Journal of Archaeological Science (2000) 27, 1139–1150

doi:10.1006/jasc.1999.0537, available online at http://www.idealibrary.com on

Patterns of Diagenesis in Bone I: The Effects of Site

Environments
Christina M. Nielsen-Marsh*
Fossil Fuels and Environmental Geochemistry, University of Newcastle, Newcastle upon Tyne NE1 7RU, U.K.

Robert E. M. Hedges
Research Laboratory for Archaeology and the History of Art, 6 Keble Road, Oxford OX1 3QJ, U.K.

(Received 9 May 1999, revised manuscript accepted 19 November 1999)

We have measured a suite of ‘‘diagenetic parameters’’ for several populations of archaeological bones buried in a
number of northwest European sites since the last Ice Age. These are: the structural damage due to microbes; changes
in bone micro-and macro-porosity; protein content; a measure of the crystallinity of hydroxyapatite as reflected in the
phosphate infra-red spectrum peak splitting; and a measure of the carbonate content as given by the ratio of carbonate
to phosphate infra-red absorption peaks. The results provide a database that clearly characterises the patterns, which
are often site-dependent, of diagenetic change in buried bone. Our main conclusions are that individual site hydrology
appears to have a strong influence on the outcome of bone preservation and that porosity is the most effective single
diagenetic parameter which both determines and reflects the preservation of bone in the burial environment.
 2000 Academic Press

Keywords: BONE, DIAGENESIS, HYDROLOGY, POROSITY, PROTEIN, HISTOLOGY.

Introduction measures corresponds directly to a simple diagenetic

process.

B
ones recovered from archaeological sites are
unquestionably altered over time. Alteration
occurs at all scales, from molecular loss and
substitution, through to crystallite reorganization, po-
rosity and microstructural changes and, in many cases,
to disintegration of the complete unit (Nelson et al.,
1986; Weiner & Bar Yosef, 1990; Sillen & Parkington,
1996; Sillen & Morris, 1996; Wright & Schwarz, 1996;
Nielsen-Marsh & Hedges, 1999). The burial environ-
ment largely determines the preservation of archaeo-
logical bone (White & Hannus, 1983; Piepenbrink,
1989; Stephan, 1997). The main aim of this paper is the
identification of those aspects of the environment that
may dominate the course of diagenesis, an important
task for the understanding of archaeological bone
preservation. A subsidiary aim is to show that different
environments have different qualitative effects on
diagenetic alteration.
We attempt to describe the course of diagenesis
using several measurable parameters of bone, which
are experimentally accessible and relate to processes
expected to operate during bone degradation. This
builds on earlier work (Hedges & Millard, 1995;
Hedges et al., 1995); however, as yet, none of these
*To whom correspondence should be addressed.
1139
0305–4403/00/121139+12 $35.00/0
Approach
This paper describes the results of a study which aimed
to build on the work discussed in Hedges et al. (1995).
Hedges et al. left unresolved questions concerning
the suitability of their measurements for studying
diagenetic alteration, as well as the exact relationship
between the burial environment and bone survival.
We have looked for relationships between selected
diagenetic parameters to determine the extent to which
they form a coherent pattern, and then investigated
how this pattern is altered in bone from different sites
and burial environments. Included in the Results and
Discussion section of this report are considerations of
results obtained from our laboratory data presented in
previous publications. These are the investigations into
the mechanisms behind mineral alteration (in particu-
lar crystallinity increases; Nielsen-Marsh & Hedges,
1997) and the role that site hydrology plays on bone
mineral dissolution and therefore bone survival (Pike
et al., in press).
The archaeological sites used in this study are de-
scribed in Appendix I. Each site has been chosen to
produce an overall database, which spans a time frame
 2000 Academic Press
1140 C. M. Nielsen-Marsh and R. E. M. Hedges

Table 1. Bones studied

No. of
Site Period Environment Species samples

Yarnton Bronze Age Palaeochannel silts Large mammals 6

Neolithic Muddy gravels 6
Anglo-Saxon Quartz/sand gravels 5
Romano British 6
Iron Age 6
Mid Bronze Age 6
Brean Down Beaker Clay/silt Large mammals 4
Early Bronze Age Sand/silt 4
Mid Bronze Age 8
Late Bronze Age Silt 10
Bercy Neolithic Channel bed Aurochs 6
Channel edge 4
Dry bank 2
Poundbury Roman Chalk graves Human 14
Gough’s Cave Pleistocene Cave Equus ferus 13
Piddington Roman Loam Large mammals 12
Human 7
Wharram Percy Mediaeval Chalk graves Human 11
Overton Down 32 years Turf Sheep 2
Chalk 2

of the last 12,000 years collated using bones from
a range of environments within northwest Europe.
Four main sites, Yarnton, Bercy, Brean Down and
Poundbury, provide the core data for this paper, but
the database is supplemented by less systematic data
from other sites. Sites in more extreme environments,
however (especially high temperature), have not been
included. Yarnton, Bercy and Brean Down contain a
range of locally differing environments. The bones
discussed in this report are summarized in Table 1.
Bone samples have been taken from long bones
(femurs have been used where available) of either
humans or large mammals. Where human bone was
collected, only skeletons from non-coffined burials
were sampled. If femurs were either not available or
unidentifiable, mandibles were sampled instead. The
samples were sand-blasted to remove surface dirt, then
sections were cut using a hand-held diamond wire saw,
and bone powder was obtained using a hand drill. The
bones collected for this project represent a wide range
of preservational standards, and sampling varied ac-
cording to the state of the preservation. In some cases,
removing ‘‘sections’’ proved to be very difficult, even
with a hacksaw blade, as the bones were very strong
and were consequently hard to cut; other samples were
extremely ‘‘crumbly’’ or soft and were relatively easy to
saw. Not all the bones had survived intact, and in a lot
of cases only fragments remained. For each sample at
least 350 mg was required to carry out all the necessary
diagenetic measurements.
In tandem with the data collected from archaeologi-
cal samples, controls (fresh bovine bone and deprotein-
ated bovine bone) were subjected to the same analyses.
The method for deproteination used here has been
adapted from Termine et al. (1973). Modern bovine
bone fragments of approximately 400–500 mg were
placed into a stoppered round-bottomed flask with
hydrazine hydrate, 98% (NH2NH2.H2O); 15–20 ml of
hydrazine was used for every 1 g of bone. The bone
was left in solution for 1 h at room temperature. After
1 h, the hydrazine was decanted off and a further
15–20 ml of fresh hydrazine was poured into the flask,
which was then incubated at 70C for 1 h. Further
hydrazine changes were made at 1, 10, 14 and 32 h
periods, keeping the temperature at 70C. The final
hydrazine wash was removed and the sample was then
serially diluted with absolute ethanol, replacing the
reagent at 30 min intervals with 1:1, 3:1 and 7:1 abso-
lute ethanol:hydrazine hydrate mixtures, and finally
three times with absolute ethanol. The samples were
dried in an oven at 110C.
Deproteinated bone has been included to act as a
control, because deproteination with hydrazine affects
the diagenetic parameters in much the same way, as
does prolonged burial.
We have also carried out laboratory experiments in
which fresh and archaeological bone has been sub-
jected to artificially accelerated degradation by (mainly
acidic) reagents at 20C, in order to investigate the
effects of different chemistries and of hydrological
regimes on bone preservation. The salient results have
been published elsewhere (Nielsen-Marsh & Hedges,
1997; Pike et al., in press), but where relevant are
included in the discussion.
 Patterns of Diagenesis in Bone I: The Effects of Site Environments 1141
Description and Measurement of Diagenetic
Parameters
The parameters measured here mainly correspond to
those first reported in Hedges et al. (1995), but
measurement methods have been improved since then
and an additional parameter (C/P) has been added.
Experimental details where previously unpublished
techniques have been used, or where familiar protocols
have been altered are described below. Otherwise the
relevant publications are cited.
Histology (HI)
Histological preservation is determined by the histo-
logical index (HI), a simple graded system of 0–5 as
measured and described in Hedges et al. (1995). This
system has been shown to correlate well (r>0·9) with
more rigorous quantitative methods of analysing
backscattered electron images in scanning electron
microscopy (Gardner, pers. comm.). Despite the rich-
ness and subtlety of such images, virtually all alteration
appears to consist of ‘‘destructive foci’’, first described
by Hackett (1981), where such foci differ mainly in
their extent and distribution. The circumstantial evi-
dence that the ‘‘foci’’, which in general take the form of
tunnels along the osteonal axes, are caused by micro-
biological attack (fungi and/or bacteria) is compelling
(Hackett, 1981; Bell, 1990; Child & Pollard, 1991;
Child, 1994, 1995; Garland, 1987).
Protein content (% N)
This is based on a measure of nitrogen content in
powdered whole bone, and has been shown to reflect
accurately the quantity of collagen in the bone, pro-
vided the % N is greater than about 0·4% by weight
(i.e. more than 10% of collagen survives). A CHN
analyser (Europa ANCA Roboprep) was used to de-
termine the nitrogen content on 10 mg of whole bone
powder, with an error of approximately 10% of the
value. Earlier work, based on the quantity of collagen
extractable from bone, was subject to larger errors,
depending on the quality of the surviving collagen and
the extraction method.
Porosity measurements
Changes in porosity necessarily relate to changes in the
physical structure, which may be due to variations in
either the organic or inorganic components. Measure-
ments described here are based on the water content in
bone in equilibrium with a controlled water vapour
pressure; microporosity (i.e. the volume of bone retain-
ing water up to a water vapour pressure of 75% of
saturation, corresponding to a nominal pore radius
of d4 nm), and macroporosity, which includes pores
of radii >4 nm. A problem with this method is its
inability to measure pore size distributions for pores
larger than about 20 nm radius, and only the total
porosity can be given. It is now clear that ‘‘mesoporos-
ity’’, which is measurable from mercury intrusion
porosimetry (Nielsen-Marsh & Hedges, 1999), is very
much altered during diagenesis, and provides a power-
ful new tool to examine porosity changes. For this
study, however, only results from water sorption
analyses are available.
Porosity measurements have been adapted from the
protocol described by Hedges et al. (1995) and were
carried out on sections of bone weighing approxi-
mately 200 to 300 mg. The samples were oven dried
overnight at around 100C. They were weighed to a
precision of 0·0001 g: each sample was weighed three
times. The samples were then placed in preweighed
containers and left to equilibrate in a thermostatted
humidity chamber (kept at 25C) for 5 days. The
environment inside the chamber was controlled using a
given molarity of sulphuric acid. To obtain a relative
humidity of 75%, 3·5  sulphuric acid was used and the
atmosphere was measured using a hygrometer. After
5 days the samples were again weighed to obtain their
new weight at 75% RH. The samples were then satu-
rated by evacuation in distilled water to give a relative
humidity of 100% RH and reweighed. The three
measurements recorded at these atmospheres (0, 75
and 100% RH) were used to obtain values for the
total porosity of bone and the distribution of macro-
porosity and microporosity. The following equations
were used to calculate the total, macroporosity and
microporosity values:
Total porosity, macroporosity and microporosity are
defined as the volume of water taken up per gram of
bone sample (cm3 g
1). Using this definition, macro-
porosity and microporosity are independent of each
other: the mass of a solid remains unchanged if
macropores are cut into it—this is not true for volume
porosity (cm3 cm
3), where macro- and micro-
porosities are inter-dependent.
Crystallinity (splitting factor, SF) measurements
Hedges et al. (1995) measured crystallinity increases
using XRD. In this study we used the index based upon
the splitting of the IR phosphate doublet at 567 and
603 cm
1, the splitting factor (SF) which corresponds
to a generalized ‘‘degree of crystallinity’’ of bioapatite
(Termine & Posner, 1966; Weiner & Bar Yosef, 1990;
Sillen & Parkington, 1996). The SF was determined
1142 C. M. Nielsen-Marsh and R. E. M. Hedges

Table 2. Pearsons correlation coefficients r for diagenetic parameters for all sites

Parameters HI %N Microporosity Macroporosity SF

%N 0·77 — — — —
Microporosity 0·49 0·78 — — —
Macroporosity
0·47
0·55
0·39 — —
SF
0·60
0·67
0·28 0·60 —
C/P 0·36 0·38 0·01
0·14
0·46

Table 3. Pearsons correlation coefficients r for diagenetic parameters for Yarnton

Parameters HI %N Microporosity Macroporosity SF

%N 0·81 — — — —
Microporosity 0·78 0·87 — — —
Macroporosity
0·36
0·58
0·34 — —
SF
0·67
0·84
0·82 0·62 —
C/P
0·63 0·71 0·87
0·12
0·77

using the simple measurement and algorithm described
by Weiner & Bar-Yosef (1990). FTIR has advantages
above XRD for this study: only a very small amount of
sample is required (1 mg), preparation that ensures
accurate results is easier, and the carbonate content
can be assessed. The method of preparation of KBr
pellets was as described in Nielsen-Marsh & Hedges
(1997). Single SF measurements are estimated to be
replicable to 0·1 and measurements reported here
were made in triplicate.
Holocene and all from Europe). They exhibit both
general and site-specific patterns. Average values for
diagenetic parameter measurements from each site
used in this study can be found in Appendix II.
General patterns
Table 2 displays the correlations found between the
diagenetic parameters across all the sites studied
here. In many cases, intra-correlations are closer than
inter-correlations, as shown in Table 3 (Yarnton).

Carbonate content (C/P) and % calcite
The carbonate/phosphate ratio is measurable to about
5% precision from the IR spectrum of archaeological
bone. Here the method employed is similar to that
reported by Wright & Schwarcz (1996). Altered bones
may show both elevated and depleted C/P ratios in
comparison with fresh bone. As for SF, the C/P values
are based on triplicate measurements. C/P is not a
direct measurement of CO2
3 content in the bone, but
correlates well with CO2

3 in bone as measured by CO2
liberated on acid dissolution (Wright & Schwarcz,
1996; Nielsen-Marsh, 1997).
FTIR also detects calcite (>3%) in the sample
(Nielsen-Marsh, 1997), represented by peaks at 1435,
875 and 713 cm
1. While peaks at 1435 and 875 cm
1
are contributed to by other CO2
3 forms (original

1
CO2
3 and diagenetic CO2

3 ) the peak at 713 cm is
due only to the presence of CaCO3 in the bone
spectrum and has been used to estimate the quantity of
calcite present.
Results and Discussion
The results are collated from 134 bones (both non-
human and human) from eight sites (seven in the
Histology. Taken overall, the pattern of histological
alteration confirms the bimodal distribution observed
in Hedges et al. (1995). The majority of the archaeo-
logical bones were either well preserved (HI:5/4) or
poorly preserved (HI:0/1), scores 2 and 3 account for
around 20% of the total number of bones.
The variation in histological preservation that can
exist within a site and between sites is large. Certainly
Mediaeval sites are as likely to contain bones of
predominantly poor histological structure as those that
are 5000 years old. The only younger site studied,
Overton Down, showed excellent histological preser-
vation after 32 years, except for one cooked bone (see
Appendix II, Table 5), implying that collagen integrity
is important for bone preservation. We found that the
histologically better preserved material generally came
from either fully waterlogged environments or rela-
tively dry sites where water levels were predominantly
static (e.g. palaeochannels or caves, Figure 1). Histo-
logically altered bones were most commonly found in
burial environments where fluctuating water levels
would have been recurrent (as in gravels near flood-
plains, see Yarnton, Neolithic samples, Appendix II,
Table 4).
The questions, whether bones buried with flesh, or at
depth, or after being cooked, are significantly different,
 Patterns of Diagenesis in Bone I: The Effects of Site Environments 1143

5 0.08
0.07

Microporosity (cm3 g–1)

4
0.06
3 0.05
HI

0.04
2
0.03
1 0.02
0.01
0
0.0 2.0 4.0 6.0 0.00
0.0 0.2 0.4 0.6 0.8 1.0
%N
Macroporosity (cm3 g–1)
Figure 1. Relationship between histological preservation (HI) and
protein content (% N) for bones buried in different hydrological Figure 2. Porosity alterations in archaeological bones (all sites). ,
environments. , Fluctuating; , static; , modern bone; , All bones; , modern bone; , deproteinated.
deproteinated.

5.0
cannot be clearly answered from our data. We did not
notice significant differences in the preservation of 4.0 Protein loss line
human bone in comparison to animal bone, but, given
the wide range of variation, the data for comparison
3.0
are insufficient. However, it is clear that histological
%N

damage to human fleshed bone is normally not less

2.0
than with animal bone (although the proportion of
cooked animal bone is not known, but must be
appreciable). 1.0

Protein content. Protein content strongly correlated 0.0

with the HI, although there were exceptions. The
correlation applies both for bones within a site and
across sites. The overall correlation coefficient was 0·77
(Table 2), which is greater than that reported in
Hedges et al. (1995), the change probably due to the
more precise measurement of protein in this study.
The exceptions comprise bones which possess good
histological preservation, but which are low in protein.
While there are no high protein bones possessing poor
preservation (see Figure 1), bones with apparently
very poor histology (HI=0) have an average of 20%
remaining protein, but can still contain up to 40%.
It is hard to avoid concluding that for most bones
(Figure 1) the loss of protein is a result of microbio-
logical action, which also causes histological alteration.
For bones with low protein and high histology scores,
however, it would appear that collagen is being lost
through a different process—such as chemically medi-
ated hydrolysis (Collins et al., 1993). Protein content
also correlates strongly with both microporosity
(r=0·78) and SF (r=
0·67), implying that, in general,
as collagenous material is lost from the bone, there is a
reorganization within the microstructure—which nec-
essarily affects the microporosity and therefore the
crystallinity of the bone. Protein loss is not, therefore,
independent of porosity, crystallinity or microbiologi-
cal attack as stated by Hedges et al. (1995).
Porosity. The earlier reported data (Hedges et al.,
1995) showed porosity changing during diagenetic
0.00 0.02 0.04 0.06 0.08
3 –1
Microporosity (cm g )
Figure 3. Relationship between microporosity and protein loss
measured in modern bone before and after deproteination. ,
Modern bone.
alteration in a correlated way (microporosity decreas-
ing, macroporosity increasing) from modern bone
values. Our present data refine this picture (Figure 2),
but show very clearly that the porosity changes are
(probably directly) related to protein loss. This is
demonstrated by measuring porosity changes for fresh
and increasingly deproteinated bone treated with
hydrazine hydrate (Figure 3). Bones that are histologi-
cally well preserved but have lost collagen fall on the
same trend (Figure 7), implying that histologically
observable alteration has no obvious independent ef-
fect on porosity. The pore size distribution is altered,
however, and characteristic structures, not seen in the
microporosity:macroporosity measurements reported
here, can be observed in the mesoporosity range using
mercury intrusion porosimetry (Nielsen-Marsh &
Hedges, 1999).
The increase in macroporosity is not surprising,
given that the protein content of a bone, amounting to
some 22% by weight, occupies 40% by volume. The
decrease in microporosity is less expected. One expla-
nation, that, as protein is removed, the inorganic
structure remaining is rearranged into a coarser
structure, receives support from the fact that the SF
1144 C. M. Nielsen-Marsh and R. E. M. Hedges
increases whenever protein is removed or lost. This is
consistent with the observed correlation coefficient
(r=0·60) between macroporosity and SF.
But microporosity–macroporosity relationships by
no means follow the expected trend determined by
protein content alone, and deviations from this trend
appear to be site (and therefore presumably environ-
ment) dependent. They are discussed in greater detail
in the following section, which describes site-specific
patterns. Some of the bones we have measured exhibit
much greater porosity increases than can be explained
by protein loss, and presumably this is due to dissolu-
tion of the mineral matrix, determined predominantly
by the hydrological environment surrounding the bone
(Pike et al., in press; Nielsen-Marsh et al., 2000).
Splitting factor. There is a general correlation between
increasing SF and overall diagenetic alteration (r=0·60
and 0·67 for macroporosity and protein content, re-
spectively), which implies that crystallinity increases
are a fundamental part of the diagenetic process.
Crystallinity increase during diagenesis of bone has
been noted by many observers (Hassan et al., 1977;
Weiner & Bar Yosef, 1990; Bartsiokos & Middleton,
1992; Wright & Schwarcz, 1996; Sillen & Morris,
1996). How this happens is not well understood. The
laboratory study by Nielsen-Marsh & Hedges, (1997)
concluded that the major contribution of SF increases
measured in archaeological bone was due to recrystal-
lization as opposed to the dissolution of the smallest
crystallites (although this must play some minor role).
Evidence in support was the increasing SF for both
fresh and deproteinated bone in various dissolution
reagents, long after measured weight loss had become
stable. Also, porosity measurements obtained from
archaeological bones in other studies indicate that a
redistribution of pore size occurs during diagenesis that
is not explainable by dissolution alone (Hedges et al,
1995; Nielsen-Marsh & Hedges, 1999).
C/P measurements. In archaeological bone the carbon-
ate peak observed in IR spectra often includes ad-
ditional carbonate. In a calcitic environment, this
CO2
3 is usually deposited as CaCO3 on the surface
and within cracks and pore spaces of the bone, but can
also be present via adsorption/exchange on to the
surface of the bioapatite (Krueger, 1991). It may also
be incorporated into the crystal lattice (presumably via
dissolution and recrystallization processes), replacing
either biogenic CO2
3 or PO3
4 (LeGeros, 1981; Lee-
Thorp et al., 1989; Koch et al., 1997). Bones containing
calcite often possess much higher C/P values than those
where CO2
3 is adsorbed or incorporated into the
mineral lattice, but such calcite is distinguished from
these other forms by the presence of the 713 cm
1
peak in the IR spectrum, where it is possible to detect
calcite at levels >3% (weight CO2
3 to bioapatite)
(Nielsen-Marsh, 1997). Low values of C/P indicate
diagenetic loss of carbonate, possibly during rearrange-
0.5
0.4
0.3
C/P
0.2
0.1
0.0
0.01 0.03 0.05 0.07
3
Microporosity (cm g ) –1
Figure 4. Relationship between microporosity and carbonate
content (C/P) for Yarnton samples (). , Modern bone;
, deproteinated.
ment or dissolution of the mineral phase. Values
similar to fresh bone may indicate both processes.
Archaeological bones that do not contain appreci-
able calcite, but with near fresh bone values for pro-
tein, porosity and SF, more often than not possess
higher C/P values than modern bone, indicating that
external carbonate from groundwater is being incor-
porated onto bioapatite crystallite surfaces over time.
For bones that have suffered greater alteration, which
includes loss of microporosity, the carbonate content
is lower than for fresh bone. It is probable in these
cases that the dissolution/recrystallization processes,
coincident with protein loss, reduces surface area and
eliminates carbonate.
In Tables 2 and 3, notable differences for Yarnton
can be seen in the correlation coefficients for CO2
3
with the other parameters. CO2
3 that is substituted
into the bioapatite lattice will directly affect lattice size,
strain and stability (LeGeros, 1981; Newesley, 1989),
whereas more superficial inclusion via deposition of
calcite or adsorption onto the bioapatite surface will
not. These latter inclusions will ‘‘mask’’ the relation-
ship between SF and porosity, which exists for struc-
tural CO2
3 content. This can be seen by the very low
correlation coefficients between C/P and other dia-
genetic parameters for most sites in Table 2. For
Yarnton, however, there are very high correlation
coefficients. Yarnton bones possess no appreciable
quantities of calcite and the most plausible reason for
such high correlations is that any diagenetic CO2
3
present has been incorporated into the lattice, or
biogenic CO2
3 has been lost with mineral dissolution.
A plot of C/P against microporosity illustrates this
point (Figure 4).
Since CO2
3 substitution in the lattice influences
crystallinity (LeGeros, 1981; Newesley, 1989), a rela-
tionship between SF and C/P would be expected on a
compositional basis alone. The C/P versus SF relation-
ship observed here (Figure 4), however, has a lesser
slope than that observed by Newesley (1989) in exper-
iments substituting CO2
3 for PO3
4 in the apatite
 Patterns of Diagenesis in Bone I: The Effects of Site Environments 1145

1.0 5

0.8 4
Newesley (1989)
0.6 3

HI
C/P

0.4 2

0.2 1

0.0 0
0.0 2.0 4.0 6.0
2.0 2.5 3.0 3.5 4.0 4.5
%N
SF
Figure 6. Relationship between histological preservation (HI) and
Figure 5. Comparison between the relationship of crystallinity (SF)
protein content (% N) for Bercy () and for all other bones. , All
and carbonate content (C/P) in samples from Yarnton ( ) and that
sites; , modern bone; , deproteinated.
observed by Newesley (1989) (– – –).

lattice. It could be argued that SF would be increased, 5.0

both by the diagenetic loss of CO2
3 and by other
diagenetic effects which would increase ‘‘crystallinity’’; 4.0
this may explain the greater slope (Figure 5).
3.0
%N

Patterns specific to particular sites and burial

environments
The general outline describing diagenetic alteration
given above can now be modified in the light of
observations from individual sites.
Histology. The relationship between histological
change seems to vary less between sites than other
forms of diagenetic alteration. In all cases, histological
attack appears to depend on the nature of the burial
, Bercy;
site in a way not fully clarified.
However, we note the following. There is consist-
ently poor histology for: Poundbury, humans interred
in chalk which have undergone wetting and drying
cycles; Piddington, human burials which have under-
gone wetting and drying cycles; Yarnton, pits dug into
a gravel flood plain, compared to bones excavated
from palaeochannels; Bercy, bone buried on the dry
bank (these should be compared to bones from Bercy
which have been immersed in the river or saturated
sediment and which have much higher HI values).
There is consistently good histology for: cave sites,
Gough’s cave, Abri Pataud (Hedges et al., 1995); Brean
Down, bones have been excavated from contexts of
sandy deposits well above the water table and therefore
dry. The sand deposits are also ‘‘sterile’’, in the sense
of being extremely low in organic matter content;
saturated sites, i.e. subsites at Bercy and Yarnton, are
both channel locations.
Protein content. Of the four main sites, Bercy is the
most depleted in protein for the given state of histo-
logical preservation (Figure 6). This may be because it
is older and/or because most of the bones have been
totally immersed in water. These histological indices
indicate that alternative mechanisms for protein loss
2.0
1.0
Protein loss line
0.0
0.0 0.2 0.4 0.6 1.0
Microporosity (cm3 g–1)
Figure 7. Evidence for characteristic ‘‘site’’ patterns: variation from
protein loss line observed in bones from different sites:
, Yarnton; , Brean Down; , Modern bone; , deproteinated.
additional to microbiological attack exist. This is also
true for bones with good histology but depleted protein
content, although such cases are rare for most sites,
including those considered here. These alternative
mechanisms have been considered in Nielsen-Marsh
et al. (2000).
Porosity. The most informative relationship here exists
between microporosity and protein content. While
most sites give data points that conform to the
microporosity/macroporosity relationship of labora-
tory deproteinated bone (Figure 3), some possess indi-
vidual trends (Figure 7). There is a general tendency
for the drier sites (e.g. Brean Down, and certain
contexts in Yarnton) to have lower microporosity, and
the wetter sites (e.g. Bercy) to have higher microporos-
ity relative to the protein loss line. This seems to be
true for bones of medium–good histological preser-
vation and so is probably not due to different modes of
microbiological alteration, but perhaps rather to
groundwater interaction.
High macroporosity (Figure 8) implies dissolution of
bone mineral, and is particularly striking for the set of
Neolithic bones excavated from gravel pits at Yarnton
1146 C. M. Nielsen-Marsh and R. E. M. Hedges
5.0
4.0 Protein loss line
3.0
%N
2.0
1.0
0.0
0.0 0.2 0.4 0.6 0.8
Macroporosity (cm3 g–1)
Figure 8. Relationship between protein content (% N) and macro-
porosity for all sites in comparison to protein loss line. , All sites;
, modern bone; , deproteinated.
(see Appendix II, Table 4). Their very high macro-
porosity may reflect cooking before burial, facilitating
extensive microbiological attack. In fact, most bones
have lower macroporosity than predicted, even for
high protein bone (Figure 8).
It should be noted that macroporosity and micro-
porosity seem relatively unaffected by calcite content.
For Poundbury, the 50% or so bones observed to have
excess calcite show no obvious difference in their
porosity compared with bones that have little or no
calcite deposition.
C/P and SF measurements. Our data on both these
parameters do not reveal strong site-dependent pat-
terns. The range in C/P and SF tends to parallel the
general range in other diagenetic parameters; however,
the large and rapid change in SF at Overton Down
(turf) is notable (see Appendix II, Table 5). Since no
signs of histological alteration and associated protein
loss are observed after 32 years of burial, it is sur-
prising that in this period a greater change to the
mineral structure has occurred than for Brean Down
or Poundbury. Possibly it reflects very active water
movement (recharge) resulting from the deposition of
bones close to the surface.
diagenetic parameters broadly appear to reflect that
process. Here, the hydrological environment appears
to be particularly important, as demonstrated by the
histological results (Figure 1) and by the difference in
preservation between bones excavated from different
environments at Bercy and Yarnton (Appendix II).
The relationship between protein content and micro-
morphological preservation is shown to be particularly
close. A second important relationship, in that a simi-
lar pattern is seen for bones from very varied environ-
ments, is that between the porosity of bone at the
1.0 macroporous level and at the microporous level. Third,
while we find that bone crystallinity and carbonate
content certainly alter in characteristic ways, they show
no clear single functional relationship to the observed
changes in bone microstructure.
Previously reported laboratory-based experiments
on the alteration of bone diagenetic parameters under
well-defined but chemically extreme conditions showed
that, during dissolution, bone crystallinity increases
(Nielsen-Marsh & Hedges, 1997). This is probably due
to simultaneous recrystallization, since simple loss of
the least crystalline components could not account for
the magnitude of the change. The manner of the
recrystallization seemed to be strongly dependent on
the composition of the aqueous solution, which has
implications for bone buried in soils, where the com-
position of ions will vary according to many factors,
including pH and redox potentials. The presence of
protein in the bone had a major effect on the rate of
dissolution and other structural alterations, which has
been supported by the archaeological data reported
here. The precise role of protein in the protection of
bioapatite integrity and further, more drastic, crystal-
linity increases cannot be inferred, however, from
crystallinity measurements alone. The results obtained
by Nielsen-Marsh & Hedges (2000) have important
implications for current decontamination protocols.
No single parameter suffices to describe the degree of
preservation of a bone, but poor preservation can be
recognized quantitatively as a constellation of altered
values, as follows:
 Histology Index (HI) altered from a fresh bone value
of 5 to below 3;
Conclusions 
The measurements reported here greatly extend pre-
vious datasets of diagenetic descriptions of archaeo-
logical bone, with regard to density of sampling and
precision and accuracy of measurement. Even so, this
dataset is still limited to bones buried in the temperate
oceanic climate of northwest Europe during the
Holocene (and mainly the last 5000 years).
An overall increase in macroporosity and decrease in
microporosity, such that the ratio of macroporous
volume/microporous volume changes from about 1
to more than 3 [we believe that alterations in bone
microstructure, especially resulting from biological
attack, are most detectable at the level of changes
in ‘‘mesoporosity’’, as measured by mercury intru-
sion porosimetry (Nielsen-Marsh & Hedges, 1999)];
We find overall patterns in the set of diagenetic  % N altered from a weight % of >4 to <2 (often to
parameters measured which are much clearer than <0·5 %);
those previously reported (Hedges et al., 1995). Our  Splitting Factor (SF) increased from 2·6 to beyond 3·0;
results suggest that there is a single uniform pro-  An alteration of the value of C/P from 0·36 to
cess that can describe the chemical and structural less than 0·3, or, in buried soils rich in calcium
deterioration of archaeological bone and that all the carbonate, to beyond 0·4 or 0·5.
 Patterns of Diagenesis in Bone I: The Effects of Site Environments 1147
Populations of bones buried in different sites can be
compared in the light of the characteristic patterns of
diagenetic parameters. Within the general trends along
the well or badly preserved axis, site-specific patterns
can be noted (see, for example, Figure 7). We have
attempted to investigate how well these are explained
by taking into account the site hydrology. In a general
way, a close relationship can be seen (bones are much
better preserved when the water content of their en-
vironment changes very little), but we have not been
able to show in detailed terms how a given bone
porosity results from the presumed hydrological his-
tory of burial. These issues have been considered in the
context of idealized models of hydrologically driven
chemical processes by Pike et al. (in press). These
authors found that although applying the results from
experimentally investigated models to archaeological
contexts was problematic, the importance of ground-
water in terms of its movements, pH, and ionic
composition could be effectively demonstrated at a
quantitative level. They found a significant difference
in the rate of dissolution and the effect on porosity
between environments where water movement is
limited (cave sites, saturated sites) and where it fluctu-
ates frequently (well-drained sites). In spite of their
results, however, it is still not clear how bone porosity
is related to hydrology, probably because, although
porosity determines the way in which a bone will
interact with its burial environment, many of the
changes are influenced by the level of protein content
and/or the microbiological alteration of the bone struc-
ture. There still remains, then, a gap between predic-
tions of diagenetic change derived from simple models
based on hydrological and geochemical processes, and
measurements of diagenetic change of bones from
actual sites.
Acknowledgements
We acknowledge with thanks all those who provided
samples for this research. CMN-M was funded by
NERC. The majority of this work was submitted as
part of a D.Phil. Thesis.
Appendix 1
Yarnton
Yarnton is located on a Thames floodplain in
Oxfordshire and was a site of occupation for approxi-
mately 6000 years (Hey & Bradley, 1994; Hey & Muir,
1997). Animal bones have been recovered from a
variety of environments in contexts on the floodplain
and also from higher ground. The bones sampled
for this project come from the whole extent of the
occupation (6000–800 ).
The site lies on an alluvial aquifer, consisting of sand
and gravel deposits, the base of which rests on imper-
meable Jurassic Oxford Clay. Much of the aquifer is
overlain by alluvial mud consisting of a 65:35 clay:silt
composition, suggested to be Iron Age or younger
which rests on Pleistocene gravels. The site can be
divided into low-lying areas (the floodplain) and higher
ground (Worton Rectory farm); these two different
areas rest on different types of gravel based soils. The
floodplain rests on the first terrace gravels and the
higher ground rests on second terrace gravels. The first
terrace is mainly Devensian in age, whereas the older
second terrace gravels were probably laid down during
a cold stage of the Upper Pleistocene.
Neolithic and Bronze Age bones have been recov-
ered from the floodplain, the Neolithic bones from pits
located on gravel islands and the Bronze Age bones
from palaeochannels. From Worton Rectory farm
bones from the Bronze Age through to the Anglo-
Saxon period have been recovered, which have all
come from the same burial environment. Both the
bones from Worton Rectory farm and the pits located
on the floodplain are expected to have undergone
fluctuating water conditions. The bones located in the
palaeochannels, however, have probably remained
waterlogged for the majority of their burial. The fol-
lowing list summarizes the samples taken for analysis.
Floodplain: palaeochannels, Bronze Age; gravel is-
lands, Neolithic. Worton Rectory Farm: Anglo Saxon,
Romano British, Iron Age and mid Bronze Age.
Brean Down
The Bronze Age coastal sand defence, Brean Down, is
a carboniferous, limestone sandcliff lying to the south
of Weston Bay and Weston-Super-Mare in Somerset,
England, and projects 1·6 km into the Bristol Channel
(Bell, 1990). Animal bones excavated from Units 8a,
6a, 5b and 4b (Beaker, Late, Middle and Early Bronze
Age, respectively: 3700–900 ) have been sampled for
this study. The units refer to an outline of the post-
glacial sequence defined in the early report from Brean
Down and have been used to define the bone samples
studied (ApSimon et al., 1961).
The layers consist primarily of sand and silt, al-
though Units 6a and 5b have a greater proportion of
sand than Units 4b and 8a. Because of the stratigraphy
at Brean Down, the movement of water within the
contexts is expected to be relatively homogeneous.
Hydrological influences may not be as important as
other environmental considerations, such as Ca2+ con-
tent, or level of bacterial activity within the burial
sediment.
Bercy
Bercy is a Neolithic settlement approximately 6000
years old, associated with the Chaussean culture. The
site is located on the dry bank of a palaeochannel,
situated along the right hand bank of the river Seine in
East Paris, France. More than 10,000 animal bones
were excavated at Bercy from a wide variety of species,
1148 C. M. Nielsen-Marsh and R. E. M. Hedges
both domestic and wild. For this study, however, only
auroch bone was sampled, since Bercy provides the
opportunity to examine diagenesis in bones of the same
species, buried for similar time periods, where the
environments in which they were buried are the only
differing conditions. The bones have been collected
from three different environmental zones: the bottom
of the river channel (bones totally immersed in the
sand/mud sediment), the channel edge (bones subjected
to periods of total immersion), and from the dry
riverbank (bones deposited in alluvial silts).
The bones taken from the dry bank are expected to
have been influenced by a fluctuating hydrological
regime, due to periodic wetting after precipitation and
occasional flooding of the Seine. Those bones found on
the edge of the channel will have been subject to
fluctuations in the river level throughout the year, but
will probably have been semi-waterlogged/saturated
for the majority of the time. Similarly, the bones from
the bottom of the palaeochannel will have lain in
waterlogged/saturated conditions. Analyses of soil
samples recovered from the bank have shown that the
soil is alkaline, at a pH level of 8·2.
Poundbury
The bones sampled for this project were selected from
the Iron Age crouch burials, which are documented in
Volume 2 of the Poundbury volumes (Farwell &
Molleson, 1993). The sample set also included a burial
from the Roman period. No form of coffin was used in
any of the burials.
At Poundbury the archaeological deposits represent
a mixture of chalk and plough soil caused by erosion
processes. In the Neolithic and the Bronze Age con-
texts, there is a higher proportion of original red
non-calcareous soils. The samples studied here were
excavated from the later soils, which possess increas-
ingly larger calcareous fractions. The bones used in this
study were excavated from the calcareous loam. The
graves were cut into the chalk soil at a depth typically
about 65 cm and this is reflected in the high percentage
of calcite present in the bones (10%) (see Appendix
2). Being buried near the surface, the bones have
probably experienced fluctuating water conditions as
the soil is recharged with each rainfall, which then
drains down through the chalk aquifer, or is lost via
evaporation at the surface.
Wharram Percy
In the late 14th century the village of Wharram Percy
consisted of 30 houses. By 1500 the village had joined
the ranks of approximately 3000 Mediaeval villages
which have failed to survive to the present day. Exca-
vations were carried out on the areas to the north and
west of the church where burials ceased after the
desertion of the village in the early 16th century.
Around 600 burials were excavated comprising about
1000 individuals. There were several superimposed
layers of graves, and later burials had disturbed many
earlier ones. The main part of the village lies on a
plateau where the subsoil is chalk, and chalk fragments
are very common in the soil profile. The soil pH
readings are accordingly alkaline, with a mean value of
8·1. The site is well drained and the bones will probably
have experienced similar conditions to those that have
been described at Poundbury.
Piddington
The Romano-British Villa excavated at Piddington,
Northamptonshire, is located 300 ft above sea level on
the side of a shallow valley. The site lies on boulder
clays, which in turn rest upon the limestone, which is
part of the Jurassic Succession. The whole of the area is
overlain by recently ploughed and rolled arable land.
Because of its position, on seasonally flooded alluvium,
the site will probably have experienced fluctuating
water levels. Samples collected for this study represent
both human burials and animal remains. All samples
have come from the same burial environment.
Gough’s Cave
Gough’s Cave, situated in Cheddar, Somerset, is a Late
Upper Palaeolithic human occupation site and field-
work has confirmed the existence of contemporaneous
Late Glacial Interstadial (12,000–13,000 ) mammal
fauna dominated by Equus ferus, from which all of the
samples for this study were taken (Currant et al.,
1989).
All of the material recovered from the most recent
excavations was excavated from an upper fine gravel or
underlying red silt, both with abundant sharp lime-
stone clasts. Because this is a cave site it is difficult to
determine what the water action will have been over
the period of burial; however, the bones are likely to
have experienced unpredictable and sporadic water
fluctuations. For the majority of their burial the
bones are likely to have been dry, but any occasional
flooding of the cave will have altered these conditions
temporarily.
Overton Down
Overton Down is an experimental earthwork, built in,
1960, situated between Marlborough and Avebury in
Wiltshire and was built in, 1960 (Bell et al., 1996). The
primary objective of this long-term experiment was to
bridge the gap in the understanding of the degradation
processes occurring to materials over relatively short
periods, and those which occur over much longer
timescales, which concern archaeologists. The material
buried at Overton (bone, textiles, coins etc.) is exca-
vated at designated time periods and examined for
diagenetic changes. A progressive scale is used for the
 Patterns of Diagenesis in Bone I: The Effects of Site Environments 1149

Table 4. Average values of diagenetic parameters for bones sampled (modern and modern deproteinated bones are included for comparison)

Porosity (cm3 g
1)

CaCO3
Site Context Total Macro Micro %N SF C/P (%) HI

Modern — 0·127 0·067 0·060 4·8 2·7 0·36 — 5

Deproteinated — 0·312 0·280 0·031 0·6 3·0 0·33 — 5
Yarnton Bronze Age (FP) 0·147 0·089 0·058 4·1 2·7 0·40 <3 5
Neolithic (FP) 0·687 0·655 0·033 0·1 3·4 0·30 <3 0
Anglo Saxon (WRF) 0·164 0·121 0·043 2·3 2·9 0·32 <3 3
Romano British (WRF) 0·146 0·112 0·0349 1·4 3·1 0·24 <3 0
Iron Age (WRF) 0·1346 0·106 0·028 0·9 3·3 0·24 <3 1
Mid Bronze Age (WRF) 0·315 0·286 0·029 1·3 3·3 0·22 <3 0
Brean Down Unit 4b 0·229 0·192 0·036 1·4 3·2 0·41 <3 1
Unit 5b 0·114 0·073 0·041 2·9 2·9 0·37 <3 4
Unit 6a 0·087 0·039 0·047 4·0 2·9 0·37 <3 5
Unit 8a 0·144 0·119 0·026 1·0 3·2 0·27 <3 0
Bercy Channel bed 0·156 0·102 0·055 2·8 3·0 0·38 <3 5
Channel edge 0·117 0·058 0·059 3·2 2·8 0·39 <3 5
Dry bank 0·318 0·284 0·034 0·7 3·3 0·31 <3 0
Poundbury Chalk graves 0·328 0·295 0·034 1·4 3·2 0·40 8–10 1
Gough’s Cave Cave 0·195 0·141 0·054 3·6 2·9 0·51 <3 5
Piddington Alluvium soil 0·322 0·281 0·041 1·3 3·2 0·31 <3 0
Wharram Percy Calcareous soils 0·268 0·226 0·042 2·4 3·2 0·34 <3 2

Table 5. Values of diagenetic parameters for Overton Down bones

Porosity (cm3 g
1)

CaCO3
Samples Context Total Macro Micro %N C/P SF (%) HI

OD. 3171 Turf 0·342 0·278 0·065 4·4 0·28 3·3 <3 5
OD. 3160 (cooked) Turf 0·402 0·347 0·052 4·3 0·22 3·4 <3 1
OD. 3109 Chalk 0·247 0·180 0·067 4·8 0·35 2·8 <3 4
OD. 3090 (cooked) Chalk 0·181 0·118 0·063 4·4 0·38 2·9 <3 5

excavation dates, so after 1, 2, 4, 8, 16, 32, 64, and 128
years, material has been and will be recovered.
The early stages of burial may be disproportionately
more important for diagenetic processes, but this is
not yet known. Four samples from the Overton Down
earthwork have been sampled for this study. The
nature of the experiments means that material
available for investigations is relatively limited and
samples need to be used to their maximum potential.
The same diagenetic measurements have been taken as
for archaeological samples, but because of the small
sample set the conclusions drawn from the results must
be relatively conservative. Nevertheless, the chance to
work on this material has added a new dimension to
this study and allowed comparisons to be drawn
between the archaeological material and the very early
stages of diagenesis.
Overton Down is unique, in that the precise con-
ditions at burial have been recorded and subsequently
monitored with each successive excavation: this is
impossible to do over archaeological timescales. A
detailed description of the burial environment and
analyses carried out on all objects recovered during the
excavation in, 1992 is described in Bell et al. (1996), but
the bones will have experienced fluctuating water
regimes with rainfall and subsequent drainage and/or
evaporation.
Because of the nature of this experiment, it was only
possible to obtain two samples from each environment.
Bones were sampled from chalk and turf layers, one
sample from each having been cooked prior to burial.
Appendix 2
Average values obtained from the analyses performed
on the samples are found in Table 4. Only two samples
from each environment at Overton Down were ob-
tained for analysis, which included bones cooked prior
to burial; therefore the results for these bones have
been placed in a separate table (Table 5).
1150 C. M. Nielsen-Marsh and R. E. M. Hedges
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