 bone splinters representing nine indeterminate bones have been
recovered. Proportional representation of splinters when added to the sample of 73 identified
bones is thus 19·5% compared with 13·4% in the SH human assemblage (see above).
One aspect of the carnivore damage on the recent assemblage differs from the SH fossil
assemblage. The punctures on spiral breaks in the recent assemblage are no larger on robust
elements like humeri and femora than they are on weaker elements. This contrasts with the
SH human assemblage, where the robust elements have larger punctures on the spiral breaks.
Foxes seem to be unable to produce spiral breaks in adult sheep bones such as humeri and
femora, and the breaks observed on the recent sheep humeri and femora were all on juvenile
or very small individuals. Puncture sizes on these spiral breaks were the same size as those on
more fragile bones like radii and ribs. Human bones are larger and more robust than sheep
bones, and the larger punctures on adult human femora and humeri in the SH assemblage
could not have been made by small canids. This indicates the presence or activity of another
carnivore producing this damage.
Other aspects of the fox-scavenged sheep assemblage differ from the SH sample because of
differing preservational conditions. For example, the sheep bones are exposed in the open, and
they constitute an attritional assemblage with a weathering profile: 43 bones are not weathered
at all, 21 are weathered at Stage 1, five at Stage 2, three at Stage 3, and one at Stage 4
(weathering stages as defined in Behrensmeyer, 1978). Most of the bones were not broken, but
15 out of the sample of 73 had spiral breaks, which is 21% compared with 153 spiral breaks
on the 197 comparable bones (i.e., humerus, radius, femur, tibia, ribs) in the SH human
assemblage, or 78%. Several of the spiral breaks had some degree of rounding as a result of
heavy chewing. No transverse breaks have been observed in the sheep assemblage, the oldest
bones of which are now 16 years after death and most no more than 8–12 years (Andrews &
Armour-Chelu, 1997).

Carnivore activities
Considering the scavenging of foxes on modern sheep carcasses, we may say that fox tooth
marks are generally less than 4·0 mm in diameter. Based on the size range of fox tooth marks
(1·1–3·7 mm), we divided the size ranges of carnivore damage on the SH assemblage (taken
           209

40
(a)
35
Percentage of punctures
on bone surface 30
25
20
15
10
5
0
0.0–0.5

0.6–1.0

1.1–1.5

1.6–2.0

2.1–2.5

2.6–3.0

3.1–3.5

3.6–4.0

4.1–4.5

4.6–5.0

5.6–6.0

6.1–6.5

6.6–7.0

7.1–7.5
Size class (mm)
30
(b)
25
Percentage of punctures
on spiral breaks

20

15

10

0
0.0–0.5

0.6–1.0

1.1–1.5

1.6–2.0

2.1–2.5

2.6–3.0

3.1–3.5

3.6–4.0

4.1–4.5

4.6–5.0

5.1–5.5

5.6–6.0

6.1–6.5

6.6–7.0

7.1–7.5

Size class (mm)

Figure 11. (a) Size distribution of surface pits for the same bones as in Figure 10 but without differentiation
into elements, comparing the SH assemblage from Atapuerca with a present-day assemblage of sheep
scavenged by foxes in Wales. (b) Size distribution of punctures on spiral breaks for the same samples. (/)
Neuadd; (.) Atapuerca.

from Table 4) into three groups (Figure 12). The group from 1·1–4·0 mm is considered to be
the tooth mark range of foxes, but it may also include chewing marks of a larger carnivore.
The other two groups are the very small tooth marks and the largest ones. The former group
(0·1–1·0) is considered most likely to be the result also of fox activity, while the latter group
(4·1–>7·6) is identified exclusively as damage from large carnivores. The anatomical elements
have been ordered (from one to 18) following the flesh and marrow consumption sequence of
small and very small mammal long bones (according to Blumenschine, 1986).
We have also found different patterns between anatomical elements and size of tooth marks
(Figure 12). As indicated by Blumenschine (1986), defleshed marrow bones (femora, tibiae,
humeri and radii, together with head contents) are most frequently available to scavengers. On
the contrary, pelvis, and lumbar vertebrae are chewed by predators in early stages. Therefore,
the largest carnivore is considered to have had priority access to the human bodies of SH,
because hindquarter (axial parts) are affected by their teeth in higher proportions than is
210      

30
(a)

20

10

0
f)

f)

f)

f)

f)

f)

)
5f

6f

8f
(1

(2

(3

(4

(7

l(8
p(

b(

d(
Pv

Fm

Tb

U
R
Sc

R
Sc

H
100
(b)
80

60

40

20

0
f)

f)

)
Sc )

Sc )

)
f/1 m
f/1 m

f/1 d
f/1 b

5f

6f

8m
4m

5m

6m

8m
f/1 l
(1

(3

(8 U

(8 R
(4 T

(7 H
(2 F

p(

b(
Pv

50
(c)
40

30

20

10

0
(1 Fm
Pv

b
(1 Tb

(1 Hm

(1 d
)

(1 Ul
)

R
Sc

)
5m
4m

R
Sc

6m

8m

8m

Figure 12. Size distribution of carnivore tooth marks linking size of features with skeletal elements. (a)
Carnivore features 4·1–7·6 mm in diameter; (b) carnivore features 1·1–4·0 mm diameter; (c) carnivore
features 0·1–1·0 mm diameter. Anatomical elements are ordered according to the consumption sequence of
Blumenschine (1986). f, Flesh; m, marrow.

present in other elements, and the smallest size of chewing-marks (considered to be foxes) are
absent on these elements. Fox activity, therefore, is consistent with scavenging rather than
predation, not surprising considering the relative sizes of humans and foxes.
           211

Figure 13. (a) Vertebra of bear from SH with a puncture mark matching a bear canine from the same site.
(b) Vertebra of bear from SH with the broad impression of a multi-cusped tooth. Note the three flat cusp
prints which accord well with bear molars.

Figure 12(a) shows the biggest size of chewing marks. These marks follow, from high to low
abundance, the consumption sequence for flesh ingestion, with highest percentages on pelvis
and lowest on ulna and radius. Figure 12(c) shows the higher abundance of the smallest
chewing marks on long bones that are most frequently available to scavengers (Blumenschine,
1986), and absence of marks on pelvis and sacrum, available for initial consumers. Figure 12(b)
follows the consumption sequence less well because this range includes chewing marks of foxes
mixed, to some extent, with those of the large carnivore.

Discussion
The distribution of body parts of the human assemblage in SH cave differs from the
background scatter of bones found in natural conditions today (Behrensmeyer, 1983; Lyman,
1994). Such dispersed bone was shown to have approximately equal numbers of skulls,
vertebrae, forelimb elements and hindlimb elements. It also differs from the bone distributions
in areas of accumulation of predation kills, what Behrensmeyer (1983) calls a ‘‘predation
patch’’. These are characterized by high frequencies of vertebrae, and this has also been found
with the fox-scavenged bone distribution in Wales (Andrews, 1995), where vertebrae are
usually the last skeletal elements to remain at death sites. In both cases the difference in
distribution can be attributed to the removal of limbs by scavengers. In contrast to these
patterns, the SH human assemblage is dominated by hindlimb elements, followed by forelimb
elements, with vertebrae and skulls relatively infrequent, and this is the pattern present in
hyaena den assemblages (Behrensmeyer, 1983).
The size and nature of many of the carnivore tooth marks on the SH assemblage are
consistent with scavenging by small to medium-sized canids. Scavengers of this size would be
unable to transport whole or partial human bodies, and so they can be discounted as possible
212      

agents of accumulation of the human bones. They would be able, however, to carry off and
totally destroy disarticulated smaller bones such as ribs and vertebrae and foot bones, and their
presence could account for the loss of these bones, and of the skeletal remains of infantile
individuals, which are only represented by a few teeth in the SH assemblage.
The evidence for another carnivore producing modifications to the SH human assemblage
is based on the larger size of some of the tooth marks both on the bone surfaces and on spiral
breaks. This pattern is not matched in the recent fox-scavenged assemblage, for which there
is a distinct upper limit (of around 4 mm) in minimum diameter of tooth marks. This indicates
the presence of a larger carnivore inflicting damage on the human assemblage, although it was
evidently one that produces minimal damage to the bone since there are so few split shafts
(common in hyaena assemblages, Binford, 1981) and the numbers of marks are low. It is also
probable, however, that some of the smaller pits and scoring may have been caused by this
larger predator. Possible candidates for this include the large felids P. leo fossilis and Panthera
gombaszogensis, and the cave bear U. deningeri (Cervera, 1992), but hyaena action is excluded
because of the low frequency of split shafts in the human sample.
Cave bear remains are extremely abundant in SH. At the present time, bears may scavenge
ungulate carcasses (Clevenger, 1991), and in addition to their main diet of vegetation and
insects, bears scavenge during the whole year with a maximum in summer (12% of their diet
by volume) and a minimum in winter (5%). However, according to descriptions of bone
modifications produced by different types of carnivores (Haynes, 1983), bears may gnaw across
cancellous tissue in a similar way to hyaenas and wolves, but typical damage appears as
rounding of broken edges (Haynes, 1983). This is not consistent with observations on the SH
human assemblage. As seen above, rounding observed on the SH fossils are produced by
movement of the bones in the sediment, because rounding affects both spiral and transverse
breakage, particularly the latter.
According to Haynes (1983) bears do not usually gnaw heavily on bones before the soft
tissue has dried or been removed. Bears may not leave tooth mark scores on the shaft surfaces,
but, when they do, they produce scores that ‘‘appear similar to rodent gnaw marks: short and
parallel, shallowly etched, straight score lines’’ (Haynes, 1983: p. 169). This feature has not
been observed on the human fossils of SH. In contrast to this, bear fossil bones from Sima de
los Huesos show clear tooth marks very likely made by bears. Pits and puncture marks on fossil
bear bones are frequent on the surface (category a) ranging up to 10·4 mm diameter, as well
as, on articular surfaces (category c) and on intact bone edges (category h). Scoring (category
b) is not as frequent as seen in human bones, and this is consistent with Haynes’ observations.
Further, bear canines and molars from Sima de los Huesos closely match the pits and puncture
marks on bear bones (Figure 13). This evidence agrees with modern bear behaviour.
According to Kurt (1976), cannibalism is more frequent among true bears than among any
other group of mammals.
Contrary to the evidence for carnivore damage on fossils of bears, the marks on human bones
do not match with bear teeth of the SH assemblage. Human fossils have tooth scores in the
compact tissue of the diaphysis which are, as described by Haynes for felids, perpendicular to
the element’s long axis, and shallow but rather sharply incised. The basic identifying charac-
teristic of large cat gnawing is, according to Haynes’ observations, the rough and irregular
marking on the epiphysis, that are wide, deep and countable, inflicted by the cheek teeth. Both
modifications have been observed on the SH human fossils. It is also noteworthy that carnivore
fossil bones in the Sima deposits have no evidence of carnivore damage, and it has been
observed that most felids, especially lions, never eat carnivores (Rudnai, 1971; Schaller, 1972).
           213

Figure 14. Human bones broken by fall of 13 m. Top left, three femora; bottom left, two humeri, radius and
ulna; top right, the remains of a single skull; bottom right, two mandibles.

Modifications like those produced today by hyaenas or large canids have not been observed
in the SH fossil assemblage. This is particularly evident in the near absence of split shafts that
are so characteristic of hyaena assemblages (Villa & Mahieu, 1991). The larger tooth marks
observed on the human fossils of SH are most similar to modifications made by felids, more
especially by lions [Figure 12(a)]. Lions closely follow the consumption sequence described by
Blumenschine (1986, 1995; see Schaller, 1972; Rudnai, 1971; Kitchner, 1991), with all parts
of the prey being eaten except for rumen contents, horns and bones. They chew and swallow
edges of ribs, pelvis, scapula and the nasal bone of herbivores, and may split some bones
(Rudnai, 1971; Schaller, 1972). Rudnai (1971) observed that young lions eat bones, including
teeth. This might also account for the loss of infantile bone remains. There is some evidence
that lions characteristically disembowel large prey and drag the digestive tract away and bury
it but this is a variable behaviour pattern that may explain why human ribs of SH assemblage
(Figure 12) have large, medium and small tooth marks.
Leopards produce a pattern of bone modification similar to that of lions (Bailey, 1993). They
typically eat prey the size of impalas until all edible parts are gone, including large parts of the
ribs and axial skeleton, parts of the skull and even sometimes the hooves. They leave the leg
bones and between half and a quarter of the axial skeleton. The vertebral ends of ribs are also
left, but smaller prey or immature individuals are consumed in their entirety. The resulting
bone assemblages therefore consist of leg bones, fewer foot bones and skulls, and fewer still
vertebrae and ribs.
214      

A common behaviour of felids, also frequent in lions, is to carry their prey away for some
distances (from a few metres to more than 400 m) before feeding begins (Rudnai, 1971;
Schaller, 1972; Kitchner, 1991; Kruuk & Turner, 1967; see also Bailey, 1993 for similar data
for leopards). The purpose of this behaviour is to avoid other predators and scavengers which
may be attracted to the kill. It is also common that when too much food is caught for one meal,
large felids may store several individuals for later (they can eat 18–30 kg of meat in one meal).
At the present time leopards are known to accumulate bones of their prey in caves (Simons,
1966; Brain, 1969, 1981), and the cave lion may have done likewise, although we have no
evidence that this was the case. Felids may also remain in the place where they have stored
their kill for days until finishing it or to preserve their prey from scavengers (Kruuk & Turner,
1967; Rudnai, 1971; Schaller, 1972). However, although lions consume all edible portions,
especially if a group feeds together, they also can abandon a considerable amount of meat for
scavengers (Schaller, 1972). All these features are consistent with the feeding sequence that
appears in SH.
There are many accounts of man-eating tigers and lions (Kingdon, 1977; Bourlière, 1955;
Schaller, 1972). Individual predators have been known to prey on up to 140 people in the
course of few months, and there is one case of 20 people killed by a pride of five lions, with the
young animals learning from the old (Kitchner, 1991). It is conceivable that lions could have
incorporated humans in their diet at that time and be the source of the humans of SH, but this
is at best circumstantial evidence that they were the primary agents of accumulation. The fact
that only humans are found in the site, with no herbivores, would indicate a degree of
specialization that would be unusual for present-day felids, and the high number of individuals
contrasts with data on modern hunter–gatherer groups, which indicates that average
population sizes do not exceed one individual per square kilometre (Hassan, 1981). The extent
of the hunting areas (Schaller, 1972) and/or the length of time needed to accumulate such a
large sample would require that lions drag their prey several kilometres for them to accumulate
their victims in a single den.
Accumulation of humans by other humans was formerly proposed by one of us (Y.F.J.). This
hypothesis considers that other humans brought dead bodies to this site as the result of a still
unknown behaviour and were thrown into the SH chamber down the 13 m shaft, which
provides the present access to the SH chamber (Arsuaga et al., 1990; Garcia et al., 1997). To
gain some insight into the possible consequences of such a process, we performed an
experiment using human bones to determine the degree of breakage by dropping them from
a 13 m height on to a hard concrete surface. This does not simulate exactly the effect of intact
bodies dropping from this height, but it would seem likely that the damage to bones
unprotected by flesh would be greater than would be the case for bones in carcasses. In the
event, most of the bones did not break or were damaged in any way, but some of those that
did break are shown in Figure 14. Two skulls used in the experiment were smashed into small
pieces except for the frontal region and parts of the cranial vault. This is a pattern of breakage
very similar to that seen in the SH assemblage. Three out of six femora broke with spiral
breaks on the shaft of one specimen and breakage of the neck two others (Figure 14). Two out
of six humeri broke, one with a jagged break of the shaft and proximal end. One fibula shaft
also broke, but no other bones. Altogether the proportion of complete bones was 82%, a very
different figure from the 3% in the SH fossil assemblage. In so far as this experiment is
relevant, it does not support the hypothesis that the human sample accumulated in this way,
and there is no direct evidence of human involvement other than the fact that only humans are
present (other than carnivores) to the exclusion of herbivores.
           215

There is also little evidence in support of the SH chamber acting as a natural trap. The
skeletal part representation is not consistent with the preservation of whole animals, and the
removal of some elements by scavengers would seem to exclude the possibility that it was a
trap of any kind. In addition, natural traps are not normally highly selective as to species,
although it is possible that an accumulation of humans in such a trap could have attracted the
carnivores also present. It is strange, in this case, that the carnivore remains do not show any
sign of damage by other carnivores except for the bears, which have not been modified either
by large felids or small canids but appear to have tooth marks made only by other bears.
A possible scenario can be formulated that takes much of this evidence into account. There
may have been a nearby source of human bodies left exposed after death by the local human
population to which large felids had access. Lions are known to scavenge recent kills, and they
may have carried some of the bodies (or parts of bodies) into the cave to protect them from
more aggressive scavengers. Absence of hyaena or wolf chewing marks, in contrast to the
presence of felid chewing marks, suggest that the bodies were adequately protected from the
former species (both known from Galeria, a site with similar age, 400 m from SH). The most
abundant carnivore damage on the human bones was from small canids which indicates that
these animals gained access to the bodies, probably after the large felids finished eating. These
carnivore activities may have occurred in some other part of the cave, with the residual human
bones transported into the SH cavern by mud flow, producing post-depositional breakage and
rounding of the bones; or alternatively, if access was possible to the SH chamber they may
have occurred in place, with the post-depositional changes occurring as a result of sediment
movement within the chamber.

Conclusions
(1) The SH fossil assemblage contains abundant human fossil bones together with bears and
some carnivore fossil bones. No herbivore remains have yet been found.
(2) The high abundance of human remains found in SH, lack of herbivores and the strange
association between humans, bears and carnivores make this site exceptional and leads to the
suggestion that the accumulating agent for the human remains were other humans.
(3) There are no stone tools, no human-made cut marks on any of the fossil remains, and
no evidence from the human population structure that would support human involvement in
the bone accumulations. There is no evidence of human occupation of the cave.
(4) The proportions of skeletal elements in the SH fossil assemblage is typical of present-day
carnivore assemblages. There are carnivore tooth marks on over half the human assemblage,
with two sizes of carnivores indicated on these fossils: a lion-sized carnivore that had priority
access to the human bodies, and a fox-sized canid which scavenged the remains left by the
former. These modifications are not present on any of the carnivore bones.
(5) Tooth marks made by cave bears have been observed only on bear bones. They have
not been observed on either the human bones or on the smaller carnivores.
(6) There is no evidence that lions were or were not the cause of death of the human
individuals found in SH. The large carnivore could have been the agent of secondary
accumulation or re-location, scavenging recently dead corpses either in the cave or transport-
ing them into the cave, with preference given to hindlimbs and perhaps also skulls.
(7) Scavenging by small canids is the main process of surface modification of the SH human
bones. The evidence of carnivore damage produced by a fox-sized scavenger on more than
half the human bones indicates heavy scavenging pressure, and the comparative rarity of fox
216      

remains in the deposits suggests that the foxes were not trapped in the cave. If they were able
to get in and out of the cave, it also explains the absence of many of the ribs and vertebrae that
are under-represented in the human assemblage. The absence of postcranial remains of very
young individuals may be attributed to scavenging pressure.
(8) There is no evidence from the breakage of the human bones that they entered the cave
by being dropped down the 13 m vertical shaft. Breakage is greater than can be attributed to
a fall of this magnitude.
(9) Post-depositional rounding on the broken ends indicates movement with or within the
sediment some time after deposition (recent caving activity can be excluded as the agent of
abrasion). In the absence of excavation data, we cannot verify the nature of the geological
agents. However, the great difference of tooth marks observed on humans and bears could
suggest that human and carnivore remains were subject to different taphonomic processes.
(10) At this stage, therefore, it is not possible to come to any conclusion as to how the
human fossil assemblage accumulated. Further work is needed to verify the human skeletal
proportions, the population structure of the human population, and the distribution of human
bones in the sediments to resolve this issue.

Acknowledgements
We are grateful to T. Torres for finding the first human fossils, and to E. Aguirre, who always
promoted taphonomic studies and for starting the research and excavation project at
Atapuerca. Thanks are extended to J. L. Arsuaga and to the Sima de los Huesos team,
especially I. Martínez and N. García, for facilities given during the analysis of this study. We
have benefited from discussions with Ana C. Pinto, J. Rodríguez, J. C. Díez, J. M. Bermúdez
de Castro and J. L. Arsuaga. We are grateful for the help given by L. Humphrey in analysing
the data and to R. Lee Lyman and one of the anonymous referees for severe but positive
criticism that has done much to improve this paper. Part of the documentary material was
made by the photo units of the Natural History Museum of London and the Museo Nacional
de Ciencias Naturales. This research was made possible by a grant from the L. S. B. Leakey
Foundation.

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