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Differential skeletal preservation at Windover Pond Causes and consequences.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 119:15–26 (2002)
Differential Skeletal Preservation at Windover Pond:
Causes and Consequences
Christopher M. Stojanowski,1,2* Ryan M. Seidemann,3 and Glen H. Doran2
1
Department of Anthropology, University of New Mexico, Albuquerque, New Mexico 87131
Department of Anthropology, Florida State University, Tallahassee, Florida 32306
3
Paul M. Hebert Law Center, Louisiana State University, Baton Rouge, Louisiana 70803
2
KEY WORDS
taphonomy; bone preservation; skeletal remains; prehistoric Florida
ABSTRACT
In this paper, we evaluate the causes of
differential skeletal preservation in the Windover Pond
skeletal series (8BR246). We collected data on sex and age
for approximately 110 individuals, and calculated a preservation score for each individual based on the presence of
80 skeletal landmarks. Our research questions evaluated
the relationship between bone preservation and individual age and sex, and between the presence of preserved
brain material and skeletal preservation, and the effects
of burial location on bone preservation. The results indicate variability in average preservation for the sample
(␮ ⫽ 0.53, SD ⫽ 0.22) with an apparent lack of sex-specific
(P ⫽ 0.79) or age-specific (P ⫽ 0.37) differences in preservation. The relationship between brain and skeletal preservation (P ⫽ 0.15) was not significant. The horizontal
distribution of burials was not significantly correlated
with skeletal preservation (north: r ⫽ ⫺0.10, P ⫽ 0.93;
east: r ⫽ 0.09, P ⫽ 0.45); however, vertical depth was a
significant predictor of preservation (r ⫽ ⫺0.31, P ⫽
0.005), indicating that skeletal preservation decreased as
burials were located closer to the ground surface. The
observed variability in preservation scores may be related
to the partial drying and resubmergence of the uppermost
burials for the last few millennia. Comparison of Windover element-specific survival rates with previous analyses based on terrestrial samples (Galloway et al. [1997]
Forensic taphonomy, Boca Raton: CRC Press; Waldron
[1987] Death, decay and reconstruction, Manchester:
Manchester University Press; Willey et al. [1997] Am J
Phys Anthropol 104:513–528) affirms the relationship between element weight or density and bone survival. The
unique taphonomic context of our study sample effected
little change in bone deterioration processes. Am J Phys
Anthropol 119:15–26, 2002. © 2002 Wiley-Liss, Inc.
The deleterious effect of marginal preservation on
demographic parameters is often stated as a concern
in demographic studies (Angel, 1969; Jackes, 1992;
Johansson and Horowitz, 1986; Konigsberg and
Frankenberg, 1992, 1994; Milner et al., 1989; Moore
et al., 1975; Paine, 1989; Paine and Harpending,
1996; Roth, 1992; Weiss, 1973). Sample representativity is affected by many factors, of which only a few
relate directly to bone preservation (Boddington,
1987). Boddington (1987) considered postdepositional disturbance and deterioration to be primary
taphonomic concerns; however, additional factors
include incomplete archaeological sampling, spatial
variability, and age-, sex-, or class-specific differential burial treatment.
The mechanics of differential bone preservation
(both between individuals and among elements of a
single individual) are fairly straightforward. Von
Endt and Ortner (1984) stated that the surface area
of an element is directly related to the speed at
which water-leaching results in the destruction of
the collagen and mineral matrix. Therefore, since
spongy or cancellous bone has a greater surface
area, one would expect these areas of the skeleton to
deteriorate first. The porous and less dense vertebra, ribs, sternum, and hand and foot bones are
generally those exhibiting marginal preservation
(Boddington et al., 1987; Nawrocki, 1995; but see
Waldron, 1987 for lumbar vertebrae). Dense areas of
bone such as long bone shafts tend to preserve better
(Galloway et al., 1997; Waldron, 1987; Willey et al.,
1997).
Density and porosity can also be used to explain
the differential preservation of elements between
individuals. Since bone mineral content increases
through childhood and adolescence, peaks in midlife, and declines with senescence (Riggs and Melton, 1986; Specker et al., 1987; Stini et al., 1992),
age-specific preservation biases may be expected.
Pathological conditions also may affect bone preservation in a similar manner where osteoclastic processes (osteoporosis) promote bone deterioration,
and osteoblastic processes (osteophyte formation)
©
2002 WILEY-LISS, INC.
*Correspondence to: Christopher M. Stojanowski, Department of
Anthropology, Florida State University, Tallahassee FL 32306.
E-mail: cstoj1@cs.com
Received 5 July 2001; accepted 10 January 2002.
DOI 10.1002/ajpa.10101
Published online in Wiley InterScience (www.interscience.wiley.
com).
16
C.M. STOJANOWSKI ET AL.
promote bone preservation (Nawrocki, 1995). Differences in body size due to sex may exhibit similar
effects on bone condition (Walker et al., 1988).
Therefore, in marginally preserved assemblages,
one would expect an underrepresentation of subadults, elderly adults, and females due to the diminished resiliency of less dense bone to taphonomic
factors such as soil acidity (Gordon and Buikstra,
1981; Walker et al., 1988; Walker, 1995) and geological factors such as soil expansion and shrinkage
(Boddington, 1987).
Although clearly demonstrable in poorly preserved samples, these results may be slightly overstated for samples of marginal, but not terrible,
preservation (Buikstra, 1997). While scholars agree
that age effects are real and have been demonstrated in actual case studies (Boddington, 1987;
Gordon and Buikstra, 1981; Henderson, 1987), the
lack of subadults in a skeletal assemblage may be
more the result of burial practices and poor archaeological recovery techniques (Saunders, 1992; Saunders et al., 1995). Despite the study by Walker
(1995), sex and general body size effects are not
supported by other case studies (e.g., Boddington,
1987; Nawrocki, 1995).
In this study, we examine the causes of differential skeletal preservation observed in the Windover
skeletal series, an Early Archaic period mortuary
pond located near the Atlantic coast of Florida. We
test a number of hypotheses regarding the causes of
differential preservation in this sample. We first test
the hypothesis that females are less well-preserved
than males, and that subadults and elderly adults
are less well-preserved than middle-aged adults. We
also evaluate differential deterioration of individual
elements and specific landmarks on a given element.
Our hypothesis is that lighter and less dense elements will display more fragmentation and lower
overall preservation than heavier, denser elements.
In addition, for a given element, those features located near or on less dense or lighter areas of bone
will preserve less frequently than features located
near denser, heavier bone sections. Because brain
preservation was a remarkable aspect of this sample, we also evaluate the relationship between presence of brain material and skeletal condition. Finally, we test two hypotheses regarding the effects
of burial location on skeletal preservation. We first
propose that burials physically located near the
pond margins are less well-preserved than those
located near the pond center. Finally, we test the
hypothesis that burials located higher in the stratigraphic column are less well-preserved than those
buried deeper in the stratigraphic column.
MATERIALS
Study materials
The Windover site (8BR246) is located in Brevard
County, Florida, approximately 8 km west of Cape
Canaveral. The site consists of a long-standing pond
which served as a cemetery for the Windover population. The pond bottom is composed of multiple
strata of anaerobic peat, whose deposition resulted
from the accumulation of decomposing plant material for the last 10,000 years (Doran, 2002). Burials
were placed in shallow depressions in the peat and
were often wrapped in fabric, covered in wood debris, and intentionally staked to the pond floor (Doran, 2002). Based on the presence of preserved brain
matter, it appears that water had filled the pond at
the time interments were occurring; however, it is
uncertain whether the burials were placed below the
water table or merely within saturated soil during
seasonal minima in water elevation (Doran, 2002).
Excavations were conducted by Glen Doran and
David Dickel over a 17-month period between 1984 –
1986. Subsequent radiocarbon dating of human
bone, wooden stakes, peat, and bottle gourd remains
placed burial activities in the Early Archaic period
(after Milanich, 1994), from 8,120 – 6,990 years BP
(Doran and Dickel, 1988a). Most burial activity may
have commenced between 7,330 –7,100 radiocarbon
years BP; however, the maximum range of dates
suggests an approximate 1,000-year interval (Doran, 2002).
The remains of 168 individuals were excavated
from the pond (Doran et al., 1990), 58 of which were
identified in backhoe spoil from construction activity. The functional sample size for most analyses is
therefore around 110 individuals (Doran, in press).
Ages range from infant to 65⫹ years, with approximately half (52%) the sample consisting of subadults
(Hauswirth et al., 1991). The ratio of males to females was approximately equal, suggesting that all
demographic groups are represented in the skeletal
assemblage.
Ninety-one human brains were recovered from the
site, and more than 37 individuals were recovered
with associated fabric material (Doran et al., 1990;
Hauswirth et al., 1991). The preserved brain and
organic plant material seems to suggest excellent
preservation of the associated human remains.
However, the preservation of skeletal material has
been variously described as good, excellent, or simply variable (Doran et al., 1986; Purdy, 1991; Tuross
et al., 1994).
Taphonomic context
At the time of cemetery use, the area consisted of
a wooded environment which flanked the pond margins. The pond itself is bowl-shaped, with a deeper
center portion ringed by a much shallower and
gently sloping ledge near the edge of the pond. It
appears that burial locations were selected based on
water levels at time of burial. Although much of the
pond circumference had been utilized, excavations
in some areas produced little human material (Doran, 2002).
Because of the basin-shaped pond bottom, the
depth and locations of stratigraphic levels vary
across the site. The idealized stratigraphic sequence
DIFFERENTIAL WINDOVER PRESERVATION
17
Fig. 1. Idealized stratigraphic sequence of Windover Pond site. (From Doran, 2002: Figure 10.1. Reprinted courtesy of the
University of Florida Press.)
18
C.M. STOJANOWSKI ET AL.
is presented in Figure 1; however, peat strata
tended to be thinner near the pond margins and
thicker and more compacted near the pond center
(Doran, 2002). These thickness and compaction differences created a context in which two individuals
of similar depth from the ground surface may not
necessarily be located in the same stratigraphic
level. Likewise, the position of each burial in relation to strata interfaces may differ by horizontal
location. Vertical control over burial position, therefore, has more to do with distance from the ground
surface.
The majority of burials were located in the lower
red/brown peat stratum, with a few burials located
in the upper red/brown peat stratum. The lower
stratum was characterized as woody to leafy, and
was deposited in a woody marsh environment beginning around 8,000 BP and terminating around 7,000
BP. The pH of the lower red/brown peat is 5.9 (Doran and Dickel, 1988b). The upper red/brown peat
consists of more woody debris and was deposited
from 7,000 – 6,000 BP, with a pH of 6.6 (Doran and
Dickel, 1988b). The highest stratum at the pond is a
fibrous black peat with a pH of 6.1 (Doran and
Dickel, 1988b). In comparison to other nearby ponds
(with an average pH of 2.9), the pH of the Windover
site is unusually neutral (Hauswirth et al., 1991).
This, combined with high sulphur levels, highly
mineralized water, and an anaerobic peat environment, is believed to be responsible for the preservation of organic remains (Hauswirth et al., 1991).
In this paper we focus on natural explanations for
differential preservation, and therefore must verify
that postdepositional disturbance, incomplete archaeological sampling, spatial variability in burial
position, and age-, sex-, or class-specific differential
burial treatment are not important factors to consider for the purposes of this paper. Postdepositional
disturbance was observed sporadically and was the
result of two unrelated processes. The first was a
general slippage of skeletal elements toward the
deeper center of the pond. This process was variable,
however, and tended to be more severe on the southern edge of the pond, where the margin ledge was
steeper (Doran, 2002). However, it is important to
note that burial density was limited in this unit of
the excavation, and this downslope movement of
bones did not significantly disperse or mix elements
from multiple burials. The majority of burials were
located on the north side of the pond where, due to a
much gentler slope, burial disturbance was not observed (Doran, 2002). Rodent disturbance was rare;
only 6 of 10,000 bone fragments displayed evidence
of gnaw marks (Doran, 2002). Root disturbance was
also limited and largely restricted to the uppermost
burials.
It is more difficult to verify that incomplete recovery or differential burial treatment did not affect the
representation of elements in this sample. However,
there was no apparent age- or sex-specific clustering
of individuals within the site, such that for any
given unit of recovery, all demographic groups were
equally represented (Doran, 2002). Excavation procedures at Windover were designed to maximize
floral and faunal recovery, and all burial fill was
wet-screened. The difficulty even in wet screening
the peat necessitated the use of 1/4 and 1/8 inch
mesh sizes; there was no discernible difference in
the type of material recovered in each screen size,
suggesting representative recovery with the 1/4 inch
mesh size (Doran, 2002).
METHODS
Age and sex were estimated by Glen Doran and
Dave Dickel, following standard osteological procedures (Bass, 1995; Buikstra and Ubelaker, 1994;
Ubelaker, 1989), and were verified by Christopher
Stojanowski with consistent results. Horizontal provenience data were recorded based on the location of
the pelvis for each individual in the analysis. In
addition, each burial was located within the stratigraphic column by recording the distance of the pelvis from the mean annual meters above sea level.
Despite the fact that a number of previously developed quantitative methodologies exist for recording skeletal preservation (Boddington, 1987; Gordon
and Buikstra, 1981; Milner, 1983; Nawrocki, 1995;
Seidemann, 1999; Walker et al., 1988, 1996; Walker
and Snethkamp, 1984; Walker, 1995; White, 1992),
none allowed the specificity we desired. We therefore developed a new scoring methodology specifically for this project. Rather than categorizing a
particular skeleton as poor, fair, or good, we generated a list of skeletal and dental landmarks that
could be converted into a proportion indicating skeletal completeness. Because one impetus for doing
this research was to evaluate the demographic integrity of the Windover sample, a list of landmarks
and measurements with demographic importance
was well-suited to our needs. We chose 35 paired
and 10 unpaired features for a total of 80 observations per individual (see Table 1).
Each feature was determined to be either
“scorable/present” or “unscorable/deteriorated” for
each individual in the sample. For our purposes,
deterioration is defined as a postdepositional, natural taphonomic modification of bone morphology
that reduces the utility of an element or feature in
sexing or aging the individual from whom the bone
derives. Pathological and cultural modifications of
an element that reduce its utility in sex and age
assessment are disregarded and are assigned a
“present” score.
Because teeth serve an integral role in many osteological analyses of marginally preserved skeletal
samples, their inclusion in this analysis was necessary, though problematic. We treated each arcade as
a single, independent unit where the arcade score
can vary between 0 –1. For adults, each tooth is
valued as one sixteenth (1/16) of the total possible
score, resulting in the subtraction of 0.0625 from the
perfect score of 1 for each missing or damaged tooth.
DIFFERENTIAL WINDOVER PRESERVATION
19
TABLE 1. Cranial and postcranial features scored with API methodology
Feature
Cranium
Superciliary arch
Supraorbital margin
Mastoid process
Temporal lines
Root of zygomatic
Gonial angle
Orbital shape
Nuchal crest
Mandibular profile
Frontal profile
Mental eminence
Palate size and shape
Femur
Length
Head diameter
Neck diameter
Midshaft circumference
Bicondylar breadth
Popliteal length
Tibia
Length
Midshaft circumference NF1
Proximal epiphysis breadth
Distal epiphysis breadth
Fibula length
Humerus
Length
Vertical head diameter
Transverse head diameter
Distal epiphysis breadth
Ulna length
Radius length
Scapula
Height
Breadth
Glenoid cavity length
Spine length
Clavicle length
Pelvis
Greater sciatic notch
Auricular surface
Ischial spines
Ilium blade orientation
Ventral arc
Pubic symphysis
Sternum
Manubrium length
Mesosternum length
Sacrum body:wing ratio
Maxillary dentition
Mandibular dentition
Bilateral
Reference
Yes
Yes
Yes
Yes
Yes
Yes
Yes
No
No
No
No
No
Bass, 1995
Buikstra and Ubelaker, 1994
Buikstra and Ubelaker, 1994
White and Folkens, 1991
Morse et al., 1983
Schwartz, 1995
Schwartz, 1995
Buikstra and Ubelaker, 1994
Schwartz, 1995
Morse et al., 1983
Buikstra and Ubelaker, 1994
Schwartz, 1995
Yes
Yes
Yes
Yes
Yes
Yes
Thieme, 1957
Dwight, 1905; Stewart, 1979
Seidemann et al., 1998
Black, 1978; DiBennardo and Taylor, 1979
Olivier, 1969
Pearson, 1917–1919
Yes
Yes
Yes
Yes
Yes
Olivier, 1969
Isçan and Miller-Shaivitz, 1984a,b
Isçan et al., 1994
Isçan et al., 1994
Ubelaker, 1989
Yes
Yes
Yes
Yes
Yes
Yes
Thieme, 1957
Krogman, 1962
Krogman, 1962; Dittrick, 1979
France, 1983
Ubelaker, 1989
Ubelaker, 1989
Yes
Yes
Yes
Yes
Yes
Krogman, 1962
Krogman, 1962
Dwight, 1894; Stewart, 1979
Olivier, 1969
Thieme, 1957
Yes
Yes
Yes
Yes
Yes
Yes
Buikstra and Ubelaker, 1994
Weaver, 1980
Schwartz, 1995
Schwartz, 1995
Phenice, 1969
Buikstra and Ubelaker, 1994
No
No
No
No2
No2
Jit et al., 1980
Jit et al., 1980
Anderson, 1962
Buikstra and Ubelaker, 1994
Buikstra and Ubelaker, 1994
1
NF, nutrient foramen.
Despite the fact that both sides of the dentition are scored in the methodology, the individual tooth scores are collapsed into a single
unit score for each arcade; therefore, dental scores are not treated in a bilateral manner.
2
Any cultural or pathological alterations (attrition,
congenital absence, premortem loss, carious destruction, calculus formation, dental mutilation, or
ablation) were disregarded. Resorbed or abscessed
alveolar regions were not scored as deterioration as
defined above. A tooth still located in the jaw was
scored as “present” if at least 50% of the crown area
was intact. For loose teeth, at least one complete
root and at least 50% of the occlusal surface area
must be observable to be considered “present.”
The dental scoring procedure for subadults is, by
its very nature, more complicated and somewhat
more subjective than for adults. Although the scor-
ing criteria are identical, most subadult arcades contain a combination of deciduous and adult teeth, the
latter of which are often partially formed and/or
erupted. To score immature arcades, we determined
the eruption age of each individual (see Ubelaker,
1989, his Fig. 71) and then generated a completeness score based on the expected number of teeth
present for an individual of that age. For individuals
less than age 5 years, the expected number of teeth
would be 10; missing or damaged alveolar areas
would result in the subtraction of 0.1 per tooth. For
individuals in the lower end of this age range (⬍9
months), no teeth would be erupted and we simply
20
C.M. STOJANOWSKI ET AL.
TABLE 2. API descriptive statistics grouped by sex and
presence of brain matter1
considered the completeness of the alveolar regions.
Complications arise with the eruption of the first
permanent molar. For example, at 6 years, the expectation is for 12 teeth per arcade, a pattern that
generally remains consistent through 11 years. For
these individuals, a missing tooth would result in
the subtraction of 0.08 per tooth. However, different
teeth would be expected at different intervals in this
age range. Furthermore, a missing deciduous tooth
would not be scored as “absent” if the corresponding
adult replacement were present. Because individuals in the 6 –11-year range are the most difficult to
generalize, common sense guided most of our scoring
decisions. After the second adult molar erupted, the
effective number of expected teeth was 14 until the
individual reached full adult dental status.
After the entire suite of traits is scored, an individual score (IS) is computed for each burial. The IS
is simply the sum of “present” features plus the
individual maxillary and mandibular scores divided
by 80, the total number of observations. The IS is
always a positive number between 0 –1, and the
average IS is the average preservation index (API)
for the sample. The API, therefore, serves as a quantitative, semicontinuous measure of preservation.
1
min, minimum; max, maximum; 95%⫾, 95% confidence interval
bounds; SE (m), standard error of the mean; SD, standard deviation.
Statistical methods and research methodology
RESULTS
To test for demographic biases, we calculated the
preservation scores by sex and by age groups (0 –5,
6 –10, 11–15, 16 –20, and 21–35 males; 21–35 females; 36 –50 males; 36 –50 female; 50⫹ male; and
50⫹ female). If skeletal decay is a relatively continuous process, then a significantly lower API score
indicates that individuals from the left tail of that
demographic group’s distribution are absent from
the archaeological record. Parametric t-tests and
nonparametric Mann-Whitney rank tests were used
to analyze sex-specific data. To analyze data by age,
we utilized ANOVA and the nonparametric equivalent Kruskal-Wallis test. We also calculated the
Pearson product-moment correlation coefficient between IS and the point estimate for the age of each
individual. These correlation coefficients were tested
against zero, using post hoc Bonferroni alpha protection.
To evaluate the relationship between element
weight and density and bone preservation, we calculated the frequency of preservation for each trait
listed in Table 1. This was accomplished by dividing
the number of “present” features by the total number of observations (for the entire sample) for each
trait. To determine an overall average completeness
score for each element with multiple traits (cranium, femur, and tibia), we averaged the individual
feature scores. For example, for a hypothetical element with three distinct scoring features (each with
frequencies of 50%, 75%, and 100%), the overall
element score would simply be the average of these,
or 75%.
We investigated the relationship between brain
preservation and skeletal condition using a t-test
The preservation scores for the Windover sample
are presented in Tables 2 and 3. There is surprisingly little variation in mean API score regardless of
age or sex. Ignoring subdivision by sex or age provides a measure of the average preservation score
for the entire sample. The API was 0.53 (n ⫽ 109,
SD ⫽ 0.22), indicating that the Windover sample is
fairly well-preserved, with 53% completeness for the
features we scored. However, as supported by
Tuross et al. (1994), the range of scores is suggestive
of the degree of variability seen in skeletal preservation in the Windover materials. The overall range
of individual preservation varies from 0 –95%.
Using the data presented in Table 2, we examined
the relationship between estimated sex (or general
body size) and API score. A two-sample t-test (t ⫽
0.26; df ⫽ 71; P ⫽ 0.79) and the Mann-Whitney test
(P ⫽ 0.77) failed to reach significance. We submitted
the data in Table 3 to a one-way ANOVA and found
no significant difference in preservation between
age groups (F ⫽ 1.09; P ⫽ 0.37). The Pearson correlation coefficient between age group and API equals
0.31, indicating that age group explains 9% of the
variability in API score. The Kruskal-Wallis analysis of variance test was also not significant (P ⫽
0.33, assuming chi-square with 9 degrees of freedom). The correlation between age and preservation
was not significant (r ⫽ 0.10, P ⫽ 0.30). Therefore,
we conclude that the demographic integrity of the
Windover sample is intact: individuals of both sexes
and all ages display, on average, similar preservation proportions.
In Table 4, we present the average preservation
scores and average element weights (whole bones)
Sex
n
Min
Max
Mean
95%⫹
95%⫺
SE (m)
SD
Brain
Male
Female
Unknown
Yes
No
38.0
0.16
0.94
0.54
0.61
0.47
0.04
0.22
29.0
0.19
0.95
0.52
0.59
0.44
0.04
0.21
34.0
0.03
0.91
0.48
0.56
0.40
0.04
0.22
86.0
0.0
0.96
0.55
0.59
0.51
0.02
0.20
23.0
0.04
0.91
0.48
0.59
0.36
0.06
0.26
and a nonparametric Mann-Whitney test. To analyze the relationship between horizontal and vertical burial location and preservation score, we calculated Pearson correlation coefficients for the
following comparisons: API vs. north coordinate,
API vs. east coordinate, and API vs. meters above
sea level (MASL). The correlations were tested
against zero with post hoc Bonferroni alpha protection.
21
DIFFERENTIAL WINDOVER PRESERVATION
1
TABLE 3. API descriptive statistics for age cohorts
n
Min
Max
Mean
95%⫹
95%⫺
SE (m)
SD
1
0–5
6–10
11–15
16–20
20–35,
male
20–35,
female
35–50,
male
35–50,
female
50⫹,
male
50⫹,
female
10.0
0.22
0.76
0.42
0.57
0.26
0.07
0.21
14.0
0.04
0.86
0.50
0.64
0.36
0.07
0.25
7.0
0.48
0.83
0.48
0.67
0.29
0.08
0.21
6.0
0.49
0.88
0.64
0.78
0.49
0.06
0.14
10.0
0.20
0.83
0.61
0.75
0.48
0.06
0.19
6.0
0.22
0.88
0.57
0.79
0.35
0.09
0.21
15.0
0.25
0.89
0.50
0.58
0.41
0.04
0.16
12.0
0.26
0.93
0.60
0.73
0.47
0.06
0.20
15.0
0.12
0.96
0.58
0.72
0.44
0.07
0.25
9.0
0.31
0.91
0.49
0.63
0.34
0.06
0.19
min, minimum; max, maximum; 95%⫾, 95% confidence interval bounds; SE (m), standard error of the mean; SD, standard deviation.
TABLE 4. Windover API scores compared against
average element weights (grams)
Element
Average API
Average weight1
Cranium
Femur
Tibia
Pelvis
Humerus
Scapula
Ulna
Fibula
Radius
Clavicle
Sternum
0.84
0.58
0.49
0.39
0.47
0.22
0.55
0.27
0.58
0.30
0.11
514
255
154
113
92
41
38
35
31
15
13
1
Average weights in grams based on data published by Lowrance
and Latimer (1957, their Table 1).
derived from a modern reference sample (Lowrance
and Latimer, 1957). The correlation between weight
and preservation is statistically significant (r ⫽
0.77; P ⫽ 0.005), indicating that the least well-preserved elements were, in general, also the lightest.
For example, the cranium is the best preserved unit
(84%), followed by the femur, radius, ulna, and tibia,
respectively. The least well-preserved elements
were the scapula (22%) and sternum (11%), two of
the lightest and least dense elements in the body. A
consideration of differential preservation within a
single element produces similar results (Table 5).
For example, the scapular body is the thinnest portion of the skeleton and presented the lowest overall
preservation score in the sample (scapula height
API ⫽ 2%). However, the dense glenoid cavity (64%)
preserved at levels higher than most other noncranial traits, with the possible exception of long bone
shaft fragments. In general, the data in Table 5
confirm previous analyses documenting a relationship between bone density and preservation (see
Galloway et al., 1997; Mays, 1991; Spennemann,
1992; Waldron, 1987).
In Table 2, we also present data on the relationship between brain and skeletal preservation. These
data indicate little difference in preservation score
between those individuals with preserved brains
and those without. A t-test grouped by brain preservation (t ⫽ ⫺1.46; df ⫽ 107; P ⫽ 0.15) and the
Mann-Whitney test (P ⫽ 0.22) were not significant.
However, a different pattern emerged when cranial
scores were tabulated independently of mandibular,
maxillary, and postcranial scores. The data in Table
6 indicate a stark contrast between postcranial and
cranial/dental preservation. As expected, the dental
remains exhibited the highest preservation score,
with the mandibular dentition exhibiting slightly
higher (though not statistically significant) completeness scores than the maxillary remains. Cranial preservation was comparable to both dentitions,
however, as indicated in Table 7. Only the maxillary
and cranial scores were significantly correlated (r ⫽
0.67; P ⬍ 0.0001). In terms of preservation of brain
tissue, significant correlations were found between
preservation of brain tissue and cranial preservation (r ⫽ 0.43; P ⬍ 0.0001), and maxillary preservation (r ⫽ 0.52; P ⬍ 0.0001). Parametric significance
tests adjusted for familywise error indicated significant differences between those individuals with
brains and those without when considering the cranial (P ⬍ 0.0001) and maxillary (P ⬍ 0.0001) scores;
the mandibular (P ⫽ 0.02) and postcranial scores
(P ⫽ 0.28) were not significantly different when Bonferroni adjustments were made. These data suggest
that the remarkable preservation of brain tissue is
related to how well the brain was encased by the
cranial architecture, an unsurprising result.
Finally, we examined variability in preservation
across the site and through the stratigraphic column. As the results indicate (Table 8), north and
east burial coordinates are not correlated with preservation. However, the correlation between API and
meters above sea level is significant (r ⫽ ⫺0.31; P ⫽
0.005), indicating that preservation decreased as
meters above sea level increased.
DISCUSSION
In this paper, we tested several hypotheses regarding the causes of differential preservation in the
Windover sample. The sample exhibited a wide
range of preservation scores, indicative of the degree
of preservation variability. As a whole, preservation
for the entire sample was approximately 53%. Of the
factors intrinsic to the bones themselves, we found
little evidence for age- or sex-specific differences in
preservation score. This result supports the notion
of Buikstra (1997) that the demographic effects of
poor preservation may only be experienced in the
most marginal situations. Although the lack of a sex
correlation is not surprising and confirms the results
22
C.M. STOJANOWSKI ET AL.
TABLE 5. Percentage of observable features for Windover sample1
Feature
Cranial paired
Superciliary arch
Supraorbital margin
Mastoid process
Temporal lines
Root of zygomatic
Gonial angle
Orbital shape
Postcranial paired
Femur
Length
Head diameter
Neck diameter
Midshaft Circum at NF
Bicondylar breadth
Popliteal length
Tibia
Length
Midshaft Circum at NF
Proximal epiphysis breadth
Distal epiphysis breadth
Fibula length
Humerus
Length
Vertical head diameter
Transverse head diameter
Distal epiphysis breadth
Ulna length
Radius length
Scapula
Height
Breadth
Glenoid cavity length
Spine length
Clavicle length
Pelvis
Greater sciatic notch
Auricular surface
Ischial spines
Ilium blade orientation
Ventral arc
Pubic symphysis
1
Left
(%)
Right
(%)
89
88
91
85
92
92
49
93
93
93
82
94
89
48
53
53
71
90
32
57
56
44
74
88
25
53
48
80
29
44
29
46
83
24
37
26
46
38
32
65
53
54
44
37
37
75
57
61
2
9
64
13
27
2
13
59
17
32
63
59
36
29
28
22
69
59
42
35
33
25
Feature
(%)
Cranial unpaired
Nuchal crest
Mandibular profile
Frontal profile
Mental eminence
Palate size and shape
82
90
92
89
66
Postcranial unpaired
Sternum
Manubrium length
Mesosternum length
Sacrum
Ratio of body:wings
Dental traits
Mandible
Maxilla
13
9
32
92
86
NF, nutrient foramen; Circum, circumference.
TABLE 6. API descriptive statistics for skeletal
and dental subgroupings1
n
Min
Max
Mean
95%⫹
95%⫺
SE (m)
SD
Crania
Maxilla
Mandible
Postcrania
102.0
0.0
1.0
0.77
0.83
0.71
0.03
0.29
102.0
0.0
1.0
0.80
0.87
0.74
0.03
0.33
98.0
0.0
1.0
0.90
0.94
0.86
0.02
0.20
102.0
0.0
0.95
0.41
0.46
0.36
0.02
0.24
1
Min, minimum; Max, maximum; 95%⫾, 95% confidence interval
bounds; SE (m), standard error of the mean; SD, standard deviation.
of independent analyses from other samples (Boddington, 1987; Buikstra, 1997; Henderson, 1987;
Nawrocki, 1995), given the range of variability and
the seemingly mediocre API for this sample, it is
somewhat surprising that an age-specific bias was
not evident.
Another result from this study was the lack of
correlation between overall skeletal preservation
and the presence or absence of preserved brain material, suggesting that factors responsible for brain
preservation were unrelated to skeletal preservation. We did, however, find a significant relationship
between cranial condition and brain preservation,
suggesting that the cranium protected brain material from the physical and chemical agents responsible for its deterioration in some individuals. The
relationship between brain preservation and cranial
condition was also evident based on the distribution
of brain matter by age; in general, subadults had
fewer intact brains than did adults, a fact attributable to the thinner cranial architecture of immature
individuals (Doran, 2002).
The relationship between preservation and burial
location perhaps provides the best explanation for
differential preservation at the site. We first examined the relationship between horizontal location
and preservation score. There were two possible reasons why position within the cemetery could have
affected skeletal condition. First, some evidence suggests that the site was utilized for brief periods of
DIFFERENTIAL WINDOVER PRESERVATION
TABLE 7. Pearson correlation coefficients and associated
p-values for skeletal subgroups
Crania
Maxilla
Mandible
Postcrania
Brain
Crania
Maxilla
Mandible
Postcrania
Brain
1.0
1.0
0.68
0.00
0.02
0.85
0.20
0.05
0.43
0.00
1.0
1.0
0.19
0.08
0.05
0.66
0.52
0.00
1.0
1.0
0.19
0.06
0.07
0.48
1.0
1.0
⫺0.16
0.13
1.0
1.0
TABLE 8. Pearson correlation coefficients and associated
p-values for API vs. provenience data
API
North
East
Depth
1
API
North
East
MASL1
1.0
1.0
⫺0.01
0.93
0.09
0.45
⫺0.31
0.01
1.0
1.0
0.74
0.00
0.00
0.98
1.0
1.0
⫺0.05
0.68
1.0
1.0
MASL, meters above mean sea level.
time over a span of 1,000 years. If groups maintained distinct burial locations, then the placement
of a burial within the pond may be indicative of the
date it was placed there. If the site was used over a
long period of time, then potentially some skeletons
have had an additional 1,000 years to deteriorate. A
second explanation was a difference in soil chemistry or soil compaction across the site. If pH varied by
location, then we expected skeletal condition to vary
likewise. However, we found no indication that preservation varies in any consistent way across the site.
Although unit-specific pH data are not currently
available, the distribution of radiocarbon dates may
explain the lack of correlation between preservation
and horizontal distribution. One of the more problematic aspects of the Windover site concerns establishing a sequence of burial activities. This results
from the fact that for any given segment of the pond,
a very early and a very late date can be found within
a few meters of one another (Doran, 2002). This may
relate to the opportunistic fashion in which final
burial location was chosen, a decision possibly based
on kin affiliation but also seasonal variation in water depth (Doran, 2002). Assuming that the water
table affected placement (for logistical reasons),
then two individuals who died in different seasons of
the same year could be buried at much different
locations in relation to the pond edge. If the water
level were low, one would expect the individual to be
buried closer to the pond center; likewise, if the
water level were high, one would expect a burial to
be closer to the shallower pond margins. The same
pattern would hold for individuals who died hundreds of years apart: due to water levels in the pond
23
at time of interment, two individuals of vastly different dates could be buried in close proximity.
Differences in vertical stratigraphic position provide another potential source of variation in skeletal
condition. As with horizontal distribution, there
were several ways in which vertical position could
have affected bone preservation. The most obvious
was the change in soil pH in different stratigraphic
levels. Because the soil was more neutral closer to
the ground surface, the expectation is for improved
preservation in burials located higher in the column
(Gordon and Buikstra, 1981; Nawrocki, 1995). However, working against this is the factor of seasonal
variation in water depth. If water depth varied after
bodies were interred, burials closer to the ground
surface would experience more frequent periods of
partial drying and subsequent resubmergence, a
process that hastens bone deterioration. Our analyses indicate that burials located closer to the modern-day ground surface were less well-preserved,
despite the soil pH being more neutral. This supports the latter notion that preservation at Windover Pond may be related to the periodic partial
drying of the burial area (Tuross et al., 1994).
Despite the unique anaerobic burial context, we
found a significant correlation between element
weight or density and skeletal preservation, a result
supported by previous analyses (Galloway et al.,
1997; Mays, 1991; Spennemann, 1992; Waldron,
1987). The consistency of the deterioration process is
demonstrated in Table 9, in which we present element- or feature-specific preservation rates based on
three comparative samples (Galloway et al., 1997;
Spennemann, 1992; Waldron, 1987). For each sample, the traits are ranked in order of completeness,
and information on the absolute preservation of
each feature is provided. In Table 9, we reproduce
the preservation scores from Tables 4 and 5 for ease
of comparison.
The data of Spennemann (1992) are derived from
a recently eroded historic cemetery in the Marshall
Islands that had been disturbed by exceptional tidal
activity and wave action. The taphonomic context is
typified by a high-energy, littoral environment with
a relatively short interval for deterioration (Spennemann, 1992). For both samples, the cranium was the
best-preserved element, and the scapula the leastpreserved. In addition, the pattern of representation
for the least well-preserved elements was identical:
the clavicle is slightly better than the sacrum, which
is slightly better than the scapula. Of the long bones,
the femur and tibia were the best-preserved, followed by the humerus; the preservation of the fibula, radius, and ulna was somewhat variable between samples. In general, the pelvis was less wellpreserved than the long bones.
The data of Waldron (1987) were derived from a
Romano-British terrestrial cemetery dating to the
second to fourth centuries AD. Whereas Spennemann (1992) reported whole bone representation,
Waldron (1987) reported preservation frequencies
24
C.M. STOJANOWSKI ET AL.
TABLE 9. Comparative preservation data from archaeological and modern reference samples1
Spennemann (1992), trait (%)
[Windover %]
Waldron (1987), trait (%)
[Windover %]
1. Cranium (100%) [85%]
2. Tibia (75%) [49%]
3. Femur (69%) [58%]
4. Humerus (63%) [47%]
5. Fibula (50%) [27%]
6. Pelvis (44%) [39%]
7. Radius/Ulna (38%) [58%/55%]
8. Clavicle (25%) [30%]
9. Sacrum (13%) [32%]
10. Scapula (0%) [10%]
1. Sciatic notch (70%) [63%]
2. Auricular surface (64%) [59%]
3. Prox femur (52%) [60%]
4. Mastoid process (59%) [91%]
5. Dist humerus (56%) [65%]
6. Dist femur (52%) [32%]
7. Glenoid fossa (50%) [64%]
8. Dist tibia (48%) [44%]
9. Prox humerus (47%) [35%]
10. Prox tibia (44%) [29%]
11. Pubic symphysis (31%) [22%]
12. Manubrium (23%) [13%]
13. Mesosternum (22%) [9%]
14. Scapular body (11%) [6%]
1
Galloway et al. (1997), trait (density)
[Windover %]
1.
2.
3.
4.
5.
6.
7.
8.
Femur midshaft (0.65) [90%]
Tibia midshaft (0.48) [80%]
Femur neck (0.32) [71%]
Dist humerus (0.27) [65%]
Femur head/Dist tibia (0.24) [53%/44%]
Prox humerus (0.20) [35%]
Dist femur (0.18) [32%]
Prox tibia (0.16) [29%]
Prox, proximal; Dist, distal.
for various bone segments and features. We attempted to correlate the landmarks of Waldron
(1987) with our own, and only report those traits
that correspond reasonably well. The percentage
representation data agree well between samples,
particularly for the most poorly preserved features.
In fact, the last four entries in column 2 of Table 9
correspond exactly in both samples, both in terms of
relative preservation and in terms of their rank
within the respective sample. It is clear that the
pubic region, the sternum, and the scapula are the
first to deteriorate. Although there are minor differences in trait representation percentages, several
comparative frequencies hold up reasonably well:
distal humerus ⬎ proximal humerus; distal tibia ⬎
proximal tibia; and sciatic notch and auricular surface ⬎ pubic symphysis. However, despite the overall similarity in the lists, two features differ rather
significantly in representation, the mastoid process
and the distal femur. The reasons for this are difficult to discern.
Finally, we compared the representation of long
bone fragments from the Windover sample to density measures derived from a prepared sample of
modern humans (Galloway et al., 1997). The comparison of these data produced the best correspondence and, in fact, the ranked density measures are
perfectly correlated with the ranked preservation
scores (Table 9, column 3). Midshaft sections of the
long bones preserved the best, followed by the distal
humerus and proximal femur. Interestingly, the
within-element relationships seen in the comparative data of Waldron (1987) are also supported by
the density measures: distal humerus ⬎ proximal
humerus; distal tibia ⬎ proximal tibia; and proximal
femur ⬎ distal femur.
Overall, the comparative data suggest that the
process of bone deterioration is relatively constant
and consistent, regardless of the exact taphonomic
context. The skull is the best-preserved unit, followed by long bone shafts, the denser epiphyseal
long bone ends, and finally the fragile sternum, clavicle, and scapula. For the pelvis, the denser auricu-
lar and sciatic region is better represented than the
pubic region (Telmon et al., 1993). Despite the
uniqueness of Windover’s taphonomic context, the
results from this study seem applicable to most
other human skeletal assemblages, including those
subjected to hydrodynamic sorting processes in a
saline-rich, high-energy coastal zone (Spennemann,
1992).
In summary, the analysis of skeletal taphonomy
for the Windover sample has accomplished several
goals. We quantified the average and range of preservation scores for the entire sample, and determined that age and sex were unrelated to skeletal
condition, as was the position of each burial within
the pond. However, preservation did decline in burials that were located closer to the modern ground
surface. As the water level in the pond fluctuated
over the last 8,000 years, burials highest in the
stratigraphic column suffered the deleterious effects
of drying and resubmergence, which resulted in deterioration of the least dense skeletal regions. No
demographic effects were discernible, and it seems
likely that the Windover sample is representative of
the population that contributed the burials.
ACKNOWLEDGMENTS
We thank Clark Spencer Larsen and three anonymous reviewers for their helpful comments on the
manuscript. Any errors or omissions remain our sole
responsibility.
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skeletal, causes, preservation, differential, windover, consequences, pond
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