J. Anat. (1988), 161, pp.

1-35 1
with 6 figures
Printed in Great Britain

Analysis of the dental morphology of Plio-Pleistocene hominids

V. Maxillary postcanine tooth morphology
B. A. WOOD AND C. A. ENGLEMAN
Hominid Palaeontology Research Group, Department of Human Anatomy and Cell
Biology, The University of Liverpool, P.O. Box 147, Liverpool L69 3BX
(Accepted 23 January 1988)

INTRODUCTION
The first four papers in this series investigated the mandibular postcanine dentition
of Plio-Pleistocene fossil hominids. Each paper used detailed metric data and precise
assessments of the morphological traits of either the crowns (Wood & Abbott, 1983;
Wood, Abbott & Graham, 1983; Wood & Uytterschaut, 1987) or roots (Wood.
Abbott & Uytterschaut, 1988) to investigate and define the distinctive characteristics
of four taxonomic categories of early hominids. In each study, the same variables and
criteria were used to assess the affinities of specimens whose taxonomic designation is
less certain and, where appropriate, they were used to develop phylogenetic,
hypotheses.
The morphometric data for the tooth crowns were obtained either from
measurements of the original specimens or from macrophotographs of the occlusal
surface of the tooth crowns. Total crown and component cusp areas were measured
by planimetry and morphological traits were scored using definitions either specially
devised or adapted from previous publications (e.g. Kraus & Furr, 1953; Ludwig,
1957; Hanihara, 1961). Fissure patterns were recorded using measurements based on
reference points which were adapted from Biggerstaff (1969). Multivariate as well as
univariate analytical methods were employed, specifically Principal Components
Analysis, for the cusp area data, and Procrustes (Gower, 1975) and Principal
Coordinates Analysis (Gower, 1966) for the fissure pattern data.
This paper, which completes the series, concentrates on the crown morphology of
the maxillary postcanine teeth, but also contributes some new data about the root
systems of these teeth as well as reviewing existing evidence about their root form.
Metrical information about the roots of the maxillary postcanine teeth was not
gathered systematically in the way it was for the mandibular postcanine teeth. This
was because of the difficulty involved in obtaining reliable radiographic images of
maxillary as opposed to mandibular teeth. In other respects, the methods of data
collection and analysis are broadly similar to those used for the mandibular teeth,
except that metrical information about fissure pattern has not been included. The
reasons are twofold. First, the results of such analyses of the mandibular molars and
premolars suggested that there is considerable overlap between the information
contained in the fissure pattern and cusp area data (Wood et al. 1983; Wood &
Uytterschaut, 1987). Indeed, in no case did the fissure pattern data of an unknown
specimen indicate affinities that had not already been suggested by analysis of the cusp
areas. Secondly, the methods used to record fissure pattern data in the previous study
required a standard and reliable method for orientating each occlusal photograph with
2 B. A. WOOD AND C. A. ENGLEMAN

Table 1. List of Plio-Pleistocene hominid maxillary molar teeth from East African
sites, together with their taxonomic category
Taxonomic
Specimen number Ml M2 M3 category
Site: Koobi Fora
KNM-ER 733E L 1 - - EAFROB
KNM-ER 807 R 1 - 1 Unknown
KNM-ER 808G R 1 - - Unknown
KNM-ER 1171H R -1 - EAFROB
KNM-ER 1171G L -1 - EAFROB
KNM-ER 1590L R 1 - - EAFHOM
KNM-ER 1590 L 1 1 - EAFHOM
KNM-ER 1804 L 1 1 - Unknown
KNM-ER 1805 R 1 1 1 Unknown
KNM-ER 1805 L 1 1 1 Unknown
KNM-ER 1808H R -1 - EAFHER
KNM-ER 1813 R - -1 Unknown
KNM-ER 1813 L 1 1 1 Unknown
KNM-ER 3733 R -1 - EAFHER
KNM-ER 3733 L 1 1 - EAFHER
Site: Olduvai Gorge
O.H. 5 R 1 - - EAFROB
O.H. 5 L 1 - - EAFROB
O.H. 6 R 1 - - EAFHOM
O.H. 13 R 1 1 1 EAFHOM
O.H. 13 L 1 1 1 EAFHOM
O.H. 15 R - -1 Unknown
O.H. 15 L -1 Unknown
O.H. 16 R 1 1 1 EAFHOM
O.H. 16 L - 1 1 EAFHOM
O.H. 21 L 1 - - Unknown
O.H. 24 R 1 - - EAFHOM
O.H. 24 L 1 - 1 EAFHOM
O.H. 30 R 1 - 1 EAFROB
O.H. 30 L 1 - 1 EAFROB
O.H. 39 R 1 1 - Unknown
O.H. 39 L - 1 - Unknown
O.H. 44 R 1 - - Unknown
Site: Chesowanja
C.H. 1 R 1 1 1 EAFROB

respect to a reference grid; the irregular shape of the outline of the maxillary molars
made such consistent orientation difficult to achieve.

MATERIALS AND METHODS

A total of 139 maxillary molar teeth (50 M1s; 42 M2s; 47 M3s) (Tables 1, 3) and 79
maxillary premolar teeth (44 P3s; 35 P4s) (Tables 2, 4) from at least 98 individuals were
studied. These figures represent the maximum number of specimens in the analysis but
for many of the detailed studies the loss of reference points through wear or damage
often results in smaller samples. The criteria for inclusion in the study were the same
as those for the mandibular teeth (Wood & Abbott, 1983, p. 199). Specimens were
placed in one of five informal taxonomic categories or were classed as 'unknown'.
Details of the formal taxa subsumed into each category are given in Wood & Abbott
(1983, pp. 198-199).
 Hominid maxillary postcanine teeth 3
Table 2. List of Plio-Pleistocene hominid maxillary premolar teeth from East African
sites, together with their taxonomic category
Taxonomic
Specimen number P3 P4 category
Site: Koobi Fora
KNM-ER 733D L I EAFROB
KNM-ER 802I L 1 EAFROB
KNM-ER 808C R 1 - Unknown
KNM-ER 1506 R 1 1 Unknown
KNM-ER 1590 R 1 1 EAFHOM
KNM-ER 1590 L I I EAFHOM
KNM-ER 1804 L I I Unknown
KNM-ER 1805 R I 'Unknown
KNM-ER 1805 L 1 1 Unknown
KNM-ER 1813 L 1 1 Unknown
KNM-ER 3733 R 1 EAFHER
KNM-ER 3886A R 1 - Unknown
Site: Olduvai Gorge
O.H. 13 R 1 1 EAFHOM
O.H. 16 R 1 1 EAFHOM
O.H. 16 L 1 1 EAFHOM
O.H. 24 L 1 EAFHOM
O.H. 30 L 1 EAFROB
O.H. 39 R 1 1 Unknown
O.H. 39 L 1 1 Unknown
Site: Chesowanja
C.H. 1 R 1 1 EAFROB

Linear measurements of the crowns of the teeth were made using vernier callipers
with tips specially machined to allow them to be inserted between teeth still in situ in
the upper jaw. Measurements were recorded to the nearest 0.1 mm and were made on
at least two occasions by BAW. The mean value of the measurements was used. The
average genuine measurement error (as opposed to obvious transcription errors) was
2 %. Mesiodistal and buccolingual diameters of both premolar and molar teeth were
measured according to the method proposed by Korenhof (1960) (Figs. 1, 2) and used
by Tobias (1967). In worn teeth allowance was made for the missing enamel (see
below). In addition to computing uncorrected, and corrected, crown base area from
the products of the linear measurements, the crown shape index (BL/MD(C) x 100)
was calculated for each intact crown.
The occlusal photographs, which provided the images from which measured crown
base, and individual cusp, areas were taken, were prepared according to the protocol
given in Wood & Abbott (1983, p. 199). The boundaries of each cusp (Figs. 3, 4) were
marked on the photographs by tracing the course of the primary fissures and by
locating reference points which are based on those suggested by Biggerstaff (1969). In
those maxillary molars that possessed an additional cuspule its area was divided
between the adjacent main cusps in the manner of the previous study. For those teeth
which showed significant interproximal wear, the outlines of the original mesial and/or
distal crown borders were reconstructed with reference to both the overall shape of
the preserved crown and the extent and orientation of the interproximal wear
facets.
Cusp areas were measured using a fixed-arm planimeter. The average of three
readings was taken and the value reduced to original size by dividing by the square of
4 B. A. WOOD AND C. A. ENGLEMAN

Table 3. List of Plio-Pleistocene hominid maxillary molar teeth from southern

African sites, together with their taxonomic category
Specimen Taxonomic Specimen Taxonomic
number MI M2 M3 category number Ml M2 M3 category
Site: Swartkrans Site: Sterkfontein
SK 13 R 1 1 1 SAFROB TM 1511 R - 1 SAFGRA
SK 13 L 1 - 1 SAFROB TM 1512 R I SAFGRA
SK 14L -1 - SAFROB TM 1514 L 1 SAFGRA
SK 17 R 1 - - SAFROB TM 1561 R 1 SAFGRA
SK 27 R -1 - Unknown Sts 1 L I SAFGRA
SK 27 L 1 - - Unknown Sts 8 L I 1 - SAFGRA
SK 31 R 1 SAFROB Sts 17 R 1 - SAFGRA
SK 36 R - - 1 SAFROB Sts 21 R I SAFGRA
SK 46 R 1 SAFROB Sts 22 L 1 - SAFGRA
SK 46 L 1 SAFROB Sts 28 R 1 1 SAFGRA
SK 47 R 1 1 - Unknown Sts 32 L 1 - SAFGRA
SK 47 L -1 1 Unknown Sts 37 L 1 1 SAFGRA
SK 48 L 1 1 1 SAFROB Sts 52a R I 1 1 SAFGRA
SK 49 R 1 - - SAFROB Sts 52a L I 1 1 SAFGRA
SK 49 L 1 SAFROB Sts 53 R 1 1 SAFGRA
SK 52 R 1 - - SAFROB Sts 53 L 1 1 SAFGRA
SK 55a L 1 - - SAFROB Sts 54 R 1 SAFGRA
SK 80 L 1 Unknown Sts 56 L I 1 - SAFGRA
SK 83 R - - 1 SAFROB Sts 57 L I SAFGRA
SK 83 L 1 SAFROB StW/H 6 L 1 SAFGRA
SK 98 L -1 - SAFROB StW/H 19B R 1 1 Unknown
SK 102 L 1 - - SAFROB SE 1508 R 1 - SAFGRA
SK 829 L 1 - - SAFROB Site: Makapansgat
SK 831a L 1 SAFROB MLD 6 R I 1 - SAFGRA
SK 832 L 1 - - SAFROB MLD 28 R 1 SAFGRA
SK 834 R -1 - SAFROB
SK 835 L 1 SAFROB Site: Kromdraai
SK 836 L - - 1 SAFROB TM 1517 R 1 1 SAFROB
SK 837 R -1 - SAFROB TM 1517a L I 1 - SAFROB
SK 838a R 1 - - SAFROB TM 1601 L I SAFROB
SK 870 L 1 SAFROB TM 1603 L 1 SAFROB
SK 1591 L 1 - - SAFROB
SK 3975 L 1 SAFROB Site: Taung
SK 3977 R 1 SAFROB IL I Unknown

the enlargement factor. Measurement error was 1 % or less. The areas of the whole
crown and its components, when these were available, were then recorded on
computer cards. For each tooth type, the parameters of the subsamples were
computed to check for obvious measurement or transcription errors. In addition to
the absolute value of each cusp area, the area of each cusp was expressed as a
percentage of the total. The mean, standard error and range of the cusp areas were
calculated for the four taxonomic categories, and the significance of any differences
was assessed using Student's t-test based on pooled variances and a two-tailed model.
Cusp size was expressed both as an overall value, and by expressing the area of each
cusp as a percentage of the measured (corrected) crown base area (MCBA). The cusp
areas of all the teeth with more than two cusps were then compared individually using
the principal components generated from a covariance matrix (PCm) and by studying
the Mahalanobis distances between the means of taxonomic categories, and the
distances between those means and each 'unknown' specimen. The SPSS canonical
variates analysis programme was used to compute posterior probabilities to estimate
the affinity between each 'unknown' specimen and the taxonomic categories.
 Hominid maxillary postcanine teeth 5
Table 4. List of Plio-Pleistocene hominid maxillary premolar teeth from southern
African sites, together with their taxonomic category
Taxonomic
Specimen number P3 P4 category
Site: Swartkrans
SK 13/14R 1 1 SAFROB
SK 13/14 L 1 1 SAFROB
SK 24 L 1 - SAFROB
SK 28 L -1 SAFROB
SK 33 R 1 - SAFROB
SK 48 R 1 - SAFROB
SK 49 R -1 SAFROB
SK 52 R 1 1 SAFROB
SK 52 L 1 1 SAFROB
SK 55a R 1 - SAFROB
SK 55a L 1 - SAFROB
SK 65 L 1 1 SAFROB
SK 74c R -1 SAFROB
SK 821 L 1 - SAFROB
SK 822 L 1 - SAFROB
SK 823 R 1 - SAFROB
SK 824 L -1 SAFROB
SK 825 L -1 SAFROB
SK 1590 R 1 1 SAFROB
Site: Sterkfontein
TM 1511 L 1 1 SAFGRA
TM 1512 R 1 - SAFGRA
Sts 1 R 1 - SAFGRA
Sts 1 L 1 - SAFGRA
Sts 30 R - 1 SAFGRA
Sts 47 R 1 - SAFGRA
Sts 52a R 1 1 SAFGRA
Sts 52a L 1 1 SAFGRA
Sts 55 R 1 - SAFGRA
Sts61 R 1 1 SAFGRA
StW/H 45 R -1 SAFGRA
StW/H 73 L 1 1 SAFGRA
Site: Makapansgat
MLD 6 R -1 SAFGRA
MLD 11/30 R 1 - SAFGRA
MLD 23 L 1 - SAFGRA
Site: Kromdraai
TM 1517 L 1 1 SAFROB

In addition to the metric data described above the presence and expression of three
morphological traits were scored for each molar tooth. These were the crista obliqua,
the Carabelli complex and the distal cuspule. The former was scored according to
Korenhof and the definitions of the three subtypes are given in Table 5; the expression
of the Carabelli trait was scored according to a scheme given in the same Table. This
scheme is not a direct adaptation of classifications designed for modem human
populations (see Kieser & Van der Merwe, 1984, for a review). Although they found
Dahlberg's to be the most reproducible scheme, most manifestations of the Carabelli
complex in early hominids take the form of an enamel shelf and the classification used
here reflects this tendency. The distal cuspule was simply scored as present or absent;
the form of a typical distal cuspule is shown in Figure 5. Occlusal wear of the molar
teeth was scored according to a nine point scale adapted from Murphy (1959) and
described in Table 5.
6 B. A. WOOD AND C. A. ENGLEMAN

I KNM-ER 1804
LP4 (reversed)
I
M
M.D. estimated

B L

B.L. maximum
Fig. 1. Diagram showing the linear measurements used in the study as they would be taken on a
typical left maxillary premolar.

IM
SK 13
M.D. I RM'

B
L

estimated

D
Fig. 2. Diagram showing the linear measurements used in the study as they would be taken on a
typical right maxillary molar.
 Hominid maxillary postcanine teeth 7
M

KNM-CH 1
RP3

B L

D
Fig. 3. Diagram to illustrate the measured areas of the major cusp components of a typical right
maxillary premolar.
M

SK 839
RM'

B L

D
Fig. 4. Diagram to illustrate the measured areas of the major cusp components of a typical right
maxillary molar.

A single morphological trait, the expression of the median longitudinal fissure, was
scored for the premolar crowns (Table 9). Occlusal wear of the premolar crowns was
assessed according to the first five points on the scale given for the molars (see
above).
RESULTS
Crown base area
A statistical summary of computed (uncorrected and corrected) and measured
(corrected) crown base areas for each of the major taxonomic categories is presented
8 B. A. WOOD AND C. A. ENGLEMAN
Table 5. Definitions of the scores of morphological traits and occlusal attrition
Crista obliqua (molars only - after Korenhof (1960))
1 A continuous crest between the tip of the protocone and the tip of the metacone, possibly with a
vertical notch in the locality of the longitudinal main groove, but never fully cut through by the latter
2 A similarly situated crest, which however, is distinctly interrupted by the longitudinal main groove
3 A total absence of the crista obliqua. The lingual ridge of the metacone and the distobuccal ridge of
the protocone have not developed more strongly than the adjacent ridges on the protocone and
paracone
Carabelli's trait (molars only)
0 Absent
1 Pit or groove at the mesiolingual corner of the crown
2 Grooves and ridges running from the lingual groove to the mesiolingual corner
3 Definite shelf of enamel related to the protocone
Median longitudinal fissure (premolars only)
1 Fissure is deep and uninterrupted
2 Fissure is evident, but interrupted by enamel ridges leading from the main cusps
3 No fissure evident
Occlusal attrition (premolars 1-4 only)
1 Unworn
2 Enamel wear only, no dentine exposure
3 Dentine exposed on one cusp only
4 Dentine exposed on two cusps
5 Dentine exposed on three cusps
6 Dentine exposed on four cusps, but the dentinal areas still discrete
7 Two dentinal areas coalesced, leaving two free areas
8 Three dentinal areas coalesced, leaving one free area

in Table 6, and individual values for teeth other than those in the taxonomic categories
are given in Table 7. In all cases computed (corrected) crown base area (CCBA)
overestimates the size of a tooth crown if its measured (corrected) crown area (MCBA)
is taken to be the more accurate estimate. The degree of this overestimation varies in
relation to the irregularity of the crown outline. In general, the percentage differences
between the two measurements were greater for the premolar than for the molar teeth.
Within the former, the values for the two 'robust' australopithecine taxonomic
categories were the most discrepant. Otherwise there were no other systematic trends,
indeed the smallest (11 %) and largest (44 %) discrepancies for individual teeth were
both for M2s belonging to the SAFGRA taxonomic group. As was the case with the
mandibular postcanine tooth crowns, there is more overlap between the two major
taxonomic categories from Southern Africa (SAFROB and SAFGRA) than between
the two taxonomic categories from East Africa, even though there is evidence that the
latter samples are both synchronic and sympatric (Walker & Leakey, 1978; Leakey,
Leakey & Behrensmeyer, 1978). In fact, the only overlap in MCBA between the ranges
of EAFROB and EAFHOM is by 6 mm2 for P3.
The size order of the crown areas of the premolars and molars is variable. In all four
major taxonomic categories the mean MCBA values for P4s exceeded that of the P3s.
The percentage differences are 4% for EAFHOM (N = 4), 14% for SAFGRA
(N = 10), 22% for SAFROB (N = 17), and 10% for EAFROB (N = 2). The molar
MCBA size order is M1 < M2 < M3 for SAFROB and SAFGRA; in EAFHOM and
EAFROB it is M1 < M3 < M2. The percentage difference in MCBA between M1 and
M2 is around 15% for all four taxonomic cafegories.
Crown shape
Crown shape has been expressed in the form of an index (BL/MD(C) x 100) and the
parameters for the maxillary postcanine dentition are set out in Table 8. Shape index
 Hominid maxillary postcanine teeth 9
M

/X \ 0 SK 829
X
^ \ LM1 (reversed)

B L

D
Fig. 5. Diagram to illustrate the form of a typical distal cuspule of a left maxillary molar.

values for each taxonomic category have been compared within premolars and
molars; pairwise comparisons which are significantly different are referred to at the
foot of Table 8. What is evident from these comparisons is that the shapes of the
crowns of the postcanine maxillary dentition of the three australopithecine taxonomic
categories are similar. The category with the exceptional shape is EAFHOM, which
shows a tendency for teeth to be mesiodistally elongated in the anterior part of the
postcanine tooth row (i.e. P3, P4, and M1). It is noteworthy that although there is no
significant difference in shape between the maxillary premolars of EAFROB and
SAFROB, the trend for the former to be relatively narrower than the latter, as noted
in the mandibular premolars (Wood & Uytterschaut, 1987), is repeated here.
Morphological traits
The expressions of the median longitudinal fissure, together with the occlusal wear
scores, for the maxillary premolar tooth crowns are given in Table 9. The main
difference is not between taxa, but between tooth types, with nearly all P4s having a
deeply incised fissure. In the sample of P3, teeth belonging to the two 'robust'
taxonomic categories are more likely to have a definite fissure than the non- 'robust'
forms.
The distributions of the scores for the three traits analysed on the maxillary molars
are given in Table 10. It is evident that for two of the traits there is variation along the
tooth row as well as between taxonomic categories. The distributions of the crista
obliqua show no consistent pattern between taxonomic categories. A well-marked
uninterrupted crest is more likely to be found in M's of the East African taxa, whereas
in the australopithecines from Southern Africa, the crest is more likely to be
interrupted. The expression of Carabelli's complex is similarly variable, but in M's
there is evidence of a difference between its appearance in EAFROB and EAFHOM.
The small sample of EAFROB all have the trait in its groove and ridge form, whereas
10 B. A. WOOD AND C. A. ENGLEMAN

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 Hominid maxillary postcanine teeth 11
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12 B. A. WOOD AND C. A. ENGLEMAN

Table 7. Crown base areas of Plio-Pleistocene hominid maxillary postcanine teeth

other than those in the main taxonomic categories (all areas in mm2)
Computed Computed Measured
(uncorrected) (corrected) (corrected)
P3
KNM-ER 808 R 110 110
KNM-ER 1506 R 98 99 79
KNM-ER 1804 L 176 182 148
KNM-ER 1805 R 87 92
KNM-ER 1805 L 94 98 86
KNM-ER 1813 L 92 92 76
KNM-ER 3733 R 107 109 77
KNM-ER 3886 R 161 161 121
O.H. 39 R 104 104 75
O.H. 39 L 101 101 76
P4
KNM-ER 1506 R 109 111 77
KNM-ER 1804 L 195 209 169
KNM-ER 1805 L 103 108 81
KNM-ER 1813 L 98 98 77
O.H. 39 R 108 108 75
O.H. 39 L 96 96 74
Ml
KNM-ER 807 R 177 177
KNM-ER 808 R 169 169 133
KNM-ER 1804 L 233 244 202
KNM-ER 1805 R 173 177 142
KNM-ER 1805 L - 156
KNM-ER 1813 L 156 156 131
O.H. 21 L 146 146 121
O.H. 39 R 140 140 110
O.H.44 R 159 159 126
SK 27 L 185 185 149
SK 47 R 189 189 126
Taung L 173 173 139
M2
KNM-ER 1804 L 285 293 241
KNM-ER 1805 R 201 201 161
KNM-ER 1805 L 173 173
KNM-ER 1808 R 156 156 122
KNM-ER 1813 L 164 164 130
KNM-ER 3733 R 186 186 139
KNM-ER 3733 L 164 164
O.H. 39 R 162 162 123
O.H. 39 L 160 160 123
SK 27 R 201 201 145
SK 47 R 195 195 148
SK 47 L 206 206 163
Stw/H 19B R 174 177 152
M3
KNM-ER 807 R 147 147 114
KNM-ER 1805 R 195 195 148
KNM-ER 1805 L 199 199 154
KNM-ER 1813 R 159 159
KNM-ER 1813 L 148 148
O.H. 15 R 229 229 185
O.H. 15 L 225 225
SK80 L 181 188
Stw/H 19B R 181 186 152
 Hominid maxillary postcanine teeth 13
in most EAFHOM specimens there is no apparent evidence of the Carabelli trait in
any form. The distribution of the distal cuspule, even among the small samples of
MIs, serves to distinguish the taxonomic categories. All nine 'robust' MIs have a distal
cuspule, whereas none of the three EAFHOM teeth, and only two out of four M's
belonging to SAFGRA, do so.
Cusp areas - premolars
A statistical summary of the absolute and relative cusp areas for the taxonomic
categories, and the individual values for the unknown specimens, are given in Tables
11 and 12. When the relative cusp sizes are compared statistically (Table 13), it is
notable that both the P's and P4s of EAFROB have relatively larger buccal cusps than
the other three taxa; the small sample sizes probably explain the reason why the
differences between EAFROB and EAFHOM means do not achieve statistical
significance. As all the premolars examined have no more than the two main cusps,
multivariate analysis of these data is not appropriate.
Cusp areas - molars
Univariate analysis
Statistical summaries of the absolute and relative main cusp areas of the specimens
in each of the four taxonomic categories are given in Tables 14 and 15, and the
absolute and relative cusp areas of the individual specimens are given in Table 16.
When the mean values for the relative cusp areas of the M's of the four taxonomic
categories are compared using Student's t-test, the number of statistically significant
comparisons does not exceed the number that would be expected by chance alone
(Table 17). For M2s and M's, the number of significant differences does exceed that
expected by chance alone, but the distribution of these differences has no obvious
systematic interpretation. One tendency worth remarking upon is the relatively small
hypocone in the M2s and M's of EAFHOM. There are no consistent differences in the
relative size of the protocone either between the two 'robust' taxa, or between the
' robust' and non-' robust' categories. This is in contrast to the results obtained from
the mandibular molar teeth which showed that in the two 'robust' categories, the
distal cusps consistently made a relatively greater contribution to the MCBA (Wood
et al. 1983).
Multivariate analysis
The percentage variance, and the eigen-vector scores of the first and second
principal components (PCm I and II) of the absolute and relative molar cusp areas are
given in Table 18. For the absolute cusp area data, the similarities in the size and sign
of the individual cusp area eigen-vectors suggest that the PCmI of these data is heavily
size-dominated (Jolicoeur & Mosimann, 1960). Inspection of the absolute and relative
cusp area plots shows that there is very little distinction between the four major
taxonomic categories. Empirically, the 'best' separation between taxa is seen on the
MI absolute cusp area plot (Fig. 6) but even so, there is still considerable overlap
between taxa.
DISCUSSION
A general feature of these results is the contrast between the extent of taxonomic
differences as seen in separate examinations of the mandibular (Wood & Abbott,
1983; Wood et al. 1983; Wood & Uytterschaut, 1987) and the maxillary postcanine
dentition. Lucas & Luke (1984) have likened the mandibular tooth crowns to 'pestles'
14 B. A. WOOD AND C. A. ENGLEMAN

Table 8. Crown shape indices (BL/MD(C) x 100) of hominid maxiliary postcanine

teeth
X N S.D. Min. Max.
Premolars
P3
EAFROB 127 2 12 7 118 136
SAFROB* 140 15 6-2 132 150
SAFGRA 135 12 8-2 116 145
EAFHOM 128 5 2-9 125 132
P4
EAFROBt 136 2 07 135 136
SAFROB 142 12 48 133 151
SAFGRA 137 8 6-8 126 148
EAFHOMI 128 6 39 123 132
KNM-ER 808 P3 R 142 KNM-ER 1813 P3 L 140
KNM-ER 1506 P3 R 143 KNM-ER 1813 P4 L 135
KNM-ER 1506 P4 R 134 KNM-ER 3733 P3 R 132
KNM-ER 1804 P3 L 142 KNM-ER 3886 P3 R 143
KNM-ER 1804 P4 L 143 O.H. 39 P3 R 120
KNM-ER 1805 P3 R 144 O.H. 39 P3 L 122
KNM-ER 1805 P3 L 146 O.H. 39 P4 R 112
KNM-ER 1805 P4 L 129 O.H. 39 P4 L 124
Molars
Ml
EAFROB 109 5 7-7 99 116
SAFROB§ 112 14 49 101 121
SAFGRA 107 9 4-8 98 113
EAFHOM 103 8 3-4 98 108
M2
EAFROBI1 106 3 4-3 102 110
SAFROB 112 8 6-4 102 123
SAFGRA 111 13 3-3 104 117
EAFHOM 118 5 6-6 110 125
M3
EAFROB 105 1 105 105
SAFROB 114 18 5.3 103 123
SAFGRA¶ 115 12 51 107 122
EAFHOM** 120 5 5.7 112 125
KNM-ER 807 Ml R 105 KNM-ER 1813 M3 R 118 O.H. 39 M2 L 115
KNM-ER 807 M3 R 113 KNM-ER 1813 Ml L 108 O.H. 44 Ml R 97
KNM-ER 808 Ml R 100 KNM-ER 1813 M2 L 118 SK 27 M2 R 107
KNM-ER 1804 Ml L 111 KNM-ER 1813 M3 L 118 SK 27 Ml L 101
KNM-ER 1804 M2 L 114 KNM-ER 3733 M2 R 108 SK 47 Ml R 105
KNM-ER 1805 Ml R 102 KNM-ER 3733 M2 L 114 SK 47 M2 R 104
KNM-ER 1805 M2 R 114 O.H. 15 M3 R 113 SK 47 M2 L 102
KNM-ER 1805 M3 R 102 O.H. 15 M3 L 124 SK 80 M3 L 124
KNM-ER 1805 M2 L 105 O.H.21 Ml L 105 StW/H 19B M2 R 108
KNM-ER 1805 M3 L 107 O.H.39 Ml R 104 Stw/H 19B M3 R 105
KNM-ER 1808 M2 R 114 O.H. 39 M2 R 111 Taung MI L 116
* SAFROB and EAFROB mean values for P3 are significantly different at P < 0 05.
SAFROB and EAFHOM mean values for P3 are significantly different at P < 0-001.
t EAFROB and EAFHOM mean values for P4 are significantly different at P < 0 05.
$ EAFHOM and SAFROB mean values for P4 are significantly different at P < 0-001.
EAFHOM and SAFGRA mean values for P4 are significantly different at P < 0 05.
§ SAFROB and SAFGRA mean values for Ml are significantly different at P < 0 05.
SAFROB and EAFHOM mean values for Ml are significantly different at P < 0-001.
1 EAFROB and SAFGRA mean values for M2 are significantly different at P < 0 05.
EAFROB and EAFHOM mean values for M2 are significantly different at P < 0-05.
SAFGRA and EAFHOM mean values for M2 are significantly different at P < 0-01.
** EAFHOM and SAFROB mean values for M3 are significantly different at P < 0 05.
 Hominid maxillary postcanine teeth 15
Table 9. Scores for occlusal wear and the expression of the median longitudinal
fissure for the four major taxonomic categories, and the individual cases, of Plio-
Pleistocene hominid maxillary premolars
Median
Occlusal wear* longitudinal fissure*
1 2 3 4 1 2 3
P3
EAFROB 0 2 0 0 1 1 0
SAFROB 3 8 3 1 10 4 0
SAFGRA 1 7 4 0 6 6 0
EAFHOM 2 0 3 0 1 2 1
P4
EAFROB 1 1 0 0 2 1 0
SAFROB 2 7 3 0 11 0 0
SAFGRA 0 4 4 0 4 1 0
EAFHOM 2 2 2 0 4 1 0

Median
Specimen number Occlusal wear* longitudinal fissure*
P3
KNM-ER 808 R 1 1
KNM-ER 1506 R 2 1
KNM-ER 1804 L 4
KNM-ER 3886 R 2 1
O.H. 39 R 1 2
O.H.39 L 1 2
P4
KNM-ER 1506 R 2 1
KNM-ER 1804 L 4
KNM-ER 1813 L 3 1
O.H. 39 R 1 2
O.H.39 L 1 2
* See Table 5 for explanation of the code.

which work in the fixed 'mortars' provided by the maxillary dentition. One of the
conclusions of this series of papers is that, at least in the early hominids examined
therein, there is more intensive selection for details of mandibular tooth form than
for their maxillary counterparts.
Tobias (1967, p. 145) has previously emphasised the need for a carefully defined
measurement system in order to ensure comparability in the measurement of teeth
with a crown outline as irregular as those of maxillary molar and premolar teeth.
Whereas the mandibular molar and premolar dimensions in previous studies (Wood
& Abbott, 1983; Wood & Uytterschaut, 1987) were all within 5 % of the average
values quoted for a similar sample by White, Johanson & Kimbel (1981), the
discrepancy between the average values for the maxillary postcanine teeth cited in that
paper, and in the present analysis, is greater. The average discrepancy is 6%, with
most of the higher percentages being related to the premolar teeth.
Premolar crown morphology
Swindler (1976) has reported that maxillary premolars belonging to extant members
of the Hominoidea are consistently bicuspid teeth. The paracone (that is the buccal
16 B. A. WOOD AND C. A. ENGLEMAN
Table 10. Scores for occlusal wear and the expression of three morphological traits in
the four major taxonomic categories, and the individual cases, of Plio-Pleistocene
hominid maxillary molars
Crista Carabelli Distal
Occlusal wear* obliqua* complex* cuspulet
1 2 3 4 5 8 1 2 3 0 1 2 3 P A
Ml
EAFROB 2 0 0 1 0 0 3 0 0 0 0 3 0 3 0
SAFROB 1 5 3 5 1 0 3 7 2 2 3 0 2 6 0
SAFGRA 0 1 3 2 2 0 0 3 0 1 2 0 2 2 2
EAFHOM 1 2 0 4 1 0 7 1 0 4 1 0 0 0 3
M2
EAFROB 2 1 0 0 0 0 0 1 2 0 0 3 0 3 0
SAFROB 1 7 0 0 0 0 0 4 3 0 2 1 0 6 1
SAFGRA 0 10 1 2 0 0 0 4 6 0 2 0 8 8 4
EAFHOM 1 2 2 0 0 0 2 2 0 0 1 1 1 3 0
M3
EAFROB 3 0 0 0 0 0 0 3 0 0 0 2 0 3 0
SAFROB 5 10 1 1 0 0 1 7 7 6 7 0 2 12 0
SAFGRA 3 7 0 0 0 0 0 1 5 1 1 3 3 7 0
EAFHOM 5 0 0 0 0 0 1 3 0 0 2 0 1 2 0

Specimen Crista Carabelli Distal

number Occlusal wear* obliqua* complex* cuspulet
Ml
KNM-ER 807 R 4 1 A
KNM-ER808 R 2 1 - A
KNM-ER 1804 L 8
KNM-ER 1805 R 4
KNM-ER 1805 L 4 -
KNM-ER 1813 L 5 1 0 A
O.H. 21 L 2 1 1 P
O.H.39 R I
O.H.44 R I 1 0 P
SK27 L 2 2 1 P
SK47 R 3 1 p
Taung L 1 2 0 P
M2
KNM-ER 1804 L 5
KNM-ER 1808 R 2
KNM-ER 1813 L 2 3 0 P
KNM-ER 3733 R 3 1 0
KMN-ER 3733 L 2
O.H. 39 R 1 P
O.H. 39 L 1 2 0 P
SK27 R I I I A
SK47 R 2 1 1 P
SK47 L 2 1 1 P
Stw/H 19B R 5 3
M3
KNM-ER 807 R 2 3 0 P
KNM-ER 1805 R 2 3 1 P
KNM-ER 1805 L 3 2 0
KMN-ER 1813 R 2 3 0
KNM-ER 1813 L 2 3 0
O.H. 15 R 2 0 P
O.H. 15 L 2 0
Stw/H 19B R 3 3 1 A
* See Table 5 for explanation of the code. t P = present; A = absent.
 Hominid maxillary postcanine teeth 17
Table 11. Absolute and relative cusp areas of early hominid maxillary premolars by
taxonomic category.
Buccal cusp (absolute) Buccal cusp (relative)
X N S.D. Min. Max. X N S.D. Min. Max.
P3
EAFROB 70-0 2 18-4 57 83 60-6 2 6-7 55-9 65A4
SAFROB 56-4 14 5-7 49 66 53-0 14 2-2 49-6 55-9
SAFGRA 46-7 12 4-7 40 53 53-1 12 2-9 47-1 564
EAFHOM 50-5 4 13-2 38 69 55-6 4 6-1 49.4 63.9
P4
EAFROB 68-5 2 4-9 65 72 54-3 2 4-6 511 57-5
SAFROB 63-2 12 3-9 58 72 49-0 12 2-4 44-3 52-8
SAFGRA 48-0 7 2-4 45 52 47 9 7 1.9 45-5 50 5
EAFHOM 47-8 5 5-1 42 56 51-0 5 1[5 48-9 52-8

Lingual cusp (absolute) Lingual cusp (relative)

X N S.D. Min. Max. X N S.D. Min. Max.

P3
EAFROB 44 0 2 14-1 43 45 39-0 2 7-3 33.9 44-1
SAFROB 49-6 14 5.5 42 60 46-6 14 2-1 43-1 49-6
SAFGRA 41-7 12 3.9 33 47 47-4 12 2-9 43-6 52 9
EAFHOM 39.3 4 12-6 38 41 44-4 4 6-1 36-1 50 7
P4
EAFROB 58 0 2 15-6 47 69 45 3 2 52 41-6 48-9
SAFROB 66 5 12 7-3 59 79 51-4 12 2-2 48-0 55.7
SAFGRA 51 9 7 3-1 47 56 51-7 7 1-6 49*5 53.5
EAFHOM 46-0 5 60 39 54 49-0 5 2-0 46-1 51.1
The sample sizes do not always accord with the numbers of specimens listed in the Materials and Methods
section. The damaged teeth which have been excluded are listed below.
P3: SAFROB -SK 1590 R P4: EAFROB KNM-ER 733 L
-

EAFHOM KNM-ER 1590 R

- SAFGRA StW/H 45 R
-

EAFHOM KNM-ER 1590 L

-

Table 12. Absolute and relative cusp areas for hominid maxillary premolars other than
those in the taxonomic categories
Buccal cusp Buccal cusp Lingual cusp Lingual cusp
(absolute) (relative) (absolute) (relative)
P3
KNM-ER 1506 R 40 50-6 39 49.4
KNM-ER 1813 L 40 52-6 37 48-7
KNM-ER 3733 R 44 57-1 33 42-9
KNM-ER 3886 R 61 50-4 60 49-6
O.H. 39 R 43 57.3 31 41-3
O.H. 39 L 43 57-3 34 44.7
P4
KNM-ER1506 R 32 41-6 44 57-1
KNM-ER 1813 L 39 50-6 38 49-4
O.H. 39 R 40 53.3 35 46-7
O.H. 39 L 38 51-4 36 48-6
18 B. A. WOOD AND C. A. ENGLEMAN

Table 13. Between-group comparisons ofpremolar relative cusp areas: significance

values based on Student's t-test, using pooled variances and a two-tailed model
Buccal cusp
EAFROB SAFROB SAFGRA EAFHOM

P3
EAFROB
SAFROB < 0-01
SAFGRA < 0-05 N.S.
EAFHOM N.S. N.S. N.S.
P4
EAFROB -
SAFROB < 0-05
SAFGRA < 0 05 N.S.
EAFHOM N.S. N.S. < 0 05

Lingual cusp
EAFROB SAFROB SAFGRA EAFHOM
P3
EAFROB -
SAFROB < 0 01
SAFGRA < 0-01 N.S.
EAFHOM N.S. N.S. N.S.
P4
EAFROB
SAFROB < 0-01
SAFGRA < 0 05 N.S.
EAFHOM N.S. N.S. < 0-05

main cusp) is apparently always larger than the protocone (or lingual main cusp) in
hylobatids, Pan, Gorilla as well as in Homo sapiens; only in Pongo are the main cusps
reported to be subequal in size (Swindler, 1976). Mahler (1973) and Swindler (1976)
also record a trend in the relative size of the crown areas of the two maxillary
premolars with the mean values of the P3 crown area being consistently larger than
those of the equivalent P4s. Thus, it is reasonable to conclude that bicuspid crowns,
a dominant paracone and P3 exceeding P4 in size, are most likely to be among the
primitive traits of maxillary premolars for the African ape/human clade.
Robinson (1956), as usual, provides a reliable and comprehensive guide to the
morphology of early hominid dental remains. He comments on the relative
homomorphy among hominid maxillary premolars, stating that " there is consequently
not a clear distinction between prehominid [i.e. australopithecine] and euhominid [i.e.
Homo] maxillary premolars" (Robinson, 1956, p. 68). However, he did note some
systematic differences between the main samples of 'robust' and 'gracile' australo-
pithecines from Southern Africa. He noted that within the Swartkrans sample there
was particular evidence of talon development in the P4s compared to the P3s. This was
associated with a more deeply incised distobuccal groove and a buccal cuspule, the
latter being found in 12 out of 19 of the Swartkrans sample of P4s. Robinson
commented that the discrepancy in crown morphology between P3 and P4 was more
marked in the Swartkrans teeth than in the substantial, but smaller, Sterkfontein
samples; the sites of Makapansgat and Kromdraai contributed too few specimens to
allow reliable conclusions to be drawn. An associated distinguishing feature of the
 Hominid maxillary postcanine teeth 19
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20 B. A. WOOD AND C. A. ENGLEMAN

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 Hominid maxillary postcanine teeth 21

Table 16. Absolute and relative cusp areas for hominid maxillary molars other than
those in the major taxonomic categories
Protocone Metacone Hypocone Paracone
Absolute Relative Absolute Relative Absolute Relative Absolute Relative
Ml
KNM-ER 808 R 41 30 8 34 25 6 27 20-3 30 22-6
KNM-ER 1813 L 36 27 5 31 23 7 31 23-7 34 26-0
KNM-ER 3733 L 45 32
O.H. 21 L 38 31 4 27 22-3 29 24-0 26 21-5
O.H. 39 R 33 300 26 236 21 191 31 28-2
O.H. 44 R 34 27 0 36 28-6 27 21-4 28 22-2
SK 27 L 45 30-2 37 24-8 31 20-8 36 24-2
SK 47 R 36 28-6 33 26 2 31 24-6 25 19-8
Taung L 39 28-1 38 27-3 30 21 6 31 22-3
M2
KNM-ER 1804 L 80 332 40 16-6 48 199 73 303
KNM-ER 1813 L 47 362 22 16-9 30 23-1 30 23-1
KNM-ER 3733 R 46 33 1 29 20 9 27 19 4 38 27-3
KNM-ER 3733 L 42 - 26 27
O.H. 39 R 34 27-6 28 228 24 195 37 30-1
O.H. 39 L 35 285 25 20-3 23 18-7 38 309
SK 27 R 54 37-2 31 21 4 17 11-7 43 29-7
SK 47 R 43 29-1 34 23-0 40 27-0 31 20-9
SK 47 L 49 30-1 36 22 1 42 258 37 22-7
StW/H 19B R - 32 21-1 34 22-4
M3
O.H. 15 R 65 35-1 37 20-0 46 24-9 39 21-1
StW/H 19B R 49 32-2 26 17-1 35 23-0 43 28-3

* EAFROB
O SAFROB
A SAFGRA
4 A EAFHOM
* Unknown specimens
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A 00
AA A A A
-1 OH 21 LE *SK 47R A,
A
A A 0
A
-2
-3 I I I i
-3 -2 -1 0 2 3
PC
Fig. 6. Plot of the first (PC I) and second (PC II) principal components generated from the covariance
matrix of the absolute cusp area data for early hominid M's.
 22 B. A. WOOD AND C. A. ENGLEMAN

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 Hominid maxillary postcanine teeth 23
' robust' australopithecines in the Swartkrans sample was the larger crown area of the
P4s compared to'the P3s.
Nearly two decades later Sperber (1974) surveyed a similar sample of australo-
pithecine teeth. In some cases he made observations about morphological traits not
considered by Robinson (1956). He was also able to provide more precise metrical
descriptions of any differences in crown morphology by measuring the mesiodistal and
buccolingual diameters of individual cusps and the talon. His data showed that,
despite the larger overall crown size of the Swartkrans teeth, the buccal cusp (or
paracone) made a consistently smaller contribution to the total crown area than it did
in the 'gracile' australopithecine sample from Sterkfontein (e.g. P3: SK = 39 %; Sts
= 51 %; P4: SK = 30 %; Sts = 45 %). He confirmed the tendency for talon formation
in the P4s of Swartkrans, and also showed that P4s were generally broader than P3s in
that sample. Corruccini (1978) has previously used a set of seven linear measurements
to characterise the crown of P3s. The results of his canonical analysis (shape only)
suggest that, far from being distinctive, a sample of six australopithecine teeth drawn
from several sites could not be separated from a modem human sample (Corruccini,
1978, p. 493).
The more accurate measure of crown area used in the present study confirms that
there are differences in overall size between the P3s and P4s of A. robustus (from
Swartkrans), and those of A. africanus (from Sterkfontein) (Table 6). However, the
present study differs from Sperber (1974) in the way it interprets the relative sizes of
the main cusps in the two Southern African taxa. The results in Table 11 suggest that
contra the figures from Sperber (1974) quoted above, in this analysis the buccal cusp
is marginally the larger cusp in both the SAFROB and SAFGRA taxonomic groups.
While there are small differences in the way the cusps are defined in the two studies,
these are unlikely to account for such large discrepancies. The most reasonable
interpretation must be that when crown areas are more accurately measured, the
patterns of taxonomic variation are actually different from those arrived at when
crown areas are estimated by the cruder method of calculating the product of the
length and breadth of an irregular outline. It is noteworthy that a larger buccal cusp
element was part of the hypothesised primitive condition for the African ape/human
clade. The EAFROB sample is small, but its values scarcely overlap with the
EAFHOM group in overall crown area, or in both absolute and relative cusp areas
(Tables 7, 11). The results suggest that the greater size of the teeth of EAFROB is due
to selective expansion of the buccal cusp (Table 11).
The occlusal wear scores show that dentine exposure is more common in SAFGRA
premolars than in the SAFROB sample (Table 9). This may reflect genuine differences
in attrition rates of enamel associated with different masticatory mechanisms in the
two groups (Grine, 1981), but it may also be related to the allegedly thinner enamel
in 'gracile' australopithecines (Robinson, 1963, p. 392). The more deeply incised
median longitudinal fissure in SAFROB (Table 9) accords with Sperber's (1974)
earlier findings.
The size discrepancy between P3 and P4 crown area, noted by Robinson (1956) and
Sperber (1974), is also borne out by the results of this study. What is especially
noteworthy is the demonstration that these more precise measures of crown area only
serve to emphasise the size discrepancy, which is 22 % for SAFROB and 14% for
SAFGRA (P4-P3/P3 x 100). The smaller samples for EAFROB and EAFHOM
provide evidence for much less P3/P4 size discrepancy in these taxa; figures are,
respectively, 10% and 4 %.
24 B. A. WOOD AND C. A. ENGLEMAN

Molar crown morphology

The extant Hominoidea show nearly as much uniformity in maxillary molar
morphology as they do in their maxillary premolar morphology (Korenhof, 1960;
Swindler, 1976). All taxa have four cusped teeth, with Homo sapiens showing the
greatest tendency for reduction in cusp number. This is most strongly expressed in the
distal part of the molar row. The paracone and protone are the largest of the four
cusps in all the extant forms, and an uninterrupted postprotcrista, or crista obliqua,
usually connects the protocone and metacone. All the extant taxa have either a proto-
conal cingulum, or varying incidences of a remnant of it, known as the Carabelli complex
(Robinson, 1956; Korenhof, 1960; Frisch, 1965; Tobias, 1967). The molar size order
in extant apes is M1 <M2 > M3; only in modem humans is it M1 > M2 > M3.
Robinson (1956, p. 96) prefaced his discussion of the comparative maxillary molar
odontology of the 'gracile' and 'robust' australopithecines of Southern Africa by
stating that "the crowns of the maxillary molars of Paranthropus and Australopithecus
are built upon fundamentally the same plan". He concluded that the main difference
between the two taxa was the absolute size of the crown. Other specific points of
contrast he listed were: (a) the relative size of M2 and M3, with the Swartkrans sample
having an M1 < M2 < M3 size order; (b) the stronger expression of the Carabelli trait
in 'gracile' australopithecine teeth; (c) a more marked buccal groove in Paranthropus,
and (d) the relatively greater contribution of the protocone to the total crown area in
the Australopithecus sample. In addition, while he noted several minor points of
difference in detailed enamel morphology he could find no evidence that the crista
obliqua was differently expressed in the two taxa. The value of a later analysis of
hominid maxillary molar morphology by Korenhof (1960) is limited because his
comments are based on a comparatively small sample (just twelve teeth), together with
photographs of casts from the collection of von Koenigswald (Robinson's study was
based on more than one hundred australopithecine maxillary molar crowns). Sperber
(1974) investigated a sample equivalent to Robinson's in size, and also found
considerable overlap in size and shape between his australopithecine samples. There
were, apparently, no significant differences in cusp area between the fossil taxa, but
Sperber did confirm the size dominance of the third molar in the 'robust'
australopithecine tooth row. By, yet again, providing evidence for the stronger
expression of the Carabelli trait in Australopithecus Sperber confirmed an earlier
observation that "the Carabelli complex and its degree of incidence differs quite
clearly in the two genera" (Robinson, 1956, p. 96).
The results of the present study confirm that there is considerable overlap in crown
area, both measured or computed, between the 'gracile' and 'robust' australopithecine
samples from Southern Africa. However, these results do show a much greater, and
statistically significant, difference in overall size between the EAFROB and EAFHOM
samples (Table 6). There is also variation in the molar size order; the SAFROB sample
is the only one in which there is strong evidence for a M1 < M2 < M3 sequence. In the
remaining taxa, M3 is either smaller than M2 (EAFHOM and EAFROB), or they are
subequal in size (SAFGRA). However, contra Sperber (1974), the present study does
suggest that the shape of M1 crowns does differ between 'gracile' and 'robust' forms
in both East and Southern Africa, with, in each case, the 'gracile' samples (SAFGRA
and EAFHOM) having buccolingually narrower MI crowns (Table 8). In this study,
evidence for taxonomic variation in the expression of the Carabelli trait comes from
the M2s and M3s of SAFROB and SAFGRA; Sperber (1974) also noted that the
contrast in expression was greater in the distal molar row. A substantial Carabelli
complex is more common in all three maxillary molars of EAFROB compared to
 Hominid maxillary postcanine teeth 25
EAFHOM; the presence of distal cuspules on M' seemingly characterises the two
'robust' australopithecine samples (Table 10). As noted previously, what variation
there is, in both absolute and relative cusp areas, shows little, or no, systematic
bias.
Maxillary postcanine root morphology
The maxillary premolar roots of most extant non-human primates are three-rooted,
being supported by two buccal and one lingual root (Duckworth, 1923; Colyer, 1936;
Peyer, 1968; Scott & Symons, 1974). In a recent radiographic study, Abbott (1984)
confirmed that more than 90 % of her samples of Gorilla, Pan and Pongo maxillary
premolars are three-rooted. In modern Homo sapiens the evidence for root number for
P3 is confusing. Some studies have indicated that the single-rooted condition occurs in
up to 90 % of samples (Pedersen, 1949), whereas two roots have been recorded in up
to 83 % of other samples (Shaw, 1931). Abbott (1984) found that more than 80 % of
her relatively small, mixed, sample of modern human P3s were single-rooted. Three-
rooted P3s are evidently rare in modern human populations. There is less dispute
about the root form of modern human P4s; they are usually single-rooted. This
evidence would suggest that the most plausible hypothesis for the primitive condition
for maxillary premolar root form in the African ape/human clade is three roots. It
is thus of interest that although all 15 maxillary molar teeth attributed to
Australopithecus afarensis from Hadar are said to be two-rooted, the flattened 'single'
buccal root usually has two root canals (Ward, Johanson & Coppens, 1982). The few
maxillary premolars recovered from Laetoli have either two, or three, roots (White,
1977, 1980).
According to Abbott (1984) the form and number of roots of maxillary molars in
extant non-human higher primates follows the generalised eutherian mammal pattern,
that is, two buccal roots and one lingual root (Gregory, 1920, 1921; Butler, 1941;
Clark, 1971). Partial root fusion has been occasionally noted in non-human primates
(Remane, 1921), but it is well-documented in modern human molars (Campbell, 1925;
Drennan, 1929; Shaw, 1931; Nelson, 1938; Visser, 1943; Abrahams, 1946/7;
Pedersen, 1949; Selmer-Olsen, 1949). Fusion of the distobuccal and lingual roots is the
most frequent fusion pattern seen in M1. In M2 fusion most commonly involves the
mesiobuccal and lingual roots, whereas in M3 neither pattern predominates (Abbott,
1984).
All of those writers who have noted or studied the root form of the maxillary
premolars of the Southern African australopithecines (Broom & Schepers, 1946;
Broom & Robinson, 1952; Robinson, 1956; Sperber, 1974) have recognised a
difference in frequency of two- and three-rooted teeth between the 'gracile' and
'robust' taxa. Robinson (1956) reported that 15 of 20 P3s from Swartkrans were three-
rooted, but in some of these the two buccal roots are partially fused. In contrast, 11
out of 13 P3s from Sterkfontein were two-rooted, with just one (Sts 54) three-rooted,
and a single (Sts 47) one-rooted tooth. Eight out of 10 Swartkrans P4s were recorded
as three-rooted, whereas all 10 Sterkfontein P4s had only two roots. Tobias (1967)
noted that the maxillary premolars of OH 5 were three-rooted. Robinson (1956, p. 66)
is explicit in regarding the three-rooted state of maxillary premolars as the primitive
condition, whereas Abbott (1984) suggests that the root form of 'robust'
australopithecine P3 and P4s may be, in detail, more like that of the maxillary molars
than the three-rooted premolars of extant higher primates. Thus, she interprets their
three-rootedness as a derived trend towards 'molarisation' of the maxillary premolars.
Sperber had made a similar point when he wrote "the robust australopithecine P4s
exhibit a trend towards molarisation by the frequent appearance of a talon on their
crowns and triple roots" (Sperber, 1974, p. 60).
2 ANA 161
26 B. A. WOOD AND C. A. ENGLEMAN

Table 19. List of specimens of East African early hominids for which there is
information about maxillary premolar root form
P3
KNM-ER 1590B L ?Three roots
KNM-ER 1805 R ?Two roots
KNM-ER 1813 L Two roots
P4
KNM-ER 733D L Three roots
KNM-ER 1506B R One or two roots
KNM-ER 15901 L Two or three roots
KNM-ER 1804 L Three roots
KNM-ER 1813 L Two roots

Table 20. Summary list of 'unknown' specimens

P3 P4 MI M2 M3
KNM-ER 807 1 1
KNM-ER 808 1 1
KNM-ER 1506 1 1 - -
KNM-ER 1804 1 1 1 1 -
KNM-ER 1805 2 1 2 2 2
KNM-ER 1813 1 1 1 1 2
KNM-ER 3886* 1
O.H. 15 2
O.H.21* 1
O.H. 39 2 2 1 2 -
O.H. 44* 1
SK27 1 1 -
SK47 1 2 -
SK 80t 1
StW/H 19B 1 1
Taung 1
* Indicates isolated teeth.
t The data for this specimen did not allow affinities to be suggested on the basis of measured crown base
area.

When compared with the richness of the data for the isolated teeth from the
Southern African cave sites, there is a paucity of reliable information from the more
intact cranial material from Koobi Fora and Olduvai. The interpretations of premolar
root form that can be made are given in Table 19, and, where relevant, these will be
referred to in the next part of the discussion.
Affinities of 'unknown' specimens
Sixteen of the 98 individual specimens included in this study were not assigned to
one of the four main taxonomic categories; these teeth are listed in Table 20. Their
exclusion from the main taxonomic groups was because they consisted of dental
remains alone (e.g. KNM-ER 808), or were teeth embedded in fragmentary specimens
(e.g. KNM-ER 1804), or because they belonged to specimens whose taxonomic
affinities have been the subject of debate (e.g. KNM-ER 1805 and 1813). Five
measurements, or categories of observations, have the potential to provide information
about affinities. Four of these, measured crown base area (MCBA), crown shape,
 Hominid maxillary postcanine teeth 27

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30 B. A. WOOD AND C. A. ENGLEMAN
absolute and relative cusp area, have had their potential information deduced in the
same way. The raw data were assembled into a covariance matrix from which
Canonical Variates have been extracted using Canonical Variates Analysis (CVA).
The SPSSX statistical programme includes the facility to compute posterior
probabilities and these have been used to calculate the percentage likelihood that a
specimen should be assigned to a particular taxonomic category. Two designs were
used to examine the affinities of each of the regional specimen groups. In the first of
these, East African specimens were 'offered' the choice between EAFHOM and
EAFROB; these results are presented for maxillary premolars and molars in Tables
21 and 22. The second design for the East African specimens added the choice of
SAFGRA; an asterisk identifies the specimens whose affinities were altered by this
addition. The first of the two designs for Southern African specimens offered the chloice
between SAFGRA and SAFROB; these results are also presented in Table 22. The
addition of EAFHOM constituted the second regional design, and if this affected
affinities, it too is indicated by an asterisk in Table 22. The efficacy of the test was
examined by applying it to the taxonomic categories themselves. Affinities have only
been recorded on Tables 21 and 22 if 66%, or more, of the 'test' specimens were
correctly classified. Likewise, affinities of the 'unknown' specimens have only been
recorded when the percentage likelihood was 66%, or more. The fifth category of
observations, the incidence of morphological traits, was not susceptible to this type of
analysis, nor, with one or two exceptions, were the traits particularly taxonomically
discriminating. Thus, reference to the value of traits will only be made when relevant
to a particular specimen; no general affinities have been recorded. The specimens will
be considered in numerical order of reference number within each site collection.
The first, KNM-ER 807, comprises the three right upper molars of an adult
dentition (Leakey & Wood, 1973); M2 and M3 are embedded in a fragment of maxilla,
but it is not well enough preserved to be helpful for taxonomy. The MCBA of M3 (114)
is below the range for EAFHOM, and well below that of SAFGRA. There is no
evidence of Carabelli's complex, nor of a distal cuspule. These items of evidence,
together with its small size, would suggest that the original assignment of these teeth
to Homo (Leakey & Wood, 1973) is still soundly based.
The next specimen, KNM-ER 808, is an immature dentition which includes the
right MI germ included in this analysis. The affinities of this tooth are mixed (Table
22). Its small size would suggest strong affinities to EAFHOM, but its absolute and
relative cusp areas and the presence of a well-marked Carabelli complex suggest
otherwise. The small size may be accounted for by its immaturity, but it would be
unusual for teeth at its stage of growth to enlarge significantly. It must be concluded
that M1 morphology can give no firm indication of the affinities of this specimen.
The specimen KNM-ER 1506 comprises part of the right side of the body of a
mandible, the crowns of M1 and M2, together with two maxillary premolar teeth. The
initial description commented upon the combination of small size and high mandibular
robusticity values (Leakey, 1973, p. 171), and it was regarded as an example of a small-
bodied (presumably female) 'robust' australopithecine. However, information
provided by the morphology of the two mandibular molar teeth suggested that the
affinities of the specimen are with Homo and not Australopithecus boisei (Wood et al.
1983). The balance of the evidence of the two maxillary premolar crowns also suggests
hominine affinities. While its P3 crown is large, the overlap in overall size between
taxonomic categories is relatively great (Table 21). The crown area of the P4 is a more
reliable indicator (Table 21), and its assignment is firmly to EAFHOM.
The maxillary fragment, KNM-ER 1804, which bears two premolars and two
 Hominid maxillary postcanine teeth 31
molars, is badly damaged. When it was first described (Leakey, 1974), KNM-ER 1804
was not attributed to a specific taxon, but the excessive size of both the premolar and
molar crowns has determined its high percentage affinity for the EAFROB group
(Table 22).
Although the skull KNM-ER 1805 provides evidence about cranial, mandibular
and dental morphology, its affinities have been sufficiently enigmatic to sustain
attributions to Homo erectus (Howell, 1978; Wolpoff, 1978), Homo africanus (Olson,
1978) and Australopithecus boisei (Tobias, 1980). However, the evidence of the cranial
base (Dean & Wood, 1982), the mandible (Chamberlain & Wood, 1985), the size of
the mandibular molar teeth (Wood & Abbott, 1983) and now the evidence of its
maxillary postcanine dentition (Tables 21, 22) suggest that its affinities lie with Homo
and not with A. africanus, or A. boisei. The only conflicting evidence is the shape of the
MI crowns (Table 22), which suggest affinities with EAFROB. However, a level of
discrimination of only 75 % for the test sample exposes the relatively poor
discriminating power of M2 shape.
The cranium KNM-ER 1813 is also an enigmatic specimen. It has been assigned to
both A. africanus (Walker & Leakey, 1978) and to either a defined (Howell, 1978), or
an undefined (Wood, 1985), species of Homo. The evidence provided in this study is
not presented in a way that would allow discrimination between early Homo taxa, but
the affinities of the premolar and molar teeth (Tables 21, 22) are clearly with Homo and
not with the SAFGRA group. A similar conclusion was reached on the basis of cranial
base morphology (Dean & Wood, 1982), but a morphometric study of the face
(Bilsborough & Wood, 1988) showed some phenetic resemblances to A. africanus.
KNM-ER 3886 comprises a maxillary molar (which has not been identified in more
detail) and a right P3 crown. The latter's size alone (Table 7) would suggest an
attribution to EAFROB, and this conclusion is not weakened by the counter proposal
suggested by the relative cusp area. It is apparent from Tables 11 and 21 that
attributions based on this criterion are not soundly based, for barely two thirds of the
reference group are correctly allocated.
Details of the Olduvai specimens included in this analysis can be found in Leakey
(1971, 1978). These remains come from Beds I and II of Olduvai Gorge, either in situ,
or as surface finds, and their firm attribution to EAFHOM indicates that their
affinities must be sought in early Homo, specifically in H. habilis, H. erectus or
perhaps a third Homo taxon (Wood, 1985). In a summary of Olduvai hominids,
Leakey (1978) allocated OH 15 to Homo sp., OH 21 and 44 to H. habilis and regarded
OH 39 as cf. H. habilis. Tobias & von Koenigswald (1964) and Tobias (1965)
suggested that OH 15 may belong to Australopithecus, but the results of this analysis
strongly suggest that this is not the case and Rightmire (1980, p. 229) has previously
concluded that "an alternate assignment to Homo sp. cannot be ruled out".
The results of the affinities of the 'unknown' specimens from Southern African sites
have been included more for record than in the belief that they constitute
authoritative evidence. This caveat is introduced because of the generally low
percentage levels of correct attributions for the Southern African reference samples
(Table 22). It is not the first time that hominine affinities have been suggested for
SK 27 and SK 47. The former is a juvenile cranium. Clarke (1977b, p. 48) listed eight
features of SK 27 which, he believed, suggested affinities with Homo; " relatively small,
narrow molars" were listed as his fifth feature. Olson (1978) drew attention to features
of the bony cranium and teeth of SK 47 which he regarded as hominine. However,
Dean & Wood (1982) concluded that, despite its immaturity, the cranial base of
SK 47 was closer to the 'robust' australopithecine than to the Homo pattern. The
32 B. A. WOOD AND C. A. ENGLEMAN
balance of the morphometric evidence gathered in the course of this study suggests
affinities between SK 27 and 47 and the 'gracile' australopithecines. However, the
presence of distal cuspules on the molar crowns is apparently a 'robust'
australopithecine trait.
The M3 of SK 80 makes up part of the so-called 'composite cranium' which is
usually referred to by the specimen number of the largest fragment, SK 847 (Clarke,
Howell & Brain, 1970; Clarke & Howell, 1972). A detailed analysis of this cranium
concluded that these remains should be assigned to an unnamed species of Homo
(Clarke, 1977 b), but others have regarded it as conspecific with A. robustus (Wolpoff,
1974) or the Sterkfontein hypodigm of A. africanus (Olson, 1978). The only datum
available for the SK 80 M3 in this study is the computed crown area of 188. This lies
within the range of SAFROB and SAFGRA, but just above the range of EAFHOM.
Little can be contributed to the taxonomic debate on the basis of this evidence, but it
must be said that it would represent a large M3 for a Homo cranium.
The single maxillary molar from Sterkfontein, StW/H 19B, was attributed to
SAFGRA, but by the slenderest of margins. The Taung MI is aligned with SAFGRA
by its size and with SAFROB by its shape. It is yet further evidence that this juvenile
skull, which is the type specimen of A. africanus, shows a mixture of 'gracile' and
'robust' australopithecine traits, but with the former predominating.

SUMMARY
A total of 139 maxillary molar crowns and 79 maxillary premolar crowns, from at
least 98 individual East and Southern African Plio-Pleistocene hominids, has been
subjected to detailed morphometric analysis. All but 16 of the 98 specimens were
assigned to taxonomic categories identified as EAFROB, EAFHOM, SAFGRA,
SAFROB and EAFHER. The analysis was based on whole crowns and the
component cusps. While there was variable overlap between the ranges of measured
crown base area of the two Southern African taxa, there was little, or no, overlap
between the two major East African taxonomic categories. Crown shape distinguished
EAFHOM from the three other australopithecine taxa, especially for P3, P4 and M1.
Of the non-metrical traits, the expression of Carabelli's complex and the incidence of
a distal cuspule discriminate best between the categories. Analysis of the absolute and
relative cusp area data shows that the major taxonomic distinction in relative cusp
area is in the premolars, in which it is apparent that EAFROB are distinguished by
their larger buccal cusps.
The principal conclusions of the assessment of the specimens in the 'unknown'
category is that the postcanine dentitions of a skull, KNM-ER 1805, and a cranium,
KNM-ER 1813, are closest in size and shape to EAFHOM. There is no dental
evidence to suggest that these specimens should be assigned to A. africanus, the formal
taxon making up the SAFGRA category.
We thank the custodians of all the fossil collections which were included in this
analysis, and the Directors, Trustees and staff of those institutions in which the
collections are housed, for permission to study the remains and for generous assistance
and hospitality while the work was being undertaken. Particular thanks are due to
Richard Leakey, the Trustees of the National Museums of Kenya and the Government
of Kenya, for allowing B.A.W. to make a detailed study of the Koobi Fora remains.
Susan Abbott's contribution to data collection was a crucial element of this study;
 Hominid maxillary postcanine teeth 33
her assistance is gratefully acknowledged; Hilde Uytterschaut helped with the
multivariate analysis. This research was supported by the N.E.R.C.

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