Journal of Archaeological Science (1996) 23, 175–181

Fish Heads, Fish Heads: An Experiment on Differential Bone

Preservation in a Salmonid Fish
Patrick M. Lubinski
Department of Anthropology, University of Wisconsin, 5240 Social Sciences, 1180 Observatory Drive,
Madison, WI 53706, USA

(Received 24 March 1994, revised manuscript accepted 7 June 1994)

An experiment modelling the effects of cooking and soil pH on salmonid bone survival was completed using cleaned
Lake Whitefish bones. The experiment tested for preservation differences between head parts and vertebrae and
between raw, boiled, and moderately burned bone under both acidic and alkaline conditions. Elements were submerged
in aqueous solution for 24-h periods, removed, dried, weighed, and re-immersed. Resulting weight and element loss
curves for acidic conditions suggest that (1) head parts are destroyed more quickly than vertebrae, and (2) element
destruction increases with heating intensity. Under alkaline conditions, broadly similar trends were observed, but at
reduced rates. ? 1996 Academic Press Limited

Keywords: SALMONID ARCHAEOFAUNAS, TAPHONOMY, BONE DIAGENESIS, SKELETAL PART

REPRESENTATION, COOKING, PH.

Introduction fish skeletal elements as a result of different responses

of elements to heating and chemical attrition.
s with mammal assemblages, measures of the There is considerable experimental evidence for

A relative frequencies between anatomical parts,

such as %MAU (Binford, 1984) may provide
useful information in the interpretation of archaeologi-
cal fish assemblages (e.g. Stewart, 1989; Belcher, 1992;
Butler, 1993). However, there are a large number of
possible explanations for any observed pattern, be-
cause there are a number of processes that might affect
skeletal parts differentially. Potential sources of differ-
ential representation in fish body parts include meth-
odological bias (in recovery, sampling, identification
and aggregation), differential disposal (during butch-
ery, transport, preparation and discard), and differen-
tial preservation (before deposition as a result of fish
bone structure, processing, ingestion and subaerial
weathering; and after deposition as a result of biologi-
cal, mechanical and chemical attrition). The most
important sources of differential skeletal part represen-
tation will vary among archaeological contexts; how-
ever, the preservation of all fish remains will be
strongly affected by the thermal history of the remains
and the chemistry of their disposal context. While there
has been experimental examination of several other
mechanisms of fish bone destruction (e.g. Jones, 1986;
Nicholson, 1992; Butler & Chatters, 1994), thermal
history and disposal context chemistry have not
been the subject of much detailed examination (cf
Richter, 1986). The experiments described in this paper
were designed to determine if there are any natural
tendencies towards differential preservation between
175
0305-4403/96/020175+07 $12.00/0
differences in preservation potential between skeletal
elements (e.g. Binford & Bertram, 1977; Lyman, 1984;
Von Endt & Ortner, 1984; Butler and Chatters, 1994).
Skeletal parts may be destroyed differentially because
they vary considerably in size and structure (shape,
density, tissue structure, porosity, etc.). Size and struc-
ture determine a bone’s inherent mechanical resistance
to physical destruction and the surface area it presents
to chemical action. For bony fishes, one might expect
that thin, flat cranial bones would be destroyed more
quickly than cylindrical vertebrae (Rojo, 1987: 210;
Wigen & Stucki, 1988: 106) because the cranial bones
appear to have less mechanical strength and more
surface area. Additionally, cranial elements are lower
in volume density (measured in g cm "3) than vertebrae
(Butler & Chatters, 1994).
Heating of bone is known to weaken the organic
phase (DeNiro et al., 1985; Richter, 1986) which gives
bone tissue some of its resilience and chemical stability
(Currey, 1990). Heat-induced loss of the organic
phase will increase the porosity of the mineral phase
(McCutcheon, 1992: 354), which will provide more
surface area to chemical action, as well as making the
bone more susceptible to mechanical breakage. For
these reasons, one might expect that moderately
burned elements would be destroyed more quickly than
unburned elements under archaeological conditions
(Stewart, 1989: 172). However, bones subjected to
more intense, high temperature burning may be more
? 1996 Academic Press Limited
176 P. M. Lubinski

Table 1. Summary of experiments

Initial Fish Fork

Beaker Solution Processing Bones used weight* (g) specimen no. length† (cm)

Experiment 1:
A Acid Boiled 27 Head, 51 trunk 3·881, 6·956 3 51
B Acid Boiled 27 Head, 51 trunk 2·818, 4·730 4 46
C Acid Burned 27 Head, 51 trunk 1·436, 2·241 5 38
D Acid Burned 27 Head, 51 trunk 2·909, 3·178 7 47
E Base Boiled 27 Head, 51 trunk 3·580, 6·320 2 50
F Base Boiled 27 Head, 51 trunk 1·986, 3·297 8 44
G Base Burned 27 Head, 51 trunk 2·199, 3·424 9 47
H Base Burned 27 Head, 51 trunk 1·657, 2·545 11 39
Experiment 2:
A Acid Raw 10 Vertebrae 0·703 6 41
B Acid Boiled 10 Vertebrae 0·877 6 41
C Acid Burned 10 Vertebrae 0·656 6 41
D Base Raw 10 Vertebrae 1·096 10 45
E Base Boiled 10 Vertebrae 0·706 10 45
F Base Burned 10 Vertebrae 0·703 10 45

*After processing.
†Fork length is a measure of fish size from the tip of the snout to the fork of the tail.

stable, because the overall shrinkage and crystal re-
organization which take place at high temperatures
(Shipman et al., 1984) lower a bone’s surface area. The
stability of intensely burned bone has been demon-
strated in several recent experimental studies (Knight,
1985; Linse & Burton, 1990: 7).
The chemical environment of deposition is widely
known to affect skeletal element preservation, princi-
pally through the pH of the soil solution (e.g. Gordon
& Buikstra, 1981; White & Hannus, 1983). Since
conventional wisdom holds that bone is destroyed in
acidic environments, the following experiments were
designed to test for differential bone destruction under
acidic conditions. Additionally, the study models
alkaline conditions because recent experiments have
shown that fish bone is destroyed in alkaline burial
environments as well (Linse & Burton, 1990; Linse,
1992).
Table 2). For Experiment 2, vertebral columns from
two fish were removed and split into three sections
each to compare preservation under raw versus boiled
versus burned heating states. For each segment, 10
vertebrae (five with spines and five without spines)
were selected.
All bones used in the experiments were cleaned with
a toothbrush and dental pick to remove any adhering
connective tissue. Skeletal elements were prepared in
three different heating states; raw, boiled, and burned.
Raw specimens (vertebrae only) were soaked in cool
water, then separated and cleaned. Boiled specimens
were boiled in tap water for 1 h and then cleaned.
Burned specimens were boiled for 1 h (to facilitate
cleaning), cleaned, dried, then heated in a muffle fur-
nace at 250)C for 2 h. This oven temperature and
duration were chosen to remove a portion of the
Table 2. Elements selected for Experiment 1

Methods Part Region Element and number

The study utilized Lake Whitefish (Coregonus clupea-
formis), an important prehistoric and modern commer- Head* Orbital Frontal (2)
Otic Posttemporal (2)
cial fish species in the Great Lakes (Heidenreich, 1978: Basicranial Parasphenoid (2)
382, Smith, 1979; Cleland, 1982). This species is a Oromandibular Maxilla (2), dentary (2), articular
member of the family Salmonidae, which also includes (2), quadrate (2)
Hyoid Hyomandibular (2), urohyal (1),
chars, ciscos, salmons, and trouts. Ten filleted whitefish ceratohyal (2), opercular (2),
carcasses, ranging from 38 to 51 cm in fork length (tip preopercular (2), subopercular (2),
of snout to fork of tail), were obtained from a local fish interopercular (2)
market. These were used for two experiments, which Pectoral Cleithrum (1)
Total head=27
are summarized in Table 1. Experiment 1 utilized head Trunk† Vertebral Thoracic vertebra (31), precaudal
and trunk elements from eight fish carcasses to com- vertebra (6)
pare preservation of head versus trunk elements and Caudal Caudal vertebra (14)
Total trunk=51
boiled versus burned elements. From each specimen,
27 of the most easily recognized and robust head *Head terms (except pectoral) after Wheeler and Jones (1989:
elements and 51 of the vertebrae were selected to Table 7.1).
represent the head and trunk regions respectively (see †Trunk terms after Cannon (1987: Table 2).

–2
–3
–4
log H2(PO4)– or H(PO4)2–
–5
–6
–7
–8
–9
3 4 5 6 7 8
pH
Figure 1. Solubility of hydroxyapatite (after Lindsay 1979: figure
12.10).
organic phase (DeNiro et al., 1985: 5) without causing
significant splitting, shrinkage or crystal reorganization
(Shipman et al., 1984). Additionally, this temperature
was chosen to mimic the core temperature of a small
open campfire, such as the experimental fire monitored
by Shokler (1991).
Two solvents were used in the experiments, one an
acidic solution with a pH of 4 and the other an alkaline
solution with a pH of 10. Aqueous test solutions
were prepared using mixtures and pH values given by
Robinson (1976: 133–134). The solutions used were
0·05  potassium acid pthalate with a pH of 4·008, and
a buffered solution of borax (sodium tetraborate) and
sodium hydroxide with a pH of 10·0. These strong pH
values were chosen not to mimic archaeological con-
ditions, but rather to produce rapid results in the
laboratory. The operating assumption is that increased
acidity and alkalinity over a short, experimental time
span can model gross patterns of preservation bias that
would be obtained over archaeological time spans
under more moderate conditions. These pH values also
were chosen to bracket neutral pH (7) and the pH at
which bone mineral is the most stable. Hydroxyapatite,
the primary mineral of bone, is the most stable (least
soluble) under slightly alkaline soil conditions at about
pH 7·9 (Figure 1). By choosing solution pH values to
either side of the values at which bone is probably the
most stable, it was expected that the experimental
solutions might serve as general models for acidic and
alkaline conditions.
Differential Bone Preservation 177
Bone units were placed along with test solution in
beakers sealed with Parafilm (to prevent evaporation)
and immersed in solution for twelve 24-h periods. For
Experiment 1, head and trunk units from each fish
were placed into the same beaker with 150 ml of
solution. In Experiment 2, each group of 10 vertebrae
was placed into a separate beaker with 50 ml of solu-
tion. Test solutions were checked with indicator paper
daily to detect any pH fluctuations. Beakers were
rinsed and filled with fresh solution on days 4 and 9
and on any day when the pH changed by a factor of
1·0. In this way, the experiment models an open system
with a continuous ion supply rather than a closed
system in which the bone might come into equilibrium
with the beaker solution.
After each 24-h immersion period, the bones were
removed, dried, counted, weighed to the nearest mg,
and re-immersed. Experiment 1 bones were oven dried
at 80)C for 1 h while Experiment 2 bones were air dried
in order not to cook the raw bone. Total drying time
from bone removal to re-immersion (including oven
drying, counting, and weighing) averaged 3·8 h. The
raw bone was inadvertently placed in the drying oven
9 for 10 min on day 1 and should therefore be considered
‘‘minimally roasted’’ rather than raw.
Bone decay was measured in terms of both element
loss and weight loss per day of immersion. Element
loss is directly relevant to zooarchaeological research
because bone counts are the basic analytic units
used by most faunal analysts (Grayson, 1984). While
weight loss may be directly relevant to researchers who
employ weights as a basic analytic unit (e.g. Wing &
Brown, 1979), it is used here primarily as a proxy
measure of preservation potential.
Weights were obtained using all fragments identifi-
able to element each day. Unidentifiable fragments
were not included. Due to this practice, bone loss in
the experiment can result from chemical weathering
or from fracture brought about by wet/dry cycles
and handling, which I believe approximate com-
mon conditions of preservation and archaeological
recovery.
Cumulative weight and element loss curves were
used to discern any patterns in decay magnitude on a
given day or in decay rates over a number of days. All
data points are included on the weight loss versus time
graphs, except those representing absolute bone gain
(negative % bone loss) due to bone moisture. These
points were rejected because (1) they are theoretically
impossible and (2) the random error is strongly non-
homogeneous, so that the ‘‘true’’ relationship must be
a regression line or curve very near or above the
collected data points. There is much more random
error to overweighing than to underweighing, since
bone wetness may increase weight dramatically (in one
case, a 0·225 g gain, which represents 29·1% of total
weight), while instrumental error for the balance is
only &0·0005 g (0·06% of total weight). Clearly, the
rejection of negative bone loss values does not remove
178 P. M. Lubinski

100 60

50
80
Cumulative weight loss (%)

Cumulative weight loss (%)

40
60
30
40
20

20
10

0 0
0 2 4 6 8 10 12 0 2 4 6 8 10 12
Time (days) Time (days)
Figure 2. Experiment 1 acidic solution weight loss. (/) boiled; Figure 3. Experiment 1 basic solution weight loss. (/) boiled;
(-) burned; (——) head; (– –) trunk. (-) burned; (——) head; (– –) trunk.

the effect of bone moisture. There is no way to discern
the effect of moisture on any particular weight value.
However, the ‘‘dip’’ patterns seen on several graphs at
day 4 indicate excess bone moisture on those days
relative to the surrounding days. This reflects in-
adequate drying, so drying time was doubled and oven
temperature was increased to 90)C beginning on day 5.
Results
The results (in terms of weight loss) of Experiment 1
bones under acidic conditions are shown in Figure 2.
The graph shows that burned head units clearly decay
the fastest and boiled trunk units decay the most
slowly. Burned trunk elements and boiled head ele-
ments lose weight at about the same rate. Burned heads
have the largest weight loss on any given day and have
the fastest decay rate based on the curve slope. To
quantify the rates of destruction, linear regressions
were calculated on the values for days 5–12 weight loss
for each fish specimen (Table 3). Based on the regres-
sion slopes, heads are destroyed 1·5–2·5 times as fast as
vertebrae among specimens with the same heating
state. Also, comparing like skeletal parts, burned
units are destroyed 1·4–2·5 times faster than their
corresponding boiled units.
Under alkaline conditions, weight loss curves show a
similar pattern of burned head part destruction as
under acidic conditions (Figure 3). Burned head units
(the two upper curves) are destroyed the most quickly,
but all boiled parts and burned vertebrae appear to
decay at about the same reduced rate. Days 5–12 linear
regressions (Table 3) show burned head elements
are destroyed 1·9–4·2 times faster than burned trunk
elements or boiled elements.
The patterns obtained from element loss, under both
acidic and alkaline conditions, are broadly similar to

Table 3. Linear regressions for Experiment 1

Specimen no. Part (Treatment) r2 Slope Specimen no. Part (Treatment) r2 Slope

Cumulative weight loss (days 5–12, fixed zero intercept):*

Acidic solution: Basic solution:
3 Trunk (Boiled) 0·97 1·40 2 Trunk (Boiled) 0·69 1·18
4 Trunk (Boiled) 0·79 2·09 8 Trunk (Boiled) 0·46 1·10
3 Head (Boiled) 0·89 3·13 2 Head (Boiled) 0·68 1·09
4 Head (Boiled) 0·94 3·45 8 Head (Boiled) 0·51 1·39
5 Trunk (Burned) 0·95 3·54 9 Trunk (Burned) 0·94 1·02
7 Trunk (Burned) 0·95 2·92 11 Trunk (Burned) 0·95 1·19
5 Head (Burned) 0·05 7·91 9 Head (Burned) 0·79 2·68
7 Head (Burned) 0·93 6·37 11 Head (Burned) 0·93 4·31
Cumulative element loss (days 0–12, fixed zero intercept):
Acidic solution:
5 Trunk (Burned) 0·71 0·53
5 Head (Burned) 0·93 3·92

*Days 1 to 4 omitted from weight loss regressions because of inadequate drying in that period.
All correlations are significant at the 0·05 level except no. 5 head weight loss (its slope was not used for rate comparisons). The non-linearity
of this curve is presumed due to moisture-related weighing error.
 Differential Bone Preservation 179

50 increasing intensity of heating from minimally roasted

to boiled to burned conditions. Under alkaline con-
ditions, the burned vertebrae were destroyed the
Cumulative element loss (%)

40 most quickly, but the minimally roasted and boiled

vertebrae decay at about the same rate. In terms of
element loss, the burned vertebrae in acidic solution
30 (the only bone unit to show any element loss) lost three
of the 10 vertebrae by day 12.
20

Discussion and Conclusions

10
Figure 6, depicting the total weight loss at the conclu-
sion of Experiment 1 under each test condition, can
serve as a summary of the results of the experiments.
0 2 4 6 8 10 12 The general patterns in this graph are representative of
Time (days) all experimental results.
Figure 4. Experiment 1 acidic solution element loss for fish no. 5 Before discussing the overall patterns of bone
(burned). (——) head; (– –) trunk. destruction and implications of the study, several
limitations should be noted. First, the experiments
obviously are not directly analogous to archaeological
80 conditions, because the pH and wet/dry cycles in the
study are more extreme than at most archaeological
sites. For this reason, the specific values obtained in the
Cumulative weight loss (%)

60
40
20
0 is important to note that fish bones vary markedly
0 2 4 6 8 10
Time (days)
Figure 5. Experiment 2 acidic and basic solution vertebrae weight
loss. (*) raw; (/) boiled; (-) burned; (——) acidic solution; (– –)
basic solution.
those obtained from weight loss. By the conclusion of
the experiment, burned units had lost up to 44% of
their elements, but boiled units had not lost any
elements. Additionally, all burned fish lost at least 18%
of their head elements, but only one of the four lost any
trunk elements. Figure 4 provides an example of ele-
ment loss for the only specimen that lost vertebrae (one
of the burned fish under acidic conditions). Days 0–12
linear regression slopes (Table 3) suggest the head loses
elements at 7·4 times the rate of the vertebral column.
Weight loss curves for Experiment 2 vertebrae are
shown in Figure 5. As one might expect, the elements
in acidic solution collectively show greater weight
loss than those in alkaline solution. In each solution
group, burned bones show the greatest loss. Under
acidic conditions, bone destruction is more rapid with
experiment and given in Figure 6 are not directly
applicable to archaeological conditions and should not
be used as ‘‘correction factors’’ for archaeological
assemblages. However, if one assumes that fish bone
reacts in a similar manner from moderate to intense
acidity, alkalinity and wet/dry cycles, then the direc-
tions of preservation bias observed in this experiment
should be appropriate analogues for Lake Whitefish
bone decay under archaeological conditions.
Second, while the results of these experiments might
be applicable to species other than Lake Whitefish, it
12
between taxa (Gregory, 1933; Cannon, 1987; Wheeler
& Jones, 1989; Colley, 1990). Significant variations in
size, shape, and composition will result in different
head versus trunk element destruction patterns and
may result in different burned versus unburned
patterns. While one might expect that all salmonid
fishes would share the general whitefish pattern,
salmon (Oncorhynchus spp.) may, for example, exhibit
a less-marked difference in destruction between skeletal
parts because they have more robust mouth parts than
the Lake Whitefish used here. For non-salmonid fishes,
the head versus trunk element destruction pattern may
be very different than Lake Whitefish, because white-
fish and other salmonids have highly cartilaginous
skulls compared to other fishes (Gregory, 1933).
Experimental results differ between alkaline and
acidic conditions. In acidic solution, the effect of
heating state on a given skeletal part is increasing
destruction from minimally roasted to boiled to mod-
erately burned conditions. These results imply that
there will be a bias against the recovery of moderately-
heated fish bones in archaeological sites. (Note that,
based on other experiments (Knight, 1985; Linse &
180 P. M. Lubinski
Head-acid
Trunk-acid
Head-base
Trunk-base
0 20
Weight loss (%)
Figure 6. Experiment 1 total weight loss at conclusion of experiment (day 12). ( ) burned; ( ) boiled.
Burton, 1990), intensely burned bone may not follow
this pattern.) This may be significant if burning is the
common fate of fish bone at an archaeological site. It
may also be significant for faunal analysts who use
burning to distinguish cultural versus natural bone
accumulations or to infer cooking technique. Under
alkaline conditions, the effect of heating state is far less
clear. There is a strong bias against burned head
elements, but vertebrae (in any heating state) and
unburned heads appear to have the same preservation
potential in alkaline archaeological sites. In light of
these results, it would seem wise for faunal analysts to
consider burned and unburned fish elements separ-
ately because of their different natural potential for
destruction at both acidic and alkaline archaeological
sites.
As for differential preservation between skeletal
parts, Experiment 1 indicates that, for a given heating
state under acidic conditions, head elements are de-
stroyed more quickly than vertebrae. These results
imply that there will be a bias against the recovery of
head elements relative to vertebrae in acidic archaeo-
logical sites, which has important implications for
zooarchaeological studies. The bias should be taken
into account when, for example, employing fish
skeletal part representation measures (such as %MAU)
to infer butchery patterns or bone transport. Under
alkaline conditions, the pattern is again far less simple.
Head element are destroyed more quickly than
vertebrae amongst burned specimens, but there is no
strong preservation bias amongst boiled specimens.
The fact that these experiments indicate different
patterns of destruction under acidic and alkaline con-
ditions has an additional implication for archaeofaunal
comparisons. When comparing assemblages from very
different sediment pH values, it would be prudent
to limit comparisons to like body parts and heating
states (for example unburned vertebrae to unburned
40 60 80
vertebrae) in order to avoid attributing simple differ-
ences in preservation to culturally-meaningful patterns.
These experiments involved only one species under a
few pH and heating states. Further experiments might
broaden the focus and consider other fish taxa and
conditions. I, nonetheless, hope that these experiments
have provided useful information for a more rigorous
interpretation of archaeological fish remains.
Acknowledgements
Earlier versions of this paper were presented at the
58th Annual Meeting of the Society for American
Archaeology and written for a seminar given by J. M.
Kenoyer at the University of Wisconsin-Madison. The
author is indebted to Goeden’s Fish Restaurant for
providing the fish used in the experiment, and to J. H.
Burton of the Laboratory of Archaeological Chemistry
for allowing access to facilities. J. H. Burton, V. L.
Butler, C. P. Lipo, C. J. O’Brien, M. J. Schoeninger,
and an anonymous reviewer provided helpful com-
ments. I am particularly appreciative of the many
suggestions and thoughtful comments given by W. R.
Belcher and M. E. Madsen.
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