Characterisation of microbial attack on archaeological bone

M.M.E. Jans
a
*, C.M. Nielsen-Marsh
b
, C.I. Smith
c
, M.J. Collins
d
, H. Kars
a
a
Institute of Geo- and Bioarchaeology, Faculty of Earth and Life Sciences, Vrije Universiteit, De Boelelaan 1085, 1081 HV Amsterdam,
The Netherlands
b
NRG, Drummond Building, University of Newcastle, Newcastle upon Tyne NE1 7RU, UK
c
Museo Nacional de Ciencias Naturales (CSIC) C/Jose Gutierrez Abascal 2, 28006 Madrid, Spain
d
BioArch, The Kings Manor, University of York, York YO7 1EP, UK
Received 9 January 2003; received in revised form 8 May 2003; accepted 21 July 2003
Abstract
As part of an EU funded project to investigate the factors inuencing bone preservation in the archaeological record, more than
250 bones from 41 archaeological sites in ve countries spanning four climatic regions were studied for diagenetic alteration. Sites
were selected to cover a range of environmental conditions and archaeological contexts. Microscopic and physical (mercury
intrusion porosimetry) analyses of these bones revealed that the majority (68%) had suered microbial attack. Furthermore,
signicant dierences were found between animal and human bone in both the state of preservation and the type of microbial attack
present. These dierences in preservation might result from dierences in early taphonomy of the bones.
2003 Elsevier Science Ltd. All rights reserved.
Keywords: Archaeological bone; Diagenesis; Microbial attack; Mercury intrusion porosimetry; Histology
1. Introduction
Bone is used extensively as a source of information in
archaeological research as it is usually the only animal
or human tissue preserved, though often it is altered or
destroyed as well. With the development of biomolecu-
lar analysis, the signicance of understanding bone
preservation has increased. One of the earliest types of
alteration is microbial attack. Yoshino et al. [42] ob-
served evidence of microbial alteration in bone 5 years
post-mortem, indeed Haversian canals were (still) lled
with bacteria. Bell et al. [5] found evidence of microbial
attack in bone in a forensic study, as soon as 3 months
post mortem, though this sample was recovered from a
predator scat.
Biological alteration in bone is in most cases caused
by fungi [31], bacteria [2,14,23], or cyanobacteria in
marine environments [4]. Wedl rst described microbial
attack on mineralised tissue in 1864, nding tunnels of
approximately 8 m in diameter in sections of teeth
exposed to untreated well water and in fossil reptile
teeth. This type of tunnelling is dened as Wedl or
centrifugal tunnelling by Hackett [14]. These tunnels
range in diameter from 5 to 10 m and under electron
microscopy [31] appear empty with well-dened calcied
walls, implying that collagen and mineral are both
resorbed by the fungi.
Three types of microscopical focal destruction
(mfd)described by Hackett as linear longitudinal,
budded and lamellateare generally assumed to be
caused by bacteria. These mfd can be distinguished
histologically by morphology; size, shape, the presence
of a hypermineralised rim and the presence of a lamel-
late content. Under light microscopy these mfd show a
granulous or brillar content, but higher magnication
shows that their interior actually consists of small pores
[2,23,40]. Mercury intrusion porosimetry (HgIP) shows
an increased volume in pores with diameters in the
0.110 m region, with the largest increase observed at
approximately 0.6 and 1.2 m [35,40], consistent with
the average diameter for bacteria (0.5 m [18]). Indeed,
Baud and Lacotte [2] argue that the small pores that are
visible at electron microscope level are the mineralised
* Corresponding author. Tel.: +31-20-4447295; fax: +31-20-6462457
E-mail address: miranda.jans@falw.vu.nl (M.M.E. Jans).
Journal of Archaeological Science 31 (2004) 8795
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remains of bacteria, in which case they would not form
the interconnected porosity observed by Turner-Walker
et al. [40].
We report here some results from a European study,
which attempted to identify the key contributory factors
to bone deterioration. Biological alteration of archaeo-
logical bone is a widespread phenomenon apparently
not limited to a single burial environment [21]. Its eect
on bone preservation is important as it can accelerate
degradation by increasing the bone porosity [35] and
simultaneously reduces chances of success for bio-
molecular research through loss or contamination of
target molecules [9]. Two methodological approaches,
mercury intrusion porosimetry (HgIP) and histology,
are combined for the rst time in this project. While
histology has been widely used previously to characterise
microbial attack (e.g. [3,11,14]), HgIP makes it possible
to analyse a larger sample of bone and provides
quantitative results [40].
2. Materials and methods
Samples of animal and human bone and related
soil were taken from excavations in Sweden, the
Netherlands, the United Kingdom, Italy and Turkey,
spanning four climatic regimes (Mediterranean, Conti-
nental, Maritime (coastal) and Subarctic) (Appendix A).
The 41 sites were chosen on the basis of well-dened
geological and archaeological context and documented
environmental parameters such as ground water table,
annual rainfall and land use history. Sites ranged from
the Neolithic to early modern times, and sampled fea-
tures included burials in con burials, stone lined graves
and rock-cut graves, settlement layers, refuse pits and
postholes. In addition, physical anthropological or
zooarchaeological data were gathered where possible
(full details are available in Kars and Kars [28]). If
possible long bones (preferably femur) were sampled.
The samples for the various analyses were always taken
from adjacent locations on the bone (preferably the
mid-shaft area of long bones).
2.1. Mercury intrusion porosimetry (HgIP)
Samples of approximately 0.51.5 g were cut from
cortical bone (usually femur) using a hacksaw. Samples
were dried for at least 48 h in an oven at 60 (C and then
analysed using an automated Micrometrics Autopore II
9220 [43]. The method estimates porosity by assuming
cylindrical pores and a contact angle between bioapatite
and mercury of 163.1( [27]; for details see [40].
2.2. Histology
The sample taken for histology was a bone section
sawed to 30 m, prepared as described in previous
studies [22,26] and viewed using transmitted and polar-
ised light microscopy. The percentage of bone micro-
structure unaected by microbial deterioration was
represented by the Oxford Histological Index (HI),
developed by Hedges et al. [20] using Table 51.1 (p. 640)
of Millard [32]. The type of mfd was determined based
on the morphological description provided by Hackett
[14] (Fig. 1).
3. Results
3.1. Characterisation of microbial attack
Two hundred and sixty-one bone samples from
41 dierent archaeological sites were analysed using
histology and 233 samples of these using mercury
intrusion porosimetry as well. Of the samples analysed
histologically (n=261), 177 samples showed microbial
attack (68%). Only 9% of the total sample could be
considered well preserved. In 12% of the bones the
characteristic Apigliano style degradation [38] could
be observed. Apigliano style degradation is character-
ised by highly crystalline and altered mineral phases,
and loss of collagen, with no histological alteration
except small micro-ssures. It remains unclear which
degradation mechanism is responsible for this type of
preservation. Excluding these atypical bones from the
analysis brings the percentage of microbially attacked
bones to 87%. The HI distribution (Fig. 2) shows a
similar pattern as found by Hedges et al. [20] with
the majority of bones being either highly altered or
unaected.
The type of mfd was assessed for the 177 samples
showing microbial alteration. Bacterial alteration,
specically linear-longitudinal and budded tunnelling, is
the most common type of tunnelling (85% of microbially
attacked bones show bacterial attack). A total of 55
specimens showed Wedl tunnelling, 101 linear longitudi-
nal, 99 budded and 48 lamellate tunnelling (one bone
sample can simultaneously be aected by more types of
tunnelling). Most mfd were commonly observed alone as
well (Fig. 3) but this was not true of lamellate tunnelling.
Despite being morphologically very dierent to budded
tunnelling, lamellate was always found in association
with it. Wedl tunnelling (fungal) is most frequently
observed in association with linear longitudinal tunnel-
ling, but the relationship was much less exclusive than
that between budded and lamellate. In fact, linear
longitudinal tunnelling is found at similar ratios with all
types (2030, Fig. 3).
To see whether the dierences between mfd were
reected in the porosity measurement, bones were
selected that showed only one type of tunnelling, and
average porosity distributions for these groups were
compared. This was not possible for lamellate tunnelling
M.M.E. Jans et al. / Journal of Archaeological Science 31 (2004) 8795 88
since this type of mfd is hardly ever found alone.
Therefore we included averages for bone showing
lamellate mfd, regardless of other mfd present. The
averaged traces are remarkably similar (Fig. 4).
Wedl tunnelling (n=23) shows the bacterial porosity
(between 0.1 and 1.2 m) as well as an additional
porosity at approximately 4 m, which is presumably the
tunnelling visible at the light microscopy level. The
presence of the double peak is unexpected as Wedl
tunnels do not show the network of small pores found in
bacterial mfd, appearing empty both at light and elec-
tron microscopical level [31]. Interestingly, in the lamel-
late traces (n=30) the rst of the double peaks indicates
a smaller modal pore size (approximately 0.3 m) than
the other mfd (approximately 0.6 m).
3.2. Animal and human bone
Dierences were found between animal and human
bone with respect to the number of bones aected as
well as the type of attack. The dierence in the extent of
attack is visible in the average porosity traces for animal
and human bone (Fig. 5). The percentage of biologically
altered human bone was 75%, diering signicantly
from animal bone where this was 57% (
2
(1,
N=261)=7.5, P%0.01). This dierence becomes more
Fig. 1. Schematic representation of the four types of tunnelling as characterised by Hackett [14]; 1: Wedl (fungal), 2: linear longitudinal, 3: budded
and 4: lamellate tunnelling.
0
10
20
30
40
50
60
70
80
90
5 4 3 2 1 0
OxIord HI
N
u
m
b
e
r
Fig. 2. The distribution of the HI categories in the sample (n=261). The
HI ranges from 5 (well preserved) to 0 (no original microstructure left)
[20]. Indicated in grey in the HI=5 bar are the Apigliano style samples
[38].
M.M.E. Jans et al. / Journal of Archaeological Science 31 (2004) 8795 89
pronounced when considering specically bacterial
attack (i.e. ignoring fungal tunnelling; Wedl) with fully
74% of human bone being altered but only 34% of
animal bone (
2
(1, N=261)=40, 613, P%0.001).
Fig. 6 shows that bacterial attack is responsible for
the majority of biological alteration in archaeological
human bone: linear longitudinal and budded tunnelling
are both frequently observed, while lamellate tunnelling
is rarer. Fungal tunnelling is less commonly found. In
contrast, Wedl tunnelling seems to be the dominant
form of tunnelling in animal bone, closely followed by
linear longitudinal tunnelling. Lamellate tunnelling is
only present in very few samples.
Logistic regression analysis is a statistical method to
model in a binomial setting the mean of response
variable (p) (binomial proportion) in terms of an ex-
planatory variable (x) [33]. In this model we used the
proportion of bacterially altered bones as the response
variable and whether or not the bone is human as the
explanatory variable (Human=1, Animal=0). This
analysis shows that human bones are more likely to be
aected by bacterial attack than animal bones (odds
ratio: 5.4, 95% CI=3.29.3). The odds ratio is the ratio
of the odds for the two possible outcomes; the odds are
the ratio of the proportions of possible outcomes. How-
ever, bone from a complete burial is slightly more likely
to be aected by bacterial attack as is shown when
complete burial is taken as an explanatory variable
(odds ratio: 6.5, 95% CI=3.811.4). This association
of complete articulated burial and bacterial attack is
further conrmed by the fact that bacterial attack is
present in all animal bone samples from articulated
Wedl (27)
Budded (20)
Lamellate (1)
24 30
21
2
1
20
6
Lin.long. (25)
Fig. 3. Bubble chart showing the relationships between the dierent
types of tunnelling, a thick line indicating a common relationship
(n=20), a dotted line an uncommon relationship (n<10). The number
of times a certain relationship is found is indicated in the small
bubbles. The number of times a certain type of tunnelling is found on
its own is indicated between parentheses.
0.00
0.01
0.02
0.03
0.04
0.05
0.001 0.01 0.1 1 10 100
Diameter (m)
I
n
t
r
u
s
i
o
n

m
l
/
m
l
Modern
Linear longitudinal
Budded
Lamellate
Wedl
Fig. 4. Mercury intrusion porosimetry plot showing the porosity traces
found for the dierent types of tunnelling; Wedl (n=23), linear
longitudinal (n=13), budded (n=12) and lamellate (n=30). The thick,
black line represents a modern unaltered bovine bone.
0.00
0.01
0.02
0.03
0.04
0.05
0.001 0.01 0.1 1 10 100 1000
Diameter m
I
n
t
r
u
s
i
o
n

m
l
/
m
l
Animal
Human
Modern
Fig. 5. Average traces for animal and human bones (bone samples
from corrosive environments (i.e. acid and/or well aerated soils [28])
are excluded) showing clearly a larger double peak in the human
bones.
0
5
10
15
20
25
30
35
40
45
Human Animal
P
e
r
c
e
n
t
a
g
e
Wedl
Linear Longitudinal
Budded
Lamellate
Fig. 6. Bar chart showing the dierence in animal and human bone
with respect to the type of tunnelling. Values for animal bone have
been normalised to compensate the dierences occurring through
dierence in sample size.
M.M.E. Jans et al. / Journal of Archaeological Science 31 (2004) 8795 90
complete skeletons (n=9), while it is only present in 34%
of the total animal samples. The (binomial) probability
for all nine samples from complete articulated burial to
be bacterially altered (binomial distribution B(9, 0.34)) is
P%0.0001. In contrast, bone from complete burial is less
likely to be aected by fungal alteration as is shown
when fungal alteration is the response variable (odds
ratio: 0.231, 95% CI=0.1220.436).
4. Discussion
In this study we used the combination of histology
and mercury intrusion porosimetry to characterise
microbial attack. The porosity traces of all types of
bacterial tunnelling showed a double peak, with in-
creases in the pore volume at about 0.6 m and 1.2 m;
albeit the former was smaller in the lamellate tunnelling
(0.3 m). Most bones with lamellate tunnels were found
on very similar sites (large, densely occupied, medieval
cemeteries). The smaller pores may therefore reect
dierences in mineral redeposition caused by high phos-
phate concentrations occurring in the soils. From the
almost exclusive association of lamellate with budded
tunnelling it can be suggested that it is a dierent (later)
stage in microbial alteration. Either budded alteration is
a prerequisite for lamellate formation or lamellate is a
sub-component of budded tunnelling.
Disappointingly, HgIP results could not discriminate
clearly between fungal and bacterial attack. The poros-
ity traces for fungal tunnelling showed a similar doublet
peak as the traces for bacterial tunnelling. This could be
caused by bacterial attack present in the sample used for
HgIP, but not visible in the sections used for histology,
since the sample analysed with histology is much smaller
than the sample analysed using HgIP. It is also possible
that bacteria attacking bone in cooperation with the
fungi cause the presence of a bacterial signature [12].
A signicant dierence in preservation was found
between animal and human bone, which is contrary to
expectation [21]. Not only is animal bone less often
attacked, fungal attack is more common, whereas in
human bone bacterial attack is the predominant type of
attack. This picture is conrmed by previous archaeo-
logical and forensic studies (on human bone) where
Wedl tunnelling is only occasionally found [3,10,16].
There are several possible explanations for this dier-
ence. Most animal bones have a dierent micro-
structure, which enables for example species
identication on the histological level [17]. Furthermore,
Robinson et al. [36] demonstrate dierences in natural
bone porosity not only between dierent animal species
(and also between juvenile and adult individuals and
(within) dierent bone elements). As porosity can be
considered a good indicator for reactivity with ground-
water [19], the original, natural bone porosity may have
predictive value for bone survival [36]. Interestingly
the one human bone sample included in this study
showed higher porosity than the mature animal bone
samples [36].
Moreover, the burial environment of animal or hu-
man bone is dierent. Refuse or settlement layers, where
animal bone is usually found, have often a high concen-
tration of organic material, and are often anaerobic,
aiding good preservation [13,34]. Additionally, humic
factors, often present in these layers, can inltrate in
bone and act as inhibitors to collagenases, preventing
bacterial attack [34]. In contrast, graveyards may have
dormant populations of bacteria that are well adapted to
attack human bone [24]. Soil bacteria have been shown
to be able to use collagen as a growth substrate [1].
However, bones from single burials and scarcely used
graveyards, where a dormant population of conditioned
bacteria would not be expected, do not show better
preservation. Mant [30] observed in his study of 150
World War II burials that soil environment does not
play a large role in the early stages of degradation. These
are much more dependent on the condition of the body
and the burial circumstances.
Moreover, an outward-in pattern for bacterial attack,
expected when soil bacteria colonise bone, is not ob-
served [37]. Furthermore, the micro-anatomy-related
localisation of bacterial attack in bone [7] suggests that
the natural bone pores (e.g. Haversian canals) are path-
ways for invading the bone. Bone from the abdominal
area is often more aected by microbial attack, being
near to the intestines where bacterial putrefaction starts
[8]. Putrefaction occurs during the early stages of
decomposition and involves endogenous bacteria and
enzymes. Intestinal bacteria (Staphylococcus sp.,
Clostridium sp. and Escherichia coli) have been shown to
invade the body as soon as 15 h post-mortem [29]. In
soils without a large bacterial population, putrefaction
can still be quite extensive, due to endogenous bacterial
populations [25]. To inhibit bacterial degradation ex-
treme circumstances are required. Examples are the
presence of bactericidal substances near the body (like
copper, mercury or lead), extreme temperatures, rapid
desiccation (mummication), or the absence of endog-
enous bacteria for example in infants [24,25,30,41].
Another possibility is dismemberment or slaughtering
shortly after death, preventing bacteria from invading
bone tissue [39]. Most animal bone fragments will there-
fore not be aected by putrefaction, which explains the
relatively low occurrence of bacterial attack. However,
the resulting good preservation of animal bone makes it
an attractive nutrient resource for fungi present in the
soil. Saprophytic fungi are dependent on environmental
factors like the presence of oxygen and a certain level of
humidity (20%) [6]; anaerobic fungi have only been
found in the rumen of certain herbivores, where
they play a role in cellulose digestion [6]. Once the
M.M.E. Jans et al. / Journal of Archaeological Science 31 (2004) 8795 91
environmental parameters are favourable, fungal attack
becomes possible.
The results of this study show that the majority of
(human) bone is likely to be biologically altered. The
resulting increase in porosity means that this large
portion of archaeological bone is more vulnerable to
other diagenetic processes. Indeed, the relative absence
of bacterial mfd in paleontological bone shows that
biological alteration needs to be prevented if bone is to
survive into the fossil record [39]. Furthermore, biologi-
cal degradation has certain consequences for biomolecu-
lar analyses, either through contamination of samples or
complete loss of information such as aDNA (e.g. [9,15]).
5. Conclusion
Microbial attack is an important contributor to bone
deterioration. HgIP and histology can be used to detect
its eects. While HgIP analyses a larger sample volume
and gives quantitative results, histology is needed to
distinguish between bacterial and fungal attack. Bone
from complete burials (animal and human) is more
likely to be aected by bacterial attack indicating that
bacterial degradation is linked to putrefaction and the
very early stages of degradation. The majority of bone,
which entered the archaeological record as fragments
(e.g. butchered animals), is either well preserved or
shows fungal attack. Fungal alteration will occur in well
preserved bone if environmental conditions are favour-
able for saprophytic fungi and can occur at any time
during burial and following excavation. The type and
extent of microbial attack seems to be determined both
by (early peri-mortem) taphonomic factors as well as
later environmental factors.
Acknowledgements
This project has been carried out with nancial
support of the European Union, Directorate General
No. XII, in the framework of the Environment and
Climate programme (19941998), project no. ENV4-
CT98-0712. Prof. John N. S. Matthews (University of
Newcastle) is gratefully acknowledged for his help with
the statistical analysis. Two anonymous reviewers are
thanked for their valuable comments.
Appendix A. Summary of sites and samples used in this study
Site Bone element (no.) Species (no.) Soil conditions Date
GB-BIF Costa (1) Cattle (2) Clay 1000 BC
Femur (1) Human (1)
Humerus (1) Unidentied (5)
Unidentied (5)
GB-KIT Radius (1) Cattle (1) Clay 800700 BC
Unidentied (5) Unidentied (5)
GB-MIL Costa (7) Cattle (19) Anthropogenic soil AD 11001400
Humerus (2) Fallow deer (1) Silt
Mandibula (1) Pig (1) Sand
Pelvis (3) Rabbit (1) Gravel
Radius (4) Sheep/goat (3)
Scapula (2)
Tibia (2)
Ulna (2)
Vertebra (2)
GB-MOO Femur (2) Cattle (6) Clay AD 11001400
Metatarsus (3) Horse (1)
Mandibula (1) Pig (1)
Radius (2)
GB-TOR Unidentied (3) Human (3) Clay 14001000 BC
IT-API Femur (9) Human (10) Terra Rossa AD 9001400
Tibia (1)
IT-FIP Femur (2) Human (2) Clay AD 400500
Sand
Anthropogenic soil
IT-HIP Femur (1) Human (2) Silt AD 3001100
Tibia (1) Cobble
IT-LMG Femur (3) Human (3) Gravel AD 700900
Cobble
IT-MES Femur (2) Human (3) Silt 300 BCAD 300
Tibia (1)* Sand
Gravel
Cobble
M.M.E. Jans et al. / Journal of Archaeological Science 31 (2004) 8795 92
Appendix A. (continued)
Site Bone element (no.) Species (no.) Soil conditions Date
IT-SDO Femur (2) Human (2) Anthropogenic soil ca. AD 13251350
IT-SEG Femur (3) Human (3) Clay AD 11001200
IT-SFS Metacarpus (1) Human (1) Church burial AD 1495
IT-SLM Femur (14) Human (14) Anthropogenic soil AD 9001000
Silt
IT-SPB Femur (2) Human (2) Grave cut into bedrock AD 9001400
IT-SUP Femur (1) Cattle (2) Sand AD 600700
Metatarsus (2) Sheep/goat (1)
NL-AAR Femur (2) Cattle (2) Silt 28502450 BC
Humerus (1) Human (1) Anthropogenic soil
Tibia (1) Pig (1)
NL-BOG Femur (3) Human (4) Clay 2400400 BC
Humerus (1)
NL-BOR Femur (3) Horse (2) Gravel AD 600700
Human (1)
NL-GIL Femur (7) Human (9) Sand ca. AD 1620
Humerus (2)
NL-NIJ Femur (10) Human (10) Sand AD 350650
Gravel AD 15001700
Moraine AD 16001800
NL-RAA Mandibula (3) Horse (3) Sand AD 300400
NL-UBE Humerus (1) Cattle (2) Sand ca. 2010 BC, AD 70104
Long bone (2) (Large) Mammal (3)
Tibia (1) Wild duck (1)
Unidentied (1)
Vertebra (1)
NL-VOR Astragalus (1) Cattle (3) Clay AD 8001300
Calcaneus (1) Horse (1) Silt
Metacarpus (1) Mammal (1)
Radius (1)
Unidentied (1)
NL-YPE Femur (13) Human (13) Sand 40003500 BC
SE-AHU Femur (28) Human (31) Sand AD 12001500
Humerus (3) Moraine
Anthropogenic soil
SE-CAR Cranium (2) Cattle (1) Sand AD 0400
Femur (1) Human (4)
Mandibula (1)
Radius (1)
SE-CLE Metacarpus (3) Cattle (8) Anthropogenic soil AD 15001600
Radius (3) Clay
Tibia (1) Silt
Vertebra (1) Sand
SE-GOT Femur (9) Human (10) Gravel AD 32002300
Tibia (1) Cobble
SE-HAG Femur (7) Dog (1) Moraine AD 9001100
Humerus (1) Human (8)
Mandibula (1)
SE-HUS Long bone (2) Cattle (1) Moraine AD 0800
Unidentied (1)
SE-LIN Femur (1) Human (1) Silt ca. 3000 BC
SE-LOV Unidentied (4) Human (4) Sand AD 8001000
Moraine
Gravel
SE-NOR Long bone (1) Cattle (4) Anthropogenic soil ca. AD 1500
Metatarsus (1) Sheep/goat (1) Silt
Sacrum (1) Sand
Vertebra (2)
SE-SAL Humerus (1) Cattle (4) Silt 1000200 BC
Metacarpus (2) Moraine
Radius (1)
SE-ULL Humerus (1) Cattle (1) Clay AD 10001500
Mandibula (1) Pig (1) Silt
Sand
M.M.E. Jans et al. / Journal of Archaeological Science 31 (2004) 8795 93
(continued on next page)
Appendix A. (continued)
Site Bone element (no.) Species (no.) Soil conditions Date
SE-VAD Costa (1) Cattle (2) Clay AD 14001500
Cranium (2) Large mammal (2) Silt
Mandibula (1) Sand
SE-VAL Femur (3) Human (4) Moraine AD 9001100
Tibia (1)
SE-VIS Mandibule (1) Dog (1) Sand 600300 BC
SE-VNG Mandibula (1) Cattle (1) Clay AD 0400
Pelvis (1) Pig (1) Silt
Sand
SE-VUO Cortical bone (3) Large mammal (4) Sand ca. 4000 BC
Long bone (4) Moose (1)
Tibia (1) Unidentied (4)*(3)
Unidentied (1)
*Bones are charred, cooked or cremated.
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