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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 53285-295 (1980)
Indonesian “Solo” (Ngandong) Endocranial
Reconst ructions: Some Preliminary
Observations and Comparisons With
Neandertal and Homo erectus Groups
RALPH L. HOLLOWAY
Deparhnent ofAnhiv&y,
Columbia Universi&,New York, New York 10027
Endocasts, Brains, Asymmetry, Hominids, Platycephaly, PaKEY WORDS
leoanthropology, Petalias, Handedness
ABSTRACT
Latex rubber brain endocasts of Solo I, V, VI, X, and XI crania
were prepared in order to determine accurate volumes by H,Odisplacement and
morphological description. The new volumes are not markedly different from
previous estimates that had been done by mensuration formulae. The mean
volume is 1,151 ml. The endocast surfaces, while devoid of gyral and sulcal relief
patterns, do show a preponderance of left-occipital-right-frontal petalia patterns,
suggesting right-handedness. Metrical observations support the viewpoint that
the Solo crania are more closely related to earlier Indonesia Homo erectus than
to Neandertals.
In 1971 and 1972, through the courtesy of
Professor R. von Koenigswald, this author had
the opportunity to endocast the interiors of all
the Solo crania directly by liquid latex rubber.
The purpose of that work was to gain as much
endocranial detail as possible, to make complete reconstructions to enable accurate volume determinations, and to compare these
with previously published estimates.
The literature on the Solo crania, a t least
with regard to endocranial volumes, is confusing. Weidenreich’s (’51) monograph on the
subject was published posthumously and does
not include discussion of the volumes or the
methods used to calculate them. Instead, these
appear in Weidenreich’s (‘43a) classic monograph on the skull of Sinanthropus, but the
methods are not precisely described, except for
the Sinanthropus crania, where endocasts
were made and direct water displacement was
used to obtain the volumes. The discussion of
the Solo endocranial volumes on p. 116, Table
XXII (cm 74, p. 1121, is in the context of
external cranial dimensions, L, B, H, and the
Lee-Pearson formula. Oppenoorth‘s (‘37) estimates were very definitely formula-based, as
were those of Dubois (’37) in which Froriep’s
(‘11) methods were applied. Weidenreich‘s values in Table XXII (see Table 1) differ from
those of both Dubois and Oppenoorth, but the
differences are not discussed in any detail.
Morphological description has so far been lim-
ited to Ariens-Kappers (’36) paper, which
briefly described the endocranial portions of
Solo IV and V.
This paper will deal mostly with the volume
determination and some of the major morphological characteristics and their mensuration and will offer some suggestions regarding
taxonomic affinities.
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MATERIALS AND METHODS
Making the casts
Liquid latex rubber (ADMOLD #3280)’ was
used to endocast all of the Solo cranial pieces.
Only crania I, V, VI, X, and XI are reported
here, owing to incompleteness of numbers 11,
111, VII, and VIII and the excessive distortion
and poor internal condition of specimen IX.
Each of the specimens required somewhat
different treatment but, by and large, the
overall methods were similar, as described in
detail elsewhere (Holloway, ’78). Basically,
each cranium was checked for cracks, which
were filled with plasticene. Six or seven coats
of liquid latex were poured into each specimen,
1
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Obtainable frum Adhesive pmduds Gorp., 1660 Bwne Ave., New
York,NY 10060.
Not for citation without permission of author.
Received October 25,1979 accepted March 20, 1980.
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0002-948318015302-0285$02.300 1980 ALAN R. LISS, INC.
285
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R.L. HOLLOWAY
TABLE I . Comparisons of cranial mpacities for the solo endocash*
~
Solo crania
solo I
solo v
Solo VI
solo IX
solo x
solo XI
~~
Author
Dubois (‘37)
Oppenoorth (‘37)
Weidenreich (‘43a)
1,143 cc (2.5%)
1,160 cc (-1.0%)
1.316 cc (5%)
l;l60 cc (12.7%)
1,035 cc (13%)
1,225 cc (2.1%)
1,035 cc (-2.1%)
1,135 cc
1,055 cc (16.7%)
1,060 cc (2.6%)
1,087 cc (6.8%)
*Avg. = 1,151 oc;S.D. = 99; Var.= 7,881.36C.V.
N.A. = Not Available
Present study
1,172 cc
1,251 cc
1,013 cc
N.A.
1,231 cc
1,090 cc
= 8.6%.
each coating dried by means of a hair dryer
(warm air) before the next was introduced.
Next, the specimens were placed into an oven
and dried a t 50°C for 4 hours. After these
cooled down to room temperature the cured
rubber endocast shells were detached from the
endocranial surfaces, with the exception of
crania VI and XI, and were immediately fitted
back into the cranial portions. This was done
to ensure easier separation of the endocasts
from the crania after plaster retainers were
built inside the crania. In the case of crania
VI and XI, which are the most complete and
which include basal cranial portions, both
were filled with liquid plaster of paris under
water, so as to stabilize dimensions. This process can only be done with complete shells,
which maintain enough rigidity to avoid the
problems of buckling and distortion that occur
with incomplete (basal portions missing)
shells.
Crania I, V, IX,and X all have their basal
portions missing; thus the stabilizing of the
endocast shell had to be done by “piece-moulding” removeable sections within the crania.
Once the pieces (about six or seven each
crania) are dried, they are removed from the
endocast shell, bonded together, and the endocast latex shell is fitted over them. In this
way the original dimensions of the situ endocast are maintained with minimal distortion.
This was checked, incidentally, by comparing
landmark distances on the convex surface of
the endocast with the corresponding measurements on the original endocranial surface. No
distortions greater than 1%were found. The
original cranial portions were not broken to
extract the endocasts.
Reconstructions
None of the Solo crania are fully intact, and
to achieve the most reliable volume estimates,
reconstruction of the basal portions is necessary. This was done by using plasticene, by
allowing the more complete basal portions of
crania VI and XI to serve as shape templates
for reconstructing the bases of I, V, and X
(Figs. 1-51, and by following the curvature of
the present portions to fill in the missing
sections. (See Holloway, ’78, for further discussion of this approach.)
Volume determinations
Volume measurements were done by water
displacement in two separate ways. First, each
endocast was made water-tight by covering
any gaps between plasticene and latex wjth a
thin film of latex. Using a large beaker (about
3,500 cc) fitted with a narrow tube, the beaker
was first filled to overflow and allowed to
drain. Next, the endocast was slowly but fully
immersed, and the resultant run-off was collected in another pre-weighed beaker and consequently weighed, giving a n immediate volume, since the specific gravity of water is 1.
Five determinations were made for each endocast, and the average was calculated.
In the second method, each endocast was
weighed in air to the nearest 0.1 gram. Next,
the endocast was weighed in water (nearest
0.1 gram) by suspending the endocast from
the rim of the weighing pan into a large
plastic tank of water. Unless Archimedes
(N.D.) was mistaken, the loss in weight exactly equals the volume of the endocast. This
method provides a narrower range of values
and was the method finally adopted.
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SOLO ENDOCRANIAL RECONSTRUCTIONS
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Fig.1. Photographs of the reconstructed endocast of Solo I. First row, left to right, frontal and occipital views; second
row, lateral views; third row, dorsal and basal views. In each view, the endocast has been oriented in a plane passing
through frontal and occipital poles.
288
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R.L. HOLLOWAY
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Fig. 2. Recomtructixl endocast of Solo V. (Dem-iptionas in Fig. 1.)
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SOLO ENDOCRANIAL RECONSTRUCTIONS
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Fig. 3. Reconsh-uctedendocast of Solo VI. (Description as in Fig. 1.)
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R.L. HOLLOWAY
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Fig. 4. Reconstructed endocast of Solo X. (Description as in Fig. 1.)
SOLO ENDOCRANIAL RECONSTRUCTIONS
291
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Fig. 5.
Recastructed end&
of Solo XI. (Description as in Fig. 1.)
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R.L. HOLLOWAY
TABLE 2. Some metrical observations on the Solo endocasts
Max. length
solo
endocasts
L
(a)
R
Max.
height
(cm)
Max.
breadth
(a)
16.4
Solo v
Solo vl
solo x
solo XI
Mean:
S.D.
17.7
16.1
17.4
16.6
16.84
0.680
4.04
C.V. ( 2 )
Height
Length
Height
Breadth
~
~~
Solo I
Breadth
Length
16.3
13.5
10.4
0.826
0.636
0.770
17.7
16.2
17.4
16.6
16.84
0.680
4.04
or less
13.3
12.6
13.4
12.3
13.02
0.536
4.1
10.9
9.8
10.6
10.45
10.43
0.402
3.8
0.751
0.780
0.770
0.741
0.774
0.033
4.3
0.616
0.607
0.609
0.629
0.619
0.0127
2.05
0.819
0.778
0.791
0.849
0.801
0.032
3.99
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Metrical studies
There is, to this author’s mind, little point
to a comprehensive list of linear measurements on these endocasts, given their distortion in various regions, as mentioned above.
Table 2 provides the three basic measurements of anterior-posterior length for both
right and left endocast halves, maximum
breadth, height, and the respective indices.
Maxjmum lengths were measured with spmxhng
calipers to the nearest mm, using light ink
marks first placed on the left and right frontal
and occipital poles and found through visual
inspection. Maximum breadth measurements
are only approximate and are usually located
on the superior and posterior portion of the
temporal lobe, more or less where the middle
meningeal vein is evident; maximal measures
are never on the parietal surface of these
endocasts. Height was measured by stretching
a metal tape across the temporal poles and
measuring to the mid-point of the tape from
vertex, in the mid-sagittal plane. Vertex was
marked on the endocast as the most superior
projection of the dorsal surface once the endocast had been aligned in a horizontal plane
through frontal-occipitalpoles.
apparent, without undue speculation. There
does exist a right-frontal-left-occipitalpetalia
combination. Only the anterior division of the
menigeal vessel is present. The sagittal sinus
appears to lead into a more or less equal
division of transverse sinuses, although I
would favor a right-sided pattern on the basis
of an apparent greater thickness of the initial
flux of the right transverse.
Solo XZ. As in Solo VI this endocast is
complete with a basal portion, except for a
somewhat larger inferior rostral bec portion
than was seen in VI. This endocast is also
more distorted than VI, with a deeper right
temporal pole and a skewed left frontal area
on the dorsal surface. Again no sulci or gyri
are apparent except perhaps in the left frontal
lobe, within the complex of “Broca’s area,”
which is more pronounced on the left side.
There is a right frontal-left occipital petalia
combination. The meningeal vessles are somewhat more clearly retained on this endocast
than on VI, showing both anterior and posterior branches, which divide immediately superior of the styloid foramen. The posterior
region is not clear enough to determine with
preciseness whether the sagittal sinus drained
RESULTS
to right or left transverse sinus, although I
would be inclined to favor the right side.
Morphological observations
Both endocasts VI and XI have undamaged
Solo VZ. This endocast is by far the most rostral portions, and these both served as the
complete, missing only a small portion of the templates for reconstructing the basal porrostral bec, which according to Jacob (personal tions of the remaining Solo endocasts.
communication) has been more recently
Solo I. The basal portion is missing, from
cleaned out and is quite deep. There is but
slight distortion, which appears limited to the just inferior to the frontal poles and back to
left temporal region, in its superior portion. A the mid-foramen magnum region, being more
small addition of approximately 10 cc of plas- extensively present on the right side. The
ticine would provide the requisite symmetry endocast is distorted in the region of the right
to match the right side. No sulci or gyri are sylvian region and in the posterior temporal
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293
SOLO ENDOCRANIAL RECONSTRUCTIONS
(and inferior parietal) lobes on the left side. right occipital region. Sulcal and gyral patNo gyral or sulcal markings are safely inter- terns are not evident. The question of petalias
pretable. Again, as in VI and XI, there is a is difficult, given the breakage of the cranium
right frontal-left occipital petalia which can- in the right occipital portion and its subsenot be accounted for by distortion but as a quent distortion. There appears to be a clear
part of its “natural” asymmetry. There is also left frontal petalia, but it cannot be stated
a right transverse sinus lead from the superior with absolute certainty that there was a right
sagittal sinus, although it is not distinctly occipital petalia. The left frontal petalia does
visible.
not appear to be any artifact of distortion but
rather a “natural” asymmetry. The sagittal
Solo V. In this largest of the endocasts the sinus drained toward the right transverse sinbase is missing from well inferior of the frontal us.
poles to mid-cerebellar lobes, although both
right and left temporal lobes are more extenVolumetric determinations
sively present than in I. The distortion is
Table 1provides the volume determinations
mostly related to the right side, resulting in
a collapsed lateral aspect of the temporal lobe from this study and compares them to previand initial portion of the sylvian fissure. There ously published values. The values reported
are no fissurational patterns worth mention- here are by the second method, i.e., Aring. There is, as in each of the cases above, a chimedes’weight loss method, since it is more
right frontal-left occipital petalia combina- accurate with less variation between meastion; but it is mainly demonstrable on the left uring trials. In addition, the parenthesized
occipital region not by the posterior projection values indicate the percent of deviation of any
of the occipital lobe, but by its thickness from results from those previously published. The
midline to lateral extent. The right transverse average of the five solo endocranial volumes
sinus appears to receive the major supply from is 1,151mm, with an S.D. (standard deviation)
the sagittal sinus. The left Broca cap is par- of 99 ml and C.V. (coefficient of variation)
ticularly pronounced; the right cap is some- equal to 8.6%.
The percent difference figures indicate that
what smaller, even though more pointed
through damage to the original cranium in in those cases in which reconstruction of the
basal portions is not substantial, the present
that area.
values are very close to those predicted by
Solo X . This endocast is roughly as com- linear measurements and formula, i.e., Solo
plete as V above, but the temporal lobes, VI and XI. Where basal reconstruction is more
particularly on the right side, are more pres- substantial, as in I, V, and X, the formulaent than they are in V and show the same based methods probably underestimate the
tucked-under aspect as those in the complete height dimension, and thus the percent differVI and XI endocasts.There appears to be some ences are positively higher using reconstructslight depression and thus distortion in the ed endocasts. In any event, the volume deterright and left temporal lobes, immediately minations presented herein do not differ
posterior to the Sylvian fissure, and in the strikingly from those previously published.
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TABLE 3 . Comparing Endocranial Indices
Solo endocasts
(N = 5)
Homo ereetus
Indonesia
(N = 6)
Homo sapiens
neandertnlensis
(N = 5)
Mean
e
h
length
S.D.
C.V.%
len&
S.D.
C.V.
Mean
height
breadth
S.D.
C.V.
0.774
0.033
4.3
0.619
0.013
2.05
0.801
0.032
3.99
0.735
0.007
0.95
0.616
0.016
2.58
0.838
0.017
2.06
Mean
M
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0.793
0.039
4.9
0.673’
0.027
3.97
0.789l
0.072’
9.12l
‘ The only available Neandertal endocasts were La Chapelle, La Ferrassie, La Quina,Neandortal,and Qebel Ihroud I. The Neundertnl
specimen has no base, thus height was not available, and the relevant indices are based on N
=
4.
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R.L. HOLLOWAY
As the section on morphological description
indicated, the temporal poles are characteristically tucked under, as are those in other
endocasts for Homo erectus (non-African), and
the reconstructions of the basal portions for I,
V, and X followed such a pattern, which
should, I believe, add to the accuracy of the
volume determinations.
Tables 2 and 3 provide the endocranial
lengths, breadths, and heights for the Solo
endocasts, and compare #em with selected
Homo erectus endocasts from Indonesia and
European Neandertals. The results emphasize
only two things in my opinion: 1)the endocasts
are clearly platyencephalic, yielding height/
length indices most similar to the Indonesia
specimens; 2) the endocasts show considerable
homogeneity, given the low S.D.s and very
low C.V.S. The figures are published herein,
should they be useful in some other context to
other authors.
DISCUSSlON
As a group, the endocasts are homogeneously devoid of sulcal and gyral relief, with minor
exceptions in the frontal lobes. The endocasts
show little, if any, significant shape differences, each of them being considerably platycephalic (Table 2). That 80% (four out of five)
show a combination of right frontal-left occipital petalias is interesting and possibly significant when combined with the observation
that the third inferior frontal region, “Broca’s
area,” is well developed on all endocasts, more
often on the left than right sides. The precise
significance of these findings is unclear, although LeMay (’76, ’77) and Galaburda et al.
(‘78) have suggested that the petalia combinations are possibly indicative of both handedness and hemispheric dominance with regard to language. From a study in progress,
this writer is finding strong evidence for leftoccipital-right-frontal petalias in most of the
hominid endocasts studied to date. However,
preliminary observations show such a combination t o be very strong in Gorilla endocasts
also. Until these results are clarified and the
functional significance is made stronger, one
cannot discuss this problem at any further
length.’
The Solo or Ngandong crania have been
recently reappraised by Santa Luca (’78) for
their morphological affinities with both Indonesian Homo erectus and Neandertal (European and Near East) groups. His opinion that
these crania were representative of a group of
Homo erectus controverts the widely publicized views of Brace (‘67) and of Brose and
Wolpoff (‘71), who have regarded the Solo
crania as reflecting part of a worldwide Neandertal stage of human evolution.
This view has had support from the past
literature. Both Dubois (’37) and Oppenoorth
(’37) considered Solo V as the most Neandertal-like of the Solo crania, but regarded the
Solo crania a s more like the Sinanthropus
material (see particularly DuBois, ’36), a conclusion shared by Ariens-Kappers (‘36). Weidenreich’s view (‘42, p. 65) was very explicit:
“Homo soloensis differs decisively from the
European Neanderthalians and resembles Pithecanthropus in general form and specific
features so closely that he can be classified
flatly as an enlarged Pithecanthropus type.”
(See also Weidenreich, ’43b, pp. 44-45). Clark
(’64) was very cautious in his book, retaining
the possibility that the Solo crania fell within
the Homo neanderthalensis (sic: p. 83) range
of variation, but he did not discount entirely
the view of an “independent evolutionary differentiation” (p. 85) of the Ngandong population with the ancestral Australoid peoples, a
view revived by Coon (’62).While not wishing
to enjoin in controversy here, examination of
the morphological and metrical features of the
Solo endocasts leads me to conclude that the
Solo group is clearly more closely aligned with
Homo erectus than Neandertals. Table 3 provides a comparison of the endocast measurements and indices for the Solo, Homo erectus
(Indonesian), and Neandertal groups, all based
solely on available endoca~ts.~
The platycephalic condition of the Solo endocasts is clearly
much more like that of the Indoneasian materials and furthermore, the location for measuring maximum breadth is on the parietal
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* Indeed,the petalia question needs more investigation and extension into pongid endocast samples. In some cases, fmntoaxipital
petalias are not discernible from the usual dorsal view, when the
endocast is oriented in a plane passing h u g h frontal and occipital
poles, but does bemme discernible when viewed fmm the basal
aspect. In some cases,the A-P petalia is not apparent but is m k e d
in its lateral aspect. Thee variations q u i r e more study than they
have received thus far.
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As the S.Dsand C.V.s make apparent,all three p u p are fairly
homogeneous in their metrical pmperties. In the cases of both the
Solo and IndonesianHomo erectm etxlocasts, some small part of this
homogeneitycan probably be attributedto reconstruction techniques,
where the most complete endocasts act as templates to build up
missing sections (basal) on those less complete. This latter observation should not, however,deemphasize their considerable homogene
ity
lobe in Neandertals (La Quina excepted) and
is wholly temporal on both Solo and Indonesian Homo erectus endocasts.
Further studies On these
endocasts are
in progress, which utilize a stereometric and
comparative framework for measuring roughly 170+ points on the dorsal surface and, after
each measurement is corrected for allomekY,
or scaling, the residuals are examined by a
variety of multivariate techniques, particularly discriminant analysis. So far the results
(tentative) suggest much stronger relationships with both the Indonesian and Chinese
Homo erectus endocasts and less strong than
with European Neandertals.4
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SOLO ENDOCRANIAL RECONSTRUCTIONS
295
Coon, C.S. (1962) Theorigin of Races. New York AE.XI
A. a o p f .
Dubois, E. (1936) Racial identity ofHomo soloensis Op
penoorth (including Homo modjokertensis von Koenigswald) and Sinnnthropus pekemsis. Davidson Black.
Konin. Akad. Van Wetemchappen, 39:l-6.
Dubois, E. (1937) Early Man in Java and P z t b n t h m
$~’!~~.~i~~~&,~~; $yLgFIYMan.
Froriep, A. (1911) Ueber die Bestimmung der Schadelcapacitat durch Messung und Berechnung. 2. Morphol.
Anthropol. Bd. XIII: 347-374
~
;
g
~
science,199;85%856.
Holloway, R.L. (1978) The relevance of endocasts for
studying
brain ev~htion.In Noba& CR. (.XI):
Sensory Systems of Primates. New York: Plenum Press,
pp. 181-200.
LeMay, M. (1976) Morphological asymmetrics of modern
ACKNOWLEDGMENTS
man, fossil man, and non-human primates. In Harnad,
H.D. SkMiS, and J. L~IUXW&~ (d):
OriginS and
1 am very grateful to Dr. G.H.R. “on KO- S.R.7
evolution of language and speech. Ann. N.Y. Acad. Sci.
enigswald, Senckenberg Institute, Frankfurt,
2&):34%360.
Federal Republic of Germany, who gave me m y , M. (1977) Asymmetries of the skull and handd
t J.N
~e ~~
lsci.
. 32:243-253.
the permission and encouragement to endosoloensis
W3.F. (1937) “he Place of
cast the original specimens, and for his has- %PenCJ”fi,
among fossil man. In M a c C d y , G.G. (ed):Early Man.
pitality and generous offer of facilities. I also
Philadelphia: J.B. Gpphcotta.,
pp. 349-360.
thank Dr. Teuku Jacob for his cooperation and santa LUB, A.P. (1978) A reexamination of p r e m e d
Neandertal-like fossils. J . of Hum. Ewl. 7:619-636.
permission to study the other Homo erectus
F. (1942) Early man in Indonesia. Far
materials in Indonesia, and for his interest Weidenreich,
Eastern Qwuterly Nov. pp. 58-65.
and advice. This research was supported in Weide-i&,
F. (&3) heskull of,yimnthropus p h i nensis: A comparative study on a primitive hominid
part by NSF grants GS 2931-1 and BNS
skull. Palaeontologica Sinica, new ser. D, No. 10, whole
7911235 and in part by a Guggenheim Fellowseries No. 127, pp. 1-484.
ship, for which I am most grateful.
Weidenreich,F. (1943b) The “Neanderthal man” and the
ancestors of “Homo sapiens”. Am. Anthrupol. 45:39-48.
LITERACITED
Weidenreich, F. (1951) Morphology of Solo Man. Vol. 43
Ariens-Kapprs, C.U. (1936) The endocranial casts of
(Part 3), Anthropology Papers of the American Museum
of Natural History, New York.
the Ehringsdorf and Homo soloensis skulls. J. Anat.
71:61-76.
Brace, L.C. (1967) The Stages of Human Evolution. En4 It would be premature to disclLss thee tentative results in any
glewood Cliffs, New Jersey: F’rentice-Hall, Inc.
detail, as they are O ~ supportive
Y
and are independent Of the
B
~D.s.,~ad,M.H. wolp0ff (1971) ~~l~ upper
morphological a d metrical aspeds d i x d herein. When discrimlithic man and late
tools, Am. Anthropol. inant analyses are done, the small Solo group (N = 5 ) tends to be
73:115694.
classified either in its own group or with either the Chinese or
Clark, W.E. LeGm (1964) The Fossil Evidence for HU- Indonesian Homo erectus groups, rather than in the Neandertal
man Evolution. Chicago: University of Chicago Press.
group.
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