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Craniofacial variation and population continuity during the South African Holocene.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 134:489–500 (2007)
Craniofacial Variation and Population Continuity During
the South African Holocene
Deano D. Stynder,1,2* Rebecca Rogers Ackermann,1 and Judith C. Sealy1
1
2
Department of Archaeology, University of Cape Town, Cape Town, South Africa
Laboratoire d’Anthropobiologie, Université Paul Sabatier, Toulouse, France
KEY WORDS
South Africa; Later Stone Age; Khoesan; craniofacial morphology
ABSTRACT
We assess craniometric variation in 153
individually dated human crania from South Africa with
the aim of investigating genetic continuity/discontinuity
during the Holocene. Evidence from the archaeological
record is used to pinpoint likely episodes of genetic discontinuity. Craniometric data are then used to assess
the likelihood of genetic change having occurred. Two
periods of possible genetic discontinuity are identified: i)
c. 4,000 BP, when an increase in overall population size,
shifts in site organization and diet, and reduced mobility,
were accompanied by reductions in stature; ii) c. 2,000
BP, when the herding of domesticates and the use of pottery vessels were introduced into the region. Results
indicate that there was a decrease in cranial size and
concomitant size-related changes in craniofacial shape
between c.4,000 BP and 3,000 BP. This was followed
almost immediately by a recovery in craniofacial size
and a return to pre-4,000 BP craniofacial shape at
c. 3,000 BP. This recovery continued gradually, extending
into the herder period without any major shifts in morphology at 2,000 BP. It is suggested that the fluctuations
in craniofacial size/shape were related to changes in
environmental factors. Results obtained are consistent
with long term continuity in South African Later Stone
Age populations during the Holocene. Am J Phys
Anthropol 134:489–500, 2007. V 2007 Wiley-Liss, Inc.
The question of population continuity/discontinuity during the South African Holocene has been the focus of anthropological research for well over a century. It has been
investigated from cultural as well as biological perspectives. Early archaeological studies invariably attributed
the various cultural and subsistence changes apparent in
the Holocene archaeological record to frequent episodes of
population migrations/replacements (e.g., Goodwin and
Van Riet Lowe, 1929). Similarly, early physical anthropological work ascribed Holocene craniofacial variation, particularly variation in cranial size, to population migrations/replacements (Meiring, 1937; Louw, 1960).
Advances in archaeological research over the last 30
years have led to a better understanding of South Africa’s Holocene archaeological record. Archaeologists have
been able to securely establish the chronology and nature of shifts in artefact frequencies and styles, the composition of food remains, settlement patterns, and economic activities. In contrast to previous work, recent
studies generally attribute most of the shifts in the Holocene archaeological record to in situ developments
(Deacon, 1984a,b). The major exception concerns the
means by which herding was introduced into the region
(Smith, 1983; Kinahan, 1994–1995). Interestingly, there
are many similarities between aspects of the Holocene
archaeological record and the cultural records of ethnographically recorded Khoesan peoples, suggesting long
term cultural continuity in southern Africa.
Although the archaeological evidence is largely suggestive of population continuity during the Holocene, it is
not conclusive. The question of population continuity/discontinuity is more directly addressed by an analysis of
the physical remains of Holocene people. Unfortunately,
there has been a dearth of recent research on Holocene
cranial remains, with the last comprehensive studies
published in the 1980s, prior to the availability of substantial numbers of radiocarbon dates for these collec-
tions (Hausman, 1980, 1982, 1984; Braüer and Rösing,
1989). We thus do not know to what extent the Holocene
human skeletal record supports the archaeological evidence for population continuity.
To address this issue, we present the results of a metrical analysis of a large sample (n 5 153) of individually
dated Holocene human crania from South Africa’s west,
south-west, south and south-east coasts and adjacent
coastal forelands (Fig. 1). These coastal regions contain
one of the richest Holocene archaeological and palaeoenvironmental records in southern Africa, and have the
most complete Holocene human skeletal record on the
subcontinent.
C 2007
V
WILEY-LISS, INC.
C
THE HOLOCENE ARCHAEOLOGICAL RECORD
The Holocene archaeological record occupies the last
half of the southern African Later Stone Age (LSA).
Today, we know that the LSA, which commenced at least
20,000 years ago (Mitchell, 2002), is characterized by
several innovations and temporally related shifts in
Grant sponsor: The Wenner-Gren Foundation for Anthropological
Research; Grant number: Gr. 7038. Grant sponsors: The PalaeoAnthropology Scientific Trust and The National Research Foundation of South Africa.
*Correspondence to: Deano D. Stynder, Department of Archaeology, University of Cape Town, Private Bag X3, Rondebosch 7701,
Cape Town, South Africa. E-mail: dstynder@yahoo.com
Received 13 March 2007; accepted 20 July 2007
DOI 10.1002/ajpa.20696
Published online 4 September 2007 in Wiley InterScience
(www.interscience.wiley.com).
490
D.D. STYNDER ET AL.
concordant interpretations of the pre-2000 BP LSA record, there is disagreement about the means by which
herding was introduced into South Africa as well as the
biological identity of the herders. One school attributes
the origin of herding to the migration of genetically distinct herders into South Africa at c. 2000 years BP
(Smith, 1983). The other school proposes that herding
entered the region either by the acculturation of indigenous hunter-gatherers (Kinahan, 1994–1995; Sadr,
1998), and/or via small groups of migrant herders who
were absorbed genetically into the gene pool of pre-existing populations (Elphick, 1985).
THE HOLOCENE HUMAN SKELETAL RECORD
Fig. 1. Map of South Africa, showing region from which crania were derived (enlarged section).
lithic technology, raw material use, and subsistence
activities. Prior to c. 2,000 BP, the archaeological record
indicates that people exclusively practiced hunting-andgathering. There is, however, evidence of two major
shifts in technology and subsistence strategy within this
hunting-and-gathering economy (Deacon, 1984a,b). The
first occurred at c. 12,000 BP, when the early LSA Robberg Industry, a predominantly microlithic industry, was
replaced by the macrolithic Oakhurst Complex. There
was a broadly concurrent shift in food remains, with the
focus changing from large to medium–small and small
animals. The second shift occurred at c. 8,000 BP when
there was a switch back to microlithic stone artefacts in
the Wilton Complex and continued exploitation of small
game animals. Wilton assemblages show some variation
through time, especially shifts in scraper morphology
and the relative frequencies of various types of backed
microliths. Late Wilton assemblages (c. 4,000 BP – colonial period) may show an increase in the use of backed
bladelets and points, the proliferation of adzes in certain
regions, and the introduction of ceramics and domesticates at c. 2,000 BP (Sampson, 1974). From c. 4,000 BP
onwards, there was a marked increase in archaeological
sites, particularly along the coast, suggesting an increase
in overall human population size compared with early or
mid-Holocene times. There is also more regional diversity in stone artefact assemblages: while microlithic artefact-making traditions persisted in some sites, as
described above, people in other areas shifted to making
macrolithic artefacts according to more or less standardized patterns. In the second half of the Holocene, subtle
shifts in site organization, food residues (increased exploitation of small ground game, plants, and marine
foods), and cultural artefacts, together suggest increased
sedentism, accompanied by resource intensification and
greater regional variation in material culture (Mazel,
1989; Hall, 1990; Binneman, 1996; Sealy, 2006). Shifts in
the LSA archaeological record prior to 2,000 BP,
although in many instances quite marked, occurred
gradually, as indicated by periods of transition between
industries (Deacon, 1984a,b).
During the last 2000 years, archaeological evidence
points toward the coexistence of two life ways: huntingand-gathering and herding. In contrast to the relatively
The South African Holocene human skeletal record is
poorly understood as there has never previously been a
comprehensive analysis of a large, well-dated sample of
Holocene crania. Nevertheless, based on a diverse array
of isolated studies, we are able to reconstruct the following scenario.
The sparse c. 12,000 BP–8,000 BP cranial sample suggests that Oakhurst populations resembled later LSA
populations, including historic Khoesan, in terms of facial morphology (Rightmire, 1974; Braüer and Rösing,
1989). These early populations are, however, thought to
differ from later populations in terms of their larger cranial size and higher levels of cranial robusticity (Bräuer
and Rösing, 1989).
The equally sparse c. 8,000 BP–4,000 BP cranial sample suggests that Classic Wilton period populations
exhibited little morphological divergence from Oakhurst
populations (Hausman, 1980; Braüer and Rösing, 1989).
In contrast, evidence from individual case studies of crania dating from c. 4,000 BP–2,000 BP, suggest that there
was a reduction in craniofacial size (e.g., de Villiers and
Wilson, 1982; Inskeep, 1987). Interestingly, Pfeiffer and
Sealy (2006) recorded a decrease in stature levels at this
time, suggesting that there was probably a general
decrease in overall body proportions.
By c. 3,000 BP, stature levels began to recover (Pfeiffer
and Sealy, 2006). Whether there was a concurrent recovery in cranial size was not known prior to the current
study. Little is also known about craniofacial morphology
at the onset of the herding period at c. 2,000 BP. The limited available evidence suggests that there was continuity
in craniofacial morphology (Hausman, 1980, 1982, 1984).
FOCUS AREAS IN THIS STUDY
On the basis of changes in behavior and/or skeletal
evidence, two periods can be identified when genetic discontinuity might have occurred, and when we have sufficient crania to test this hypothesis: i) c. 4,000 BP, when
an increase in overall population size, shifts in site organization and diet, and decreased mobility was accompanied by reduced body dimensions; ii) c. 2,000 BP, when
herding was introduced into the region. Significant
changes in craniofacial morphology at either of these
times may indicate population discontinuity. On the
other hand, a lack of significant change would be consistent with population continuity.
MATERIALS AND METHODS
Only adult crania possessing a full complement of
landmarks (see below) were included in this study, limit-
American Journal of Physical Anthropology—DOI 10.1002/ajpa
491
HOLOCENE POPULATION CONTINUITY
TABLE 1. South African Holocene crania used in this study
Region
West coast
South-west coast
Accession no.
Sex
SAM -AP 1446
UCT 227
UCT 429
UCT 387
UCT 164
SAM-AP 5069
UCT 445
SAM-AP 4931
SAM-AP 4867
SAM-AP 6020
SAM-AP 5035a
SAM-AP 5032
SAM-AP 5012
UCT 60
SAM-AP 6332
SAM-AP 1247a
SAM-AP 4905
UCT 94
SAM-AP 4314
SAM-AP 6075
SAM-AP 4669
SAM-AP 6074
SAM-AP 4920a
SAM-AP 5034
SAM-AP 6334
SAM-AP 6149
SAM-AP 5083
UCT 55
SAM-AP 4630
SAM-AP 4659
SAM-AP 6041a
SAM-AP 4901
SAM-AP 6264
UCT 120
SAM-AP 3053
SAM-AP 5041
SAM-AP 5035b
SAM-AP 1443
SAM-AP 1142
UCT 220
SAM-AP 6260a
SAM-AP 4636
SAM-AP 6313b
SAM-AP 5082
SAM-AP 6313a
SAM-AP 1441
UCT 134
SAM-AP 4942
UCT 436
SAM-AP 4301
SAM-AP 4299
SAM-AP 6043
SAM-AP 4300
SAM-AP 4899
SAM-AP 39
SAM-AP 5070
SAM-AP 4906a
SAM-AP 5095
SAM-AP 4627
SAM-AP 4202
UCT 167
NMB 1827
UCT 162
UCT 421
SAM-AP 6147
SAM-AP 6071
SAM-AP 6317
SAM-AP 4906b
UCT 435
UCT 343
M
M
M
M
F
F
M
M
M
M
M
M
F
M
M
F
M
F
F
F
F
M
F
F
M
M
M
F
M
F
M
M
M
F
M
F
M
M
M
M
F
M
M
M
F
M
M
M
F
F
F
M
F
M
F
F
F
F
F
F
M
F
M
F
M
M
M
F
F
F
Date (BP)
Laboratory no.
Citationa
740
1,000
1,870
2,055
2,360
2,634
2,720
3,750
590
620
620
765
812
950
980
1,180
1,210
1,270
1,319
1,330
1,333
1,360
1,364
1,390
1,400
1,440
1,490
1,680
1,775
1,815
1,824
1,892
1,950
1,960
1,990
2,010
2,011
2,050
2,090
2,100
2,120
2,130
2,140
2,150
2,161
2,170
2,210
2,220
2,240
2,250
2,294
2,295
2,304
2,440
2,448
2,573
2,635
2,660
2,665
2,673
2,695
2,815
2,880
2,895
2,920
2,935
2,970
2,977
2,980
2,985
Pta-9085
Pta-4405
Pta-8814
GrA-23218
Pta-8750
OxA-V-2066-34
Pta-5617
Pta-4827
Pta-4407
Pta-4189
Pta-4401
OxA-V-2056-35
OxA-V-2065-36
Pta-2005
Pta-8767
Pta-4281
Pta-4349
GrA-23216
OxA-V-2066-26
Pta-4186
OxA-V-2056-28
Pta-4148
OxA-V-2059-17
Pta-4771
Pta-8790
Gx-13182
Pta-926
GrA-23075
Gx-13178
OxA-V-2056-43
OxA-V-2056-27
OxA-V-2065-40
Pta-9073
Pta-5677
Pta-4411
Pta-4376
OxA-V-2055-46
Pta-2309
OxA-V-2056-32
Pta-5678
Pta-9069
Pta-4379
OxA-V-2056-47
Pta-4199
OxA-V-2055-44
Pta-4201
GrA-23226
Pta-4829
Pta-8751
OxA-V-2055-40
OxA-V-2065-46
OxA-V-2056-40
OxA-V-2065-37
Pta-4149
OxA-V-2055-43
OxA-V-2056-46
OxA-V-2065-35
Pta-4674
OxA-V-2056-34
OxA-V-2056-25
GrA-23222
GrA-23229
Pta-929
GrA-23217
Pta-8774
OxA-V-2055-42
Pta-8807
OxA-V-2056-48
Pta-5034
GrA-23221
This study
[8]
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6
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6
6
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6
6
6
6
6
6
6
6
6
6
6
6
6
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6
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6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
30
50
35
40
30
28
60
60
45
30
35
25
26
50
50
50
50
40
25
40
25
40
32
40
50
70
50
40
80
29
27
28
60
50
50
50
30
50
27
21
60
45
29
60
30
60
40
45
60
30
29
28
29
60
29
31
29
70
27
29
45
40
50
45
60
32
60
33
60
45
American Journal of Physical Anthropology—DOI 10.1002/ajpa
492
D.D. STYNDER ET AL.
TABLE 1. (Continued)
Region
South coast
Accession no.
Sex
Date (BP)
Laboratory no.
Citationa
SAM-AP 6051
SAM-AP 6319
SAM-AP 6318
SAM-AP 4974
SAM-AP 4298
UCT 112
SAM-AP 6272
SAM-AP 4692b
UCT 262
NMB 1207
UCT 583
UCT 157
A 1153
NMB 1219
NMB 1338
UCT 114
UCT 83
SAM-AP 4180
UCT 582
A 1154
UCT 70
A 1117
SAM-AP 4898
SAM-AP 1260
ALB 244(1)
UCT 75
NMB 1707
NMB 5
SAM-AP 4874
SAM-AP 6213
UCT 109
NMB 83
SAM-AP 4790
ALB 323
SAM-AP 320g
A 1166
A 1152
SAM-AP 1473
A 1127
UCT 78
SAM-AP 278g
NMB 1203
NMB 1204
SAM-AP 4312
A 1114
UCT 107
SAM-AP 34
SAM-AP 1146
NMB 82
ALB 222
ALB 301
SAM-AP 5050
A 1115
NMB 1639
SAM-AP 1878b
NMB 86
SAM-AP 5049
SAM-AP 5048
A 1172
NMB 1242
NMB 1273
NMB 1202
SAM-AP 1128
SAM-AP 1145
NMB 4
SAM-AP 1871
ALB 354
A 1112
SAM-AP 1879
F
F
M
F
F
M
M
M
M
M
M
M
F
M
M
M
M
F
F
M
M
F
M
M
F
F
M
F
M
M
M
M
F
F
M
M
M
M
F
F
F
F
F
F
M
M
M
M
M
M
M
F
M
F
F
F
M
M
F
M
M
M
F
M
M
F
F
F
M
3,190 6 50
3,200 6 35
3,310 6 60
3,363 6 34
3,380 6 33
4,445 6 50
5,830 6 80
c. 12,000
510 6 40
560 6 50
560 6 45
587 6 28
636 6 26
650 6 60
650 6 35
650 6 40
680 6 40
688 6 27
740 6 40
905 6 25
920 6 40
1,060 6 50
1,084 6 26
1,137 6 27
1,180 6 50
1,340 6 40
1,394 6 24
1,423 6 26
1,426 6 29
1,558 6 27
1,590 6 50
1,590 6 40
1,610 6 150
1,620 6 35
1,707 6 27
1,818 6 27
1,850 6 35
1,880 6 60
1,891 6 29
2,145 6 40
2,158 6 28
2,180 6 50
2,210 6 35
2,260 6 170
2,271 6 33
2,290 6 50
2,310 6 25
2,321 6 28
2,335 6 40
2,640 6 60
2,570 6 50
2,580 6 60
2,588 6 28
2,590 6 60
2,620 6 35
2,705 6 40
2,740 6 50
2,780 6 60
2,950 6 40
3,030 6 26
3,050 6 60
3,140 6 50
3,156 6 33
3,210 6 70
3,236 6 33
3,310 6 60
3,340 6 60
3,355 6 45
3,440 6 60
Pta-2969
Pta-8752
Pta-8741
OxA-V-2055-48
OxA-V-2055-41
Pta-2003
Pta-9082
–
GrA-23221
Pta-8755
Pta-8760
OxA-V-2055-45
OxA-V-2065-47
Pta-8804
GrA-23711
GrA-23654
GrA-23072
OxA-V-2056-23
Pta-7178
OxA-V-2066-33
GrA-23074
Pta-8727
OxA-V-2056-37
OxA-V-2066-28
Pta-8587
GrA-23069
OxA-V-2064-53
OxA-V-2064-49
OxA-V-2056-45
OxA-V-2065-39
GrA-23656
GrA-23227
Pta-2163
Pta-8578
OxA-V-2056-24
OxA-V-2056-33 A
Pta-8757
Pta-8773
OxA-V-2066-36
GrA-23241
OxA-V-2065-43
Pta-8783
Pta-8744
Pta-2164
OxA-V-2055-51
Pta-6815
Pta-6599
OxA-V-2065-44
GrA-23228
Pta-8636
Pta-8684
Pta-7927
OxA-V-2065-48
Pta-6965
Pta-2145
GrA-23657
Pta-7934
Pta-7924
GrA-23647
OxA-V-2064-50
Pta-6942
Pta-8801
OxA-V-2055-49
Pta-2284
OxA-V-2064-48
Pta-2273
Pta-8680
GrA-23232
Pta-2283
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(continued)
American Journal of Physical Anthropology—DOI 10.1002/ajpa
493
HOLOCENE POPULATION CONTINUITY
TABLE 1. (Continued)
Region
South-east coast
Accession no.
Sex
UCT 161
SAM-AP 31
SAM-AP 32
NMB 1640
A 1124
A 1139
NMB 1275
UCT 180
SAM-AP 4182
SAM-AP 5055
UCT 156
A 2226
A 2227
ALB 131
F
M
M
F
M
F
M
M
M
M
M
M
M
M
Date (BP)
3,451
3,576
3,754
4,240
4,320
4,800
4,850
6,180
6,811
6,995
10,110
800
1,150
4,700
6
6
6
6
6
6
6
6
6
6
6
6
6
6
26
30
35
70
32
50
60
70
36
50
80
50
50
60
Laboratory no.
Citationa
OxA-V-2064-54
OxA-V-2065-34
OxA-V-2055-47
Pta-8792
OxA-V-2056-42
Pta-8816
Pta-6986
Pta-3718
OxA-V-2056-26
OxA-V-2065-42
GrA-23223
Pta-8728
Pta-8819
Pta-5979
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a
Refers to the source of the radiocarbon date, as follows: 1 (Morris, 1992); 2 (Sealy and Pfeiffer, 2000); 3 (Pfeiffer and Sealy, 2006);
4 (Sealy, 1989); 5 (Sealy and van der Merwe, 1988); 6 (de Villiers and Wilson, 1982); 7 (Morris et al., 2004–2005); 8 (Hausman,
1980); 9 (Patrick, 1989); 10 (Albany Museum register); 11 (Sealy, 2006).
b
A date has been obtained on postcranial bone from SAM-AP 4692, but has not been published, and is not available for citation.
However, reliable sources report that it is c. 12,000 BP.
TABLE 2. Cranial landmarks used in this study
and their descriptions
Landmark no.
Landmark
Landmark description
1
2
B
1/2BN
3
4
5
6
7
8
9
10
11
12
N
NS
PR
D
ZYO
FMO
ZYM
PTP
P
1/2BL
13
14
15
16
17
AST
L
O
BA
BOC
18
19
H
TSP
20
MXT
Bregma
Halfway along
Bregma to Nasion arc
Nasion
Nasospinale
Prosthion
Dacryon
Zygoorbitale
Frontomalare orbitale
Zygomaxillare
Pterion posterior
Porion
Halfway along Bregma
to Lambda arc
Asterion
Lambda
Opisthion
Basion
Basioccipital-sphenoid
synchondrosis
Hormion
Temporal-sphenoid junction
at petrous
Maxillary tuberosity
ing our sample to n 5 153 (Table 1). Adult status and
sex was determined on the basis of a combination of cranial and, when available, postcranial morphological characteristics (Buikstra and Ubelaker, 1994; White and Folkens, 2000). Information about geographic location was
acquired from museum registers (see Stynder, 2006). All
crania analyzed, except one (SAM-AP 4692), have been
securely radiocarbon (14C) dated, many specifically for
this study. Following Sealy and Pfeiffer (2000) and Mitchell (2002), dates were left uncalibrated for consistency
with other discussions of the South African LSA.
Three-dimensional coordinates of 20 cranial landmarks
were recorded on the left side of the cranium (Table 2)
using a MicroscribeTM 3-D digitizer and InScribe-32 software (Immersion Corp., San Jose, CA). Most landmarks
are type I according to the criteria of Bookstein (1991);
four are type II (P, O, BA, MXT) and two are type III (1/
2BN, 1/2BL).
Analytical approaches employed consist of Principal
Components Analyses (PCA) of untransformed (preserving size) linear distances, Mahalanobis distances (D)
calculated from untransformed linear distances, and
Generalized Procrustes Analyses (GPA) of coordinate
data. Male and female crania were analyzed separately
in those analyses where sex-related morphological differences would have influenced results.
PCA was employed to investigate principal differences
in cranial form throughout the Holocene. The computation of the PCs in the PCA was done via the correlation
matrix. Prior to analysis, coordinate data were transformed into linear distances, and a subset of 48 variables
was selected for further analyses (Table 3). These were
chosen to cover all major regions of the cranium without
redundancy.
Mahalanobis distances (D) were calculated between
individual crania (Van Vark and Schaafsma, 1992) using
a covariance matrix calculated directly from the 153 crania which make up the study sample. Means and standard deviations of Mahalanobis distances (D) were calculated for temporally bound categories to measure levels
of homogeneity and heterogeneity between samples.
GPA was used to quantify and visually examine
change in (centroid) size and size-related shape (allometry) through time. In this study, allometry was used in
the sense of Bookstein (1991), where allometry is shape
change that is related to an increase in size. With the
aid of APS software version 2.41 (Penin et al., 2002;
Berge and Penin, 2004), an allometry value for each cranium was calculated using a multivariate regression in
which the independent variable was centroid size, and
the dependent variables were principal components of
shape (PCS) scores calculated from Procrustes residuals.
In this study, enough PCS scores were included in each
analysis to account for more than 90% of shape variance.
The significance of this relationship was then tested
using an F test of significance. The resulting variable of
this multivariate regression is called the ‘‘common allometric shape vector’’ (Penin et al., 2002). Points along this
American Journal of Physical Anthropology—DOI 10.1002/ajpa
494
D.D. STYNDER ET AL.
TABLE 3. Subset of linear distances selected
for further analyses
No.
Distance
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
B-1/2BN
B-N
B-PTP
B-1/2BL
1/2BN-N
1/2BN-PTP
N-NS
N-D
N-ZYO
N-FMO
N-PTP
NS-PR
NS-ZYO
NS-ZYM
PR-ZYM
PR-H
PR-MXT
D-ZYO
D-FMO
ZYO-FMO
ZYO-ZYM
FMO-ZYM
FMO-PTP
ZYM-PTP
ZYM-P
ZYM-MXT
PTP-P
PTP-1/2BL
PTP-AST
PTP-L
P-AST
P-TSP
P-MXT
1/2BL-L
AST-L
AST-O
AST-BA
AST-TSP
L-O
O-BA
BA-BOC
BA-H
BA-TSP
BOC-H
BOC-TSP
BOC-MXT
H-MXT
TSP-MXT
vector were plotted against radiocarbon date to visualize
the magnitude of allometric change through time.
RESULTS
Primary cranial form differences through time
Figure 2 provides PCA plots of male and female samples (geographic regions combined). In both cases, the
most positive values along PC 1 represent relatively
large crania characterized by long/broad faces, prognathic upper-facial regions, and long/low frontal bones. The
most negative values represent small crania characterized by short/narrow faces, retracted upper-facial
regions, and short/high frontal bones. The most positive
values on PC 2 represent crania which exhibit increased
facial dimensions relative to neurocranial dimensions.
The most negative values represent crania which exhibit
reduced facial dimensions relative to neurocranial
Fig. 2. Plot of PC 1 and PC 2 of a principal components
analysis based on (a) male crania and (b) female crania.
dimensions. In both male and female samples, there is
considerable overlap in crania from the selected periods.
Nevertheless, some patterning is discernible. In the male
sample which covers most of the Holocene (Fig. 2a), the
distribution along PC 1 indicates that some of the largest crania in the sample postdate 2,000 BP. Most pre4,000 BP crania are also large. In terms of shape, these
large crania are characterized by long/broad faces, prominent upper facial regions, and long/low frontal bones.
By contrast, most c. 4,000 BP–2,000 BP crania are
smaller, particularly in comparison to post-2,000 BP crania, and are characterized by short/narrow faces,
retracted upper facial regions, and short/high frontal
bones. Similar patterns are discernible in the female
sample (Fig. 2b). Along PC 2, most male post-2,000 BP
crania exhibit positive values, indicating a morphology
characterized by increased facial dimensions relative to
neurocranial dimensions. On the other hand, most pre2,000 BP crania, particularly 4,000 BP–2,000 BP crania,
exhibit negative values, indicating a morphology characterized by facial dimensions that are decreased relative
to neurocranial dimensions. Again, similar patterns are
discernible in the female sample (Fig. 2b).
From the above, it is evident that size and size-related
shape are important aspects of morphological variation
during the Holocene. Below, we investigate these aspects
in more detail.
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HOLOCENE POPULATION CONTINUITY
Fig. 3. Centroid size versus radiocarbon date, post-4,000 BP
(sexes combined). Lines represent linear regression model: solid
line for the south coast, dashed line for the south-west coast.
Cranial size through time
In Figure 3, the centroid sizes of crania (sexes combined) dating to the last 4000 years are plotted against
radiocarbon date. Regression lines are fitted to the samples from the south-west and south coasts (i.e., those
with the best spread through time) to assist with their
interpretation. In the case of the south coast sample, the
regression line (r2 5 0.171, F 5 12.58, P 5 0.001) indicates a subtle; yet significant increase in cranial size.
The south-west coast sample also exhibits a subtle
increase in size; however, this increase is not significant
(r2 5 0.025, F 5 1.66, P 5 0.202). Small sample size during the last 1,000 years (when some of the largest crania
occur in the south coast sample), may account for this
lack of statistical significance. Small sample size also
prevents us from determining whether there was an
increase in cranial size along the west and south-east
coasts. The positions of crania from these regions in the
plot, are however, not inconsistent with a broad geographic increase in cranial size. When we lump crania
from all geographic regions, we find that there is a significant increase in cranial size in the sample as a whole
(r2 5 0.093, F 5 14.322, P 5 0.0002). This increase is
gradual with no fluctuations at 2,000 BP.
Figure 4 displays a combined plot of male and female
centroid sizes against radiocarbon date; this time with
the pre-4,000 BP sample included (all male). Interestingly, many pre-4,000 BP crania are quite large compared with those just postdating 4,000 BP. A MannWhitney U test (P 5 0.029) indicates that there is a significant size difference between crania in the pre-4,000
BP sample and those dating between 4,000 BP and 3,000
BP. Whether large cranial size was the norm prior to c.
4,000 BP cannot be answered conclusively on the basis
of this small pre-4,000 BP sample. Although it is possible
that the sample is biased toward large crania, previous
observations of large cranial size (Rightmire, 1974; Braüer and Rösing, 1989), and indirect evidence of increased
stature levels in early populations (Pfeiffer and Sealy,
2006), suggest that cranial size was probably relatively
large at the time. Nevertheless, the reduction in cranial
size during the mid-Holocene (observable in the males)
should be viewed as preliminary until a substantially
larger pre-4,000 BP sample can be included.
495
Fig. 4. Centroid size versus radiocarbon date, all 153 crania
(sexes combined).
In Figure 5, the timing of the reduction and subsequent increase in cranial size is further explored (geographic regions combined). It is evident that median
male cranial size is lowest between 4,000 and 3,000 BP
(Fig. 5a). In the female sample, which only covers the
last 5,000 years, median cranial size is also lowest at
this time (Fig. 5b). Despite a general increase in cranial
size after 3,000 BP, large standard deviations in centroid
size indicate substantial variation in cranial size during
this period.
Allometric change through time
In Figure 6, the allometric shape vectors are plotted
against radiocarbon date for post-4,000 BP males and
females. In both samples, there are gradual increases in
allometric shape vector values following lows between
4,000 BP and 3,000 BP. In the male sample (Fig. 6a),
regression lines are fitted to the samples from the
south-west and south coasts. The regression statistics
indicate that the increases in allometric shape vector
values are significant in both samples (sw coast: r2 5
0.119, F 5 4.48, P 5 0.04; s coast: r2 5 0.269, F 5 13.22,
P 5 0.001). The regression statistic for the male sample
as a whole is also significant (r2 5 0.15, F 5 13.37, p \
0.001). In the case of the females (Fig. 6b), the increase
in the allometric shape vector values of the south-west
coast sample is not statistically significant (r2 5 0.026, F
5 0.82, P 5 0.37). However, the increase is significant in
the case of the south coast sample (r2 5 0.221, F 5 6.52,
P 5 0.018). As in the males, the regression statistics for
the females as a whole is significant (r2 5 0.08, F 5
5.07, P 5 0.03).
Shape differences associated with the allometric shape
vector are visualized in the sketches adjacent to the
graphs. In both cases, crania with the most positive values along the allometric shape vector are characterized
by projecting upper facial regions, relatively long/broad
faces and long/low frontal bones. Crania with the most
negative values along the allometric shape vector exhibit
nonprojecting upper-facial regions, relatively short/narrow faces and short/steep frontal bones. Most of the
shape changes related to allometry occur in the face.
Apart from minor changes in the frontal (most of which
American Journal of Physical Anthropology—DOI 10.1002/ajpa
496
D.D. STYNDER ET AL.
Fig. 5. Box plots of (a) male centroid size for the last 12,000 years and (b) female centroid size for the last 5,000 years. The vertical line in the centre of the box marks the median of the sample. The length of each box represents the range within which the central
50% of the values fall, with the box edges at the first and third quartiles. The whiskers extend to the highest and lowest values of the
interquartile range. In the male sample, time is divided into two 2,000 year intervals for the pre-5,000 BP sample, four 1,000 year
intervals for the 5,000 to 1,000 BP sample, and a 500 year interval for the post 1,000 BP sample. In the female sample, the asterisk
represents a value falling outside of the lowest value of the interquartile range for the 3,000 to 4,000 BP sample. Time is divided into
four 1,000 year intervals for the 5,000 to 1,000 BP sample, and a 500 year interval for the post-1,000 BP sample.
are influenced by the position of nasion) and posterior
neurocranial height, the neurocranial shape remains relatively stable between small and large crania.
Figure 7 presents a plot of the allometric shape vector
for the entire male sample against radiocarbon date. As
expected, this plot broadly mirrors the pattern of male
cranial size change during the last 12,000 years. On the
basis of the distribution of the individual crania, it is
apparent that prior to c. 4,000 BP and after c. 2,000 BP,
cranial morphology is dominated by long/broad faces,
retracted maxillae, pronounced upper facial regions, and
low/long frontal regions. The neurocranium shifts
slightly upwards in posterior aspects, but length remains
largely constant. Between c. 4,000 BP and 2,000 BP, cranial morphology is dominated by short/ narrow faces,
pronounced maxillae, retracted upper facial regions, and
steep frontal regions. The neurocranium shifts slightly
downwards in posterior aspects, but again, length
remains largely constant.
Interindividual Mahalanobis distances (D)
Table 4 presents the mean interindividual Mahalanobis distances (D) for four temporal categories (regions
combined). In the male sample, the pre-4,000 BP sample
which encompasses the widest temporal span, exhibits
the highest mean interindividual distance (D). In both
male and female samples, the lowest mean interindividual distances (D) occur between 4,000 BP and 3,000 BP,
when cranial size was at its lowest. After 3,000 BP, there
is a gradual increase in mean interindividual distances
in both samples. Overall, the minimal change in these
mean values indicates that levels of craniofacial homoge-
neity did not change much throughout the duration of
the Holocene.
DISCUSSION AND CONCLUSIONS
Results indicate that there was a subtle late mid-Holocene fluctuation in the craniofacial size of South African
LSA coastal inhabitants. Because of the small pre-4,000
BP sample, it is not possible to pinpoint with confidence
when the initial reduction in size occurred, however, it is
most apparent between c. 4,000 BP and 3,000 BP. Reductions in overall cranial size were accompanied by allometric shape changes that primarily affected the shape
of the face and frontal region. Whereas pre-4,000 BP crania exhibited projecting upper facial regions, relatively
long/broad faces and long/low frontal bones, crania dating to between c. 4,000 BP and 3,000 BP were primarily
characterized by nonprojecting upper-facial regions, relatively short/narrow faces and short/steep frontal bones.
By c. 3,000 BP, cranial size began to recover, increasing
in a linear fashion into the last 1,000 years. In addition
to the increase in size, primary craniofacial shape
shifted back to the pre-4,000 BP pattern.
In terms of chronology, the fluctuation in craniofacial
size corresponds almost exactly with a fluctuation in stature levels of LSA people along South Africa’s south-west
and south coasts (Pfeiffer and Sealy, 2006). This correspondence suggests that the observed fluctuation in cranial size was probably part of a general fluctuation in
body size along these coasts. On the basis of a relatively
large pre-4,000 BP postcranial sample, Pfeiffer and Sealy
(2006) were able to fix the initial decline in stature to c.
4,000 BP.
American Journal of Physical Anthropology—DOI 10.1002/ajpa
HOLOCENE POPULATION CONTINUITY
497
Fig. 6. Plots of individual allometric shape vector values for the last 4,000 years versus radiocarbon date. In the graphs, lines
represent linear regression models: solid lines for the south coast, dashed lines for the south-west coast. In the illustrations, grey
represents the mean shape, and black represents shape change relative to the mean. (a) Males: 40 PCS scores (98.78% of total
shape variation) are included in the analysis. The result of the F test for the regression between centroid size (independent variable) and the 40 PCS scores (dependent variable) is significant (r2 5 0.71, F 5 2.40, P < 0.003). (b) Females: 29 PCS scores (96.27%
of total shape variance) are included in the analysis. The result of the F test for the regression between centroid size (independent
variable) and the 29 PCS scores (dependent variable) is significant (r2 5 0.68, F 5 2.09, P < 0.024).
Reductions in craniofacial size during the Holocene are
not unique to South African populations. Similar diachronic trends have been documented in other Holocene
populations around the globe, including Europe (Sardi
et al., 2004), North Africa (Carlson, 1976; Carlson and
Van Gerven, 1977; Sardi et al., 2004), Australia (Brown,
1992), East Asia (Brown and Maeda, 2004), and SubSaharan Africa (Henneberg and Steyn, 1993). However,
this is the first time that a similar reduction, albeit temporary, is recorded in South African Holocene populations.
Interestingly, reductions in cranial and postcranial
dimensions occurred at a time when there was an overall
increase in population size, reductions in mobility levels
and a greater emphasis on group identity amongst LSA
South Africans inhabiting the study region. These
changes were accompanied by a general shift toward
resource intensification that included the consumption of
a broader range of food types (Mazel, 1989; Hall, 1990;
Binneman, 1996). In other parts of the world such as the
Near East, similar processes of resource intensification
had led to the emergence of agriculture (Bar-Yosef and
Meadow, 1995).
It has been suggested that the switch to agriculture
may in turn have contributed to the reductions observed
in human cranial size. In particular, the greater emphasis placed on softer carbohydrate staples and cooked
foods, may have led to a relaxation of stress on the facial
skeleton, causing reductions in cranial size (Carlson,
American Journal of Physical Anthropology—DOI 10.1002/ajpa
498
D.D. STYNDER ET AL.
Fig. 7. Plot of individual male allometric shape vector values (all crania) against radiocarbon date. In the illustrations, grey represents the mean shape, and black represents the shape change relative to the mean. To calculate the allometric shape vector, 45
PCS scores (99.42% of total shape variation) are included in the analysis. The result of the F test for the regression between centroid size (independent variable) and the 45 PCS scores (dependent variable) is significant (r2 5 0.68, F 5 2.09, P < 0.007).
TABLE 4. Means and standard deviations of individual
Mahalanobis distances (D) for four time-bound categories
Male
\2,000 BP
3,000 BP–2,000 BP
4,000 BP–3,000 BP
12,000 BP–4,000 BP
Female
Mean
Std. dev.
Mean
Std. dev.
9.815
9.607
9.474
9.999
1.046
0.823
0.895
0.933
9.709
9.345
8.698
–
0.949
1.107
0.748
–
1976; Carlson and Van Gerven, 1977). Although the process of intensification along South Africa’s coast encompassed some dietary shifts, these were not of the order
seen in early agricultural communities. The process of
mid- to late-Holocene intensification in South Africa did
not result in the emergence of local forms of agriculture.
Instead, the domestic animals associated with herding
and the crops associated with farming had their origins
in regions further north on the continent, and/or on other
continents. These subsistence strategies, and the use of
earthenware cooking vessels, only became a factor in the
South African socio/economic landscape after c. 2,000 BP.
Prior to this time, the types of food available to Holocene
people, as well as methods of food preparation, remained
relatively constant. Although people increased their consumption of marine foods, small animals and plant foods,
these foods were previously consumed. It is thus unlikely
that biomechanical adaptations related to changes in diet
and food preparation would have led to the observed
reductions in craniofacial size.
An increase in disease load and nutritional insufficiencies, results of increased population size and
greater sedentism during the agricultural period, have
also been proposed as possible causes of skeletal size
reductions during the Holocene (Macchiarelli and Bondioli, 1986). It is well known that those cells that are
responsible for the development of dental and skeletal
tissue are easily disrupted by negative environmental
influences when tissue is being formed (Larsen, 2002).
In children, this disruption may lead to arrests in long
bone growth, hampering the achievement of full growth
potential (Ruff, 2002). The effects of negative environmental influences, particularly nutritional insufficiencies on craniofacial form, are less well-known. On
the basis of the studies carried out on nonhuman primates (Pucciarelli et al., 1990), nutritional insufficiencies
may lead to reduced cranial size and changes in cranial
form, particularly with respect to reductions in facial
dimensions. This is similar to the pattern of cranial
change observed in this study.
Despite some evidence for reduced mobility, South
African pre-2,000 BP groups did not reach the levels of
sedentism apparent in early agricultural communities. It
is thus not surprising that LSA skeletons exhibit no
skeletal evidence of the types of infectious diseases that
commonly affected early agricultural communities
(Pfeiffer and Crowder, 2004; Pfeiffer, 2007). Chronic
stress indicators such as cribra orbitalia, porotic hyperostosis, enamel hypoplasia and growth arrest lines do
however occur (Patrick, 1989; Pfeiffer, 2007). These conditions are usually cited as indicators of nutritional and/
or pathogenic stress in past population. According to
Pfeiffer and Sealy (2006), populations living during the
latter half of the Holocene may have been under
increased nutritional stress due to a growth in population and resultant competition for scarce resources. They
suggest further that the observed reductions in stature
may have been due to this increase in nutritional stress
(Pfeiffer and Sealy, 2006).
If nutritional insufficiencies played a role in skeletal
reductions, the reason for the recovery in size at c. 3,000
BP is not clear. Pfeiffer and Sealy (2006) proposed that
the observed increase in stature may indicate that
people solved nutritional problems within the existent
hunter-gatherer economy. Although the nutritional
insufficiency hypothesis has merits, there is currently
little skeletal evidence to support it other than reduction
in size. What is required is a systematic analysis of
chronic stress indicators across the Holocene which
American Journal of Physical Anthropology—DOI 10.1002/ajpa
HOLOCENE POPULATION CONTINUITY
would provide data about dietary-related health through
time.
Even if nutritional insufficiencies are not at the root of
cranial and postcranial size reductions, the likely cause is
probably environmental in nature. It is unlikely that
gene flow caused the observed reduction in craniofacial
size. Gene flow into a population generally increases phenotypic variability (González-Jozé et al., 2007). In our
study sample, mean interindividual Mahalanobis distances (D) for both male and female samples indicate that
morphological variability was at its lowest between 4,000
BP and 3,000 BP, when craniofacial size was at its smallest. Although internal genetic change cannot be discounted at this stage, most of the available evidence suggests that the observed skeletal reductions were plastic in
nature. It is significant that the primary shape change
occurred in the viscerocranium, while the neurocranium
and basicranium—two regions with high levels of heritability—remained relatively stable. This observation is
consistent with previous studies which showed that shifts
in environmental factors are more likely to influence the
viscerocranium while leaving the neurocranium and basicranium relatively unaffected (Wood and Lieberman,
2001; González-Jozé et al., 2005). In contrast to the more
long term consequences that genetic changes, particularly
gene flow, may have on phenotype (Martı́nez-Abadı́as
et al., 2006), skeletal changes due to shifts in environmental factors are more likely to exhibit a quick reversal
if there is a change in circumstances (Van Wieringen,
1986; Eveleth, 1994; Ruff, 2002). Significantly, following
the observed reductions in overall body dimensions, we
see a concurrent recovery in stature (Pfeiffer and Sealy,
2006) and cranial size as well as a shift back to the general craniofacial shape which characterized pre-4,000 BP
populations. The fact that there was a reversion to the
earlier morphological state strengthens the suggestion
that the initial reduction in skeletal size was probably
due to shifts in environmental factors.
In contrast to changes in craniofacial form between c.
4,000 BP and 3,000 BP, there were no dramatic changes
at c. 2,000 BP. Although cranial size was generally
greater, the increase in cranial size did not commence at
2,000 BP but was instead a culmination of the recovery
that began at c. 3,000 BP. The higher mean interindividual Mahalanobis distance (D) values for the post-2,000
BP sample, also appears to be part of a general increase
in inter-individual variation after 3,000 BP. In the light
of studies linking a reduction in craniofacial size to the
consumption of domesticates and cooked food, the fact
that there was no reduction in cranial size at 2,000 BP
is quite surprising, particularly since isotopic data is
consistent with the inclusion of domesticates in diets
(Pfeiffer and Sealy, 2006; Sealy, 2006).
The results of this study favor population continuity in
South African LSA populations during the Holocene.
Although there was a fluctuation in craniofacial form
sometime during the late mid-Holocene, this was likely
plastic in nature and possibly linked in some way to the
process of intensification which occurred at the time.
This study demonstrates that cranial size may fluctuate
within a relatively short time span. Significantly, variation in craniofacial size was often used in early studies
to argue for population change in South African Holocene populations. Our results confirm that variation in
cranial size and facial morphology cannot always be
used as reliable indicators of genetic change in South
African Holocene populations.
499
ACKNOWLEDGMENTS
The authors would like to thank the curators of the
osteological collections at Iziko Museums of Cape Town,
the Department of Human Biology at the University of
Cape Town, the Albany Museum in Grahamstown, the
National Museum in Bloemfontein, and the Department
of Anatomical Sciences at the University of the Witwatersrand, for facilitating access. The South African Heritage Resources Agency (SAHRA) provided the permits to
date the crania used in this study. Clark Larsen and two
anonymous reviewers provided useful comments which
improved this paper.
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