Correlation of cranial and mandibular prognathism
in extant and fossil hominids
1
2
Fred Spoor , Meave G. Leakey & Louise N. Leakey
1
3
Evolutionary Anatomy Unit, Department of Anatomy & Developmental Biology, University College London,
Gower Street, London, WC1E 6BT, United Kingdom e-mail: f.spoor@ucl.ac.uk
2
3
Division of Palaeontology, National Museums of Kenya, Nairobi, Kenya, and Department
of Anatomical Sciences, Stony Brook University, New York, USA
Koobi Fora Research Project, Nairobi, Kenya, and Department of Anthropology, Stony Brook University,
New York, USA
This pilot study investigates the correlation between mandibular symphysial orientation and both
craniofacial and subnasal prognathism in modern humans, chimpanzees, gorillas, and a range of other
catarrhine species. The purpose was to assess the potential value of using the degree of prognathism as a
parameter that might relate isolated crania and mandibles in the Plio-Pleistocene hominin fossil record
of Africa. It is found that angles expressing cranial and mandibular prognathism are intraspecifically
correlated in modern humans from pre-industralised populations, but not in the African apes, or
interspecifically among catarrhine species. Hominin fossils investigated in this comparative context
broadly follow the pattern of correlation shown by modern humans, with some potentially interesting
differences that warrant further investigation. In all, the results suggest that the degree of prognathism has
little practical value in determining associations between isolated hominin mandibles and crania.
INTRODUCTION
The association of isolated mandibles and crania in the fossil
hominin record can be problematic if multiple species are
known to be present. Lack of consensus involving unassociated
type specimens results in varying hypodigms, and complicates
discussions of taxonomy and phylogeny. An example is the
interpretation of the African Late Pliocene and Early Pleistocene
hominin record not attributable to Paranthropus. The species
names Homo habilis, H. rudolfensis and H. ergaster are frequently
used with the relatively complete and emblematic crania
KNM-ER 1813, KNM-ER 1470 and KNM-ER 3733 in mind.
However, their respective affiliations rest with the correct
association with two type mandibles, OH7 (H. habilis) and
KNM-ER 992 (H. ergaster). The latter specimen has been linked
with a range of different crania, and the currently widely
accepted association with H. erectus-like specimens, such as
KNM-ER 3733, only came with the discovery of KNM-WT
15000, which preserves both its cranium and mandible (see e.g.
Wood, 1991). More recently, the discovery of the maxilla
OH65 has reopened the debate as to whether H. rudolfensis is a
junior synonym of H. habilis (Blumenschine et al., 2003; Tobias,
2003). This is because the edentulous type cranium of H. rudolfensis, KNM-ER 1470, could actually be the same species as OH7
(including the type mandible of H. habilis). If this is the case
then smaller early Homo specimens such as OH13 and
KNM-ER 1813 perhaps represent an as yet unnamed taxon.
Modern human skulls show reduced prognathism of the
mandibular symphysis and the cranial aspect of the face,
including the subnasal area. This could suggest that these
forms of prognathism are correlated among hominids, and that
human evolution is characterised by a process of joint, and
integrated change from a more prognathic to a more orthognathic facial morphology. If correct, this phenomenon could have
the potential to relate fossil crania and mandibles of the same
taxon. Indeed, the more vertically-oriented symphysis of the
Australopithecus bahrelghazali type mandible KT12/H1 has been
said to indicate a more orthognathic face (Brunet et al., 1996).
The contemporary Kenyanthropus platyops type cranium KNMWT 40000 does show such facial morphology (Leakey et al.,
2001), and this could be seen as evidence linking the two fossils.
To assess the comparative basis for associating fossil hominin
crania and mandibles using the degree of prognathism we
studied the correlation pattern in modern humans, African
apes, and a sample of additional catarrhine species. In
orthodontic surgery there is a long tradition of comparing the
degree of cranial and mandibular prognathism in pragmatic
efforts to obtain a well-occluded dentition, and a balanced
facial appearance (e.g. Ranly, 1988; Proffit, 2000; Wolford et al.,
2004). However, in human palaeontology this issue has
attracted little attention. Most recently, however, Kimbel et al.
(2004) discussed the facial morphology of A. afarensis, and
noticed the unusual combination in A.L. 444-2 and A.L. 417-1 of
a prognathic subnasal area, and a steep, vertically-oriented
mandibular symphysis.
MATERIALS AND METHODS
The sample is summarised in Table 1. The modern humans
comprise of two subsamples: (1) skulls in the collections of the
Natural History Museum (London) and the National
Museums of Kenya, representing indigenous, pre-industrial
populations from all six widely inhabited continents, and (2)
skulls in the Department of Anatomy, University College
London, without definitive provenance, but representing
populations of the last 100 years from the London area, and
possibly the Indian subcontinent. These will be referred to here
as the pre-industrial and anatomy samples, respectively. The
non-human primate specimens are from the collections of the
Powell Cotton Museum (Birchington), the Royal College of
Surgeons (London), the Natural History Museum (London),
the Anthropological Institute of the University of Zürich, and
the Departments of Anatomy and Anthropology and the Grant
Museum of University College London,. The African ape
samples include the eastern lowland gorilla Gorilla gorilla, and
representatives of all chimpanzee Pan troglodytes subspecies.
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Transactions of the Royal Society of South Africa
Vol. 60(2)
Table 1. The sample analysed in this study, giving the number of individuals (n ), the labels used in Figures 2–5, and the craniofacial, subnasal and
mandibular angles.
n (male, female)
or accession no.
Label
Homo sapiens (all)
pre-industrial
UCL-Anatomy
Pan troglodytes
Gorilla gorilla
Pongo pygmaeus
Hylobates hoolock
Hylobates klossi
Hylobates syndactylus
Cercocebus torquatus
Cercopithecus campbelli
Cercopithecus cephus
Cercopithecus diana
Cercopithecus nictitans
Chlorocebus aethiops
Erythrocebus patas
Macaca fascicularis
Macaca nigra
Macaca sinica
Macaca sylvanus
Mandrillus sphinx
Miopithecus talapoin
Papio hamadryas
Theropithecus gelada
Colobus guereza
Colobus polykomos
Nasalis larvatus
Procolobus badius
Procolobus verus
Semnopithecus entellus
Trachypithecus obscurus
Trachypithecus vetellus
48
25
23
55 (25m, 30f)
44 (23m, 21f)
5 (3m, 2f)
1f
1m
1f
1f
2f
1m
1m
2 (1m, 1f)
1f
1f
3 (2m, 1f)
1f
1m
1m
1m
1m
3 (2m, 1f)
2 (1m, 1f)
2 (1m, 1f)
2 (1m, 1f)
1f
2m
3 (2m, 1f)
1m
1m
1f
Homo erectus
Paranthropus robustus
Australopithecus africanus
Australopithecus afarensis
Australopithecus afarensis
Kenyanthropus platyops
Hominini sp.
Australopithecus bahrelghazali
Australopithecus anamensis
Australopithecus anamensis
KNM-WT 15000
SK 12
Sts 52ab
A.L. 444-2
A.L. 417-1
KNM-WT 40000
KNM-WT 8556
KT12-H1
KNM-KP 29281
KNM-KP 29283
Species
Craniofacial angle
Subnasal angle
Mandibular angle
Mean
S.D.
Mean
S.D.
Mean
S.D.
–
h, Hs
a
p, Pt
g, Gg
Po
Hh
Hk
Hy
Ct
Cc
Ce
Cd
Cn
Ca
Ep
Mf
Mn
Mi
My
Ms
Mt
Ph
Tg
Cg
Cp
Nl
Pb
Pv
Se
To
Tv
78
78
78
60
63
61
65
70
60
55
64
61
59
58
64
60
53
53
46
50
46
66
39
50
58
61
57
61
77
68
65
61
5.0
5.8
4.2
3.5
3.3
4.2
–
–
–
–
2.7
–
–
0.1
–
–
2.0
–
–
–
–
–
5.0
1.0
1.5
1.1
7.9
9.4
5.4
4.1
4.5
6.1
–
–
–
–
2.8
–
–
1.5
–
–
3.1
–
–
–
–
–
8.1
1.1
2.7
2.3
7.3
6.0
–
–
–
81
78
84
45
42
39
57
58
56
47
36
47
44
42
46
44
37
35
41
38
46
40
44
48
45
52
54
47
43
47
49
48
8.2
6.9
8.1
5.8
4.9
1.8
–
–
–
–
4.6
–
–
2.4
–
–
2.5
–
–
–
–
–
5.5
4.1
5.2
3.3
1.6
1.9
–
–
–
60
62
58
43
42
35
56
55
52
57
45
57
38
59
45
43
49
50
48
56
54
52
44
61
41
52
42
50
51
53
40
40
1
2
3
4
5
6&7
6
7
8
8
70
63
67
65
–
69
–
–
–
–
–
–
–
–
–
–
–
–
–
–
48
35
48
41
44
53
–
–
–
33
–
–
–
–
–
–
–
–
–
–
59
67
60
65
71
–
64
65
47
–
–
–
–
–
–
–
–
–
–
–
The match between mandibles and crania was carefully
checked, and the specimens have largely intact dentitions
without evidence of substantial pathology.
Five hominin fossils were included, which each preserve the
relevant cranial and mandibular areas (Table 1). Moreover, to
explore the earliest available hominin evidence the most
complete A. anamensis mandible and maxilla were considered,
even though these specimens are not of the same individual.
Lastly, we assessed the potential match of the cranium
KNM-WT 40000 with the A. bahrelghazali mandible KT12-H1, as
well as with KNM-WT 8556, a mandible from the same
geographic area, although somewhat younger in age (Leakey
et al., 2001). The original fossils were measured, apart from
KT12-H1 (cast made available by M. Brunet), and A.L. 444-2
(reconstruction made available by W. Kimbel, cross-checked
4.4
2.7
–
–
–
with the original fossils). KNM-WT 15000 and Sts 52 are
subadult, with second molars in occlusion, and the third molars
unerupted in the former, and partially erupted in the latter.
Craniofacial, subnasal, and mandibular prognathism is
quantified using the lines sellion to prosthion, nasospinal
to prosthion, and infradentale to gnathion, respectively
(Figure 1). Angles of these lines were calculated to the orientation of the relevant (maxillary or mandibular) alveolar margin
projected onto the midsagittal plane. The alveolar margin
orientation is defined as the line from the margin between the
second and third molar to either prosthion or infradentale
(Figure 1). Angles were also calculated relative to the postcanine alveolar margin, replacing prosthion and infradentale
by the margin between the canine and third premolar. However, these results are not reported here, as they are basically
Spoor et al.: Correlation of cranial and mandibular prognathism in extant and fossil hominids
Figure 1. Lateral view of a skull of Pan troglodytes, showing the landmarks employed in this
study. Unlike sellion (se) and the alveolar margin
between the second and third molars, the position of nasospinale (ns), prosthion (pr), infradentale (id) and gnathion (gn) may not be visible,
and these are therefore indicated by markers.
87
Figure 2. Bivariate double logarithmic bivariate plot of the craniofacial angle against the
mandibular angle for the individual specimens of Homo sapiens, Pan troglodytes, and Gorilla
gorilla, as well as the hominin fossils. RMA regression lines are given for H. sapiens (dashed,
total sample; solid, pre-industrial sample). Symbols listed in Table 1.
the same as obtained for the full alveolar
margin.
The angles were calculated from landmark coordinates. For the non-hominid
specimens these were recorded from threedimensional medical CT data (images
in sagittal plane with a pixel size varying
from 0.19 to 0.49 mm, depending on cranial
size). The landmarks of the hominid skulls
were taken from digital images with a focal
distance of 1–2 m, aligned to provide an
exact lateral view of the facial profile. Nasospinale, prosthion, infradentale and gnathion may not be visible in this view, and are
indicated by markers oriented in the sagittal
plane (Figure 1). Tests show that this simple
technique has a maximum measurement
error of ±1 degree for the focal and interlandmark distances used here (Wright, Figure 3. Bivariate double logarithmic plot of the craniofacial angle against the mandibular
2001).
angle, for the species means of the extant catarrhine sample, and the individual hominin
The relationship between the cranial and fossils. The RMA regression line of the pre-industrial modern human sample is shown.
mandibular angles was assessed by calcu- Symbols listed in Table 1.
lating Spearman rank correlation coeffiare given in Table 2.
cients (rrank). When correlation is significant (P < 0.05), it was
investigated whether the relationship is allometric rather
In the modern human sample the craniofacial and mandibular
than isometric by testing the null hypothesis that the slope of
angles are positively correlated, and this relationship is
the reduced major axis (RMA) regression of the logged values
allometric. The slope indicates that widening of the mandibular
is 1 (software: PAST, v1.20). Correlation was investigated intraangle (i.e. a more upright symphysis) tends to correspond with
specifically for the modern human and African ape samples,
increasingly less widening of the craniofacial angle. The two
and interspecifically for the full extant catarrhine sample.
subsamples of modern humans show markedly different
results, with a stronger correlation between the two angles in
the pre-industrial group, and no correlation in the anatomy
RESULTS
group. Moreover, for the pre-industrial sample an isometric
Sample statistics of the craniofacial, subnasal and mandibular
relationship cannot be rejected (P > 0.05). The estimated RMA
angles are given in Table 1, and Figures 2–5 show intraspecific
slope, based on raw values, is less than 1, suggesting that
and interspecific bivariate plots of these angles. Correlation
variation in the mandibular angle is accompanied by less
coefficients, and RMA regression statistics, where appropriate,
88
Transactions of the Royal Society of South Africa
Vol. 60(2)
change in craniofacial angle. However, its
95% confidence interval does include 1,
leaving open the possibility of one-to-one
variation.
The subnasal and mandibular angles in
modern humans are correlated in the preindustrial skulls, but not in the anatomy, or
combined samples. This relationship is
allometric, but with a slope larger than 1,
rather than less than 1 found for the craniofacial angle. This means that a more upright
symphysis tends to correspond with an
increasingly more orthognathic subnasal
clivus.
The craniofacial and subnasal angles are
not correlated intraspecifically with the
mandibular angle in the chimpanzee and
gorilla samples, and treating the sexes separately gives the same result. These angles
are not correlated interspecifically in the
catarrhine sample either. Excluding the derived, orthognathic modern humans from Figure 4. Bivariate double logarithmic plot of the subnasal angle against the mandibular angle
the catarrhines does not alter the result.
for the individual specimens of Homo sapiens, Pan troglodytes, and Gorilla gorilla, as well as
Bivariate plots of the cranial versus man- the hominin fossils. The RMA regression line of the pre-industrial modern human sample
dibular angles clarify how the modern is shown. Symbols listed in Table 1.
human correlation patterns compare with
the morphology of the fossil hominins, the
African apes and other catarrhines. There is
no overlap of the areas of intraspecific variation of modern humans on the one hand,
and those of the African apes on the other
(Figures 2, 4). The latter cluster above the
intraspecific RMA for the pre-industrial
modern humans, in particular in the plot of
the subnasal angle (Figure 4). This indicates
that, compared with modern humans, the
African apes are not only more prognathic
overall, but also have a less vertically inclined mandibular symphysis for their
degree of cranial prognathism. Plots of the
catarrhine species means broadly show the
same pattern (Figures 3, 5). However, for
the relationship between the craniofacial
and mandibular angles several species,
including the hylobatids and the longmuzzled papionines, fall close to the modern human regression trend (Figure 3).
Moreover, the subnasal angle of hylobatids Figure 5. Bivariate double logarithmic plot of the subnasal angle against the mandibular
and many cercopithecids fall well within angle, for the species means of the extant catarrhine sample, and the individual hominin
the range of modern humans (Figure 5).
fossils. The RMA regression line of the pre-industrial modern human sample is shown.
The hominin fossils broadly follow the Symbols listed in Table 1.
RMA regression of the pre-industrial
modern humans, falling at, or well below the lower end of the
they are also close to the other hominin specimens, apart from
modern range (Figures 2, 4). The A. afarensis and P. robustus
A. anamensis. Relative to the modern human regression the hyspecimens differ from the A. africanus and H. erectus specimens
pothetical combination of either mandible with KNM-WT
by combining a more vertical mandibular symphysis with a
40000 follows the trend of the A. africanus, A. anamensis and
prognathic cranium, in particular for the subnasal angle. This
H. erectus specimens more closely than that of the A. afarensis
difference approximately equals the maximum scatter around
and P. robustus specimens (Figures 4, 5).
the RMA shown by the modern human sample. Both the
A. anamensis maxilla and mandible are characterised by the
DISCUSSION
most prognathic morphology in the hominin sample (Table 1).
In this short study we set out to assess whether or not there is
Combined their position relative to the modern human regresa comparative basis for using the degree of prognathism in
sion is similar to that of the A. africanus and H. erectus specimens
efforts to associate isolated fossil hominin mandibles and
(Figures 4, 5). Finally, the mandibles KNM-WT 8556 and
crania. The results clearly indicate that in pre-industrial
KT12-H1 do not differ in their symphysial orientation, and
modern humans worldwide the degree of cranial and mandib-
Spoor et al.: Correlation of cranial and mandibular prognathism in extant and fossil hominids
Table 2. Correlation and regression statistics of the craniofacial and
submandibular angles against the mandibular angle. The Spearman
rank correlation coefficient (rrank) is given with its level of significance,
and the RMA slope is given with its 95% confidence interval in brackets,
and with the probability that it is 1. x, P < 0.05; xx, P < 0.01; xxx, P <
0.001; ns, not significant (P ≥ 0.05).
y-axis:
x-axis:
Craniofacial angle
Mandibular angle
rrank
0.324 x
RMA slope (log)
0.666 (±0.182)
Subnasal angle
Mandibular angle
Homo sapiens
All
Pre-industrial
p(slope = 1)
xxx
Intercept (log)
0.621
0.142 ns
rrank
0.476 xx
0.541 xx
RMA slope (log)
0.867 (±0.313)
1.774 (±0.639)
p(slope = 1)
ns
x
Intercept (log)
0.251
–1.565
RMA slope (raw)
0.841 (±0.309)
Intercept (raw)
12.466
UCL-Anatomy
rrank
0.185 ns
–0.126 ns
Pan troglodytes
rrank
–0.128 ns
–0.151 ns
Gorilla gorilla
rrank
–0.028 ns
0.069 ns
Interspecific
rrank
0.321 ns
0.279 ns
rrank
0.248 ns
0.202 ns
excl. H. sapiens
ular prognathism are correlated. However, this correlation is
absent in the sample representing industrialised society,
suggesting that changes in diet, mastication and health care
may have released constraints underlying the integration of
craniofacial and mandibular morphology. That no correlation
was found intraspecifically in chimpanzees and gorillas could
imply that such constraints emerged in the course of human
evolution, perhaps in relation to the reduction of the size and
projection of the lower face.
Interspecifically catarrhines clearly do not show a single
pattern of correlated variation in the degree of cranial and
mandibular prognathism, with aspects such as facial length
and hafting resulting in great variation between species. The
position of the African apes away from the modern human
regression trends is largely the consequence of the position of
gnathion, which is positioned posteriorly because of the
well-developed inferior tranverse torus (shelf), whereas in
modern humans it is placed particularly anteriorly because of
chin development.
The hominins investigated here do seem to follow broadly
the modern human regression trends, but without showing
a simple pattern from more prognathic to orthognathic
morphology. The H. erectus and A. africanus specimens are
similar in their facial angles, whereas the A. afarensis and
P. robustus specimens cluster separately. The latter could be
interpreted as adding a similarity involving the masticatory
system to the list of morphologies shared by these taxa (see
Kimbel et al., 2004, p. 231). However, differences between the
four taxa fall within modern human variation around the
regression trends, and additional data are required to assess
this further. It could be argued that the subadult H. erectus and
A. africanus specimens would have been more prognathic as
adults. However, this is not necessarily the case as KNM-WT
15000 is similar in craniofacial and subnasal angles to
KNM-ER 3733 (68 and 47 degrees, respectively), and Sts 52 is
89
close to Sts 5, Sts 17 and Sts 71 (range 63–68, and 44-46 degrees,
respectively).
In all, the results suggest that the degree of prognathism has
little practical value when attempting to associate isolated
hominin mandibles and crania, given the absence of an
unambiguous and consistent pattern of correlation, and the
substantial ranges of intraspecific variation. Typically, the
results indicate that the relatively vertically inclined symphysis
of KT12/H1 cannot be used to predict an orthognathic face,
because in A. afarensis and P. robustus this morphology is
associated with prognathic crania. On the other hand, if it
holds true that the fossil hominins broadly follow the modern
human pattern of correlation, it will be less likely that a more
orthognathic cranium, such as KNM-WT 40000, is associated
with a particularly prognathic mandible, as exemplified by
the A. anamensis specimen KNM-KP 29281. The findings
presented here do indicate that on a more fundamental level
the integration of cranial and mandibular morphology in
human evolution warrants further investigation. Perhaps
when the underlying biological mechanisms and constraints
are better understood it will be possible to predict what morphology is of particular importance when matching cranial and
mandibular elements in the fossil hominin record.
ACKNOWLEDGEMENTS
With this paper we salute Phillip V. Tobias, and acknowledge
his many contributions to palaeoanthropology. We thank
the National Museums of Kenya, the National Museum of
Ethiopia, the Transvaal Museum (South Africa), the Institute of
Human Origins (USA), and the museums listed in the Materials
& Methods section for access to specimens in their care. We are
grateful to Leslie Aiello, Gustl Anzenberger, Michel Brunet,
Simon Chaplin, Helen Chatterjee, Chris Dean, Heidi Fourie,
John Harrison, Jane Hughes, Nathan Jeffery, Paula Jenkins, Bill
Kimbel, Charlie Lockwood, Emma Mbua, Peter Morris, Alan
Walker and Tanwen Wright for help with various aspects of the
research. Financial support was provided by the Leakey Foundation and the National Geographic Society.
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Transactions
of the Royal Society of South Africa
Volume 60(2) — October 2005
© Royal Society of South Africa. ISSN 0035-919X
http://www.journals.co.za/ej/ejour_royalsa.html
Foreword — By D.E. Rawlings · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
Phillip V. Tobias: a biographical note — By R. Leakey · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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Aspects of Hominid Evolution
Species diversity in human evolution: challenges and opportunities
— R. Foley · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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Variation in early hominin temporal bone morphology and its implications for species
diversity
— C.A. Lockwood, W.H. Kimbel & J.M. Lynch · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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Pliocene hominid fossils from Gamedah, Middle Awash, Ethiopia
— T.D. White, B. Asfaw & G. Suwa · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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Correlation of cranial and mandibular prognathism in extant and fossil hominids
— F. Spoor, M.G. Leakey & L.N. Leakey · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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‘A tale of two taxa’
— B. Wood · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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Dental development and upper molar cusp dimensions of South African australopithecines
— S. Boccone & J. Moggi Cecchi · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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Lesser known chapters in the history of the Makapansgat hominid site
— J.M. Maguire · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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Dating of the Sterkfontein hominids: progress and possibilities
— T.C. Partridge · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
107
Prehistory in blood and bone: An essay on the reconstruction of the past from genetics
and morphology
A.G. Morris · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
111
Palaeoanthropological and molecular studies on the origin of modern humans in
China
— X. Wu· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
115
Human Genetics
Genetic prediction of common multifactorial diseases
— M. Bobrow · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
Genetics, race and medicine
— T. Jenkins · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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Human Behaviour and Culture
Remembrance of things past: amnesic syndrome with partial preservation of professional skills
— L Geffen & G. Geffen · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
Have you checked your hyoid lately? Strangulation, pathology, trauma, accident
— K.A.R. Kennedy · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
The southern African San and their rock art
— J.D. Lewis-Williams · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
Antiquity of the smoking habit in Africa
— N.J. van der Merwe · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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ààà
Phillip Vallentine Tobias: an appreciation
— F. Clark Howell · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
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Professor P.V. Tobias FRS, Hon. FRSSAf
Curriculum vitae · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
List of publications · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ·
The journal is abstracted in:
Biological Abstracts, Chemical Abstracts, Excerpta Medica, Mathematical Reviews, ISI® Web of Science.
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