A partial skull of Paranthropus
robustus from Cooper’s Cave,
South Africa
Christine Steininger*‡, Lee R. Berger* and
Brian F. Kuhn*
Introduction
A partial skull, COB 101, was identified in the collections of
the Transvaal Museum, Pretoria.1 The fossil was found in the
collections that were attributed by museum records to Cooper’s
Cave. Two other ex situ hominin specimens have previously
been described from early collecting activities—an upper third
molar attributed to Australopithecus africanus2,3 and an upper
central incisor attributed to Homo sp. indet.4 In situ hominin
specimens have been recovered from the Cooper’s D locality by
more recent excavation activities and these are currently being
studied and assessed.5
The Cooper’s Cave site is hosted in dolomites of the Monte
Cristo Formation (Malmani Subgroup, Transvaal Supergroup).
The site is situated approximately 45 km northwest of Johannes-
burg, half-way between Sterkfontein to the west and Kromdraai
to the east. Given our understanding based on recent excavations,
Cooper’s Cave comprises three fossil-rich areas, provisionally
recognized as Cooper’s A, B, and D localities, with most of the
fauna having been recovered from Cooper ’s D. Based on
biostratigraphy, the age of the excavated deposits of Cooper’s D
is estimated at between 1.6 and 1.9 million years ago (Mya).5,6
This is penecontemporaneous with Sterkfontein Members 4 and
5,7–10 Swartkrans Members 1, 2 and 3,11–13 Kromdraai A and B,12,14
and Drimolen.15,16 Radiometric dating methods are currently
being applied to the various Cooper’s Cave sites to try to obtain
more precise dates. Initial radiometric results are, however,
comparable to the biostratigraphic age-estimations (R.
Pickering, pers. comm.)
The purpose of this paper is to provide a description and
analysis of the hominin specimen, COB 101, which evinces
morphology not previously known for the Cooper ’s Cave
hominin assemblage.
Materials and methods
COB 101 was compared with original hominin specimens of
A. africanus (n = 17), Paranthropus robustus (n = 15), Homo habilis
(n = 7) and H. ergaster (n = 1), and high-resolution casts of
A. afarensis (n = 3), P. aethiopicus (n = 1) and P. boisei (n = 6)
(Table 1). Observations were aided by stereomicroscopy. All
dimensions were chord distances taken with digital callipers to
the nearest 0.1 mm. Because of the nature of the preservation of
COB 101, measurements of breadth on all taxa were taken from
the midsagittal plane. Non-metric facial parameters follow Lock-
wood17 and Rak.18 Facial measurements follow Martin and
Knußmann19 and Lockwood.17 Dental diameters were measured
following Wood.20
Description
Only the right portion of the upper face, including an upper
right third premolar, is preserved in COB 101 (Fig. 1). While the
specimen is distorted and has post-depositional damage. The
Research Letters South African Journal of Science 104, March/April 2008 143
*Institute for Human Evolution and the Bernard Price Institute for Palaeontological
Research, School of Geosciences, University of the Witwatersrand, Private Bag 3, WITS
2050, South Africa.
‡Author for correspondence. E-mail: christine.steininger@student.wits.ac.za
A partial hominin skull (COB 101) was identified in the fossil collec-
tions of the Transvaal Museum, Pretoria, attributed to the Cooper’s
Cave site in South Africa. The find represents the most complete
hominin specimen recovered from localities at this site to date.
COB 101 comprises the supraorbital, zygomatic, infraorbital and
nasoalveolar regions of the right side, and the right upper third
premolar. The specimen has undergone post-depositional distortion
that resulted in the flattening of the facial structures. Here we
describe and compare COB 101 with other hominin material from
Africa and find that this specimen shares numerous diagnostic
features with Paranthropus robustus. The discovery of COB 101
augments the number of specimens attributed to this species from
other South African sites and other Cooper’s Cave localities.
Fig. 1. Stereo-photograph of the anterior view of COB 101 (scale in cm) with detail of the occlusal view of the right P3 inset (not to scale).
supraorbital area, the right side of the glabella
and torus are preserved. The supraorbital
and zygomatic regions are slightly separated
but held together by adhering breccia. The
zygomatic has been displaced from its anatom-
ical position and is compressed against the
maxilla. The inferior part of the infraorbital
and the nasoalveolar regions are the best-
preserved areas of the specimen, with the only
damage being slight weathering. Age-related
features are not well preserved, but based on
moderate wear of the permanent premolar,
the individual is almost certainly an adult,
comparable in dental wear stage to P. robustus
specimens SK 49 and SK 57 from Swartkrans.
The temporal line of COB 101 is placed high
and is well marked on the frontal squama,
indicating the probable presence of a sagittal
crest. Recent studies on sexual dimorphism
on P. robustus suggest that a prominent
sagittal crest reaching onto the frontal bone is
consistent with larger males of this species.21
Accordingly, even though this specimen is
damaged it is possible to state that it is proba-
bly a male. The supraorbital torus is continu-
ous, similar in morphology to Clarke’s ‘supraorbital rib’
attributed to paranthropines22 On the torus, just lateral to the
glabella, a small crescent-shaped supraorbital notch changes the
relief of the torus slightly. Overall, the torus is thin, 6 mm at its
thickest vertical height and vermiculated throughout. The
superciliary eminence is absent and this trait is consistent with
other specimens attributed to P. robustus. Anteriorly, the pre-
served portion of the glabella is flat and there seems to be varia-
tion of the glabella prominence among most species included in
this study, except for P. aethiopicus and P. boisei, where the glabella
is consistently marked (Table 2).
The zygomaticomaxillary fossa of COB 101 is deep. This feature
has been noted as dominant in paranthropines and A. afarensis.18
The inferolateral orbital margin is rounded, and is similar in
appearance to that of A. africanus and P. robustus. The zygomatic
prominence is slight, located inferiorly on the specimen, close to
the masseteric origin (Table 2).
On the maxilla, the marginally distorted anterior pillar slightly
overhangs the nasal aperture. The widest portion of the preserved
anterior pillar is 10.6 mm. The anterior pillar is found in most
A. africanus specimens and all P. robustus specimens. A marked
canine–premolar eminence forms the inferior part of the ante-
rior pillar. The maxillary sulcus is preserved immediately below
the infraorbital foramen. A large canine fossa, at the same level
144 South African Journal of Science 104, March/April 2008 Research Letters
Table 1. List of early hominin specimens used in this study.
Taxon Site Age (Mya) *Specimens
A. afarensis Hadar 3.4 AL 200-1
3.18 AL 333-1, 333-2
A. africanus Makapansgat ~3.2 MLD 6, 9, 45
Sterkfontein 2.1–2.8 TM 1511, 1512, 1514
Sts 15, 53, 63, 71, 3009
Stw 13, 73, 360, 370, 391, 505
P. aethiopicus West Turkana 2.5 KNM-WT 17000
P. boisei Koobi Fora 1.66–1.76 KNM-ER 406, 732
1.83–193 KNM-ER 13750
Olduvai Gorge 1.7–1.85 OH 5
West Turkana 2.5 KNM-WT 17400
Chesawanja 1.5–1.6 KNM-CH 1
P. robustus Kromdraai 1.3–1.8 TM 1517
Swartkrans 1.5–1.8 SK 11, 12, 46, 48, 65, 79, 83, 821, 826, 845
SKW 11, 12
1.5–1.8 SKX 265
Drimolen 1.5–2 DNH 7
H. habilis Sterkfontein 2.4–2.8 Stw 53
H. habilis Koobi Fora 1.87–1.92 KNM-ER 1470, 1813
1.8–1.9 KNM-ER 1805, 3732
Olduvai Gorge 1.7–1.85 OH 16
1.7–1.85 OH 24
H. ergaster Swartkrans 1.5–1.8 SK 847
*For the East African specimens, only high resolution casts were used.
Table 2. Non-metrical character states for each hominin taxon.
Taxon n Supercilliary Temporal line Glabella Inferolateral Zygomatico- Anterior Maxillary
eminence prominence orbital margin maxillary fossa pillars sulcus
Character state: Absent Present Moderate High Flat Slight Marked Round Sharp Absent Present Absent Present Absent Present
COB 101 X X X X X X X
A. afarensis 3 1 1 1 1
A. africanus 17 3 2 2 2 3 2 2 3 1 2 14 2 4
P. aethiopicus 1 1 1 1 1 1 1 1
P. boisei 6 4 5 4 5 1 1 3 1 1
P. robustus 15 4 4 1 1 1 5 1 7 14 10
H. habilis 7 4 2 6 1 4 1 2 2 4 1
H. ergaster 1 1 1 1 1 1
Taxon n Canine fossa Maxillary Subforamen Lateral nasal Prenasal fossa Inferior nasal Anterior nasal
furrow divide margin margin spine
Character state: Absent Present Absent Present Absent Present Round Sharp Absent Present Smooth Sharp Absent Anterior Posterior
COB 101 X X X X X X X
A. afarensis 3 2 1 1 1 2 2 2 1
A. africanus 17 3 2 4 7 4 7 13 1 9 2 6 9 1 2 3
P. aethiopicus 1 1 1 1 1 1
P. boisei 6 2 3 3 2 1 1 1 2 2 3
P. robustus 15 6 6 9 4 6 10 2 4 4 13 8
H. habilis 7 1 1 1 2 1 2 1 3 1 2 1
H. ergaster 1
The number of individuals is given for each character state.
as the inferior nasal margin, faces later-
ally. A maxillary furrow, which is a
variable though prominent feature of
most A. africanus specimens, is absent
in COB 101. The subforamen divide is
present in COB 101 as a thin marked
ridge, starting just above the canine
alveolar margin that diverts laterally
slightly below the infraorbital fora-
men,thereby framing the maxillary
sulcus. The combination of the anterior
pillars and the subforamen divide, and
the lack of a maxillary furrow, is consis-
tent with most P. robustus specimens
(Table 2).
In the nasoalveolar region, the lateral
nasal margin is rounded with a deep
prenasal fossa24 behind it. The depth of
the fossa may be exaggerated by the dis-
tortion of the anterior pillars. The nasal
spine is posterior from the inferior
nasal margin. The nasoalveolar gutter is
present, sloping smoothly up into the
nasal aperture with no distinct nasal
sill present. The slight alveolar prog-
nathism observed is exaggerated by the
incisor eminences.
Though we were able to obtain some
measurements in the nasoalveolar re-
gion of the face, these measurements
provide only estimates due to the
crushed and distorted nature of COB
101. The measurements are as follows:
alveolar height is c. 23.2 mm, nasal breadth is c. 13.1 mm, and
snout breadth is c. 22.7 mm.
The third premolar crown is almost complete, but is crushed
mesiodistally and buccally. The only aspect of the tooth that is
not damaged is the lingual face and a small part of the
mesiolingual face. Part of the occlusal surface of the protocone is
missing and the occlusal surface of the paracone is partially
crushed. The occlusal surface and the mesial face are stained by
manganese. The tooth is moderately worn, with small patches of
dentine visible in the area of the protocone. This surface is worn
flat and the occlusal edges of the tooth are rounded. Although
the mesial face of the tooth is distorted and slightly crushed,
interstitial wear from the canine is still evident and the crown
has retained its oval shape. The buccal face is moderately convex
occlusocervically, and more so mesiodistally, giving the tooth a
bulbous appearance. Buccal grooves are not present. Perikymata
are visible on the mesiobuccal aspect of the tooth. The lingual
face is similar to the buccal face in its convexity, but is much
wider along the cervical margin. Because of damage, the
mesiodistal diameter was measured across the least crushed part
of the tooth, across the protocone just below the central fissure.
The estimated crown diameters are: mesiodistal c. 9.9 mm,
buccolingual c.14.5 mm, and preserved crown height c. 6.7 mm
(Table 3).
Discussion
The combination of features in the supraorbital and the
nasoalveolar region in COB 101 are similar to these morphological
areas in P. robustus (see Table 2). These features tend to be related
to robusticity and degree of development, and it is important to
note that the morphology of the nasoalveolar region is often
used to separate paranthropines from australopithecines.19,24
The presence of an anterior pillar and the lack of a maxillary
furrow are considered diagnostic features found in most
P. robustus specimens.18 In this regard COB 101 most closely
resembles two P. robustus specimens from Swartkrans (SK 48 and
SK 79).
Morphologically, the molarized upper third premolar of
COB 101 is similar to specimens attributed to P. robustus and
P. boisei. Notable features are the oval crown, large bulbous
appearance of the tooth, and the flat wear on the occlusal
surface. The mesiodistal diameter of COB 101 is well within the
specimens attributed to P. robustus (9.8 ± 0.5 mm, n = 23) and
A. africanus (9.1 ± 0.5 mm, n = 28) where the buccolingual
diameter falls within only the P. robustus range (14.2 ± 0.9 mm).
The size of the upper premolar of COB 101 indicates affini-
ties with P. robustus (Fig. 2). We hesitate, however, to assume
that these dimensions provide evidence for placement within
P. robustus. There are two reasons for caution: (1) the damaged
state of the upper third premolar means that our measurements
are estimates, and (2) mesiodistal and buccolingual diameters of
upper third premolars are recognized as being highly variable
among hominin species, with a particularly high variation in
A. africanus and P. robustus24–27 (Fig. 2).
In summary, most of the morphological evidence in the
supraorbital and the nasoalveolar regions of COB 101 is consis-
tent with other specimens attributed to P. robustus, particularly
SK 48 and SK 79. In addition, the morphology and wear of the
upper third premolar provides further evidence for associating
the specimen with the genus Paranthropus. The evidence reviewed
above suggests that COB 101 is best attributed to P. robustus.
As COB 101 is of unknown provenance, its precise association
Research Letters South African Journal of Science 104, March/April 2008 145
Table 3. Descriptive statistics for dental measurements (in mm).
Measurement Taxon n Min. Max. Mean s.d. CV%
Mesiodistal diameter COB 101 c. 9.9
A. africanus 28 7.9 10.6 9.1 0.5 5.7
P. robustus 23 9 10.8 9.8 0.5 5.3
H. habilis 10 8 9.5 8.9 0.5 5.6
Buccolingual diameter COB 101 c. 14.5
A. africanus 27 10.7 14.4 12.7 0.9 6.8
P. robustus 22 12.7 15.6 14.2 0.9 6.1
H. habilis 8 11 12.7 11.7 0.6 5.3
Fig. 2. Crown diameters of COB 101 upper third premolar compared with those of other early hominins.
with other Paranthropus material recovered from Cooper’s D, or
the Cooper’s specimens attributed to A. africanus and Homo,
remains unclear. Owing to the possibility of a temporal overlap
between latest A. africanus (c. 1.5–2.1 Mya),7–9 and earliest P. robus-
tus (c. 1.9 Mya),11,13 and considering the early dates proposed
for some of the deposits at Cooper’s,5 the presence of all three
species at this site is plausible.
We thank two reviewers for their helpful comments. Thanks also go to Francis
Thackeray, Stephany Potze and Heidi Fourie of the Transvaal Museum, Pretoria,
for their assistance with the hominin material in their care. We wish also to thank
Andre Keyser and Colin Menter for allowing us brief access to a Drimolen
specimen, DNH 7. This research has been funded by the Palaeontological Scientific
Trust (PAST) and the University of the Witwatersrand.
Received 12 October 2007. Accepted 2 March 2008.
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146 South African Journal of Science 104, March/April 2008 Research Letters
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/DAN <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>
/NLD <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>
/ESP <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>
/SUO <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>
/ITA <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>
/NOR <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>
/SVE <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>
/ENU <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>
>>
>> setdistillerparams
<<
/HWResolution [1200 1200]
/PageSize [612.000 792.000]
>> setpagedevice