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Body size among Holocene foragers of the Cape Ecozone southern Africa.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 129:1–11 (2006)
Body Size Among Holocene Foragers of the Cape
Ecozone, Southern Africa
S. Pfeiffer1,2* and J. Sealy2
1
2
Department of Anthropology, University of Toronto, Toronto, Ontario M5S 363, Canada
Department of Archaeology, University of Cape Town, 7701 Rondebosch, South Africa
KEY WORDS
Later Stone Age; hunter-gatherers; Khoesan; bioarchaeology; stature; stable
isotopes; paleodiet
ABSTRACT
Temporal and geographic variability in
adult body size can be a useful indicator of a population’s adaptation. The southern African Cape supported
foraging populations exclusively until some pastoralism
is seen, ca. 2000 BP. This paper describes and interprets
body-size patterns among foragers, as deduced from
maximum femoral lengths and femoral head diameters,
using 127 individually dated adult skeletons from the
western (70) and southern (57) regions of the Cape (64
male, 60 female, 3 sex undetermined). Estimated statures are comparable to historic Khoesan samples, but
show lower values and greater variance during the fifth/
fourth millennium before the present among both sexes
and both biomes. Variation in femoral length does not
Adult body size can be an important indicator of a population’s adaptation to its environment. Stature and weight
are ubiquitous measures of health among modern societies. Reconstructions of body size in past populations are
important to studies of the evolution of our species and
the adaptation of that species to diverse global environments. Stature can be reconstructed from the length of
major long bones, especially the femur, while the size of
joint surfaces at the hip contributes information regarding mass of the torso (Ruff, 1994; Ruff et al., 1997). Adult
stature reflects an individual’s history of nutrition, in balance with the energy demands of activity and disease,
within the context of population genotype. Stature has
been incorporated as a key component of the ‘‘health
index’’ through which biological variables are quantified
to elucidate past quality of life (Steckel, 1995; Steckel and
Rose, 2002; Steckel et al., 2002). While researchers have
explored the increased morbidity associated with the origin of agriculture (Cohen and Armelagos, 1984), the
dynamics of health among foragers are much less fully
explored. It is in this context that this study presents
information about past hunter-gatherers of southern
Africa, during millennia of foraging before the adoption of
herding (Sadr, 2003).
The Cape Ecozone (Deacon and Lancaster, 1988) comprises the western and southern coasts of South Africa
from the Cape Fold Mountains to the Indian and Atlantic
Oceans, and includes an area of coastal lowlands. The
principal vegetation type of the Cape Ecozone is the fineleaved scrub often referred to as fynbos. In terms of the
number of plant species per unit of area, it is one of the
richest plant communities in the world (Meadows and
Sugden, 1993). The southern region has a predominantly
rocky shoreline and a coastal platform with an elevation
#
2005 WILEY-LISS, INC.
correlate with diet protein, as reflected in stable isotope
(d13C and d15N) values. Positive correlations between
femoral head diameter and isotopic indicators suggest
greater body mass with more reliance on marine protein.
A decline in femoral length begins at around 4000 BP, a
time when archaeologists suggest that population
growth led to the incorporation of lower-ranked food
resources, and to reduced mobility. A clearly identifiable
linear recovery, beginning at around 3000 BP, greatly
predates the earliest evidence of pastoralism on the
Cape. Apparent problems of food sufficiency were addressed and solved, within a hunting and gathering
economy. Am J Phys Anthropol 129:1–11, 2006.
'
2005 Wiley-Liss, Inc.
of approximately 200 m. The coastal platform ranges from
2–20 km in width, and abuts the Cape Fold Mountains to
the north, which separate the Cape Ecozone from the
Karoo. The climate along the south coast is warm and
temperate, with year-round rainfall ranging between
700–1,200 mm per year (Anonymous, 2003). Coastal and
marine resources are linked to the productivity of the
Indian Ocean. Because the dominant vegetation is afromontane forest, the region is classified as forest biome
(Butchart, 1995; Rutherford and Westfall, 1986).
The western region of the Cape Ecozone consists predominantly of a broad, sandy coastal lowland with a
Mediterranean-like climate of mild, wet winters and hot,
dry summers. The average annual rainfall ranges from
less than 200 mm to slightly more than 1,000 mm per
year (Anonymous, 2003). The coastal and marine
resources of the western region are linked to the productivity of the South Atlantic Ocean. The more southerly
Grant sponsor: Social Sciences and Humanities Research Council
of Canada; Grant sponsor: National Research Foundation of South
Africa; Grant sponsor: University of Cape Town; Grant sponsor:
Palaeoanthropology Scientific Trust.
*Correspondence to: S. Pfeiffer, now at the Department of Anthropology, University of Toronto, 100 St. George Street, Toronto,
Ontario M5S 3G3, Canada. E-mail: susan.pfeiffer@utoronto.ca
Received 1 March 2004; accepted 15 September 2004
DOI 10.1002/ajpa.20231
Published online 13 September 2005 in Wiley InterScience
(www.interscience.wiley.com).
2
S. PFEIFFER AND J. SEALY
portion of the western region is classified as fynbos
biome, and the northern portion is classified as succulent
karoo biome (Butchart, 1995; Rutherford and Westfall,
1986).
The Cape Ecozone of southern Africa has been home
to foraging peoples for thousands of years. Foraging
(hunting and gathering) was the only mode of subsistence throughout the Middle Stone Age and during most
of the Later Stone Age. The transition from Pleistocene
to Holocene adaptations included a shift from reliance
on social, grazing prey to reliance on nongregarious
browsers and a variety of food sources that, while nutritious, came in smaller package sizes. Geophytes, fruits
and nuts, and tortoises were among the most important
food resources; along the coast, fish and shellfish were
key items (Deacon and Deacon, 1999; Mitchell, 2002).
Archaeological study affirmed continuity from the Holocene foragers and pastoralists of the region to living peoples of Botswana, Namibia, and South Africa who speak
Khoesan languages. Evidence for continuity comes from
the tools they made, the technological approaches they
used, and the symbolism of their rock art (Deacon and
Deacon, 1999; Mitchell, 2002). In historic times, studies of
stature documented the Khoe pastoralists to be taller
than the San foragers (Wells, 1960), but Khoesan groups
have consistently been shorter than Black African and
European average statures.
Prior studies have used smaller samples to explore
variability in body size in the Cape during the Holocene.
Wilson and Lundy (1994) published a pilot study of 45
femora from dated, prehistoric skeletons, mainly from
the western region, concluding that prehistoric statures
were comparable to historically documented statures for
Bushmen. Wilson and Lundy (1994, p. 7) further noted
‘‘some evidence of diachronic change in the statures of
both the female and male samples, with a decrease after
3000 BP and an increase after 2000 BP.’’ A radiological
study of humeri noted a temporal disjuncture in the western region as well (Smith et al., 1992). Sealy and Pfeiffer (2000), studying femora, noted a similar pattern
among skeletons from the southern region. There, it
appeared that skeletons with more terrestrial isotopic
signatures tended to be smaller, prior to 2000 BP, but
this tendency did not pertain in more recent times.
Past studies of skeletal size among Later Stone Age
foragers of the Cape suggested that perhaps the origin of
pastoralism, at around 2000 BP, provided the catalyst
for change. The current study provides a new analysis of
a substantially larger sample of dated skeletons. Using
the femur as a proxy for stature (length) and mass (head
diameter), the purpose of this work is to explore patterns
of adult body size throughout the Holocene, emphasizing
the period between approximately 4,000 and 2,000 years
ago, during which average adult body size was reduced.
The focus is on variability within Holocene hunter-gatherer societies, but this study also elucidates the extent to
which the introduction of sheep herding, ca. 2,000 years
ago, may have influenced the populations of the Cape.
The pattern of change in body size is explored particularly with regard to sources of dietary protein, as
reflected in values of d15N and d13C from bone collagen.
MATERIALS AND METHODS
Skeletons for this study are derived from that section
of the South African coastline south of 328 South and
west of 248 East, extending inland as far as the Cape
Fig. 1. Map of southern Africa, highlighting southern and
western regions from which Later Stone Age skeletons were
derived for this study.
Fold Mountain belt. This area falls today within the
Western Cape Province of South Africa (Fig. 1). The skeletal sample includes 57 skeletons from the southern
region (26 male, 30 female, 1 sex undetermined), and 70
from the western region (38 male, 30 female, 2 sex undetermined) (deVilliers and Wilson, 1982; Morris, 1992;
Muller, 2002; Patrick, 1989; Pfeiffer et al., 1999; Sealy
and Pfeiffer, 2000; Sealy, 1989; Sealy et al., 1992; Sealy
and van der Merwe, 1986, 1988) (Table 1). Most of the
southern-region skeletons come from sites that yielded
multiple skeletons, like Matjes River Rock Shelter,
Oakhurst, and Drury’s Cave. Most of the western-region
skeletons were isolated finds or small burial groupings
from coastal dunes or rock-shelter contexts. Each skeleton is both temporally and spatially identified. All but
two have radiocarbon dates based on samples of bone tissue, and the remaining two (UCT 345, Nelson Bay Cave
burial 2; UCT 390, Faraoskop) are very closely associated with radiocarbon dates from the burial context.
Among skeletons from the past 2,000 years, information
from the archaeological context, bone morphology, and
stable isotopes was used to avoid the remains of obvious
pastoralists and new immigrants to the region. The distinction between forager and pastoralist, however, may not
always be clearly marked during this time of transition.
Femora from 127 skeletons held at five curatorial
institutions in South Africa were measured, using standard osteological methods. Left femora were used, except
in cases where measures were only available from the
right side. Sex was ascertained from morphological indicators, chiefly the pelvis and cranium. In cases where indicators are equivocal, an ‘‘undetermined’’ sex is indicated.
No stature estimation formulae have been developed
that are specifically appropriate for Khoesan people.
Both the crural index and the ratio of femoral head diameter to femoral length are intermediate between published values for blacks and whites (Powell et al., 2004;
Ruff, 1994), so stature was estimated from various equations, the results of which should approximate true stature. The white and black regression equations proposed
by Feldesman and Fountain (1996), the generic femur/
stature ratio of 26.74 proposed by Feldsman et al.
(1990), and the sex-specific white and black equations
developed by Trotter (1970) are applied. When applied
appropriately, the equations of Feldesman and Fountain
(1996) are reported to have a mean absolute deviation of
BODY SIZE AMONG FORAGERS
3.3 cm; the generic femur/stature ratio has a mean absolute deviation of 4.4 cm. (Feldesman and Fountain,
1996). Error terms associated with the equations of
Trotter (1970) range from 3–4 cm.
Collagen samples were analyzed for stable carbon and
nitrogen isotopes, for the purpose of assessing paleodiet.
Variation in the composition of flora of the western and
southern regions dictates that studies of paleodiet should
ideally focus on carbon and nitrogen systems, respectively. However, Sealy (1997) demonstrated a strong covariance between individual d13C and d15N values across
both regions. Both d13C and d15N values reflect the relative importance of marine and terrestrial foods in each
individual’s diet. Therefore, both carbon and nitrogen
values are used in this study to assess possible relationships between diet and body size.
Southern-region skeletons range in age from 560–9720
BP; western region skeletons range from 620–9750 BP.
This analysis uses uncalibrated radiocarbon dates. Each
skeleton is the remains of a person who in life may have
ingested a mix of proteins from terrestrial and marine
sources. Because carbon in the marine system is less
rapidly mixed with newly formed 14C from the upper
atmosphere, a radiocarbon date will overage tissue to
the extent that collagen incorporates marine-based carbon. This is a recognized phenomenon, but there is no
system of calibration that accommodates this bias and
retains any systematic error term. Further, the events of
the Later Stone Age were identified and documented
using uncalibrated radiocarbon dates. In order to place
this study in the broader context of the Later Stone Age,
dates were not calibrated (as per Mitchell, 2002).
RESULTS
Summary statistics for maximum femoral length and
maximum femoral head diameter are provided in Table 2.
Among the 108 available pairs of osteometric values, the
Pearson correlation coefficient is 0.705 (P < 0.001), indicating that while length and head diameter covary substantially, they also contribute some different information about body size. The ratios of femoral head diameter
to maximum length are significantly different for men
(M) and women (F) (M, 9.66; SD, 0.47; F, 9.30; SD, 0.42;
t ¼ 4.22; P < 0.01), indicating that sexual dimorphism
affects both size and shape of the femur. Nevertheless,
given the near balance of sexes within the sample (64
males, 60 females, 3 sex undetermined), the sexes are
pooled in some of the subsequent analyses.
Stature estimation
Table 3 shows that estimated average statures based
on femur lengths are intermediate between those of San
(Bushmen) men and women reported by Dart (1937a,b)
and those of Khoe (‘‘Hottentot’’) men reported by
Schultze (1928). The formulae of Trotter (1970) yield taller male statures and greater sexual dimorphism. All
estimates fall within the range of relatively short adult
modern humans. Both the male and female mean estimates are below the 10th centiles for the current US
population (Anonymous, 2004a,b).
Body size through time
The sexes are combined in the exploration of temporal
variability. Both osteometric variables show similar temporal patterns of variation. As illustrated in Figure 2,
3
both relationships are best modeled by a quadratic curve
(femur length: r2 ¼ 0.106, F ¼ 6.57, P ¼ 0.002; head diameter: r2 ¼ 0.124, F ¼ 8.37, P < 0.001). The decline in
body size occurs in both biomes at around the same time.
Specimens that are older than 8000 BP have considerable influence on the strength of the relationship. If the
femoral lengths older than 6000 BP are discounted, the
relationship is strengthened (r2 ¼ 0.183, F ¼ 12.23, P <
0.001). Removing the oldest, largest femoral head diameter (UCT 374, Elands Bay, 9750 BP) reduces the
strength of that relationship, but it remains significant
(F ¼ 4.23, P ¼ 0.02). There are fewer data points prior
to 4000 BP than since 2000 BP, but with the exception of
femoral head diameter noted above, values appear to be
similar before 4000 and since 2000 BP. Between 4000
and 2000 BP, there is increased variability introduced by
some much smaller femora. The plot of femoral length
includes a relatively small number of very short femora
at around 3000 BP, such that a few anomalous individuals might be skewing the temporal pattern. However,
the plot of femoral head diameter includes more data
points in that time range, and confirms that this is a
general phenomenon.
To explore the relative magnitude and timing of this
transient period of reduced stature, the time range is
divided into 1,000-year periods, beginning with 500–
1499 BP (Fig. 3). In both the western and southern
regions, the small number of skeletons from the period
of 3500–4500 BP mark the beginning of reduced body
size, in both linear and mass dimensions.
Focusing on those 87 skeletons with 14C dates more
recent than 3000 BP, Figure 4 shows that femoral
lengths increase linearly from the beginning of the third
millennium (79 pairs, r2 ¼ 0.27, F ¼ 28.84, P < 0.001).
The increase in body size begins about 1,000 years earlier than the shift in dietary protein sources that was
associated with the initiation of pastoralism.
Body size in relation to diet
Figures 5 and 6 summarize the isotopic values for skeletons in this study. Figure 5 shows d15N plotted against
d13C for western and southern Cape sample sets. Values
are significantly correlated, in a manner similar to that
previously demonstrated for coastal regions of the Cape
(Sealy, 1997). The fitted regression equation for the southern region (54 pairs) is d15N ¼ 28.0 þ 1.05 d13C, r2 ¼ 0.59
(P < 0.001). For the western region (67 pairs), the fitted
regression equation is d15N ¼ 23.5 þ 0.65 d13C, r2 ¼ 0.47
(P < 0.001). The shallower slope of the line for the western region and the lower r2 are probably associated with
the aridity, and hence higher terrestrial d15N values,
found in parts of the western region. Because isotope
values from the two regions are not precisely comparable,
western and southern sample sets are distinguished in
the analysis below. However, general consistency between
the two regions increases our confidence in trends seen in
the separate data sets.
Figure 6 shows d13C and d15N values through time for
the two regions. These values are scattered, indicating
high interindividual variation in the diet. There are,
however, some patterns in both d13C and d15N values.
There is relatively less spread in the values in the earlier millennia, but these times are represented by rather
small sample sizes, especially from the western region.
The d15N plot shows no apparent shift in sources of dietary protein during the period when a reduction in body
4
S. PFEIFFER AND J. SEALY
TABLE 1. Cape ecosystem Holocene skeletons used in this study
13
15
Sex
Laboratory
number for
date
d C (%)
d N (%)
Southern Cape
SAM-AP 6232
ALB 282
NMB 1704
A 1055
UCT 254
SAM-AP 6219(B)
SAM-AP 6219(C)
SAM-AP 4212
SAM-AP 6216
SAM-AP 6219(A)
SAM-AP 4825
SAM-AP 4292
A 1112
SAM-AP 1878(A)
Matjes River skeleton #1
UCT 107
UCT 246
SAM-AP 34
SAM-AP 1889
SAM-AP 1893
ALB 50
NMB 1639
A 1070(A)
SAM-AP 1878(B)
A 1184 (VII)
UCT 345 (NBC)
SAM-AP 5048
NMB 1705(A)
A 1186
NMB 1241(A)
NMB 1440
NMB 1705(B)
SAM-AP 1145
NMB 1241(B)
UCT 347
SAM-AP 1871
SAM-AP 1879
NMB 1271
SAM-AP 3026
SAM-AP 1131
SAM-AP 3021
NMB 1640
UCT 209
A 1184 (VIII)
NMB 1437
NMB 1274
A 1184 (VI)
NMB not accessioned SS2
NMB not accessioned SS3
UCT 206(1)
A 1184 (B)
NMB 1312
UCT 201
UCT 199
UCT 200
UCT 202
SAM-AP 4827
13.3
17.0
12.2
13.2
15.7
13.1
12.9
15.8
13.5
14.5
14.3
16.7
15.3
13.0
13.8
13.3
12.0
15.4
11.5
11.7
11.3
12.2
14.0
12.0
15.6
12.5
12.3
12.9
17.0
12.7
15.0
13.1
12.5
15.0
16.6
12.4
11.1
12.9
12.1
13.5
12.0
12.0
12.8
14.9
13.8
13.6
14.9
13.5
14.6
12.0
15.4
13.2
13.3
14.1
12.0
13.6
13.2
11.4
8.0
9.3
12.4
9.3
12.6
13.8
8.7
13.0
12.1
14.0
8.6
13.3
14.4
13.6
17.9
15.1
12.5
17.5
16.6
16.9
15.9
12.4
15.9
8.4
14.7
16.6
14.4
8.3
13.5
11.4
13.4
15.3
12.9
12.81
16.2
17.5
13.0
16.8
13.2
15.7
16.3
13.2
10.5
13.2
13.2
9.6
13.0
13.9
14.4
11.5
13.8
11.7
12.7
16.3
12.3
14.5
F
F
F
M
M
M
M
F
M
F
F
F
F
M
F
M
F
M
M
M
M
F
M
F
F
F
M
F
F
F
F
X
M
F
M
F
M
F
F
F
F
F
M
M
M
F
M
F
M
M
M
M
F
M
F
F
M
Pta-7924
Pta-6964
Pta-6953
Pta-6958
Pta-6948
Pta-6978
Pta-2284
Pta-6950
OxA-V-2055
Pta-2273
Pta-2283
Pta-6957
Pta-7925
TO-8401
Pta-6595
Pta-6983
Pta-4348
Pta-6973
Pta-6947
Pta-6981
Pta-6974
Pta-6976
Pta-6975
Pta-4367
Pta-6962
Pta-7983
Pta-4226
Pta-3718
Pta-4354
Pta-3724
Pta-6636
Western Cape
SAM-AP 6020
SAM-AP 5035A
SAM-AP 6331
SAM-AP 1863
SAM-AP 6221
UCT 428
UCT 60
SAM-AP 4929
UCT 230
SAM-AP 1247A
15.4
14.4
15.0
10.9
16.1
15.3
14.6
17.1
16.6
15.2
12.4
14.0
14.7
13.8
12.7
14.5
14.1
13.0
13.6
12.8
M
M
F
F
F
M
M
X
M
F
Pta-4189
Pta-4813
TO-8953
Pta-4708
Pta-4356
GX-14815
Pta-2005
Pta-4823
Pta-4736
Pta-4281
Pta-6611
Pta-8580
Pta-6963
Pta-6955
Pta-6820
Pta-6631
Pta-6635
Pta-6642
Pta-6628
Pta-6619
Pta-6607
Pta-6617
Pta-6949
Pta-6592
Pta-6952
Pta-6815
Pta-6812
Pta-6599
Pta-6594
Pta-6613
Pta-8557
Pta-6965
Pta-6960
Pta-2145
Pta-6971
Date (BP)
560 6 50
570 6 50
760 6 50
1080 6 40
1270 6 50
1580 6 60
1700 6 50
1710 6 70
1840 6 60
1880 6 20
2060 6 50
2120 6 50
2160 6 50
2170 6 20
2280 6 60
2290 6 50
2290 6 60
2310 6 25
2310 6 50
2360 6 20
2380 6 45
2590 6 60
2610 6 50
2620 6 35
2750 6 60
Ca. 2500–3000
2780 6 60
2780 6 60
2880 6 60
2970 6 60
3040 6 60
3070 6 60
3210 6 70
3290 6 90
3236 6 33
3310 6 60
3440 6 60
3570 6 50
3980 6 60
4030 6 110
4030 6 60
4120 6 60
4880 6 70
4920 6 60
4940 6 70
5120 6 50
5120 6 70
5370 6 70
5390 6 70
5450 6 70
5960 6 70
5980 6 50
5990 6 70
6180 6 70
7120 6 60
9100 6 90
9720 6 140
620
630
790
800
880
940
955
1040
1110
1180
6
6
6
6
6
6
6
6
6
6
30
50
90
50
50
70
50
50
50
50
Maximum
femur
length
Femur
head
diameter
417
392
453.5
449
399
442
439
403
381
375
426
369
375
437
38
35.5
40
42
40
42
41
37
36
37
39
33
35
42
36.5
40
42
39
42
40
35
37
36.5
38
36
35
36
35
32.5
400
421
393
428
394
349
396.5
389
383
379
374
349
372
387
440
330
435
417
405
400
402
384
395
377
401
451
440
420
378
435
436
420
453
410
432
448
418
457
391
406
431
419
458
443
411
Citation5
34
38.5
39
33.5
42
37
40
36
36
37.5
36
36
44
39
43
39
40
35
41
44
39.5
42.5
37
44
40
41
40
1
This study
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
This study
1
1
2
1
3
5
1
1
1
1
1
2
1
3
2
2
1
5
This study
1
1
4
1
1
1
1
1
1
4
1
5
4
4
4
4
1
42
42
39
38
37
40.5
42
39
44
40
6,7
6
This study
6
6,7
2
2,7
6
6
6,7
5
BODY SIZE AMONG FORAGERS
TABLE 1. (Continued)
13
SAM-AP 4905
SAM-AP 4934
SAM-AP 6075
SAM-AP 5034
SAM-AP 6149
SAM-AP 6050
SAM-AP 5083
UCT 97
SAM-AP 4630
SAM-AP 6041a
UCT 120
SAM-AP 3053
SAM-AP 6083A
UCT 386
SAM-AP 6041b
SAM-AP 5041
SAM-AP 1443
UCT 396
UCT 220
UCT 390
SAM-AP 4305
SAM-AP 4309
Unaccessioned,
North of Cape Town2
UCT 385
SAM-AP 4813
SAM-AP 5082
UCT 394
SAM-AP 4308
SAM-AP 1441
SAM-AP 4720
SAM-AP 4306
SAM-AP 4304A3
UCT 169
SAM-AP 6023
SAM-AP 1157
SAM-AP 4203A3
SAM-AP 4899
Unaccessioned,
North of Cape Town4
SAM-AP 6017
SAM-AP 5075
SAM-AP 6031
SAM-AP 4943
SAM-AP 5095
UCT 427
UCT 445
SAM-AP 5091
UCT 222
UCT 162
UCT 158
SAM-AP 6051
SAM-AP 5040
UCT 373
SAM-AP 4637
SAM-AP 1149
UCT 112
UCT 248
SAM-AP 4203B
SAM-AP 5068
SAM-AP 37
UCT 374
1
15
d C (%)
d N (%)
14.7
14.7
15.0
16.0
14.4
13.9
14.5
11.8
15.4
12.0
13.5
13.6
13.4
16.8
15.7
17.9
11.8
17.7
11.5
17.2
12.5
11.7
13.3
14.0
17.0
17.1
11.3
13.3
12.3
15.0
16.1
11.3
15.8
14.2
13.7
14.0
12.7
10.2
15.5
16.5
14.1
16.3
16.1
15.5
Sex
Date (BP)
Maximum
femur
length
M
M
F
F
M
F
F
M
M
M
F
M
F
M
M
F
M
F
M
M
M
F
M
Pta-4349
Pta-4210
Pta-4186
Pta-4771
GX-13182
Pta-2855
Pta-926
Pta-4828
GX-13178
Pta-4722
Pta-5677
Pta-4411
Pta-4358
Pta-5283
Pta-4768
Pta-4376
Pta-2309
Pta-4965
Pta-5678
Pta-4660
Pta-4385
TO-8952
1210 6 50
1220 6 50
1330 6 40
1390 6 40
1440 6 70
1480 6 50
1490 6 55
1560 6 40
1775 6 80
1800 6 50
1960 6 50
1990 6 50
2000 6 50
2000 6 50
2010 6 45
2010 6 50
2050 6 50
2090 6 60
2100 6 21
ca. 2100
2100 6 45
2120 6 45
2120 6 60
Pta-5281
Pta-4204
Pta-4199
Pta-4964
Pta-4404
Pta-4201
GX-13179
Pta-4350
Pta-4656
Pta-5694
GX-13180
Pta-4217
Pta-4412
Pta-4149
GX-23455
2130
2140
2150
2150
2170
2170
2195
2210
2220
2320
2355
2420
2590
2440
2460
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
60
45
60
70
60
60
80
50
50
50
85
60
50
60
50
406
388
425
373
428
414
414
420
408.5
Pta-4293
Pta-4669
Pta-4814
Pta-4821
Pta-4674
GX-14816
Pta-5617
Pta-4724
GX-13184
Pta-929
Pta-5686
Pta-2969
Pta-4225
Pta-1754
Pta-4803
Pta-4690
Pta-2003
GX-13185
Pta-4798
Pta-4370
Pta-4353
Pta-3086
2490
2530
2560
2610
2660
2670
2720
2830
2830
2880
3190
3190
3570
3835
3880
3970
4445
4730
4760
5680
6120
9750
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
50
60
50
50
70
80
60
50
85
50
60
50
60
50
50
50
50
95
50
70
70
100
390
383
414
363
388
430
400
405
379
379
434
401
376
411
387
413
414
438
390
451
390
16.9
14.9
11.6
17.5
11.8
13.0
12.1
13.3
12.3
13.2
12.2
13.8
10.6
14.2
14.2
16.4
15.9
15.5
15.0
17.3
15.2
15.2
13.5
15.8
15.3
10.8
F
F
M
M
M
M
M
F
M
F
M
F
X
M
F
13.3
10.6
11.6
15.2
13.2
11.8
14.8
14.9
12.6
11.5
13.5
11.9
17.6
14.0
17.2
12.3
11.2
14.2
11.2
14.6
14.9
12.3
14.9
16.7
16.0
12.3
15.7
13.9
16.2
12.8
16.7
16.1
14.5
13.8
11.3
15.3
10.5
14.3
16.9
14.1
15.8
14.6
13.1
17.4
M
M
M
F
F
F
M
M
M
M
F
F
F
F
F
M
M
F
F
M
F
M
13.8
12.1
15.9
Laboratory
number for
date
420
425
407
435
408
396
436
423
432
408
427
391
431
417
401
389
389
424
402
395
413
380
432
403
400
397
Femur
head
diameter
Citation5
41
43
39
40
43
37
39
40.5
40
42
37
41
38
42
41
41
39
35
37
39
43
38
34
6,7
6,7
6,7
6
6,7
8
6,7
6
6,7
6
This study
6,7
6,7
9
6
6,7
6,7
9
6
9
6
6,7
This study
37
37
41
39
42
9
6,7
6,7
9
6,7
6,7
6,7
6,7
6,7
This study
6,7
6,7
6,7
6,7
10
40
40
40
38
39
34
39
39
37
39
34
36
40
36
41
39
40
37.5
33
36
39
44.5
37
36
38
39
47
6,7
6
6
6
6
2
2
6
6,7
6,7
This study
8
6,7
7,11
6
6
6,7
6,7
6
6,7
6,7
6
Correction of previously published value of 10.4 d15N (Sealy and Pfeiffer, 2000).
Excavated April 1998, 17 Moolman St.; Bloubergstrand, single young adult male; archaeological report on file at Department of
Archaeology, University of Cape Town.
3
Correction of previously published dates (Sealy and van der Merwe, 1988).
4
Middle-aged female found with adolescent, Melkbosstrand; excavation and analysis described in Pfeiffer et al., 1999.
5
1, Sealy and Pfeiffer, 2000; 2, Morris, 1992; 3, Inskeep, 1987; 4, Patrick, 1989; 5, Muller, 2002; 6, Sealy, 1989; 7, Sealy and van der
Merwe, 1988; 8, deVilliers and Wilson, 1982; 9, Sealy et al., 1992; 10, Pfeiffer et al., 1999; 11, Sealy and van der Merwe, 1986.
2
6
S. PFEIFFER AND J. SEALY
TABLE 2. Summary statistics for femoral dimensions, southern and western Cape samples
Southern Cape
Maximum Femoral Length
Femoral Head Diameter
Western Cape
Maximum Femoral Length
Femoral Head Diameter
Total
Maximum Femoral Length
Femoral Head Diameter
N
Males
Mean (mm)
SD
N
Females
Mean (mm)
SD
N
Sex indeterminate
Mean (mm)
SD
23
26
418.0
40.2
27.1
2.5
28
29
395.8
36.7
28.2
2.3
1
1
403.0
38.5
35
36
414.6
40.5
22.4
2.5
28
27
403.4
37.5
18.0
2.0
2
39.0
0.0
57
61
416.6
40.4
23.9
2.4
54
54
399.8
37.1
24.1
2.2
1
3
403.0
38.8
0.3
TABLE 3. Estimated stature compared to earliest documented statures of Khoe and San people
N
San (Dart, 1937)
Khoe (Schultz 1928)
Holocene skeletons
White, M and F (Trotter, 1970)
Black, M and F (Trotter, 1970)
White (Fledesman and Fountain, 1996)
Black (Fledesman and Fountain, 1996)
Generic ratio (Feldesman et al., 1990)
Mean of estimates
20
73
58
Males
Mean (cm)
SD
N
Females
Mean (cm)
SD
155.8
162.4
5.51
6.23
19
146.1
5.93
160.4
158.1
157.0
154.5
155.5
157.1
5.76
5.12
7.87
7.23
9.05
152.8
150.9
151.7
149.6
149.4
150.9
5.87
5.42
7.74
7.10
8.89
56
size begins, i.e., 3500–4500 BP. The d13C values, too, do
not show substantial directional change at this time.
However, there is a clear tendency toward lower d15N
values, i.e., less reliance on high trophic level marine
sources of protein, after 1,500 years ago, particularly in
the southern region. The d13C values become more negative during this time, indicating more terrestrial foods in
the diet, especially in the western region.
Both body size variables, sexes pooled, were compared
with both isotopic variables. To explore the possibility
that certain mixes of dietary protein yielded larger or
smaller bodies, values for d13C and d15N were partitioned into the lightest, moderate, and heaviest signals,
roughly corresponding with most terrestrial, mixed, and
most marine dietary intake. Comparisons against femur
length show tendencies for the longest femora to come
from ‘‘mixed’’ diets and the shortest femora to come from
terrestrial diets, but not significantly so. When femoral
head diameter is compared against isotopic value, there is
no apparent pattern. Therefore, more terrestrial, more
marine, or more mixed dietary sources do not correlate
with either aspect of body size when femora are considered collectively.
To integrate the temporal factor into the analysis,
body-size variables were divided into the pre-3000 BP
sample (N ¼ 39) and post-3000 BP sample (N ¼ 88).
Table 4 shows that in the earlier group, both measures
of body size correlate positively with date, and in the
more recent group, body size correlates negatively with
date. Femoral length shows no significant relationships
with d13C or d15N, in either region or time period.
Femoral head diameter shows significant, positive correlations with both d13C and d15N values in both regions
and time periods, although not in a fully consistent fashion. The correlation in the earlier time period is still significant when the largest, oldest data point (UCT 374) is
removed. A higher proportion of marine protein in the
diet leads to heavier isotopic values in both the carbon
and nitrogen metabolic pathways, so correlations with
both d13C and d15N suggest that diets with proportionally more marine protein appear to be associated with
stockier body builds, but there is no evidence for the
type of dietary protein influencing adult stature.
DISCUSSION AND CONCLUSIONS
Archaeologically derived Holocene skeletons from the
Cape Ecozone are generally of a size that is consistent
with the historically documented statures of Khoesan
peoples. The presence in this data set of one large
femoral head with a date early in the Holocene is a
reminder that variability in body size may yet be demonstrated. Many of the earliest radiocarbon dates are
associated with fragmentary skeletons, from which no
femoral measures are available. For example, a fragmentary skeleton from Drury’s Cave (SAM-AP 4208B) has a
date of 9540 6 120 BP (Sealy and Pfeiffer, 2000). Bones
that are complete enough to measure include a very
large metatarsal I. Its maximum length of 70 mm compares to a mean of 54 mm (N ¼ 32, 16 M, 16 F) derived
from other Later Stone Age skeletons of the southern
region. Given the diversity of body size among Middle
Stone Age postcranial remains (Churchill et al., 1996;
Grine, 1998; Grine et al., 1998; Pearson et al., 1998;
Pearson and Grine, 1996, 1997; Pfeiffer and Zehr, 1996;
Rightmire and Deacon, 1991), discussion of the origin
and antiquity of the characteristic Khoesan small stature will continue until more remains are recovered. The
present research demonstrates that by early in the Holocene, the dominant pattern was one of small body size.
This study demonstrated that small-bodied foragers of
the Cape experienced a period of increased variability in
growth, including some very short adult statures,
between ca. 4000 and 2000 BP. Recovery to pre-4000 BP
statures is initiated much earlier than the initiation of
some pastoralism in the region, suggesting that pastoralism does not explain the improved growth characteris-
BODY SIZE AMONG FORAGERS
Fig. 2. a: Maximum femur lengths among Holocene foragers
of southern (plus signs) and western (open circles) regions of the
Cape; curves represent quadratic regression model, solid line for
southern region, dashed line for western region. b: Maximum
femoral head diameters among Holocene foragers of southern (plus
signs) and western (open circles) regions of the Cape; curves represent quadratic regression model, solid line for southern region,
dashed line for western region.
tics. Information from measurements of femoral head
diameter suggests that reliance on relatively more marine protein supported greater body mass, but there is no
significant relationship between femoral length measurements and isotopic indicators of dietary protein sources. If
7
Fig. 3. a: Box plots of maximum femur lengths with southern (open) and western (hatched) foragers of Cape represented
separately. Each solid horizontal band represents sample median. Each box represents the interquartile range, which contains
50% of values. The whiskers extend to highest and lowest
values. Time is divided into thousand-year intervals, beginning
with 500–1499 BP. The interval to the extreme right represents
5500–10,000 BP. b: Box plots of maximum femoral head diameters, with southern (open) and western (hatched) foragers of
the Cape represented separately. Each solid horizontal band
represents the sample median. Each box represents the interquartile range which contains 50% of the values. The whiskers
extend to the highest and lowest values. Time is divided into
thousand year intervals, beginning with 500–1499 BP. The
interval to the extreme right represents 5500 to 10,000 BP.
8
S. PFEIFFER AND J. SEALY
Fig. 4. Maximum femur lengths of skeletons from southern
(plus signs, solid line) and western (open circles, dashed line)
regions of Cape, dating from 3000–500 BP, N ¼ 78.
Fig. 6. a: Scatterplots of d13C (%) for southern (plus signs)
and western (open circles) regions of Cape, from 500 to 10,000 BP.
b: Scatter plot of d15N (%) for the southern (plus signs) and
western (open circles) regions of the Cape, from 500 to 10,000 BP.
Fig. 5. Plot of d15N vs. d13C values (%) for southern (plus
signs, solid line) and western (open circles, dashed line) regions
of Cape.
the reduced statures are linked to diet, explanations
should be sought that focus on the amount of food available rather than a shift in the type of protein being
exploited.
Nutritional insufficiency and the presence of disease are
the two factors most commonly associated with a failure to
achieve predicted adult stature. Surveys of adult and juvenile skeletal remains from southern Africa, pre-2000 BP,
show very little evidence of chronic stress indicators or
infectious diseases (Pfeiffer, 2002, 2005; Pfeiffer and Crow-
der, 2004). The cause or causes of the growth-stunting
remain unknown, but they seem more likely to be based on
diet than on disease, with chronic and/or cyclical insufficiency of nutrients being most probable.
Evidence of Later Stone Age foragers in the southern
and western regions of the Cape is extensive, with good
preservation thanks to shell middens and dry rock-shelter deposits. Throughout the Holocene, population fluctuations are seen as reflecting, in part, climatic shifts
that affected the southernmost parts of the continent
more broadly (Deacon and Deacon, 1999; Sampson, 1974;
Wadley, 1993) Because of hot, dry conditions, the interior
of South Africa was largely depopulated between 8000
and 4000 BP, concentrating human activity nearer the
coastlines (Deacon, 1974).
The extent to which southern and western Cape foragers were interacting across regions is not clear (Binneman, 1996; Hall, 1990; Mazel, 1989; Sealy and Pfeiffer,
9
BODY SIZE AMONG FORAGERS
1
TABLE 4. Pearson correlation coefficients between measures of body size temporal and dietary variables
Pre-3000 south
Maximum femur
N
24
Date
0.405*
0.243
d13C
0.053
d15N
Head diameter
N
27
Date
0.450*
0.140
d13C
15
0.081
d N
Pre-3000 west
Pre-3000 total
Post-3000 south
Post-3000 west
Post-3000 total
11
0.176
0.267
0.476
35
0.364*
0.244
0.149
27
0.545**
0.317
0.052
53
0.510**
0.001
0.071
80
0.524**
0.107
0.064
10
0.711*
0.602
0.765*
37
0.537**
0.334*
0.157
29
0.335
0.464*
0.319
55
0.362**
0.036
0.000
84
0.367**
0.112
0.239*
1
Skeletal sample is divided into pre- and post-3000 BP segments.
* P ¼ 0.05–0.01.
** P < 0.01.
2000; Wadley, 1987); model building tends to approach
the regions as separate phenomena. Inskeep (1987),
working primarily from evidence provided at the stratified southern Cape site of Nelson Bay Cave, argued that
shellfish became a more regular component of the diet
after 4500 BP, and that there was a noteworthy shift
toward less standardized artifacts, greater reliance on
local stone raw material, and intensified exploitation of
local food sources ca. 3300 BP. Hall (1990) noted that, in
inland sites further to the east, people started to consume significant quantities of riverine mussels and fish
from 4300 BP. In both the eastern and western regions,
plant foods became more important in the late Holocene.
Jerardino Wiesenborn (1996) suggested that, in the west,
climate and demographic factors combined to stimulate a
dramatic move to large open-air sites which preserve
huge quantities of marine shell refuse (‘‘megamiddens’’)
in that region, ca. 3500–3000 BP. Also from the western
region, there are isolated instances of interpersonal violence in the third millennium that have stimulated discussion of unusual and unexplained community stresses
(Bahn, 2003; Morris and Parkington, 1982; Morris et al.,
1987; Pfeiffer et al., 1999; Pfeiffer and Van der Merwe,
2004).
Archaeologists working with other lines of evidence
have identified patterns of reduced mobility and group
interaction at around 4500 BP (Binneman, 1996; Hall,
1990; Mazel, 1989). Part of this change involved broaderspectrum food collection, with more reliance on plant
foods, mollusks, and/or fresh-water fish. Shellfish, fish,
and plant foods provide relatively small ‘‘packages’’ of
food, though they are predictable, reliable resources
(Hall, 1990; Parkington et al., 1988). These authors (and
others) suggest that this resulted from the need to obtain
more food to feed more people, i.e., to a nutritional challenge associated with a growing population. This pattern is
also seen in the isotopic values presented here, in which
values from the southern region show some very terrestrially oriented signatures, consistent with some people
having limited access to the coast. In the second half of the
Holocene, foragers incorporated a broader spectrum of
hunted and gathered foods, presumably because there
were more mouths to feed.
Despite variations in climate, topography, and resource availability, the western (fynbos) and southern
(forest) regions may have offered similar challenges to
Holocene foragers. In both regions, sometime prior to
4000 BP, there were changes in living conditions that
compromised human growth. The increased variability
in stature and its subsequent mitigation occur too
broadly and too quickly to reflect genetic change. Isotopic data indicate that the sources of dietary protein were
not changing dramatically, but this evidence does not
address questions of dietary sufficiency or food security.
This study is the first to suggest that the difficulties
occurred across the full expanse of the Cape. There are
suggestions that, in the second half of the Holocene,
Cape foragers were moving toward more control of their
food resources, with the fire-farming of geophytes and
the drying of mollusks being two examples from different
regions. It has been further suggested that such collective strategies could have preadapted some communities
for the adoption of pastoralism, when sheep and goats
were introduced from more northerly parts of the continent. The isotope data indicate that soon after 2000 BP,
there is a gentle shift of dietary protein sources toward
the consumption of more terrestrial (and fewer marine)
protein foods. This would be consistent with some inclusion of domestic stock at that time.
When the temporal shift in femur length was initially
noted, it was characterized as a pre- and post-2000 BP
phenomenon. The arrival of pastoralism was characterized as a solution for communities which were nutritionally challenged. This examination of a larger sample
from a broad geographic region confirms the broad geographic basis of the decline in adult body size, but
demonstrates that an increase in stature predates the
introduction of domesticated stock by about a millennium. Femur lengths increase linearly from 3000 BP
onward. Assuming that the earlier decline in average
femur length represents health difficulties among foragers of the Cape during the period of roughly 4000–3000
BP, the gradual increase in average femur length after
3000 BP represents foragers solving their food security
problem within the framework of a hunting and gathering lifestyle.
The causes and details of the fifth/fourth millennium
dietary problem and its third millennium solution are
not clear at this time. Possible demographic, climatic,
and technological causes should be explored through
diverse lines of evidence. This study outlines the evidence available from the study of adult body size. It also
demonstrates that past foraging peoples were subject to
good times and bad. Studies of past human health across
subsistence transitions, like the origin of agriculture,
tend to characterize the health of hunter-gatherers as
stable through time and space. This study illustrates
that the patterns of growth among individual huntergatherers in the Cape ecozone were far from homogeneous during the Holocene. Within the rubric of hunting
10
S. PFEIFFER AND J. SEALY
and gathering, the interplay of human numbers and natural resources led to challenges, including food
shortages, which in this case were solved without shifting to a new system of subsistence.
ACKNOWLEDGMENTS
The authors thank the curators who provided access
to the collections that form the basis of this research.
They include Lita Webley and Johan Binneman (Albany
Museum, Grahamstown), James Brink (Florisbad
Research Station, National Museum, Bloemfontein),
Kevin Kuykendall (University of the Witwatersrand),
Alan Morris (University of Cape Town), and Graham
Avery (Iziko Museums of Cape Town). Permission for
sampling for radiocarbon dating and isotope assessment
was granted by the South African Heritage Resource
Agency. The work benefited from consultation with Jay
Stock.
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