The endocranium of MLD 37/38 - New insights using Virtual Anthropology
MLD 37/38 is an incomplete but well-preserved, about 3 million years old Australopithecus africanus cranium, found embedded in two blocks of pink breccia mined in the Limeworks cave at Makapansgat, South Africa, in 1958/59 (Dart, 1959, 1962). Most of the face and parts of the frontal neurocranium are missing. Furthermore, the endocranial cavity is filled with stone matrix so that the endocranial capacity of MLD 37/38 could only be roughly estimated earlier and the endocranial morphology could not be analyzed at all with the traditional methods.
Using the virtual representation of MLD 37/38 obtained by CT scanning, segmenting the natural endocast and reference-based reconstructing of the missing frontal endocast portion, this work gives a new estimate of the endocranial capacity, reveals some features of this specimen’s endocranium that where inaccessible so far and thus adds to the knowledge of the endocranium of the species Australopithecus africanus .
The segmented endocast comprises the calvarial surface up to
approximately the coronal suture, left of endobregma missing small
portions and on the right side enclosing also parts of the frontal
lobe, both occipital lobes, both cerebellar lobes, the entire right
temporal lobe and also most of the left temporal lobe only missing very
anterior parts.
The volume of the preserved natural endocast has been determined to be
378 cm³ and the total endocranial capacity has been estimated to be 450
cm³, using Sts 5 as reference-specimen. This estimate is significantly
smaller than the first estimate by Dart (1962, 480 cm³) and larger than
the more recent estimates of Holloway (1972, 435 cm³) and Conroy et al.
(1990, 425 cm³). The sample mean of endocranial capacity for
Australopithecus africanus thus increases to 454 ± 32 cm³
(n=7).
Beside the endocranial capacity, other qualitative and quantitative
data regarding the shape of the endocast were examined. In occipital
view, the endocast is round in shape, in superior view egg-shaped. In
general, the endocast of MLD 37/38 is lower and wider than that of Sts
5, StW 505, Sts 71 and Taung. Only Sts 60 is even more lower and wider
than the endocast of MLD 37/38.
The four measured distances (projected measurements of the cranial
base, after Falk et al., 2000) for MLD 37/38 lie well within the range
of the other available Australopithecus africanus specimens.
Endocasts of Australopithecus africanus are more similar to
those of Homo , while Paranthropus endocasts are more
ape-like in these metric aspects. Because a handful of distance
measurements provide only restricted information of the shape, the
shape of endocasts should be analyzed in the future using geometric
morphometrics (see Neubauer et al., 2006).
Other qualitative aspects of endocasts are convolutional details and
asymmetries. While other natural and artificial endocasts, regardless
if real or virtual endocasts, can show detailed convolutions, no such
convolutional details could be found on the endocranial surface of MLD
37/38. Also, the very controversially discussed lunate sulcus is not
detectable on this specimen. The slight displacement of the right
parietal bone over the left causes small asymmetries of the endocast. A
small right occipital petalia can be seen.
Furthermore, endocasts give evidence of venous drainage from the
brain and arterial supply to the internal table of bone and the dura
mater. MLD 37/38 shows a well-developed lateral venous drainage system
(transverse and sigmoid sinuses) on both sides while there is no
evidence for an enlarged occipital-marginal sinus system. Except for
the juvenile Taung fossil, all known Australopithecus africanus
specimens exhibit this pattern of cranial venous drainage. In contrast
to the condition in Australopithecus africanus, enlarged
occipital-marginal sinus systems are known in high frequencies in
Australopithecus/Paranthropus robustus ,
Australopithecus/Paranthropus boisei , and Australopithecus
afarensis and have been used as a claim to robust australopithecine
membership (Tobias, 1968; Falk and Conroy, 1983; Kimbel, 1984; Falk,
1986; White and Falk, 1999; Falk et al., 2000; Holloway et al.,
2002).
Internal blood supply of the brain is not detectable on an endocast but
meningeal vessels frequently leave imprints on the internal table of
bone. Several branches of meningeal vessels could be detected on the
left side of the endocast of MLD 37/38. On the right side only
rudimentary branches can be seen because the cranium broke apart on the
boundary between fossilized bone and the natural endocast in this area.
While one branch on the left side gives evidence that the middle third
of the braincase was supplied by the external carotid artery through
the foramen spinosum, another branch on the ride side stems from the
orbit and therefore originates from the internal carotid artery.
Because statistical analysis (n= 43 apes, Chi2=18.92, df=1, p<0.001,
Falk, 1993) showed that Falk’s pattern A (blood supply by meningeal
vessels originating from the internal carotid artery with an increasing
extent of supply by the middle meningeal artery from A1 to A4) or E
(blood supply alone by the middle meningeal artery originating from the
external artery) in one hemisphere strongly predicts the same pattern
in the other, it can be concluded that there was blood supply from
both, the internal and external carotid artery to the middle third of
the braincase (pattern A2, A3 or A4).
Furthermore, the stone matrix was removed from some apertures and
foramina so that they can be followed from outside inwardly. Parts of
the right orbit were preserved within the stone matrix. The superior
orbital fissure was found to be round in shape and therefore should be
called superior orbital foramen instead. This feature in MLD 37/38
conforms to the character state in African apes, Australopithecus
afarensis , and Australopithecus africanus in contrast to
humans and robust Australopiths that show elongated, comma-shaped
superior orbital fissures. The crescent of foramina (superior orbital
fissure/foramen, foramen rotundum, foramen ovale, foramen spinosum) in
MLD 37/38 has the same morphological condition as the other
Australopithecus africanus and Australopithecus afarensis
specimens.
This work demonstrates that Virtual Anthropology, using biomedical
imaging and geometric morphometrics, reveals features of precious
fossil hominids that would otherwise be inaccessible using traditional
methods.