STUDIES I N GROWTH CHANGES I N THE SKULL

AND FACE O F ANTHROPOIDS

v. GROWTH CHANGES IN THE SKULL AND FACE OF THE ORANG-UTAN'

WILTON MARION KROGMAN

Anatomical Laboratory, Western Reserve University, Cleveland, Ohio

NINE FIGURES

CONTENTS
Introduction and material ............................................ 343
Superposition in norma lateralis : qualitative study .................... 345
A quantitative analysis: method ...................................... 348
A. The cranium ................................................. 350
B. The face .................................................... 353
Growth changes in cranium and face of the Great Anthropoids ........... 356
Summary ........................................................... 361
Literature cited ..................................................... 362

INTRODUCTION AND MATERIAL

As I have indicated in the study of growth changes in the
skull and face of the Gorilla ('30) the analysis of growth
changes in the Anthropoid skull is based upon 168 Gorillas,
133 Chimpanzees and 134 Orangs, classified and grouped
according to dental age (see Table I). Of this number 78
were graphically reproduced upon the Reserve craniostat in
norma lateralis sinistra. Selected representatives of the sev-
eral dental age groups were then superimposed and the result-
ing details of change in proportion and position noted.
In the entire aeries of craniostatic studies the analysis has
been limited t o growth changes as observed in norma lateralis
sinistra. This aspect was chosen since it undoubtedly yields
the most significant results to serve as a preliminary investi-
Studies I-V form parts of a thesis submitted to the Faculty of the Graduate
Schools of Art, Literature, and Science, University of Chicago, in partial ful-
fillment of the requirements for the degree of Doctor of Philosophy, June, 1929.
343
344 WILTON M A R I O N K R O Q M A N

gation. Future research in growth changes in the Anthropoid

cranium and face will be extended t o norma facialis and
norma basilaris.
The principle of the superposition of tracings of cranio-
static reproductions seeks to retain the advantages, while
eliminating many of the defects, of the Frankfort horizontal.
To this end the nasion-porion plane has been devised: in
practice any skull of definite dental age is oriented in the
Frankfort plane and a parallel to that plane projected
through nasion. Succeeding skull tracings are superimposed
upon this, so oriented that the nasion of each falls upon the
nasion parallel, with porion superposed.
The skulls utilized here for superposition are as follows :
1. N.M. 49972, Male. No teeth erupted.
2. Am.N.H. A(W.K.G.), Male. Deciduous il-2, ml-2, deciduous e erupting.
3. Am.N.H. L243, Male. Deciduous dentition complete ; M1 erupting.
4. Am.N.H. 17350, Male. Deciduous dentition complete; M1 erupted; M2 in
crypt.
5. W.R.U. B1024, Male. Deciduous dentition complete; M 1 erupted; I1 and
M2 erupting.
6. W.R.U. B1169, Male. Deciduous c ; ml-2; 11, M1 and M2 erupted; I2
erupting.
7. An1.N.H. 4OOC-A, Female. M3 newly erupted.

In each figure the younger skull is indicated by a solid line,

the older by a broken line. In going from figure to figure the
older skull is carried over; thus, the broken line of Figure 1
becomes the solid of Figure 2 and this relation is preserved
throughout.
TABLE I
Stages in tooth eruption serving as subdivisions f o r the present investigation
il-2 c ml-2
Stage 1 il-2 c ml-2
il-2 e ml-2 MA
Stage I' il-2 ml-2
Pml-2 M 2 erupting
Stage 'I1 G-2 M2 erupting
Stage Iv Ceruptiug Pml-2 M1-2 M3erupting
C erupting Pml-2 M1-2 M3 erupting
11-2 Cnew Pml-2 M1-2-3 (new)
v I1-2
~

Stage Pml-2 MI-2-3 (new)

11-2 C
- Pml-2_.
Stage V I 11-2 C
~

Pmi-2
 SKULL AND FACE GROWTH O F THE ORANG-UTAN 345

T H E SUPERPOSITION I N NORMA LATERALIS SINISTRA O F SELECTED

AVERAGE REPRESENTATIVES O F DENTAL AGE GROUPS
I n Figure,l skull A (W.K.G.) has been superimposed upon
skull 49972. Here, with the latter as the youngest skull
observed in the entire anthropoid series, we note for the first
time the relatively enormous amount of growth before the
eruption of the deciduous teeth. The impression is one of a
uniform increase in size in almost every respect. There has
occurred a marked general increase in the size of the cranium
with a growth in height nearly equal to that in length. The
occipital region is more full superiorly, imparting to its
inferior aspect a relative flatness. The foramen magnum has
increased in size and moved slightly up and back, and ap-
parently in keeping with this the lambdoid suture has inclined
forward. The coronal suture, however, retains the same
obliquity. The parietal bone has grown vertically, anteriorly
and posteriorly. The squamous has also extended forwards
and backwards, a trifle more at its posterior margin. The
squamous has gained height, at its anterior end only, a gain
in keeping with the increase in height of the alisphenoid,
which we have not seen in the other Great Anthropoids. The
alisphenoid has not gained in antero-posterior dimension, but
has moved forward. The fronto-sphenoid articulation, how-
ever, remains at the same level and the pterygoid is rela-
tively unchanged except in height. The forward growth of
the parietal has pushed the frontal forward and this carries
the mask with it. Hence the entire face has grown forward,
the orbit has enlarged without changing its vertical axis, and
the alveolar region has grown markedly along its whole
length. As a corollary of the forward-downward growth of
the face the zygomatic arch presents a forward growth and
a downward inclination. An interesting feature of the Orang
skull is the observable deviation of the coronal suture. This
suture is not a relatively straight line, as in the Gorilla and
Chimpanzee, but it shows a tendency, with increasing age,
for the pars temporalis and pars media to assume differing
degrees of obliquity. In the Figure under consideration it is
346 WILTON M A R I O N K R O G M A N

to be noted that the pars temporalis is almost vertical, while

the pars media is inclined backward and then upward.
It is worthy of note that the inner table of the frontal bone is
beveled in the pars bregmatica, and the outer table in the pars
temporalis, a condition which permits the center point i.e. a t
pars media, of the coronal to serve as pivot for a shearing
movement. I n several young Orang skulls it has been noted
that the continuity of the temporal ridge is broken as it
crosses the coronal suture, as though there had been a sudden
movement or a differential rate of increase, in size or position.
I n Figure 2 skull L243 is superimposed upon skull “A.”
Again there is noted a uniform proportionate increase in size,
though not so marked as before. The cranium has gained in
length and height, while the parietal and squamous bones
evince only a little growth, and the position of the foramen
magnum is unchanged. The relations of the coronal and
lambdoid sutures have not changed, though the mid-point of
the coronal suture has taken a decided backward movement.
The pterionic complex, except for an antero-posterior increase
in the alisphenoid is unchanged. The growth in the face is
most marked in the alveolar region, which has gained greatly
in height and somewhat in length. The zygomatic arch has
retained its earlier downward inclination and has grown
slightly forward. The orbito-nasal portion of the face has
grown forward, while the orbit has increased in size without
changing its vertical axis.
I n Figure 3 skull 17350 has been superimposed upon skull
L243. We see here that the cranium had already practically
attained its full size and that following this the growth
process is one of adjustment. There has been no increase in
size either vertically or occipitally. Curiously enough the
increase forwards is due to occipital expansion pushing pari-
etal forward along squamous suture, and, with it, the frontal.
With this the foramen magnum has moved upward. The
squamous also has extended anteriorly and has pushed the
alisphenoid forward and downward without gain in size. The
pterygoid has grown slightly. The face shows very little
 SKULL AND FACE GROWTH O F THE ORANG-UTAN 347

growth ; some in the orbital region, and slightly in the anterior

aspect of the alveolar portion, while the zygomatic arch has
grown forward and somewhat downward.
I n Figure 4 skull B1024 has been superimposed upon skull
17350. As before we note that there is no increase in the
size of the cranium. The slight differences in position of
sutures and the form of the temporal fossae are merely indi-
vidual characters like the angulation in the coronal suture
which reappears in B1024. The face however shows a definite
progressive change in its relation to the cranium. I n the first
place the alisphenoid has increased in width, the pterygoid
has moved forward; the orbit has changed its vertical axis
so that it now faces more nearly straight ahead; the zygomatic
arch has lost some of its downward inclination; and the lower
face has grown forward while above, superiorly it has lost
the child-like curve of the forehead. All these features are
expressive of a definite rotation forwards round a transverse
horizontal axis passing through the nasion. Heretofore it has
been growing downward and forward; it now commences to
tilt and a study of the alisphenoid in this series indicates how
this particular facial movement comes to be.
In Figure 5 skull B1169 has been superimposed upon skull
B1024. The apparent growth in cranial height is probably an
individual character associated with the difference in occiput.
In association with this change in the occipital region the
foramen magnum lies much further upward and backward in
B1169 than in B1024. The face shows further progress in the
tilt alluded to in the preceding paragraphs. The alisphenoid
has increased in size and moved forward while the pterygoid
has retained its position and added substance to its anterior
part. Continued forward movement of the frontal has car-
ried the orbit forward without affecting its size or inclina-
tion. The lower part of the face presents a decided growth
in the alveolar portion, a growth somewhat more marked
anteriorly, enhancing the impression of an upward swing.
The picture of growth in the face of the Orang at,the period
between eruption of first and second molars may be clarified
348 WILTON MARION KROGMAN

if we conceive of the interrelationship of two lines of growth,

a superior or orbito-nasal, and an inferior, or alveolar. These
may be likened to two cords extending from a common.point,
connected at their distal ends so that extension of the one will
also affect the other though to a different degree. I n the
Orang there is relatively greater growth in the alveolar
region than in the orbito-nasal, and the alveolar region tilts
forward and upward as though bridled by the lack of com-
parable growth in the orbito-nasal region. I n the Gorilla and
Chimpanzee the growth is more equal in both orbito-nasal and
alveolar portions of the face so that the mask moves steadily
forward. The end-result in the Orang reminds one of
achondroplasia, though of course there is no suggestion of
this condition being invoked. Most certainly, however, of the
three great Anthropoids the Orang most nearly presents the
concave “dish face” of the achondroplast.
I n Figure 6 skull 900 C-A is superimposed upon skull
B1169. Since we are dealing with the superposition of female
upon a male skull it is difficult to judge as t o growth in the
skull. Comparison with other skulls warrants the statement
that there is neither growth nor movement in the skull after
the eruption of second molar; the cranium becomes a unit.
We witness in the face, however, a final continuation of the
swinging process already noted. We are aware of a great dis-
proportionate growth between alveolar and orbito-nasal
regions. The alisphenoid has increased its antero-posterior
dimension as is natural in this change of form in hafting and
a great growth of the pterygoid plus a marked growth of the
alveolar region downward and anteriorly results in an accen-
tuation of the general upward trend of the antero-inferior
part of the face. To adult life the Orang carries this pecu-
liar growth pattern wherein the face seems disproportion-
ately large.
A QUANTITATIVE ANALYSIS O F GROWTH CHANGES: METHOD

The qualitative method of analysing growth changes quite

obviously limits observation to selected average skulls from
 SKULL A N D FACE GROWTH O F THE ORANG-UTAN 349

established dental age groups. As a consequence individual

skulls have served as a basis of interpretation.
I n order, then, to supplement the qualitative method we
may turn to the expression of growth changes in terms of
absolute measurements. Accordingly in this section we
undertake an analysis of the measurements of the total series
of skulls included in the present study. The measurements
have been made on ad1 the craniostatic reproductions, and, in
addition, upon the skulls in the Reserve collection.
The measurements taken are those best calculated to aug-
ment the qualitative analysis already presented. They do
not, therefore, strictly conform to the orthodox anthropo-
metric measurements, though they are based on standard
landmarks. A list and definition follows:
I. On the cranium.
A. Expression of growth i n length.
1. Nasion to opisthokranion, wherever found (NL). The
occipital crest has been included since it is, i n last
analysis, an indication of growth change.
2. Pre-porionic length of 1 (PRE-L). Determined by
the erection of a perpendicular to the Frankfort
horizontal a t porion. (This is a variant of Cam-
eron, '28.)
3. Post-porionie length of NL (POST-L).
4. Porion axis to opisthokranion as determined by 1
(PO-L) .
5. Porion axis to opisthion (PO-OPIS).
6. Porion axis to asterion (PO-AS).
7. Width of alisphenoid (AL).
8. Width of pterygoid process of the sphenoid a t its base
(PTERY).
9. Width of parietal bone. Measured as the distance
between two planes running along the lambdoid and
coronal sutures (PA).
B. Expression of growth in height.
1. Porion axis to plane of vertex (PO-VER) .
2. Porion axis to level of squamods suture. The level is
a n average, disregarding extreme variation in ser-
ration (PO-SQ).
350 WILTON MAR.ION KROGMAN

11. On the face.

A. Expression of growth in length.
1. Porion axis to nasion (PO-NA).
2. Porion axis to orbitale (PO-ORB).
3. Porion axis to prosthion (PO-PROS).
4. Porion axis to posterior margin of alveolus of canine
(PO-C).
5. Porion axis to posterior margin of alveolus of first
permanent molar (PO-M) .
6. Porion axis to most anterior point of malo-temporal
suture (PO-ZY) .
7. Posterior margin of alveolus of canine to anterior
margin of alveolus of first permanent molar ( C-M1) .
8. Prosthion to posterior margin of alveolus of first per-
manent molar (PRO-M) .
R. Expression of growth in height.
1. Nasion to prosthion (NA-PRO).
2. Height of orbit (ORBIT).
3. Orbitale t o nasion (NA-ORB).
It will be noted that such measurements as Na-Po are at
once an indication of growth in height as well as length;
further that as pointed out by Hellman ('27) many of
the measurements are more indicative of a change in posi-
tion than of growth. These considerations will be dealt with
later.
A . The cranium
A consideration of Table 11, presenting an analysis of
measurements expressing growth in length, demonstrates that
the Orang skull has grown 72.8 percent in length (NL). The
growth is limited entirely to the portion of the skull anterior
to porion (Pre-L), which presents an increase of 64.2 percent
while the post-porionic distance (Post-L) has not grown at
all. This may be compared to the Gorilla and Chimpanzee
patterns in which in the former 90 percent of gain in length
is pre-porionic, and in the latter 75 percent. I n striking con-
trast to the Anthropoid growth pattern is the human condi-
tion in which definitive skull length is attained by equal pre-
and post-porionic growth.
 SKULL AND FACE GROWTH O F THE ORANG-UTAN 351

Specifically in the Orang, and generally in the Anthropoid

skull, the region of the alisphenoid and coronal suture must
be responsible for whatever growth in length has occurred.
In the Orang the alisphenoid has increased its breadth (i.e.
length as measured antero-posteriorly) 123.3 percent while
TABLE I1
Growth of the skull of the orang in length

DENTALAGE

I
___-
AV.

105.4
NL

I % Inc. -I AV.
PRE-L

I % Inc.
~ -
Av.
POST-L
-
I
~ _
% Inc.
_ _

54.8 50.6
I1 110.3 4.7 59.3 8.2 51.0 .8
I11 123.0 10.5 73.0 23.0 50.0 - 2.0
IV 122.0 - .9 72.0 - 1.5 50.0 -
V 126.0 3.8 76.0 5.5 50.0 -
VI 140.0 11.1 90.0 18.4 50.0 -

- .__ -
DENTAL AGE
1
~ ~

Av. Inc. 9% Inc. AV. % Inc.

I 56.3 34.3 33.3 1
I1 55.7 - 1.2 35.5 3.5 ~ 32.9 i - 1.2
I11 57.5 3.4 37.0 4.2 37.0 ' 12.4

I
~

IV 59.0 2.8 44.0 18.9 35.0 - 5.4

V
VI
56.0
56.0 ~.
-- 5.1
-
42.0
42.5 ~
1 - 4.6
_ 1.2
_ 1 37.5
- ~
-
7.1

I 10.3 13.4 68.6

I1 12.4 21.5 16.5 23.0 69.4 ' 1.1
111 17.5 45.5 23.0 39.5 75.0 , 8.1
IV 16.5 - 75.0 -
V 22.0 - 75.0
I
-
VI 23.0 2.4 74.0 - 1.4

the parietal bone has added only 8 percent, a fact which

localizes growth quite definitely.
The changing proportions in the length of the skull with
respect to the relation of the anterior and posterior porionic
distances to the total length may be expressed by the follow-
X 100
ing index POST-L
- PRE-L.
352 WILTON MARION KROGMAN

Dental age Indez

I 92
I1 86
I11 68
IV 69
V 65
VI 65

The interesting fact here is that already at Stage 111, after

eruption of the first molar but considerably before the erup-
tion of the second, the skull has gained its definitive length.
A further increase is noted after the eruption of third molar,
but this is due to the preponderance of male skulls, abso-
lutely larger in their dimensions, a fact which colors many
of our averages for Orangs.
The distance from porion to point of greatest length
(PO-L) does not change, while from porion to asterion
(PO-AS) seems to present a slight increase of about 10 per-
cent, reflecting the tendency of the squamous to tilt its pos-
terior end upward, a process at once inclusive of some growth
and movement. The backward and upward trend of the fora-
men magnum, as indicated by the measurement from porion
to opisthion (PO-OPIS) presents an increase of 23.9 percent,
the greatest movement occurring just before the eruption of
second molar.
The width of the pterygoid (PTERY) presents an increase
of 73.9 percent, as contrasted with 123.3 percent in the case
of the alisphenoid. It is important that the pterygoid is vir-
tually stable shortly after the eruption of the first molar,
wheras the alisphenoid presents an increase just preceding
the eruption of third molar. The initial rate of increase of
the two is about the same. The skull of the Orang presents
less growth than that of either the Gorilla or Chimpanzee,
and, moreover, what growth is attained is very early, at the
latest between eruption of first and second molars, and prob-
ably just after eruption of the first, at a time when the perma-
nent incisors and premolars are coming in. The posterior
portion of the skull is already fixed before the appearance
of the deciduous teeth : the anterior portion finds its greatest
 SKULL AND FACE GROWTH O F T H E ORANG-UTAN 353

growth in length just after eruption of first molar and its

souice of greatest growth in the alisphenoid.
The data for a study of the growth of the skull in height
are in the following table:

___ - -

I1 71.0 - .9 -
I11 78.0 9.0 ~ 21.4
30.0 ~ 40.0
IV 76.0 -2.5 30.0 -
V
771
i
1
75.0
79.0
-1.3
5.2
1 30.0
32.0 I
-
6.7

The increase in height (PO-VER) discounting VI, which

is influenced by the crest of the male, is about 10 percent, and
is attained before the eruption of second molar. The squa-
mous (PO-SQ) increases about 40 percent in height and, as in
the Gorilla and Chimpanzee, is largely sliding up over the
parietal. I n its vertical dimension the Orang reveals the
stability common to the Anthropoids.

B. The face
I n Table 111 we have presented the measurements taken
on the face which serve as an indication of growth in length.
For the orbito-nasal region we note an increase of 56 percent
for PO-NA and 70.3 percent for PO-ORB; for the nasal
region an increase in 82.8 percent for PO-ZY; and for the
alveolar region an increase of 109 percent for PO-PRO. F o r
the superior part of the face we have an even progression,
from the orbito-nasal region to the nasal, but as we go from
the nasal region we note a relatively great velocity in the
al~7eolarregion. The upper face has completed its growth by
the end of the eruption of the incisors and premolars, whereas
the lower face presents an increase just before the eruption
of third molar. This disproportionate rate of increase serves,
in a measure, along with the disproportionate growth in the
alisphenoid and the pterygoid, to explain the pronounced
354 WILTON MARION KROQMAN

upward swing of the Orang face with progress toward ma-

turity. There is an absolute and relative disproportion which
will be more clearly seen later.
The alveolar region, the scene of greatest growth in length
of the face, now commands our attention. The distance from
TABLE I11
Growth of face of orang in length
PO-NA PO-ORB PO-PRO
DENTAL AGE -
_. -~ ~ _.
. -~
Av. 1 % Inc. AV. 1 % Inc.
I
I 59.6 54.6 i 78.7
I1 65.2 9.4 61.0 11.7 96.1 22.1
I11 73.0 12.0 76.5 25.2 118.0 24.9
IV 75.0 2.7 78.0 1.9 120.0 1.7
V 77.0 2.6 75.0 - 3.8 141.0 17.5
VI 93.0 20.8 93.0 24.0 164.5 17.4

PO-c P0.M PO-ZP

DENTALAGE ~- ~~ ~~ ~~

Av. 1 96 Inc. I Av. 1 % Inc. AV. I % Inc.

I 70.8 45.6 38.3
I1 82.5 5.2 56.4 23.7 44.8 16.8
111 103.0 24.8 76.0 33.7 53.5 17.2
IV 101.0 - 2.0 76.0 - 54.0 1.0
V 119.0 18.8 99.0 30.1 57.5 3.3
VI 140.0 17.6 118.0 19.0 70.0 21.7
___-
C-M' PRO-M
DENTAL AGE __-- - ~-

~_______
i AV. % Inc. AV. 1 96 Ine.
I 17.5 40.0- 1-__~
I1 19.3 10.3 44.6 I 11.4
111 21.0 8.8 48.0 3.1
IV 20.5 - 2.5 44.0 ' - 8.3
V 19.0 - 7.3 54.0 I 22.7
VI 20.0 5.3 59.0 9.3

molar, viz., a forward and downward movement just before

its own eruption, and just before the eruption of second and
 SKULL A N D FACE GROWTH OF T H E ORANG-UTAN 355

third molars ; indicating that most of the growth of the alve-

olar region is from back to front, i.e. pre-eruptive addition
to the length of the alveolus in preparation for the eruption
of the tooth. As in the Gorilla and Chimpanzee it is improb-
able that there is any interstitial growth between the perma-
nent canine and first molar. The observed increase is prob-
ably a corollary of the enlarged alveolus for the canine,
especially in males.
The growth of the face in height is presented in the follow-
ing table:

NA-PRO I ORBIT NA-ORB

DENTAL AQE
Av. Av. To Ine. Av.

I 31.2 23.2
I1
111 I 62.2
73.5
24.1
18.2 25.0 - 7.8

~~
IV
V
VI
1 ~~
75.0
92.3
96.0 1 ~~~
2.2
23.1
4.0
I 38.0
38.0
38.0
_ _
~ -
11.8 25.0

_ _ ~ _ ~~

Since we see that the Orang face has gained only 91.6 per-.
cent in height (NA-PRO) as compared to a gain of 109
percent in length (PO-PRO), we gain an insight as to the
mechanism of the pronounced upward swing of the face.
This may be thought of as a definitive upward trend of the
anterior-inferior portion of the face or it may be considered
as a mechanism of relief for the large lower face in the pres-
ence of the relatively slight growth of the upper part. It is
certain that, of the three Anthropoids, there is greatest rela-
tive disproportionate of the two, parts of the face in the
Orang.
The orbit of the Orang adds 21.6 percent to its height, the.
most growth in any of the great Anthropoids. In common
with the Gorilla and Chimpanzee, the Orangrface attains its
definitive height by growth inferior to the orbital margin. It
i s still a matter of growth in the nasal and alveolar regions.
356 WILTON M A R I O N K R O G M A N

GROWTH CHANGES I N THE CRANIUM AND FACE O F THE

GREAT ANTHROPOIDS
We have thus seen, from two viewpoints, which have been
termed the qualitative and quantitative, that the problem of
growth changes in the skull and face is a complicated one,
compounded of two major elements, increase of dimensions
and readjustment of proportions. I n the initial stages in all
three Anthropoids, up to shortly before the eruption of first
molar, increase in dimensions is preponderant. There is how-
ever a continuous, relatively rapid, progressive change in
proportion. The skull is conforming throughout to a funda-
mental type.
It cannot be emphasized too strongly that the posterior
moiety of the skull, the parieto-occipital and occipital regions,
is the most stable of the entire skull. This is expressed by
the relative lack of growth posterior to porion, and the very
early attainment of the maximum vertical dimension. The
inferior portion of the occipital region, as witnessed by the
upward and backward migration of the foramen magnum,
presents an area of unrest. There is a decided movement
here which may reflect an attempt a t expansion, a view upheld
by the fact that the second part of the masto-occipital suture
is patent until well along in life.
It is in the anterior portion of the skull that we find the
locus of skull growth. The tendency of the squamous to
increase in height, most marked in the Gorilla, seems to point
to an expansion of the parietal area, the squamous riding up
over the parietal as if providing a covering for a lateral
growth. The fronto-parieto-sphenoid region seems to hold
the key to growth of the skull, not only by its increase in
dimension, but by the complex adjustment to the changing
proportions of skull and face. The alisphenoid constantly
doubling its antero-posterior diameter from birth until the
completion of the permanent dentition, migrating forward in
response to the push of the temporo-parietal region, presents
us with the area of greatest growth and adjustment. It is here
we must look for a solution of cranio-facial growth changes,
and further studies must be made of this specific area.
 SKULL AND FACE GROWTH O F THE ORANG-UTAN 357

The growth of the face is, in the main, accomplished in

conformity with a pattern peculiarly its own, adjusted to the
needs of the functions of sight, respiration, and food prepara-
tion. We note, therefore, growth changes definitely linked up
with these functions : the orbital region relatively constant
after birth: the nasal region evidencing early growth: and
the alveolar region the scene of greatest growth. We note,
however, that secondarily in the face, there is a reflection of
cranial growth, such as the downward “slide” of the
Gorilla face in response to a downward inflection of the
fronto-sphenoid region. We may point, indeed, to the form
of the face as occasioned by growth forces possibly inherent
in the basilar region of the skull. And this brings us to the.
problem of the noted upward swing of the antero-inferior
portion of the face, most marked in the Orang, less so in the
Gorilla, and almost absent in the Chimpanzee.
To state the presence of a growth pattern is one thing:
to explain it, quite another. We have already noted that the
relative foreshortening of the basilar region is apparently
reflected in the face, as witness achondroplasia; we have
adduced, on the other hand, that the observed facial dispro-
portion in the Orang may be due to the differential rate of
increase between upper and lower parts of the face. Certain
it is that there seems to be a correlation between the upward
trend of the face and the marked growth in the alveolar and
orbito-nasal regions. If we regard the face and cranium as
an entity-and, in the final analysis, we must-it follows that
growth is inter-relational. Hence the growth in any given
part will find a reflection in the part most intimately asso-
ciated with it.
The growth of the alveolar region of the face of the three
great Anthropoids is a direct reflection of the dental equip-
ment. The quite regular alternation of increase in length,
increase in height, period of rest or equilibrium, is setting the
stage for the eruption of the teeth. The pre-eruptive periods
are those of greatest growth activity. We see a further
reflection of the force of the erupting tooth as growth-motive

THE AMERICAN JOURNAL OF AN.4TOJfY, VOT,. 47, NO. 2

358 WILTON MARION KROGMAN

when we observe the downward and forward movement of

the first permanent molar; at first obliquely down and an-
terior as it descends from its own crypt high in the maxilla':
then relatively straight forward in response to the push of
second and third molars, as they, in turn, come down and
forward. The growth of the posterior region of the maxilla
is thus explicable.
Let us turn now to a consideration of Figures 7 and 8 and
9. I n attempting to present as simple a picture as possible
of the very complex process of growth-changes, recourse has
been had to a diagrammatic representation of the cranium
and face of the Gorilla, Chimpanzee and Orang, at each of
the dental ages we have outlined. The measurements which
have been utilized are as follows:
1. Porion axis to posterior margin of alveolus of 1\11 (PO-M) .
2. Porion axis to prosthion (PO-PRO).
3. Porion axis to nasion (PO-NA) .
4. Porion axis to level of vertex (PO-VER).
5 . Porion axis to point of greatest length (PO-L).
6. Prosthion to posterior margin of alveolus of 191 (PRO-11).
7. Nasion to prosthion (NA-PRO).
8. Nasion to point of greatest length (NL).
The plotting, on millimeter paper, of the foregoing permits
differentiation of the cranium (PO-NA-VER-L) from the face
(PO-NA-PRO-If) ; further we may divide the face into two
areas, orbito-nasal and nasal (PO-NA-PRO) or upper, and
alveolar (PO-PRO-M) or lower. This permits of the anal
ysis of each of these areas, both with respect to growth as
an entity, aiid with respect to reciprocal relationships.
The stability of cranial height and occiput is marked. In
each figure the horizontal line at the cranial summit repre-
sents the range through which the vertex migrates about the
perpendicular erected at porion. Iii the Gorilla the range is
r+10 mm. (+ for pre-porionic o r forward migration and -
for post-porionic or backward migration) though the vertex
moves forward as the age increases. I n the Chimpanzee the
range is +lo, namely forwards only; a majority of the skulls
presenting the vertex at the perpendicular. I n the Orang the
 SKULL A N D FACE GROWTH OF THE ORANG-UTAN 359

range is +15, the vertex moving anteriorly with age as in

the Chimpanzee, though again the majority of skulls have the
vertex at the perpendicular.
I n the Gorilla there is a marked upward trend of the pos-
terior point marking the site of greatest length (DL). This
is more movement than growth as the cranium flattens occip-
itally with formation of the lambdoid crest or its more dimin-
utive homologue in the female.
As an expression of the growth of the face and its changing
relationships, the angle NA-PO-PRO has been calculated for
the several dental ages :

DENTAL AQE QORILLA CHIMPANZEE ORANQ

I 47" 10' 39" 10' 38" 30'

I1 42" 30' 40" 50' 39" 50'
111 40" 0' 40" 30' 36" 30'
IV 37" 30' 39" 50' 36" 50'
v 34O 20' 40" 10' 37" 10'
VI 35" 50' 37" 40' 30" 20'

This strikingly emphasizes the process of facial growth-

change in the Great Anthropoids. In the Gorilla the angle
decreases about 12 degrees, in the Chimpanzee only 2 degrees,
and in the Orang 8 degrees. I n the Gorilla the decrease is
regular and progressive ; in the Chimpanzee there is a funda-
mental retention of the same angle, the final decrease being
negligible, and in the Orang there is an initial decrease, with
a marked change in the angle upon the eruption of the third
molar.
Since NA is, in keeping with the stability of the orbit, rela-
tively fixed, it follows that the point PRO has been closing in,
to decrease the angle. The amount of growth concomitant
with the degree of change tends to obscure this feature in
the diagrams that have been constructed, yet the obliquity of
the line PRO-11, most marked in the Gorilla and the Orang,
is a measure of the forward and upward swing of the point
PRO, as representative of the antero-inferior aspect of the
face. And in following the point 11, as indicative of the
360 WILTON MARION KROGMAN

first permanent molar, we get a perfect trajectory of an

obliquely forward and downward movement. The interrela-
tionship of gain in height and length is likewise aptly
illustrated.
We are introduced to a further important concept of
growth-change, namely, that of primary and secondary
change. I n the Gorilla, as we record growth between the
completion of the deciduous dentition and the eruption of the
first molar, we observe a primary or basic change in pattern:
the alveolar region grows in such a manner as to present a
definitive downward obliquity. I n the Chimpanzee we also
note that at the same age-period the alveolar portion of the
face grows markedly, so that by the eruption of the first molar
the face has its final pattern. The Orang face is established
as a type by the eruption of the deciduous teeth.
As already pointed out we may express the change in face
as compounded growth originated by the dentition together
with that produced by other factors including sex. Thus in
the Gorilla, a tremendous downward growth of the alveolar
region occurs in preparation for eruption of the third molar.
This, as pointed out by Doctor Todd, may be partly “respi-
ratory” in nature. (Cf. Gundobin, ’12.) At the same period
in the Orang there is, in addition, a pronounced upward tilt
of the face. I n this, however, is almost certainly a pro-
nounced sex influence since in dental age V we have more
females, and in VI more males, represented.
Stripped to their absolute essentials the growth-changes in
the cranium and face of the Gorilla, Chimpanzee and Orang
conform to a major pattern: stability of the posterior part
of the cranium; stability of cranial height of the cranium;
greatest growth in the anterior part of the cranium, spe-
cifically in the sphenoid area; early attainment of definitive
cranial form; facial growth far more pronounced than cranial
growth; lower face growing faster than upper; general tend-
ency for the face to tilt upward, after an initial downward
and forward growth. All of these are common to the Giant
Anthropoids. As among themselves the differences are those
 SKULL AND FACE GROWTH OF THE ORANG-UTAN 361

of degree rather than of kind. The Gorilla and Orang tend to

present the same type of growth pattern, constrasted with that
of the Chimpanzee, which, as will be indicated in later studies,
is apparently intermediate between the anthropoid and human
growth-change patterns.
SUMMARY
1. The study of growth changes in the cranium and face of
the Orang is based on a craniostatic and anthropometric
analysis of 134 skulls, classified according to dental age.
2. The growth of the Orang skull in length is accomplished
solely by pre-porionic expansion, and definitive length is
accomplished shortly after the eruption of the first permanent
molar.
3. The growth of the Orang cranium in height is very slight
but the squamous gains 40 percent in its vertical diameter,
apparently sliding up over the parietal t o provide for lateral
expansion.
4. The Orang face gains more in length than in height,
and there is a differential increase in length of orbito-nasal
and alveolar regions so that the lower portion of the face,
after an initial downward and forward projection, is tilted
forwards and upwards.
5. The problem of growth changes in the cranium and face
of the Anthropoids involves an interplay of proportional
growth and readjustment. Up to the eruption of the first
molar the first factor only is present, after that the pattern
is one of growth-change.
6. The Anthropoid cranium is marked by stability of post-
porionic and height dimensions, marked growth pre-porion-
ically, pronounced facial growth, most active in the alveolar
region, and an early forward and downward growth of the
face, followed by an upward tilt, most marked in the Orang.
7. Of the giant Anthropoids the growth pattern of the
Chimpanzee is nearest that of Man.
362 WILTON MARION KROGMAN

LITERATURE CITED
CAMERON,J. 1928 Pre-porion and post-porion dimensions. Am. J, Phys.
Anthrop., vol. 12, pp. 182-191.
GUNDOBIN,N. Y. 1912 Die Besonderlieiten des Kindesalters. Berlin.
HELLMAN, M. 1927 Preliminary study in development as it affects the human
face. Dental Cosmos, vol. 69, pp. 250-269.
KROGNAN,W. M. 1931 Growth changes in the cranium and face of the Gorilla.
Bm. Jour. h a t . ,
1701.47, pp. 89-116.

Figures 1 to 3
SKULL -4ND FACE GROWTH OF THE ORANG-UTAN 36.3

4 81024-----8

Figures 4 to 6
364 WILTON MARION KROQMAN

I I I I I

I
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I I I I

- I I I ' I --u)
100 50 0 50
Figures 7 and 8
 SKULL AND FACE GROWTH OF THE ORANG-UTAN 365

I I I I

I I 1 I
b
10 100 50 0 so
Figure 9