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Basicranial anatomy of Plio-Pleistocene hominids from East and South Africa.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 59157-174 (1982)
Basicranial Anatomy of PlioPleistocene Hominids From East
and South Africa
M.C. DEAN AND B.A. WOOD
Department of Anatomy, The Middlesex Hospital Medical School, Cleuehnd
Street, London WlP 6DB
KEY WORDS
Basicranium, Hominids, Systematics
ABSTRACT
The results of a metrical analysis of the basicraniurn of 19 PlioPleistocene fossil hominid crania are presented. The sample includes crania attributed to Australopithecus africanus, Australopithecus boisei, and robustus, and
Homo erectus as well as crania whose attribution is still under discussion. These
results confirm significant differences between the cranial base patterns of the
“gracile”and “robust”australopithecines and the three crania attributed to Homo
erectus have a pattern which resembles that of modern humans. None of the crania
examined from East Africa sites have base patterns which resemble that of the
“gracile” australopithecines. The crania KNM-ER 407 and 732 have patterns
which are compatible with them being smaller-bodiedfemalesof Australopithecus
boisei; KNM-ER 1470 and 1813 have base patterns which most closely resemble
that of Homo erectus. The cranial base pattern of KNM-ER 1805 is compatible
with its inclusion in either Australopithecus boisei or Homo. When account is
taken of the immaturity of Taung, the evidence of its cranial base pattern suggests that if it had reached adulthood it would have resembled the “gracile”australopithecine crania from Sterkfontein and Makapansgat.
In an earlier paper we established a comparative metrical framework for assessing the
significance of variations in the basicranial
morphology of early hominid crania (Deanand
Wood, 1981a). Linear and angular measurements were made with the basicranium viewed
in normal basilaris, and special emphasis was
given to establishing the relative positions of
important bilateral landmarks. In this way we
hoped to be able to integrate what was already
known about cranial base flexion in the sagittal plane, with changes in the position and
orientation of bilateral structures.
The results of the comparativestudy showed
a remarkably consistent pattern of cranial
base morphology in the three pongid taxa. The
basicrania of Gorilla, Pongo, and Pan are relatively long and narrow with sagittally orient&ed petrous temporal bones and a posteriorly
situated foramen magnum. In contrast, the
cranial base in Homo sapiens is broad but compressed anteroposteriorly. The foramen magnum is more anteriorly situated and the petrous temporal bones are shorter and orientated
with their long axes more aligned to the cor-
0002-WW6902-0157
$&.OO
0 1982 ALAN R. LISS, INC.
onal plane (Fig. 1).Four hominid crania were
examined in the earlier study, all belonging to
the genus Australopithecus. Sts 5 and MLD
37/38 represented Australopithecus africanus,
the “gracile”australopithecine, and KNM-ER
406 and OH 5 sampled one of the species of
“robust” australopithecine, Australopithecus
boisei.
All four hominid crania retain a relatively
long cranial base and the overall morphology
of the two “gracile” crania is remarkably
pongidlike. However, the anteriorly positioned
foramen magnum and the more coronally orientated petrous bones of KNM-ER 406 and
OH 5 are reminiscent of the basicranial morphology of Homo sapiens.
In this second part of our investigation we
have enlarged the fossil sample from 4 to 19.
This has not only allowed us to make modest,
but important, increases in the sample sizes of
the two australopithecine taxa, but the larger
sample also includes three crania attributed to
Homo erectus. Most of the remaining fossil
W i v e d July 6,1981; accepted April 26,1982.
158
M.C. DEAN AND B.A. WOOD
A
B
Fig. 1. Skull base diagrams of A. Homo sapiens and B.
Gorilla with the landmarks used to define measurements
employed in this study. A full list of the landmarks together
with the measurement definitions are given in Dean and
Wood (1981a).
hominid crania have either been attributed to
taxa for which there are no other good examples of basicranial morphology, e.g., OH 24;
or they have been proposed as examples of
smaller-bodied females of the “robust” australopithecines, e.g., KNM-ER 407 and 732
(Leakey, 1974; Wood, 1978); or they are crania
whose affinities are sufficiently enigmatic for
their taxonomic attribution to have been the
subject of debate, e.g., KNM-ER 1470 and
1813 (Leakey, 1974; Leakey, et al., 1978;
Walker, 1976 and Wood, 1976) and SK 847
(Clarke et al., 1970; Clarke and Howell, 1972;
Wolpoff, 1971). Two other specimens from
South African sites have been treated separately: TM 1517 from Kromdraai because it is
fragmentary, and the skull from Taung because of its immaturity.
The aims of this new study are to test
whether, with a larger sample, it is still possible to discern two patterns of basicranial morphology in Austrulopithecus; to establish
whether Homo erectus crania show a consistent pattern of cranial base morphology, and to
attempt to clarify the taxonomic affinities of
the individual fossil crania by comparing their
basicranial shape with that of established
fossil groups. A comparative metrical framework for this analysis was established in an
earlier paper (Dean and Wood, 1981a). This
paper should be consulted for the detailed
results of the measurements taken on the samples of extant taxa, but they are presented in a
summarized form in Table 3. The problems of
simultaneously comparing a series of variables
are often solved by resorting, with little or no
preliminary analysis, to sophisticated multivariate analytical techniques. However, although such an analysis is in progress, we are
conscious that it cannot replace, and should
always be preceded by, careful inspection and
consideration of the original data. It is, of
course, possible to employ simpler bivariate
statistical tests of affinity, but we considered
that the small sample sizes of the fossil groups
in this study seriously limited the usefulness
of such tests. We decided therefore, to present
the data for each specimen separately, and to
159
BASICRANIUM OF PLIO-PLEISTOCENEHOMINIDS
base our interpretations on observed patterns
of overall cranial shape, and on variables which
were shown to be useful discriminators in our
earlier comparative study. We believe that
metrical data like these, from a region such as
the cranial base, and collected from extant
taxa as well as fossils could be used as part of a
cladistic analysis of fossil hominids, and such a
study, based on a series of cranial morphological features, is in progress.
MATERIALS AND METHODS
Nineteen fossil crania were examined and
measured, and except for two specimens, OH 5
and OH 9, all the observations were made on
the original fossils. The reference numbers of
the specimens, their site of origin, and the location of the original specimens are given in
Table 1. All specimens are judged to be adult,
except for SK 47 which is sub adult, and Taung
which is juvenile. Details of the comparative
extant samples of Homo, Gorilla, Pongo, and
Pun are given in Dean and Wood (1981a).
The fossils are not equally well preserved.
Five of the specimens used in the study (KNMER 406, KNM-ER 3733, KNM-ER 3883, OH 5,
and Sts 5) are sufficiently well preserved to
identify the landmarks, and it was possible to
make reliable estimates of all the measurements on these crania. Three crania (Sts 19,
MLD 37/38, and SK 47) are damaged in the left
or right infratemporal region and the position
of the damaged infratemporal crest was derived by doubling the distance from the preserved side to the midline. Five specimens
(KNM-ER407,1470,1805,1813, and OH 9)are
damaged in the region of the basioccipital and
the foramen magnum; interpretation of the
base of KNM-ER 1470 is further complicated
by distortion. Where landmarks are preserved
on one side of the cranium, measurements were
derived by doubling the distance to the midline. The cranial base is particularly poorly preserved in KNM-ER 1470; nonetheless, by using the preserved margins of the both foramina
ovale and the right carotid canal, and part of
the right petrous temporal bone, it is possible
to estimate the distances FOIFO, CCICC, and
the angle a.The positions of the infratemporal
crest (preserved on the right) and the lateral extremities of the tympanic plate can also be
estimated.
The cranium OH 24 was found in a badly distorted state embedded in a mass of calcareous
matrix, but many of the pieces of the crushed
skull have been separated, cleaned, and reassembled (Leakey et al., 1971). Despite this extensive and skilled attempt at restoration the
cranium remains slightly distorted and the
right petrous and tympanic plate are still in
their crushed position. There has also been
some reconstruction of the lateral end of the
left tympanic plate.
TABLE 1. Details o f the Plio-Pleistocene hominid fossils used in the studv
Site of origin
Koobi Fora
(East Rudolfl,
Kenya
Olduvai Gorge,
Tanzania
Sterkfontein
South Africa
Swartkrans,
South Africa
Makapansgat,
South Africa
Kromdraai
South Africa
Taung
South Africa
Museum or Collection
registration number
Institution where
original is kept
KNM-ER 406
KNM-ER 407
KNM-ER 732
KNM-ER 1470
KNM-ER 1805
KNM-ER 1813
KNM-ER 3733
KNM-ER 3883
OH 5 (Cast)
OH 9 (Cast)
OH 24
Sts 5
Sts 19
Sts 25
SK 47
SK 847
MLD 37/38
National Museum, Nairobi
National Museum, Nairobi
National Museum, Nairobi
National Museum, Nairobi
National Museum, Nairobi
National Museum, Nairobi
National Museum, Nairobi
National Museum, Nairobi
National Museum, Dar es Salaam
National Museum, Nairobi
National Museum, Nairobi
Transvaal Museum, Pretoria
Transvaal Museum, Pretoria
Transvaal Museum, Pretoria
Transvaal Museum, Pretoria
Transvaal Museum, Pretoria
Department of Anatomy,
University of Witwatersrand,
Johannesburg.
Transvaal Museum, Pretoria
TM 1517
Taung 1
Department of Anatomy,
University of Witwatersrand,
Johannesburg.
160
M.C. DEAN AND B.A.WOOD
Five crania (KNM-ER 732, Sts 25, SK 847
TM 1517, and Taung) are well enough preserved on one side only. Where any two or more
of the following midline landmarks were preserved (prosthion, staphylion, hormion, sphenobasion, basion, opisthion, or opisthocranion),
or where there were other indications of the
sagittal plane, these were used to construct a
midline, and measurements and the positions
of landmarks were derived by doubling the distance from the midline to the intact side. In
some cases the position of a landmark was inferred from the topography of nearby preserved bone.
All landmarks whose positions have been derived by reconstruction, or by mirror imaging,
are indicated by broken lines in Figures 2-6,
and measurements involving these landmarks
are put in parenthesis in Table 2. I t must be
emphasized that measurements taken on crania reconstructed in the ways outlined above
are inherently less reliable than those made on
intact specimens. However, it is our belief that
in most cases the measurements made using
reconstructed landmarks are accurate to
within a few millimetres, and it is on this basis
that these specimens were included in the
study. Nevertheless, the results should be interpreted with this important caveat in mind.
The landmarks and the measurements are
those that were listed and defined in Dean and
Wood (1981a),and the landmarks have been located in a diagram (Fig. 1)of two crania seen in
normal basilaris, one Homo sapiens and the
other a Gorilla. Measurements were made on
three occasions and the mean values were recorded. In cases of doubt the measurements
were checked by a second observer. Definitions
of the landmarks and details of the measurement technique are given in Dean and Wood
( 1981a).
surements of Homo sapiens and Gorilla have
been included at the top of each figure for comparison. The results will considered in the four
groups used in Table 2.
1. Adult fossils attributed to Australopithecus africanus (Sts5, Sts 19, Sts 25, and MLD
37/38) (Figure 2.) The skull base diagrams of all
these fossils more closely resemble the pattern
of the pongid samples than they do the modern
human sample. Overall they are small and the
bilateral structures all lie relatively close to the
midline, except for the carotid canals which are
relatively further apart than those of the
pongid taxa. Compared to the pongids the
length of the cranial base as a whole is reduced
in Sts 19,25, and MLD 37/38, but rather less so
in Sts 5, and the foramen magnum is closer to
the bitympanic line. The bitympanic width is
also reduced in comparison with the pongid
taxa, but not to the extent it is in the modern
Homo sapiens sample; this may be related to a
reduction in the size of the masticatory apparatus (Dean and Wood, 1981a).The petrous axes
are aligned more sagittally than in the modern
Homo sapiens sample, but slightly less so
than in the pongid taxa. The range for a in the
“gracile”australopithecines is 59“ to 72”. This
is comparable to the distribution of values in
Pongo ( a = 60”-71”), but it is at the lower end
of the ranges of the Pan and Gorilla samples.
The orientation of the tympanic axis is very
similar to that in the pongid taxa but the range
within the “gracile” group also overlaps the
range of the modern Homo sapiens sample.
The more sagittal orientation of the petrous
axis is the most notable contrast between the
cranial base patterns of the “gracile”australopithecines and modern Homo sapiens, and the
shortened tympanic plates and reduced skull
base length of the “gracile”australopithecines
are features which contrast with the pattern
typical of the pongids.
RESULTS
2. Fossils attributed to Australopithecus
The results of the 15 linear measurements boisei and robustus (KNM-ER 406, OH 5, and
and the two angles for the fossil sample are SK 47) (Figure 3). The cranial base diagrams of
given in Table 2. The crania are grouped in the this group of fossils are distinct from the patTable according to accepted taxonomic attri- tern of both the comparative pongid sample
bution, or treated as individual specimens, as and the “gracile”australopithecines, and they
outlined in the introduction. The parameters share some features of the cranial base pattern
for the comparative samples are given in Dean of the modern Homo sapiens sample. Overall
and Wood (1981a. Table 2); the measurement the cranial base of this group is large and parorder has been revised in the present paper to ticularly wide across the lateral extremities of
separate those which relate to basicranial the tympanic plates. The cranial base is
width from length measurements. The results shorter than that of the pongids and the anterare also presented in a series of two-dimension- ior margin of the foramen magnum is characal plots (Fig. 2-6). In each of these plots the teristically positioned well in front of the
mean values for the linear and angular mea- bitympanic line, unlike either the modern
48
44
44
47
58
57
43
61
54
57
40
(41)
(45)
(51)
44
49
(46)
(41)
(36)
48
50
42
47
65
59
51
58
57
67
53
136)
(60)
55
48
48
(44)
(50)
(38)
60
160)
70
(65)
60
61
(62)
(61)
(44)
50
(49)
50
50
61
64
(54)
66
69
76
72
78
72
(63)
67
73
69
85
83
78
89
84
92
69
68
28
114)
(25)
(20)
(20)
23
(21)
(18)
(15)
26
25
(26)
25
29
29
22
35
31
30
29
(26)
25
29
28
29
27
27
35
31
29
27
(30)
29
28
25
32
26
20
32
26
24
22
25
33
(30)
19
32
30
23
(35)
9
23
22
19
26
34
36
25
29
29
25
(25)
24
(20)
20
20
(21)
(14)
19
23
24
30
24
20
26
25
28
19
24
20
'All linear measurements in mm.
'All angles to the nearest degree.
'Measurements in parentheses are either estimates, or, in the case of widths, were derived by doubling measurements taken on one side only.
Sts 5
92
66
Sts 19
89
63
Sts 25
(90P
MLD37138
97
62
Australopithecus
KNM-ER406
132
79
(Paranthropus)
OH5
122
72
boisei and robustus SK47
100
68
Homo erectus
KNM-ER3733 120
79
KNM-ER3883 115
78
OH9
123
84
Individual fossils
KNM-ER407
102
64
KNM-ER732 (100)
KNM-ER1470 131
KNM-ER1813 103
66
KNM-ER1805 118
72
OH24
(103)
67
SK847
(98) (60)
TM1517
(118) 66
TAUNG
54
Fossil Groups
Australopithecus
africanus
Lengths
(16)
(16)
(25)
(22)
21
22
29
(28)
12
27
24
30
25
20
47
50
(56)
44
60
45
49
63
29
53
41
(43)
47
57
60
45
55
55
60
(27)
24
42
28
31
35
18
24
28
22
37
24
30
31
23
30
28
30
49
45
(56) (45)
45 46
(58) 46
38 54
41 50
(57) 42
32 50
40
95
112
108
110
115
90
112
105
(105)
50 65 103
39 59 98
(40) 72 104
48 60 93
53 44 100
55 45 102
37 45 107
57 48 104
52 55 105
60 50 108
Angles'
IT/IT- FOIFO- IT/IT
TP-TP SP-SP CC-CC FO-FO IT-IT SM-SM AP-AP FM-FM BS-OP TP-CC CC-PA SB-BS TPITP TPlTP -BS a
p
Widths
Table 2. Cranial base data for hominid fossils'
rn
8
5
0
5:
5
M
c,
F
1
0
*
G
0
"1
cd
k
d
3
:
*m
162
M.C. DEAN AND B.A. WOOD
TABLE 3. Cranial base data for comparative groups
Widths
Comparative groups
Homo sapiens
X
n = 30
S.D.
cv
Range
Pan troglodytes
n = 30
Gorilla gorilla
n = 30
-
X
S.D.
cv
Range
X
S.D.
cv
Range
Pongo pygmaeus i7
n = 30
S.D.
cv
Range
TP-TP
SP-SP
CC-CC
FO-FO
IT-IT
SM-SM
PA-PA
FM-FM
99.2
3.8
3.8
91-108
78.0
3.8
4.9
71-86
56.8
3.4
5.9
50-63
49.4
2.8
5.7
45-54
65.5
4.7
7.3
57-77
81.6
4.2
5.2
72-90
32.2
3.4
10.4
27-39
29.9
2.1
7.1
25-33
101.8
6.3
6.2
92-118
59.1
3.6
6.1
54-66
42.5
2.8
6.4
38-48
42.9
2.8
6.4
37-48
51.0
3.3
6.5
42-59
63.6
3.6
5.6
58-69
24.0
2.4
10.6
19-28
23.4
1.9
8.3
20-28
125.9
9.0
7.1
112-143
65.4
3.8
5.6
60-72
47.5
4.5
9.5
40-57
51.3
3.2
6.2
42-57
59.1
4.1
6.9
51-67
79.8
5.2
6.5
72-89
30.4
2.8
9.4
23-34
28.6
2.2
7.8
24-35
114.7
11.2
9.8
93-136
68.2
5.5
8.1
55-77
49.5
3.8
7.6
41-57
45.3
3.6
7.8
41-54
51.5
5.1
9.8
41-63
76.1
5.4
7.1
65-86
28.8
2.7
9.3
23-35
24.4
2.1
8.6
21-31
Angles
-
Lengths
BS-OP
TP-CC
CC-PA
SB-BS
ITIITTPlTP
35.0
2.7
7.8
29-40
21.9
2.1
9.7
18-25
18.6
2.3
12.0
16-23
25.5
2.8
11.2
20-31
47.3
4.1
8.8
41-55
25.9
2.2
8.3
22-32
48.0
4.7
9.7
40-58
45.7
4.8
12.7
31-55
107.0
3.3
3.1
102-114
28.2
2.9
10.1
22-34
31.8
2.9
9.2
26-36
24.0
2.5
9.3
20-29
26.3
2.4
9.1
20-31
48.8
4.2
8.6
42-56
24.6
3.3
13.2
19-29
51.8
3.8
7.4
41-58
68.9
5.3
7.7
60-78
96.0
4.9
5.1
86-105
31.8
2.5
7.9
28-37
40.5
3.5
8.6
34-46
29.8
3.1
10.2
20-37
31.4
3.1
9.9
25-36
61.6
6.6
10.6
52-75
31.3
3.8
12.1
25-38
68.3
7.8
11.4
52-85
71.9
4.9
6.8
60-81
95.0
3.0
3.2
88-102
30.7
3.0
9.8
24-35
34.7
3.8
10.9
29-41
28.4
2.9
10.3
23-35
30.2
4.4
14.5
21-37
57.9
6.7
11.6
43-70
28.0
3.7
13.1
21-38
61.5
7.1
11.6
45-75
68.1
3.0
4.4
60-71
101.0
4.5
4.4
93-108
humans or the “gracile”australopithecines and
the pongids. The orientation of the petrous
axis closely resembles that of the modern
human sample and values for this angle in the
“robust”group of fossils fall within the middle
of the range, a! = 31” - 55”, for modern Homo
supiens. The orientation of the tympanic axis
also resembles that of the modern human
group, but values for 0 for the “robust” group
fall towards the lower end of the modern
human range. In this particular fossil group
this is almost certainly due to the long tym-
FOIFOTPITP
ITIlT
-BS
a
P
panic plates reducing the value of /3. The bicarotid canal width (CC-CC)in KNM-ER 406
and OH 5 exceeds the modern human mean
value, but whereas in the Homo sapiens sample the bi-infratemporal fossa width (IT-IT)is
disproportionately wider than the other bilateral landmarks this is not the case in the “robust” group. There appears to have been little
reduction in the length of the body of the
sphenoid bone in the “robust”crania (if spenoid
length is taken as equivalent to the distance
ITIIT-FOIFO).
163
BASICRANIUM OF PLIO-PLEISTOCENEHOMINIDS
x
X
X
X
GoriIla
Homo
X
X
x
x
Sts 19
Sts 5
X
x
Sts 25
+
X
,1(
MU) 37138
I
All diagrams are to scale and are reduced to 40% of life size
II
Landmarks in Homo and
correspond to their mean value
I I I Where the position of a landmark can only be estimated, or is derived
by mirror imaging the preserved side, it is marked by a broken line
Fig. 2. Skull base diagrams of fossils attributed to Australopithecus africanus, with Homo and Gorilla for comparison.
164
M.C. DEAN AND B.A.WOOD
X
X
X
X
Homo
-
Gorilla
X
X
X
x
OH 5
KNM-ER 406
X
x
SK 47
I
All diagrams are to scale and are reduced to 40% of life size
II
Landmarks in Homo and G
M correspond to their mean value
I I1 Where the position of a landmark can only be estimated. or is derived
by mirror imaging the preserved side, it is marked by a broken line
Fig. 3. Skull base diagramsof fossils attributed to Austmlopithecus boisei and robustus, withHomo and Gorilla for comparison.
165
BASICRANIUM OF PLIO-PLEISTOCENE HOMINIDS
X
X
X
X
Homo
-
Gorilla
X
X
X
KNM-ER 3733
X
KNM-ER 3883
X
X
OH 9
I
All diagrams are to scale and are reduced to 40% of life size
II
Landmarks in Homo and Gorilla correspond to their mean value
I II
Where the position of a landmark can only be estimated, or is derived
by mirror imaging the presented side, it is marked by a broken line
Fig. 4. Skull base diagrams of fossils attributed to Homo erectus, with Homo and Gorilla for comparison.
166
M.C. DEAN AND B.A. WOOD
X
X
X
KNM-ER 407
KNM-ER 732
x
X
0
I
II
x
0
All diagrams are to scale and are reduced to 40% of life size
Landmarks in w a n d Q&Jg correspond to their mean value
i I I Where the position of a landmark can only be estimated. or is derived
by mirror imaging the preserved side,it is marked by a broken line
Fig. 5. Skull base diagrams of individual Plio-Pleistocenehominid crania from East Africa, with Homo and Gorilla for
comparison.
167
BASICRANIUM OF PLIO-PLEISTOCENEHOMINIDS
X
X
X
X
Gorilla
-
Homo
X
0
X
8
,
’.-__-’
‘
SK 847
OH 24
\ i
X
.4
r:
U
0
x
0
TM 1517
s
,
.-.
L.’
Taung
I
All diagrams are to scale and are reduced to 40% of life size
II
Landmarks in Homo and Gorilla correspond to their mean value
II I
Where the position of a landmark can only be estimated. or i s derived
by mirror imaging the preserved side,it is marked by a broken line
Fig. 6. Skull base diagrams of individual Plio-Pleistocenehominid crania from South Africa, with Homo and Gorilla for
comparison.
168
M.C. DEAN AND B.A. WOOD
KNM-ER 1470 and KNM-ER 1813
3. Fossils attributed to Homo erectus (KNMER 3733,3883, and OH 9) (Figure 4).The craniThe cranial base patterns of these two fossils
al base pattern of this group of fossils resem- resemble those of the Homo erectus group. The
bles that of modern Homo sapiens. They differ foramen magnum lies on the bitympanic line,
from the “robust” group and resemble the Ho- the angle p is low and the angle a high. The
mo sapiens sample in two ways. The first is the sphenoid bone is foreshortened, but wide
position of the foramen magnum which lies ap- across the infratemporal fossa at the base of
proximately in the bitympanic line, and not the greater wings. Two obvious differences bewell in front of it as it does in the “robust” tween these two crania are the relatively short
australopithecine fossils. The second is the sphenoid, and the smaller overall size of KNMbroadening of the sphenoid so that the bi-infra- ER 1813.
temporal fossa width exceeds the biforamen
ovale and bicarotid canal widths; this trend is,
KNM-ER 1805 and TM 151 7
however, not as marked in Homo erectus as it
These fossils combine a relatively long craniis in Homo sapiens. The petrous axis in Homo
erectus is more sagittally orientated than in al base and a foramen magnum which is posiHomo sapiens and the “robust”australopithe- tioned well in front of the bitympanic line. The
cines, with the values of a falling at the upper values for angle 0 are high, with the tympanic
end of the Homo sapiens range; in contrast the axis being orientated further forwards than it
values for /3 fall towards the bottom of the is in modern Homo sapiens, but the angles a
range for Homo sapiens. This is partly due to are comparable to the mean values for both the
the relatively long tympanic plates in Homo “robust” australopithecines and the modern
erectus which have the effect of reducing the Homo sapiens sample. The presumed position
value of /3 and accentuating the difference be- of the spheno-occipitalsynchondrosis in KNMER 1805 is some way behind the line connecttween the tympanic and petrous axes.
4. Individual Plio-Pleistocene hominid cra- ing the posterior margins of the foramen ovale,
nia (KNM-ER 407,732,1470,1805, and 1813, thus suggesting that the sphenoid body may
OH 24, SK 847, TM 1517, and Taung) (Figures have been longer than indicated by the ITIIT5 and 6). For convenience these fossils have FOIFO distance. Apart from the relatively
been divided into four groups on the basis of si- high value for angle (3, and a marked narrowing
milarities in cranial base patterns; no other im- of the biforamen ovale width (a feature which
plications are intended at this stage and the they share with KNM-ER 732), the basigroupings are not intended to be taxonomic crania of these two specimens are similar in
shape to those attributed to the “robust”
units.
australopithecines.
KNM-ER 407, KNM-ER 732, OH 24, and SK
Taung
847
Developmentally,
this juvenile specimen is
This group of fossils shows a combination of
features, some of which are characteristic of not directly comparable with the other fossil
the “robust” australopithecines, and others of hominids used in this study, and for this reawhich are seen in Homo erectus and in the son it has been considered separately. It is posmodern Homo sapiens sample. The position of sible that gingival eruption of the permanent
the foramen magnum relative to the bitympan- first molars has occurred in this skull but it is
ic line resembles that in the “robust” sample, unlikely that the first permanent molars were
but the relatively wide and foreshortened sphe- in functional occlusion. This corresponds to a
noid bone are features of Homo. The value of a developmental age of about 6 years in modern
for the orientation of the petrous axis is 49“ in man and to about 3 years 4 months in the three
KNM-ER 407,45” in KNM-ER 732,54” in OH great apes (Dean and Wood, 1981b).A separ24, and 50” in SK 847. With the exception of ate study on the growth of the hominoid craniKNM-ER 732, these values are slightly higher al base in norma basilaris (Dean, in preparathan those for the “robust” sample, and within tion) suggests that at the time of eruption of
the range for Homo erectus. There is little to the first molars the angles a and p would not
distinguish between the cranial base patterns yet have reached their adult values. Of all the
of these four fossils except that the petrous measurements used in this study, only the
angle and the sphenoid lengthlwidth ratio of length and width of the foramen magnum of
OH 24 and SK 847 are more indicative of the the Taung skull can be expected to approach
the adult values (Ashton and Spence, 1958).
pattern seen in Homo erectus.
BASICRANIUM OF PLIO-PLEISTOCENE HOMINIDS
The value of 50” estimated for angle a is already greater than the mean value for the adult
Homo sapiens sample, and it also exceeds the
values for the adult and subadult “robust”
australopithecine fossils used in this study.
During the growth of the cranial base from the
time of eruption of the first permanent molars
to adulthood an increase of about 10O occurs in
both the pongid taxa and Homo sapiens (Dean,
in preparation). If this is added to the existing
value for a,50°, then the inference is that the
petrous axis in the Taung adult wouId have
been approximately 60”. This would place it
out of the observed range of the “robust”crania
and well within the “gracile”australopithecine
range.
DISCUSSION
Differences in the degree of cranial base flexion in the skulls of hominoids have been linked
with a variety of morphological and functional
correlates. Huxley (1863)suggested that there
was a relationship between facial prognathism
and cranial base flexion, and since then Huxley
(1867),Weidenreich (1947),Bjork (1950),Scott
(1958), and Cramer (1977)have presented evidence to support such an association. Other
workers have concentrated on the effects of the
expansion of the neurocranium on cranial base
flexion during both human phylogeny and ontogeny (Moss, 1958 and Biegert, 1963). The
type of habitual posture, and the relative proportions of neurocranium and viscerocranium
have also been related to the degree of cranial
base flexion (Schultz, 1955; Clark, 1959; Du
Brul and Laskin, 1961, and Du Brul, 1977).
More recently Laitman (1977)and Laitman et
al. (1978,1979)have presented evidence to suggest that the degree of exocranial base flexion
may indicate the position of structures in the
upper part of the respiratory tree in primates.
In an earlier paper (Dean and Wood, 1981a)
we emphasized the importance of recording
variation in the shape and form of the cranial
base when it is viewed in norma basilaris as
well as in the sagittal plane. Among the problems of sagittal studies, not touched on in our
earlier discussion, is the difficulty of describing a complicated anatomical region across a
range of primate comparative groups without
a reliable set of homologous landmarks (Lestrel and Moore, 1978). For example, while nasion is a reliable indicator of the plane of the
foramen caecum (the anterior end of the cranial
base) in modern human crania, because of an
upward remodelling which occurs during
growth in many nonhuman primates it is a
169
much less reliable indicator of the position of
the foramen caecum in these groups (Ashton,
1957; Scott, 1958, 1963; Cramer, 1977). Likewise intrataxonomic variations in the degree of
upward remodelling have been shown to occur
in the region of the sella turcica (Latham, 1972;
Gould, 1978), and there is also evidence that
growth changes occurring at the spheno-occipita1 synchrondrosis may differ among
primate taxa (Sirianni and Van Ness, 1978).
In discussing the significance of the results
of this study of hominid basicrania we are, of
course, conscious that they refer to a restricted
set of metrical criteria from only one area of the
skull. However, the results of the preliminary
study of the variation of this region in a series
of samples of extant primates, established the
existence of consistent morphological patterns
which could, moreover, be plausibly related to
known patterns of cranial base flexion in the
sagittal plane. Thus it is against this comparative background, and the knowledge that the
cranial base must reflect, or respond to, a
whole series of important functional demands,
that the results should be assessed.
The additional of Sts 19 and 25 to the sample
of adult crania attributed to Aus tralopithecus
africanus has reinforced the conclusions based
on the initial observations on Sts 5 and MLD
37138. In general Sts 19 and 25 follow the pattern of MLD 37/38 in having a more foreshortened sphenoid than Sts 5. Sts 19 has both a
shortened sphenoid and the least sagittally orientated petrous axis of the group which makes
it the most “Homo-like”of the sample.
The suggeston that Sts 19 may not be a typical “gracile”australopithecine is not a new one.
When Sts 19 was first described in detail, and
referred to as skull No. 8 (Broom and Robinson, 1950).the authors referred to the “human”
proportions of the posterior cranial fossa. In
the same monograph Schepers (1950) commented that “the manner in which the cerebellum has come to be shifted forward and below
the cerebral occiput is most striking. The general arrangement is that found for the human
brain, especially in Homo sapiens.” Clarke
(1977)considered the morphology of the cranial base of Sts 19 in detail and suggested there
were several features of the morphology of the
temporal and sphenoid bones that were Homolike. He considered the possibility that Sts 19
should be excluded from Australopithecus africanus, but also noted the alternative, that the
“moreHomo-like characteristics of Sts 19 compared to the more pongidlike characteristics of
Sts 5 might be indicative only of a wide range
170
M.C. DEAN AND B.A. WOOD
of variation among the Sterkfontein Australopithecus population.” The results of this
metrical analysis suggest that the Clarke’ssecond interpretation is the more probable one.
The subadult “robust”australopithecine cranium SK47 has been included with the adult
crania KNM-ER 406 and OH 5 in this study because it is the only “robust” australopithecine
specimen from South Africa with a more or less
complete and undistorted cranial base. Despite
its immaturity Ashton and Zuckerman (1952)
noted that in SK 47 “themuscular markings of
the occipital region of the fossil are far more
pronounced than in either man, or the chimpanzee, and are as well developed as in gorillas
of corresponding age.” In addition, they comment that the inion and external occipital crest
in SK 47 are better marked than those of adolescent gorillas and chimpanzees. The results
of our metrical study suggest that, if it had
reached full adulthood, SK 47 would have come
to resemble the two adult “robust” australopithecine crania. Olson (1978) has previously
drawn attention to cranial and dental features
of SK 47 which he considers are hominine, but
these cranial base metrical data on their own
are not sufficiently discriminatory to make
any significant contribution to deciding the
relative strengths of the hominine and “robust”
australopithecine affinities of SK 47. Nonetheless some features, such as the anteriorly
situated foramen magnum, suggest a closer
association with the “robust” taxon. Howell
(1978)has maintained a separation at the species level between specimens recovered from
Kromdraai and Swartkrans, but the incomplete cranial basc of TM 1517 resembles the
cranial base pattern of the “robust” sample
used in this study much more closely than any
of them resemble the pattern in the “gracile”
australopithecines. However, it differs from
other “robust” crania in having a more elongated base (IT-ITITP-TP)and a larger angle p.
The cranial base pattern of TM 1517 also resembles those of KNM-ER 732 (a probable female of the “robust”taxon) and KNM-ER 1805.
The calvarium KNM-ER 407 and the cranium KNM-ER 732 were recovered from the
lower part of the Upper Member at Koobi Fora
in North Kenya (Leakeyet al., 1978).Both specimens show marked post-orbital constriction,
and have expanded and heavily pneumatized
mastoid regions. The mandibular fossae are
wide and extend laterally well beyond the sides
of the cranial vault. Although the specimens
are incomplete they have sufficient anatomical
areas in common to suggest that they should
be considered as belonging to the same taxon.
A preliminary assessment of KNM-ER 407
lead to the conclusion that it “probably is
either a gracile species of Australopithecus or
else a very early representative of Homo”
(Leakey, 1970).Since then Wolpoff (1978a)has
espoused the former proposition, but the latter
has received no support. From the outset,
KNM-ER 732 was considered as a possible
smaller-bodied female of Australopithecus
(Leakey, 1971), and although these reports
made no specific attribution, the clear inference was that the material should be referred to Australopithecus boisei (Leakey,
1970 and 1976). Since these preliminary pronouncements two substantial reviews of the
fossil hominid evidence from Koobi Fora concurred with the judgement that KNM-ER 407
and 732 are probable females of Australopithecus boisei (Howell, 1978; Wood 1978).
I t is clear from the cranial base diagrams
(Fig. 5) and the results in Table 2, that neither
KNM-407 or 732 have a basicranium which resembles that of the “gracile” australopithecines. The forward position of the foramen
magnum in both crania is a feature which resembles the “robust” australopithecine group,
and in view of the features of the frontal and
malar region which KNM-ER 732 shares with
specimens attributed to Australopithecus boisei (Wood,1978, in press), the most parsimonius taxonomic placement of these two specimens is within Australopithecus boisei.
The cranium KNM-ER 1470 was recovered
from the Lower Member of the Koobi Fora Formation and its age is between about 1.8 Myr.
and 2.8-3.0 Myr. (Drake et al., 1980; Gleadow,
1980; McDougall et al., 1980), and palaeomagnetic evidence suggests that it may be close to
2.4 Myr old (Hillhouse et al., 1977). Initial
assessment suggested that its affinities were
with Homo (Leakey, 1973; Wood, 1976) but
Walker argued for caution, and quite properly
stressed the features, particularly of the face,
which are shared between KNM-ER 1470 and
Australopithecus (Walker, 1976). Subsequent
reviews have favoured its inclusion in Homo
habilis (Howell, 1978) or treated it as a conspecific of the “gracile” australopithecine
material from Sterkfontein (Olson, 1978). Evidence derived from the cranial base of KNMER 1470 has to be interpreted in the light of
the distortion which has been assessed by
stereographic projection (Walker, 1981). This
indicates that the right temporal has been
pushed forwards, but not laterally. If allowance is made for these changes it gives no
cause to modify the conclusion that the hominid group KNM-ER 1470 most closely resem-
BASICRANIUM OF PLIO-PLEISTOCENE HOMINIDS
171
bles is Homo erectus. There are clear contrasts to Homo habilis, the pattern of basicranial
between the cranial base of KNM-ER 1470 and morphology in KNM-ER 1813 is strongly in
that of the “gracile”australopithecines, and the favour of its inclusion in Homo, and for it not
breadth across the infratemporal crests to be regarded as a “gracile”australopithecine.
distinguishes it from “robust” australopithe- The foreshortening of the skull base, widening
cine crania. Although the overall morphology of the sphenoid, and the angulation of tymof the cranial vault and the face of KNM-ER panic and petrous parts of the temporal bone
1470 set it apart from crania and calottes at- are all clear and unambiguous evidence against
tributed to Homo erectus, the form of its KNM-ER 1813 being placed within Australocranial base suggests that if it is assigned to pithecus africanus.
Homo habilis then the cranial base of this taxThe cranium OH 24 was found at site DK in
on had already developed some of the probable Bed I of Olduvai Gorge. The cranium was
derived features we presently associate with found in a crushed state and was skillfully reHomo erectus and Homo sapiens.
stored by Dr. R.J. Clarke. The initial descripThe skull KNM-ER 1805 was recovered from tion pointed to the similarities between OH 24
just below the Okote Tuff in the Upper Mem- and the Homo habilis skull, OH 13, from Bed
ber at Koobi Fora. Radiometric and magneto- I1 (M.D. Leakey et al., 1971). Although some
stratigraphic evidence all point to a date differences were noted between OH 24, and
around 1.6 Myr (Fitch and Miller, 1976; Brock OH 7 and 16, the report concluded that “beand Isaac, 1976). Its relatively small cranial yond doubt, the new specimen represents the
capacity, 582 m13, (Holloway, 1978), and the genus Homo as defined by Leakey, Tobias, and
combination of a compound nuchal crest and Napier and differs fundamentally from the
parasagittal crests have perhaps been features
australopithecines.” Although its inclusion in
which have led some workers to attribute it to Homo habilis has been supported by some
Australopithecus boisei (Tobias, 1980). Pre- (Holloway, 1976, 1978; Tobias, 1980, 1981).
sumbably different features prompted Howell many workers have been impressed by the sim(1978)and Wolpoff (1978b)to include ER 1805 ilarities between OH 24 and Australopithecus
in Homo erectus, and other workers to place it africanus (Leakey, 1974; Howell, 1978; Olson,
in a separate species of Australopithecus,
1978; Leakey and Walker, 1980). The cranial
either named (Olson, 1978), or unnamed base of OH 24 is, however, unlike that of the
(Holloway, 1976; Leakey 1976). The evidence “gracile”australopithecines. The petrous axes
from the cranial base is compatible with the in- are more coronally aligned and the greater
clusion of KNM-ER 1805 in either Australo- width across the infratemporal crests suggests
pithecus boisei or a species within the genus that the sphenoid in OH 24 was broader than in
Homo, but is strong evidence against its at- the Australopithecus africanus specimens. We
tribution to Australopithecus africanus.
are confident that the differences in cranial
The precise stratigraphic relationships of
base length between OH 24 and AustralopitheKNM-ER 1813 have yet to be worked out, but
cus afn’canus are not simply the result of
it is likely that it was buried in sediments in the distortion.
Since the discovery that the parts of the cralower part of the Upper Member, or the upper
part of the Lower Member, at Koobi Fora. Its nium SK 847 from Swartkrans fitted with a
small brain size, 509 m13(Holloway, 1978) and left temporal (SK 846b) and a maxillary fragthe overall shape of the cranial vault and face ment (SK 80),the affinities of this “composite”
have prompted many workers to suggest affin- cranium have been in dispute. The initial asessment of its affinities considered that it differed
ities with, or its inclusion in, Australopithecus
africanus (Holloway, 1978; Olson, 1978),while significantly from the “robust” australopithemore hominine aspects of relative tooth size, cines, and it was regarded as Homo sp. indet.
dental morphology, and tooth wear have led (Clarke et al., 1970), and a similar taxonomic
others to include it in Homo habilis (Howell, conclusion was reached after a much more de1978; Tobias, 1980).Many features of the mor- tailed analysis (Clarke, 1977). Howell (1978)
phology of the cranial vault, face, and denti- has assigned it to Homo habilis, but Wolpoff
has argued consistently against any taxonomtion convincingly preclude the inclusion of
KNM-ER 1813 in Australopithecus boisei ic distinction between SK 847 and the “robust”
(Wood,1978).Thus, with the taxonomic claims australopithecines. (Wolpoff, 1970, 1971,
1974). In his survey of nasal and mastoid morlimited to its inclusion in either Australopithecus africanus or Homo habilis the evidence phology Olson (1978)regarded the “composite”
from the cranial base is crucial. If KNM-ER cranium as a conspecific of the “gracile”
1470 and OH 24 (uide infra) do prove to belong australopithecines from Sterkfontein. In this
172
M.C. DEAN AND B.A. WOOD
present study the shape of the basicranium of
SK 847 has been shown to be unlike that of the
“gracile”australopithecines, and thus must be
counted as evidence against them being regarded as conspecific. Although the overall
pattern of basicranial morphology of SK 847
shows similarities to both Homo and the “robust” australopithecines, the relatively wide
bi-infratemporal fossa breadth suggests interesting affinities with specimens which have
been attributed to Homo.
Although the cranial base of Taung is incomplete, any information it does contain is potentially important contributory evidence for any
assessment of its affinities. In the wake of new
evidence about the dating of Taung, Tobias
suggested that serious consideration be given
to the proposal that the Taung child may be a
“late surviving member of A. robustus or A. cf
robustus” (Tobias, 1973).Tobias (1981)has recently restated his views on Taung and has reemphasized his belief that Taung has closer affinities with the “robust” than with the
“gracile”australopithecines.
The assessment of the significance of differences between adult specimens is notoriously
difficult. One of us (Dean, in preparation) has
investigated growth changes in the basicrania
of extant higher primates which have been
aged using dental criteria (Dean and Wood,
1981b).These results suggest that in the adult
equivalent of the Taung skull the petrous axes
would have been aligned at an angle of approximately 60”. This is a much more sagittal orientation than is found in the “robust” australopithecines and is inferential morphological
evidence in favour of regarding the Taung
child as a juvenile representative of Australopithecus africanus as known from Sterkfontein
and Makapansgat.
A more general feature of these results is
that, because of the similarities between the
cranial base patterns of “robust”australopithecines and Homo, it would be unwise to try to
distinguish between the two taxa solely on the
evidence of the cranial base. We have discussed elsewhere the possibility of parallel
evolution in Homo and the “robust” australopithecines (Dean and Wood, 1981a),and while
many apparently derived features have been
adduced in support of the concept of a ‘robust’
australopithecine clade (White, et al., 1981;
Rak, 1981)we submit that these similarities in
cranial base morphology between Homo and
the “robust” taxa, together with evidence of
shared patterns of tooth eruption (Clements
and Zuckerman, 1953) and jaw growth (Skin-
ner, 1978)should be given due consideration in
any analysis of hominid phylogeny.
The implications of the differences between
the cranial base patterns of the “robust” and
“gracile” australopithecines have been discussed elsewhere (Dean and Wood, 1981a)and
confirm the preliminary observations of Tobias (1967) and Du Brul (1977). There are profound differences in basicranial morphology
between Homo habilis and Homo erectus on
the one hand, and Australopithecus africanus
on the other. The reorientation and rearrangement of the cranial base which has taken place
in these early Homo taxa suggests that the
relationship between Homo habitis and Australopithecus africanus may not be as close as
some workers have claimed (Robinson, 1972;
Leakey, 1974; Olson, 1978; Leakey and
Walker, 1980). The taxonomic and phylogenetic relationships of the small brained “gracile”
crania from East Africa (e.g., KNM-ER 1813
and OH 24) remain an enigma, but the results
presented here suggest that their inclusion in
Australopithecus africanus may be premature.
The basicranial morphology of hominids recovered from Laetoli and Hadar will be crucial
evidence in any phylogenetic analysis, and
more details of well-preserved cranial specimens are awaited with interest (Olson, 1981).
We are conscious that the taxonomic and
phylogenetic relationship of fossil hominids
can be neither confirmed nor refuted on the
basis of observations made on one region of the
cranium. However, it is clear that one profitable way of analyzing the affinities of fossil
groups is to accumulate evidence from a series
of investigations dealing with a comprehensive set of anatomical regions or morphological
variables, and we submit that this study is a
contribution to such an analytical framework.
ACKNOWLEDGMENTS
M.C. Dean was in receipt of an NERC Research studentship and B.A. Wood was supported by a project grant from the NERC. We
are grateful to the Trustees of the National
Museum, Nairobi, the Transvaal Museum,
Pretoria, the Tanzanian Government, and the
Head of the Department of Anatomy, University of the Witwatersrand for allowing us access to material in their care. Our thanks go to
C.K. Brain, Alan Hughes, M.D. Leakey, R.E.
Leakey, P.V. Tobias, and Elizabeth Vrba for
providing hospitality and for stimulating and
productive discussion of many ideas included
in this paper.
BASICRANIUM OF PLIO-PLEISTOCENE HOMINIDS
We thank Elizabeth Marshall for typing the
manuscript, and we are indebted to several
referees for their careful assessment of this
paper. Some of their suggestions have been incorporated into the revised version of the
manuscript.
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