Six Lectures on Sinanthropus pekinensis and Related Problems

seems that this omission is all the more justified because the new facts instead of altering the statements made earlier on the problem concerning the descent of man oniy serve to confirm them. The most essential facts are illustrated by 55 figures compiled in Plates l-IX. For more detailed information the reader is kindly referred to former publications on the subject in question ot to such which are still

in preparation.

Franz Weidenreich Peking, February 20, 1939

BULLETIN OF

THE GEOLOGICAL SOCIETY OF CHINA. Vo{ XIX, No. 1.

MARCH 1939.

Six Lectures on Sinanthropus pekinensis and Related Problems By FRANZ WEIDENREICH (Viefung Professor of Anatomy and Hon.

Director, Cenoxoic Research Laboratory,

Peiping Union Medical College)

With lo Plates

PREFACE The six subjects selected and combined into a common title represent lectures delivered under various auspices in the United States and Europe during the spring and summer of 1938.

Lecture Ill constitutes the James Arthur Lecture for 1938 held at the American Museum of Natural History in New York, Lecture V was given before the Anthropological Section of the Deuxieme Congrès Interna-

tional des Sciences Anthropologiques et Ethnologiques in Copenhagen August 1938. New discoveries made in the interim, especially those dealing with

the various Australoplihecus types, are not included herein, since they were still unknown at the time the lectures were given. However, it

seems that this omission is all the more justified because the new facts instead of altering the statements made earlier on the problem concerning the descent of man oniy serve to confirm them. The most essential facts are illustrated by 55 figures compiled in Plates l-IX. For more detailed information the reader is kindly referred to former publications on the subject in question ot to such which are still

in preparation.

Franz Weidenreich Peking, February 20, 1939

Sinant/iropus pekinensis and his significance for the problem of human evolution For mafly years our knowledge of fossil man was restricted to almost

one group of types combined under the term of Homo primigeahus or Neanderthal man.

These types are characterized by numerous peculiarities by

which they differ from present mankind and by which they prove their more primitive nature.

in addition to that group, there existed a fossil form of

uncertain classification, namely the type represented by the famous skull cap of Trinil, which its discoverer Eug. Dubois considered to be a kind of missing

link between the anthropoids and hominids and termed it Pithecanthropus. The correct judgement of morphological characters of human remains is often impaired by too much stress being laid upon the geological conditions

and the unreserved acceptance of the idea that remains derived from older geological strata must under all circumstances reveal more primitive particularities than such from younger levels.

This belief led to the conclusion that

certain characteristics of recent man must be considered typical for bominids from the beginning of their differentiation and that the retention of the original

primitiveness would therefore represent a more or less rundamental difference between anthropoids and hominids.

The discovery of Sinanthropus pekinensis shed new light on this problem.

Indeed, it is a fact that certain anthropologists at first refused to see

more in Sinanthropus than a mere variation of the long known Neanderthal type.

However, as additional material comes to light and as more complete

studies are made of the specimens in hand, it becomes evident that Sinanthro-

Bulletin of the Geological Society of China

2

pus is a hominid type of his own., and, though. closely related to the Neander-

thal man, must be considered much more primitive than the various representatives of that group.

In other words, Sinanthropus is, morphologically

speaking, an ancestral form of the Neanderthal group and its immediate preceding phase in:the course of human evolution.

It is well known that remain of fossil man in most cases merely consist of one skeletal part or a fragment thereof and rarely refer to numerous individuals.

The Krapina remains form such an exception. An entire

skeleton which is more or less preserved is only available of La Chapelle-auxSaints. In those cases in which we only have one single specimen it, of course,

is impossible to decide whether a given peculiarity repres2nts an accidenTwo of the most important discoveries which in past decades have played a decisive role in the course taken by tal variation or a typical character.

palaeoanthropology, namely the Heidelberg mandible and the skull cap of The conditions involving the Sinanthropus Up to the present, remains of at least 40 indivi-

Trinil fall within that group. material are quite different.

duals of both sexes have been recovered, ranging from four or five years old to

a rather advanced age. It is true, that in this instance also, we are not dealing with entire skeletons but rather with skulls and teeth. However, this material is so complete and numerous that it permits not only a reconstruction

of the entire skull but also the determination to a certain extent of the range of variability.

The cranial material consists of 5 brain cases and 10 frag-

ments which belong to six additional individuals, foür larger and several smaller parts of the face, including three fragments of upper jaws, belonging to four different individuals. Furthermore, we have 14 lower jaws of all ages

and finally 150 teeth which partly represent isolated specimens or are embedcled within the jaws.

Our collection comprises all types of teeth mostly of

both sides and with several specimens of each type, only the upper deciduous

Sinanthropus and his significance in evolution

3

incisors and molars are lacking. A g1eat number of the teeth is in a completely unworn state. So as to begin with the main conclusion, we may state that the variation

of Sinanthropus peinensis embraces types of an extraordinary primitiveness

on the one hand, and on the other, those which approach the lowér limit of the Neanderthal stage, thus making the latter appear as the next morphological phase within the line of human evolution.

This fact is supported not only

by the features of the brain case but also by those of the lower jaws and It is of the greatest interest, however, that both primitive peculiarities and more advanced ones may be found to exist in different individuals as well teeth.

as side-by-side in one and the same individual.

In regard to the brain case, there are six structures characteristic for Sinanthropus. The first and the most striking one is the existence of a pair of heavy supraorbital ridges which, connected by means of a similar median

ridge, roof over the nasal bridge and are separated from the forehead by a well developed furrow (Figs. 1 and 7). With this feature Sinanthropus differs from the Neanderthal group in that the ridges there are morís restricted to the lateral orbital regions and continue directly to the forehead (Fig. I O). in chimpanzee and gorilla there is only a difference in development and location of these ridges but not a fundamental one when compared with Sinanthro-

pus (Fig. I a, b).

The second peculiarity of Sinanthropus is the remarkable lowness of the brain case (Fig. lb; 2b; 5a). The ratio between length and height is the same as in anthropoids (auric. height-length index: Sinanthropus Skull Il Locus L :49.2, chimpanzee: 51 .2). whereas within the NeandertlaT

group no skuJI shows such a low ratio, the lowest here being 54.7. A third peculiarity combined with the one just mentioned are the landmarks of the greatesi breadth which coincide with those of the bianricular breadth like ir1 great apes (Fig. 2a, b), whereas in Neanderthal and recent man these land.

Bulletin of the Geological Society of China

4

marks are situated on the superior portion of the squama of the temporal bone or on the inferior portion of the parietal bone (Fig. 2e).

in accordance with

this appearance, the lateral bones forming the walls of the brain case are turned medially in Sinanthropus (Fig. 2b) while in Neanderthal and recent

man they are located vertically or turned outward (Fig. 2c).

The fourth

peculiarity concerns the torus occipitalis which in Sinanthropus represents a fairly thick uniform ridge (Figs. 2h; 5a; 14), whereas in Neanderthal man the

torus as a whole is less developed and divided into three distinct portions of

which the middle one is reduced to a narrow bridge linking the two lateral ones (Fig. 2e). Sinanthropus in addition shows a continuous crest coursing from the lateral end of the torus down to the mastoid process (Fig. 14). At the point where it crosses the angle of the parietal bone it appears in the form

of a distinct roundish swelling (tp), a condition which has become almost completely obscure in the Neanderthal man. The fifth peculiarity concerns

the foramen magnum which is situated further backward than in Neanderthal

and recent man (Fig. 2h, c).

The sixth peculiarity finally concerns the con-

figuration of the basal part of the temporal bone, especially the course of the axis of the pyramis, which does not form a straight line like in other hominids

but is distinctly bent forward in its median portion like in anthropoids (Fig. 6).

The styloid process is lacking. 'All these 1particularities of the Slnanthropus brain case must be con-

sidered a consequence of the smallness of the cranial capacity. This capacity

varies between 850 cc and 1220 cc. The three best preserved skulls of which the capacites are directly measurable hold circa 1000 cc, as an average volume.

in contrast to Sinanthropus the average cranial capacity of recent

man amounts to 1350 cc and that of the Neanderthal man even to 1425. The smallest skull of the latter groupthe Gibraltar skullshows a capacity

tr

r

-

TABLE I Sinanthropus-

Pithe-

rrt uit it canth- NeanLoc. L Loc. L Lac, L Loc. E rorusS derthai juv 198

189

191

147

-

193

195

¿'

143 133

185

188

¿

94

85

140 130

177

183

80.5

--

107

147

199

201

¿'

81

118

104

147

198

203

/'

79

115

109

144 156

195

208

I'

90

116

100

142 146

204

206

sian

195

145 132

90

81.5 96

66.2

pelle

man8

44°

40.4 38.4

122°

68°

46°

40.9 34.5

54.7

68.5°

45.5°

36.5

700

48

36.4 28.0

113°

1325

¡07°

40.5

1620

39°

500

¡/9° ¡/0° 1325 410

59° 84°

56.3 63.8 51.3 45.4 59.8 32.5 50.0

panzee

Neanderthal group Spy L LaCha-Rhode- Recent Cl8im-

148 136

84

102

73

Jj?.

89

94

77

¿

1. Greatest length (g-op) 2. Glabeflo-inion length (g-i) 3. Greatest breadth: a. auricular b. tempero-parietal 107.5

70

Sex

4. Last frontal breadth

84

40.9

66.5°

42°

118°

70.9 52.2

51.0 .34.4 38.9

70°

72.4 39.7

43°

99°

73.1

49.2

40.3

67°

75.0 55.4

37.Z

44°

1 10°(?)

-

68.7 53.4

42.1

63°

70.7 43.1

40°

97°

69.2 39.5

60°

1000? c.1370 -

71.0 51.4 37.9 47°

1040

c.1200 c.1015 c.11OO 915

-

74° .

- -

5. Auricular-bregmatic height (p0-b) 6. Calvarial height (over glabelloinion line)

.

2) Reconstruction after Weinert. 3) Average figures of all races.

7. Cranial index 8. Length-auricular height index 9. Calvarial height index 10. Bregmatic index. (Schwalbe) 11. Bregmatic angle (Schwalbe) 12. Superior inion angle (Hg) 13. Irnon angle (ho) 14. Capacity (cc) 1) Corrected measurements.

Bulletin of the Geological Society of China

6

of 1 290 cc, that is 70 cc more than that of the largest Sinanthropus skull and

about 400 cc more than that of the smallest one.

Table I gives the main measurements of the brain case of the four Sina'ithro pus skulls compared with those within the Neanderthal group, recent

man (average) and chimpanzee.

lt is evident from these figures that Sinon-

thropus differs considerably from the Neanderthal group and distinctly approa-

ches the anthropoids represented here by the chimpanzee.

The peculiarities of the endocranium of Sinanthropus as demonstrated

by casts complete the picture with respect to the brain. The kam is small

and flat (Fig. 25b) particularly within the vertex and obelion regions. In addition, the temporal lobe is very narrow and separated from the cerebellum

by a wide and deep notch (Fig. 27a). A feature similar to the last mentioned may only be observed to occur in the Rhodesian man but not in any of the other representatives of the Neanderthal group nor in recent man

(Fig. 27 b, c).

Strikingly pithecoid is the appearance of the entire area of

the frontal pole (Fig. 22e).

The frontal lobe is small and tapering in the form of a keel-like orbital rostrum which protrudes forward and downward. This formation together with the entire pattern of convolutions and fissures closely resemble the corresponding features in the chimpanzee brain (Fig, 22b).

in recent man as well as in Neanderthal man the area of the frontal pole is broadened añd flattened and the pattern much more cómplicated (Fig. 22 e-g). The grooves of the middle meningeal artery are of special interest The rami-

fication of this vessel reveals an anthropoidlike, especially gorilla-like type

(Fig.23 a, b). The posterior main branch (r. temporalis) is the largest, longest and the most ramified of the two or three main branches, whereas the anterior branch (r. frontalis) is weak, short and poor in its ramification.

Exactly the reverse conditions exist in recent man (Fig. 23 e, d).

Nean-

'lerthal man follows the recent human type, with the exception óf the lhode-

Sinanthropus and his significance in evolution

7

sian man who displays a closer resemblance to the Sinanthropus pattern of ramification.

With respect to the facial skeleton the differences between Sinanthro-

pus and the Neanderthal group do not seem to be as striking as those of the

brain case (Fig.. 3 b). Yet the more pithecoid character is also rather pronounced here. The orbits re very deep and relatively spacious. The lacrimal fossa is 'lacking. The upper orbital fissure is narrow and short. The appearance of the lateral borders of the orbits resembles the conditions of the gorilla. The interorbital breadth is considerable. The nasal bridge

is slightly flat and broad but the pit between the forehead and the nose is rather shallow (Fig. I b). The apertura piriformis is low and wide. The zygomatic bone is high, flat and directed more nteriorly than laterally (Fig.

I b). There is evidence of a strong total prognathism of the maxilla which causes the lateral border of the apertura piriformis to be distinctly forwardly

directed (Fig. 1 b).

The anterior nasal spine is lacking and likewise a dis-

tinct threshold at the entrance to the nasal floor (Fig. 3 b).

The anterior

surface of the maxilla protrudes and is divided by a narrow groove, coursing from the infraorbital foramen toward the second premolar, into a medial and lateral portion.

The jugum of the canine represents a longish swelling sup-

porting the lateral wall of the nasal cavity (Fig. 3 b). vaulted and high.

The palate is widely The zygomatic arch is missing but from the conditions of

its connections with the skull it may be seen that the arch extended .a con. siderable distance away from the skull in a relatively low level and slightly declined in forward. direction (Fig. 1 b). The face as a whole shows that its upper part, because of the heavy and far laterally projecting supraorbital .

ridges and the similarly located cheek bones, is broad and low, whereas the breadth of the projecting alveolar part of the maxilla is relatively narrow

(Fig. 3 b). These features give the face a rather pithecoid appearance which

8

Bulletin of the Geological Society of Chino

is not attained even by the Rhodesian skull.

The great variability in (Fig. 43

b) size and pattern of the mandible, partly due to the difference in sex, renders it dîfficuit to recognize the most characteristic features.

Apparently as such.

however, must be considered the position of the digastric fossa at the basal surface, the steepness and breadth of the ramus, the narrowness of the dental

arch and the regular curvature in its frontal part. Features which the jaw of Sinanthropus has in common with the martdibles of the Neanderthal group

concern the strong inclination of the symphyseal region and the absence of a chin (mentum osseum) - Fig. 43.

The responsible factor for Sinanthropus representing a type of his own, however, among all homimds known hitherto and for his ranging within

the order of anthropoids is the character of his teeth

The sigmficance of

such a criterion is all the more important because palaeontology always considered the character of teeth as decisive for classification. The Sinanthropus

teeth as a whole are much stronger in crown and root than any teeth of the Neanderthal group and more so than those of recent man. This s especially true of the canines and curiously also of the first upper premólar.

The lower

premolars and molars are low and long (Figs. 38 and 39), while they are high

and short in Neanderthal and recent man. The persistence of a cingulum (Fig. 38 a, b; Fig: 39 c., b) and even indicatioús of stylai cusps are almost standard features. The lower molars (Fig. 32 a; Fig. 36 e) show a typical

Dryopithecus-pattern, and frequently six cusps will be found with the mcta-

conid representing 'the largest and longest cusp.

The trigonid is broader

than the talonid. The entire chewing surface is covered by wrinkles in specific formation and arrangement.

The second lower premolar has a long

talonid and a short trigonid (Fig. 39 a., o; Fig. 41 b), the latter at the same time being higher than the talonid and protruding toward the inesial side. This gives the tooth an asymmetric shape with its longitudinal axis running

Sinanthropu and his significance in evolution

9

in oblique direction from the mesiobuccal side to the distolingual one (Fig.

41 b).

The first lower premolar is provided with a small isolated lingual

cusp limited by a cingulum-like border (Fig. 40 b).

The talonid is developed

here also and thus causes the tooth to be asymmetrical.

The two premolars

which show these formations most distinctly are represented by germs and

were completely embedded within the mandible.

They resemble certain

chimpanzee premolars (Fig. 40 a.; 41 a) so closely that, if found as isolated

teeth elsewhere, their hominid affinity would have been recognizable only with great difficulty.

The Sinanthropus canines have a special character in

so far that the upper and lower differ from each other in their shape (Fig. 33).

While the upper canine is conical, the lower one rather has the appea-

rance of being an incisor with a but slightly pointed edge.

They differ from

the canines of the anthropoids (Fig. 34 a; 44 a) in that they are less massive,

less high and lese pointed than the latter.

The upper Sinanthropus canine,

however, bears a surprisingly close resemblance to the lower canine of the fossil orang of Yiinnan, especially in regard to the development of the cingulum

(Fig. 34).

The upper central incisors show a very strong lingual tubercie

which continues into several prolongations, a feature rather characteristic for the respective teeth of anthropoids.

All these peculiarities of the Sinanthro-

pus teeth are either completely lacking within the dentition of the Neanderthal

group or at best faintly indicated.

In respect to the size of the individual types the molars of Sinanthro-

pus are larger than those of chimpanzee and equal to those of orang, while

the incisors are much smaller than those of the two apes (Fig. 42). The gorilla, however, shows the same ratio in this respect as Sinanthropus, with

the only difference that the entire dentition of the former is much stronger than that of the latter.

Bulletin of th Geological Society of China

IO

A particularly primitive character of the Sinanthro pus dentiton is the

persistence oí a distinct paraconid in the first and second deciduous molar. In the latter tooth, not only the paraconid but also the entire trigonid is preserved in such a completeness as is never found in either fossil or living anth-

ropoids; however it recalls the respective conditior2s of Eocene Tarsioids

(Fig. 37). The sequence of the eruption in Sinanthropus also reveals anthropoid characteristics.

\Vithin the deciduous dentition the lower canine erupts last,

whereas in recent man jt makes its appearance before the molars.

The

second lower molar of the permanent dentition cuts the gum during the process

f eruption of the permuent teeth like in anthropoids and Neanclertha man but not after as in recent man.

Although all the peculiarities desrribed until now prove th special primitive character of Sinanthro pi's when compared with the other fossil hominids, yet his close relationship ti Neanderthal and recent man is evident

within the range o his variabity. As apparent from the figures of Table I single iudivituals approach the lower boundary of variations within the Neandei ial group so closely, especially in regard to size and form

of .ie brain case, that there is an almost continuous line leading from the sinanthropus phase through that of Neanderthal man to that of recent man. As regards direct genetical relations between these three stages, most palacoanthropologists are inclined to consider the Neanderthal group a discOntinued

branch with no bridge leading to recent man. Hrdlicka is one of the few who at all times was of different opinion. The reasons generally offered loT the alleged specialisation of Neanderthal man refer in particular to the roomi-

nes of the pulp cavities of the lower molars, the so-called "taurodontism.' lt can easily be demonstrated that taurodontism is not a speciality of Neanderthal man only but also a characteristic of Sinanthropus. Furthermore. rooiw

Sinanthropus and his significance in evolution

11

pulp cavities may he found in the same degree of development in present an-

thropoids (female orang) as well as in recent man (palaeolithic recent man, Eskimo, Bushman, Amerindians).

The question whether Sinanthro pus himself bears indications of spe-

cialisation had already been raised by Davidson Black and answered in the negative. Marcellin Boule recently admitted that Sinanthropus represented direct ancestor of recent man.

Indeed, the new discoveries have not con-

On the contrary, there is distinct evidence of the presence of several features, though tributed anything which could he interpreted as specialisation.

irrelevant from the phylogenetic standpoint, which nevertheless occur in exactly the same manner and degree of formation in recent mankind.

The first peculiarity is the development of a strong sagittal crest at the crown of all Sinanthropus skulls discovered hitherto (Figs. 2h: 3h).

Such a crest is faintly

indicated in Pithecanthropus, somewhat more distinct than in the latter in the skull of the Rhodesian man, but very pronounced in Eskimo, Northern Chinese, Amerindians, Australian and Tasmanian (Figs. 16 and 17). The second peculiarity of general human character is the persistence of a metopic suture in Skull II of Locus L (Fig. I 6a).

This suture is extraordinarily rare in anthropoids and has never been found in any of the skulls of the Neander-

thal group but is common in recent man, regardless of races. The third peculiarity is the presence of a so-called os epactale or Inca bone in three of the four Sinanthrupus skulls in which the occipital hone is preserved (Fig. 2b). This peculiarity may also be found to a certain extent within the entire

mankind of today, but in Amerindians it occurs in more than 7%. The fourth peculiarity is the formation óf the so-called torus mandibularis which is developed on three of the five adult Sinanthropus mandibles with this region prçserved.

This peculiarity is a rather common occurrence in Eskimo and

Lapps, and, to a less degree, may also be observed in Chinese, neolithic Jap-

Bulletin of ¿he Geological Society of China

I2

atiese, Icelanders, Norwegians, Dinaric types and Bushman.

Th fifth pecu-

liarity is the shovel sh3pe of both upper incisors, especially of the lateral ones,

a feature characteristic for present-day Mongolians. These facts prove that Sinanthropus not only ranges within the direct

line of evolution but also bears a certain relationship to the Mongol group of For four of the five just mentioned pcculiaities are specifically developed in branci3es of this race, whereas they constitute a mOre or recent mankind.

less rare occurrence in other.

This appearence of special Features within

certain racial groups and their absence in others, indicates.jn addition that the

differentiation of races can be traced back to a much earlier time than is generally supposed.

That Neanderthal mañ had already divided into several

racially characterized types becomes more evident with each new discovery. As I have shown ten years ago, the skulls of La Chapelle-aux-Saints, Rhodesia

and Homo soloensis (Java) undoubtedly represent three different fossil racial types which may be distinguished as the European, South African and Sunda branches of HQmQ primigenius.

Sir Arthur Keith arrived at the same con-

clusior*, namely that the existing separation of mankind into different races is

very ancient having already started before the Pleistocene period and that

each racial stock was evolved in the very territory occupied by it at thê dawn of its history.

Thus we come to the question concerning the genetic relationship between Sinonthropus and the other hominids.

Must Sinanthropus be considered the direct ancestor of the entire mankind of today? In other words, are all the other fossil hominids in the world who represent a more advanced

stage than Sinanthropus direct descendants of the latter? This question touches the essential point of the theory on human evolution. Under the dominant influence of Haeckel's interprettiora f Darwin's theory, it was generally assumed that a cradle of mankind must have existed, temporally and

Sinanthropus and his significance in evolution

13

regionally fixed, that is to say, that humn evolution originated exclusively

within a single centermonocentric evolutionwherefrom all the new types evolved from time to time.

These newcomers spread over the entire world,

thereby compelling the older types, that is the more primitive ones, to gradu..

ally retire to the remotest boundaries of the habitable world. The localisation of such a prolific center varied according to the latest discoveries, once Africa was considered to be that cradle, then Asia. This idea furthermore led to the conception that certain branches are discontinued while other, geologically

older but nevertheless more advanced ones, have no relation to more primitive forms,

All these difficulties may be overcome immediately if we consent to

suppose that the early ancestors of man from the very beginning of their appearance were dispersed over a great part of the world and developed independently from each other within the regions accidentally selected for habitation.

In such case the connecting tie between the various types would

consist only of their general tendency to develop into man, whereas local 3ifferences would be restricted merely to special features which we call racial characters. lt is well known that the latter never affect the general nature of the various branches of mankind. According to such a hypothesis poly-

centric evolution -the Far-Eastern Sinanthropus probably living during the same epoch as the already more advanced Heidelberg man of Central Europe cannot be regarded as a direct progenitor of the latter but nevertheless still as one representing an ancestral form of mankind and thereby be consubstantial with thé real ancestor of that European type. The evolution of this type would have taken place in another part of the world and at a much earlier period than in the case of Sinanlhropus.

The idea of there having existed various centers of evolution seems strongly supported by the important discoveries recently made in Java.

In

earlier discussions of the relation between Sinanthropus and Pithecanthropi,s,

Bulletin of the Geological Society of China

14 I

expressed the opinion that Pithecanthro pus and Sinanthropus represent

the same stage of evolution with but slight differences, which may be of racial character.

in addition, I considered Pithecanthropus, a slightly more advan-

ced stage in respect to the special appearance of the frontal region of the brain case.

At the same time I called attention to the fact that there is a

close resemblance between Pithecanthropus and Homo soloensis, a representa-

tive of the Neanderthal type of Java.

I reached the conclusion, that despite

the great difference in size the two types coincide in their general form and in certain special details so completely that Homo soloensis must be consider-

ed as an advanced Pithecanthropus form. The discovery of an almost complete brain case of a second Pithecanthropus skull by Dr. R. von Koenigswald gives this idea further support.

The Wadjak and Talgai man possibly

represent the earliest fossil type of recent man of this special hominid line. The theory of a polycentric evolution simply explains the incontestable

fact that present mankind also, despite the conformity in general human char-

acters, embraces such primitive groups as the Australian native or Bushman

on one hand, and those of the advanced European on the other. With reference to the Australian, it may be that we are not actually dealing with a primitive type remaining stationary for a long time but rather with one merely

having the appearance of primitiveness because it was left behind by other forms which underwent an earlier and more speedy evolution within different

regions. The general resemblance between the Australian aboriginal of today

and the Upper Palaeolithic European would then be a consequence of both representing a similar stage of evolution rather than that of an early immigration of the Palaeolithic European into the Australian continent.

The discovery of Sinanthropus peinensis finally yields a more certain basis for tracing not only the course of the sequent stages of evolution but

also of the preceding ones.

It has been demonstrated that Sinanthropus,

Sinanthro pus and his significance in evolution

15

according to his zoological character, must be ranged within the anthropoid group.

Thus, we are justified to assume that those preceding types approa-

ched the anthropoid group more than Sinanthropus himself.

lt is not permis-

sible, however, to identify the ancestor to Sinanthropus directly with either gorilla, chimpanzee and orang nor with their supposed forerunner like Dryopiliiecus, Sivapithecus or any other. The nature of the Sinanthropus denti-

tion, especially of the canines and the first lower premolar, is incompatible

with such an attempt. This, however, does not hold good for one fossil anthropoid form which has become known within recent years and is repre-

sented by two specimens, namely AustralopiLhecus Darta juvenileand Austratopithecus Brooman adult. The teeth of the juvenile Auslralopithecus resemble the Sinanthro pus teeth closer than they do those of gorilla or chimpanzee.

The canine is missing in the adult specimen but the size and

shape of its alveolus and the lack of a diastema indicates that this tooth must have been much smaller than the canines of great apes.

As is revealed by

the character of the skull Australopithecus must nevertheless be considered a real anthropoid and not a hominid. It follows from these facts that this Afri-

can type, together with Sinanlhropus, belong to a special anthropoid stock which must be distinguished from that of Dryopithecus.

The former stock

includes anthropoids with a "homomorphic" canine group, whereas the latter

The first apparently retained its primitive primate character, while the latter lost it by special

embraces such 'with a "heteromorphic" canine gn.ip. differentiation.

Thus far we have only referred to skull parts of Sinanthropus with no mention of spine and limb-bones. The reason for so doing is that very few extremity bones have been discovered until date. This scarcity is not because the bones have been dissolved in the course of time or because of our oversight but obviously because Sinanthropus himself (ailed to deposit them at

16

Bulletin of the Geological Society of China

the site of our excavation.

Until nowen fragments of spine and limb-bones

have been unearthed, namely a fragment of an atlas, the greater part of a collar bone a part of the distal end of an arm-bone, a carpal bone (os hmatum)

and quite recently six fragments of femora (Figs. 45-47; 51; 53; 54) and one of a humerus.

Notwithstanding the incompleteness of these specimens and

thé scarcity of the entire material, it is possible to state that these bones display a much more human-like appearance than the teeth and the skull of Sin-

anthropus do.

The character of the femur proves that Sinanthropus already

had adopted an erect posture (Figs. 45;47). lt must concluded from this fact

that the body of the hominids attained the stage of recent man much earlier than the skull. As is evident from the cultural remains, Slnanihropus. already knew the art of hunting and the use of fire, and was quite skilled in the

manufacturing of tools and their uses. Such an achievement, of conise. could

only have been made if Sinanthropus did not have to rely on his hands for locomotion.

lt is true, Boule voiced the opinion that the culture of Choukou-

tien does not beking to Sinanth ropas but to a more advanced hominid type, whose physical remains are still to be found. This conception merely based

upon the idea that the Sinanthropus brain is too small to be capable of pro.. ducing such an advanced cultural standard, is contradicted by the entire course and the results of the excavions. As to the size of the brain the capacities of the SinanthroPus skulls vary between 850 and 1220 cc The maximum value is considerably higher than the minimum value for recent man and is in excess of the average cranial capacities of anthropoids by more than twice their volume.

The fact that a hominid with such primitive characters in teeth and skull already had adopted an upright posture suggests that the transformation of the lower limbs, previously adapted to climbing, must have taken place much earlier than is generally supposed. From the character of the teeth it may be concluded that the divi5I0 of the anthropoids already occurred before

Sinanthro pus and his significance in evolullori

7

the differentiation of Druopilhecus took place, implying that the branching-off

of the hominid line must be dated back at least to the lower Miocene. Thereby

the frequently raised question concerning the existence of the Tertiary man looses its foothold.

For, no matter to which Pleistocene period Sinanthro-

pus may be attributed, that first differentiation occuned in the Tertiary epoch.

Although numerous essential questions remain open, yet one may be considered as solved, namely that man developed from a primate form which, from the viewpoint of zoological classification, represents an anthropoid.

In

some details he bears a closer resemblance to the gorilla, in others to the

chimpanzee or the orang.

For provided, as we believe, the recent anthro-

poids represent specialisations derived from one original common stock, the differences or resemblances between the individual forms of the anthropoids,

on one hand, and the hominids, on the other, may be due as much to the condition that one form retained common primitive characters more persistently

than others. A certain conformity, of course, must have existed among all the representatives previous to being separated. - But, on account of the com-

plete absence of adequate fossil material, it is not worth our while to deal with this question for the moment

II. The general and special charactcr of the SinanEhropu dentition When discussing problems concerning the descent of man, teeth have at all times played an important role. Yet all controversies could be reduced

to the relatively simple question, namely whether man and his ancestors represent a line of their own originating directly from an indifferent and low primate type or whether they deviated relatively late from an already higher differentiated form the organization of Which

of present anthropoids.

Speaking of anthro

corresponded in principle to that

d in this sense we must bear in mind that it implies only a certain priate group which approaches man

closer than any of the other d0, omitting the various representatives themselves of

minor differences as exist among

any one group. 1f it so happens that the hominids, for instance, resemble the chimpanzee in their essential features more than a lemur or a spider

or a baboon, we are justified in

ranging the hominids within the same order as the chimpanzee. The only question remaining open for further dise550 would be the assignment of the special position within this group

lt is known that Huxley was the first to recognize that man, according to the entire structure of his body, had to be considered the closest relative of the anthropoids. This famous Esgj5 anatomjst based his statement chiefly on the results of comparative stud of the organs of modern man. The knowedge of fossil man at that time Was rather poor, being restricted almost exclusively to the Neanderthal ma of Dusseldorf; All the discoveries since then have served to confirm Huxley'5 viewpoint and at the saine time 18

General and special character of Sinanlhropus dentition

19

demonstrated that that similarity increases in the same degree as the fossil types differ from recent man. Only the teeth did not seem to fit into this scheme satisfactorily.

The

dentition of recent man is first of all distinguished from that ofthe anthropoids by the complete lack of a diastema in the upper and lower jaw and correspon-

ding therewith by the absence of tusk-like caninès.

As to the other teeth they are relatively smaller and their pattern is considerably less complicated than is the case in all of the genera of anthropoids.

That which is known of the teeth of fossil man failed to support the idea of really striking differences from the conditions of modern man in contrast to expectations.

There were,

of course, a great number of special features showing a distinct approach to a more primitive stage but their intepretation as "pithecoid" characters remained debated.

The discovery of the Heidelberg mandible was greatly responsible

for such a skeptical attitude.

in this case a jaw of a very primitive character

is combined with relatively small teeth, the general appearance of which bears a closer resemblance to recent man than the bone would suggest.

Hence the

conclusion that the hominids had preserved the simple character of the teeth from.the very beginning of their differentiation and never passed through an anthropoid-like organisation during their evolution.

Such a viewpoint would prove erroneous in any case.

For the teeth

alone could not render invalid the evidence yielded by all the other organs. But the discovery of Sinanthropus revealed that the idea of the teeth of re cent

man being of a real primitve nature is based on the incorrect conception that a

speeimen recovered from a geologically older stratum must under all circumstances and in all its parts represent a more primitive character than such found at a younger formation. For, the particularities of the Sinanthropus teeth completely harmonize with the marked degree of primitiveness

manifested by the skull itself but the considerable' range of their variability

20

Brillelln of the Geological Society of China

at the same time proves the impossibility of drawing cogent conclusions merely

ort the basis of one single object. The Sinanthropus teeth now available number 147 specimens compos-

ing the entire permanent dentition and the lower set of the deciduous teeth. Each type is also represented by several unworn specimens consisting of germs

or teeth just erupting.

63 of the teeth are isolated while the remaining ones

are still embedded within the corresponding jaws

The character of all the teeth is the same. First of at! they are larger on the whole than the teeth of recent man. This holds good not only for the crowns but all the more for the roots, especially for the roots of the incisors, canines and premolars. Besides, a considerable difference in size is to be noted between the Sinanthropus teeth themselves which can only interpret-

ed as being due to difference in sex, the larger ones belonging to male and the smaller to female individuaJ. in all cases the patterns are much more complicated than those of recent man approaching the anthropoid tooth pattern to an astonishing extent. Taking all into account, the Slnanihropus teeth

resmble those of the anthropoids much closer than they do those of recent man, with the only exceptions of the canines and the first lower premolar.

On the other hand, there is no indication whatever that any other primate group approaches Sinanthropus closer than the anthropoids do. The gap existing between anthropoids, other Catarrhinae, Platyrrhinae and Prosimians

remnin unchanged and undiminished despite the accession of the prehominid type. The fact to be admitted however is that the Sinanthropus teeth have

retained certain primitive characters more intensely than is true of any of the

other higher primate group, particularly the arithopoid. Yet; this fact cannot affect the general rdationship between hominids and anthropoids but only their special positions within the common group.

Besides the general features just mentioned the anthropoid character is especially evident in the second lower premolar and the lower molars.

As

General and special character of Sinanthropus dentition

21

to the general shape and pattern of the premolar (Figs. 39 a, c; 41 b-d), it is 1) low and long, 2) .there is a well developed cingulum on the labial side, 3) the crown is asymmetric by the presence of a large and well developed talonid

whereas the smaller trigonid embracing the two cusps has completely shifted

to the mesial side, 4) the large and high root is furrowed on the mesial and distal sides indicating a division into a labial and lingual portion. All these traits are also characteristic for the second lower premolar of the anthropoids (Figs. 39b; 41 a).

In recent man this tooth is high and short., a cingulum i

completely missing, the crown is symmetric on account of the total loss of the

talonid and the cusps occupy the midline of the tooth, the root being rather simple (Figs. 39c1; 41 e,f).

The first and second lower molars (Figs. 32a,b; 36e) ate like the pre-

molars low and long, with a greater length than breadth. A cingulum or corresponding differentiations are rather pronounced on the labial side. The trigonid portion of the crown is distinctly broader than the talonid portion. The number of the cusps amounts to five or, in the majority of cases, to six,

three of which occupy the labial and three the lingual moiety. The metaconid is the largest cusp, larger and especially longer than the protoconid. The hypoconid is smaller than the entoconid.

The arrangement of the cusps shows

a typical asymmetric "Droypithecus. pattern?

The inner slopes of the cusps

are covered by wrinkles in typical position ad development.

The roots

are large and stout, their distal branch deviating, straighter, higher and longer

but narràwer than the mesial one; the latter terminates in two distinct tips.

All these peculiarities are characteristic for anthropid molars (Fig.

36 a, b, c). In recent man (Figs. 32c; 36f) the molars are high and short, especially the second which is quardrangular. A cingulum is missing.

The talonid portion is broader than the trigonid one. There is a distinct tendency towards reducing the number of cusps to four, especially in the second molar, the metaconid being of the size as the protoconid, occassionally even smaller.

22

Bulletin of the Geological Society of China

The arrangements of the cusps show a symmetrie "cross" -or "plus" pzLttern. Wrinkles are usually absent.

The roots are slim, the branches convergent

with both of the same size.

The teeth of the Neanderthal group occupying an intermediary posi-

tion between Sinanthropus and recent mn bear a closer resemblance to the latter's teeth than to those of the former. Notwithstanding this fact, numerous

details reveal the direction taken by the dentition in the course of human evolution. The essential tendencr is the reduction in size. This becomes manifest not only in the decrease of the height of the roots and their robustness

or the special shortening of the crowns of the premo!ars and molars but also in the reduction of number and size of the cusps and the smoothness of the chewing surface due to loss of the wrinkles. This reduction and simplification

i concomitant to the reduction of the entire masticatory apparatus represented by the decrease of the robustness of the jaws and the shortening of the muzzle

(Figs lb, C; 43 b, c). Viewed from this general standpoint the change of the dental character is in accordance with the transformation of the whole skull leading from the anthropoid-like Sinanthropus stage to Homo sapiens.

That the character of the teeth of recent man supposedly representing an originally primitive one, should have preserved everyone of those original traits

throughout all the phases of evolution turns out to be a legend,

The relative

simplicity of the teeth of modern man is a secondary acquisition.

Despite the general similarity between the anthropoid and hominid dentition, there are some differences of great significance.

These in volve, as

already mentiâend above, the canines and the first lower premolar, The anthropoid canines bear a tusk-like character, are high, sharply pointed, very

massive at the base and provided with high and strong roots (Fig. 44 a, b). In addition, there are wide gaps between the canines and the second incisor

of the.upper jaw and the canine and the first premolar of the lower jaw foi lodging the respective canines in occlusion. The size of the canines and the

General and special character of Sinanthropus dentition

23

width of the diastemata vary not only according to sex but also individually

but large canines and wide gaps present the rule. in Sinanthropus gaps do not exist (Fig. 44c) nor do crown or root of the canines, even in the maximum of their development, reach the minimum value true of anthropoids.

Besides, the form of the crowns themselves show remarkable differ-

While the cingulum in recent anthropoids is faintly developed being restricted to a narrow ridge encirching the base, in Sinanthropus it ences.

represents a broad and distinct formation with special differentiations on the

mesial and distal sides (Fig. 34b). A comparison of the proportions of canines of Sinanthropus and recent man (Fig. 34c) shows that a considerable reduction in size has taken place in the latter, suggesting that the f ore-

runner of Sinanthropus may have also been equipped with bigger canines than is true for his descendant. At any rate, I consider the differences in the entire character of the crowon sufficiently substantial to warrant a sep-

aration of the hominids from the anthropoids as far as the latter are represented by the three living types: gorilla, chimpanzee and orang.

Of special interest is the different sppearence of the upper and lower canine of Sinanthropus (Fig. 33). In recent man these teeth appear so alike as to render a distinction between them quite difficult.

in anthro-

poids, on the other hand, in addition to differences in shape and dimensions at the base of the crowns, the upper canine is higher and more robust but, as a rule, both canines seem fairly alike with regard to the configuration of the upper parts of the crown. The crown of the upper canine in Sinanthropus (Fig, 33a) is stout and pointed whereas that of the lower one (Fig. 33b) is relatively flat and shows a curved edge instead of a pointed tip,

This peculiarity gires the tooth an incisor-like aspect rather than one

of a typical canine. in addition, the special patterns are also different. It is true, both teeth show a distinct cingulum of the same character but the arrangement of the ridges and folds of the lingual surface differ in a

24

Bulletin of'the Geological Society of China

characteristic manner. In the lower canine the distribution is such that the entiie crown seems somewhat bent distaiward, while that of the upper canine appears straight. Another decisive difference between Sinanibró pus and the anthro-

poids involves the first lower premolar. This tooth shows a "sectorial" character in anthropoids (Fig. 44a, b); it is asymmetric, its mesial portion being prolongated mesial-and labiaiward and the two cusps substituted by a strong, pyramidal top with a sharp mesial edge. In -Sinanthropus the tooth (Figs. 38a; 40 b, c) is only slightly asymmetric with no prolongation and there are two distinct cusps, though the lingual one is smaller and lower than the labial. A certain variability in canines as well as in the first premolar also occurs in anthropoids with respect to the special features under discussion. In some cases an approach even to the conditions existent in Sinanthropus

may1 ie evident (igs. 34a, 40a). Before discussing the nature of the relationship between Sinanthropus and the hominids on the one hand and the anthropoids on the other, we must clarify again the meaning of the latter term. At the beginning of

this lecture I left no room for doubt that the general character of the Sinanthropus dentition is similar to that of an anthropoid when compared with the dentition of other representatives of the primate group. Notwithstanding this fact there are certain differences which necessitate a discrimination within the. anthropoid group Itself. According to my opinion, the differences in question are not of the same nature es those existent between

gorilla, chimpanzee and orang because they involve traits common to all these genera. Sinanthro pus and therewith the hominids are anthropoids with a more "homomorphic" canine group (Fig. 44c), while the other anthropoids are characterized by a more "heteromorphic" canine group

General and special caracter of Sinanthropus dentition

25

(the first premolar considered as belonging within the canine group) - Fig.

45 a, b. The question which now arises concerns the genetic relations of the

two groups. Must the homomorphic group be regarded as a late differentiation of the heteromorphic group or is the reverse true? Such a decision depends. entirely upon adequate fossil material. Some authorities take Dryopjthecus for the common ancestor of the horninids and anthropoids of

today. Since Dryopithecus and all the related fossil types already exhibit a rather pronounced heteromorphic. canine groupf the hominids would represent later differentiations or, in other words, the hominids formerly equipped with tusk-like canines and a sectorial first lower premolar must

have subsequently lost these peculiarities.. The facts revealed by Sinanthropus, however, fail to favour such a view. The general and special appearances conform, it is true, to a surprisingly high degree to the anthropoid dentition but the canine group is distinctly homomorphic. On the other hand, Australopithecus, Dart's juvenile form as also Broom's

adult one, has a dentition very similar to that of Sinanthropus, especially tÑe for the canine group since a diastema is missing in the upper jaw and the size of the upper canine not iñ excess of that of Sinanthropus Nevertheless, however, the general shape, the size of the brain case, the character of the brain and certain other characteristic features - existence of an ape-like premaxillaprove that Australopithecus cannot be consklered a real hominid. This type apparently retained some traits of the commön anthropoid stem before dividing into bominids and anthropoids in the strict sense of the word.

Another reason why I consider hominids a more primitive stock and the Dryoplthecus group as a secondarily differentiated one is the fact that the dentition of the former shows the retaining of some very primitive

26

Bulletin of the Geological Society qf China

features, whereas they have become lost in the latter. The difference in shape and pattern between the upper ar4 lower canines ¿lescribed above constiutes some of these peculiarities, especially the incisor character of the lower canine and the persistence of a welL developed cingulum in the upper

and lower canines. The con-sectorial habitus of the first lower premolar and the complete absence of diastemata point to the same direction. Of special importance in this respect is the second deciduous molar of Sinanthropus (Fig. 37b). The trigonid and its cusps, including the paraconid and the distal trigonid crest here are well preserved resembling the charac-

terè of the permanent molars of Eocene Tarsioids (Fig. 37a). The trigonid, n particular the paraconid, have not been retained in such completeness in either Dryopithecus itself (Fig. 37c) nor in any of the recent anth-

ropoids and the distal crest is more or less reduced in all of these cases (Fig. 37d). Oa the other hand, it is evident, that the special differentiations of the canine group in recent anthropoids are secondary acquisitions. This is manifest in their great variability in the degree of achievement with a distinct tendency to return to primary conditions The cingulum of the canines for instance is, as mentioned above, usually restricted to an extre-

mely narrow crest surrounding the base of the crown but, especially in chimpanzee, it occasionally occurs as a broad band ascending on the mesial and distal sides forming special features there which are quite similar

to those observed in the Sinanthropus canines. Moreover, the lower canine of a fossil orang derived from a cave in Southern China and contemporaneous to Sinanthropus exhibits, in contrast to the regular conditions

of the orang of today. a broad band-like cingulum with the same enlagement of the mesial and distal sides (Fig 34a). As to the first lower premolar the lingual cusp, usually absent in recent anthropoids, may be more or less 4eveloped and in correspondence therewith the non-sectorial character

of the tooth restored (Fig. 38b). The diastemata are also aubject to a

-

General and special character of Sinantliro pus dentition

27

great variability ranging from a very wide gap to a complete absence regardless of the size of the canines.

Taking ail these facts into account, there is no escape from the assumption that the conditions existent in hominids were originally anth-

ropoid and those observed in recent anthropoids later differentiations. However, since Dryopithecus or Sivapithecus show the same character as the latter in decisive traits, they cannot be considered direct ancestors of the

hominid line but must be regarded as already specialized (Fig. 44b). Fur.. thermore, since Dryopithecus and related types were airead fully developed in Miocene times, the division of the common anthropoid stem in hominids, on one side, and anthropoids in the strict sense (including Dryopithecus),

on the other, must be dated back to a pre-Miocene period. The relation of the common anthropoid stem to the remaining primate groups is obscur-

ed for the present on account of the deficiency in adequate comparative fossil material.

111. The phylogenetic development of the hominid

brain and its connection with the transformation of the skull A study dealing with the phylogenetic evolution of the human brain

will always incite the idea whether it may not be possible to discover morphological facts which would permit to draw conclusions also on the development of the mental faculties and the human culture. The idea of a close relation between structural peculiarities, even if

manifested in bony substance, and Dsychical qualities is not in the least a recent one. Gall's phrenology which dominated the early part of the 19th

century was based exclusively on such a conception. One hundred years later Walkhof f attempted to interpret the formation of the mental protuberance of the lower jaw as a direct consequence of the faculty of speech. From the absence of such a protuberance in Neanderthal man he deduced the lack of speaking power as far as this hominid is concerned. Another

example in this respect is Pithecanthropus erectus ta whom Ernst Haeckel

gave the additional name aldus, inferring the total absence of that faculty from the smallness of his brain cap as a whole. Boule and Anthony, in studying the endocranial conditions of the skull of La Chapelle-aux-Saints, regarded the relative smallness of its frontal part as an indication of inferior psychical powers in general and specially also of a defective faculty of

speech; in addition, they concluded right-handedness from the greater prominence of the left hemisphere of the brain. Davidson Black did the same in considering Sinanthro pus on the basis of the endocast of Skull E to

be right-handed; (rom the strong development of the left orbital operculum 2B

Phylo genetic development of The hominid brain

29

the same author ¿leduced that Sinanthro pus must have already been prov-

ided "with a cerebral mechanism for the elaboration of articulate speech."

Shelishear and Elliot Smith using the same material arrived at similar conclusions based on the same reasoning. in addition, they surmised a greater ability in motion and in particular in locomotion because of the special formation of the region of the temporal pole and the middle temporal area. As regards the fôrmation of the chin it can now be taken as proven that there is not the slightest direct relation between the general and special

structure of the mental protuberance and the faculty of speech. Thus the

question arises whether and how fa it may be justified to infer other mental qualities from special features of endocranial casts.

There is no doubt that the internal wall of the brain case represents a fair counterpart of the brain surface. But the question is whether its special feature really must be considered a direct morphological expression of definite functional qualities. When ezaniining the organs of the body

with reference to the relation between shape and function, two different types may be distinguished, namely such organs the function of which is determined by their particular form and such without those relations. The first type of organs comprises the teeth, the eye, the chest, the heart, th stomach, the bones; the seàond type includes the glands, the spleen, the voluntary muscles, etc. The teeth, for instance, serve as tools for cutting and chewing; the eye is a camera obscura equipped with a real lens. On the other hand, glands merely have the task of producing certain substances; whether their form is globular or flat, straight or curved, is irrelevant to their physiological achievement. A typical example in this regard is the liver, the special form of which is entirely conditioned by the

arrangement of the adjacent organs.

30

Bulletin of the Geological Society f China

To which category does the brain belong? Morphologically it is comuosed of cell centers and nerve tracts connecting the former. Viewed from a purely stereometrical standpoint, a globular form with the cells at the periphery and the nerve fibers in the center would be best suited for

that purpose, because in such case communication could always be maintained within the "inner line." Moreover, whenever an increase in central substance and nerve tracts is required, they can be supplied in. the simplest manner by enlarging the surface, either in the form of an

enlargement of the entire globe or by a folding of the surface and a simultaneous condensation of the center. As a matter of fact, such a principle is actually realized in the human brain.

Of course, an ideal form of a globe does not exist but there is no doubt that the brains of vertebrates during their evolution strives 1:0 change from a more cordlike form to that oí a globe (Fig. 25). This general tendency is handicapped to a certain extent by the conditions of the space which depends chiefly upon the constructive necessities of the skull. Thus, the actual state must be understood as a compromise between the principle of unrestricted growth as manifested in the brain and the hardly flexible constructive realities of the broin case. Therefore, when attempting to devote a study to the phylogenetic development of the hominid brain, we must turn our attention to three

points, namely the general increase in size, the tendency to assume a globular form and the effect of the architectural requirements upon the first and second tendencies. Whether and how far we may be able to proceed beyond the limit of these points for the purpose of finding indications which

may permit to interpret such fine structures as the special configuration of the brain surface, must be considered separately.

Ph.10 genetic development cf the hominid brain

31

The general increase in size during the phylogenetic evolution can easily be proven. The Neanderthal man, despite the primitiveness of his

brain case, when compared with that of recent man, fails to furnish evidence in support of this fact. The cranial capacity of the various specimens of that group known hitherto varies between 1296 cc (Gibraltar

skull) and 1626 cc (skull of La Ghapelle-aux-Saints) with an average of about 1425 cc. Such an average capacity exceeds even that of recent man by approximately 100 cc, when taking sexual and racial differences into account. lt is, of course, probable that the value obtained would be lower if a large number of iudividuals were available for the determination of the cranial capacity. At any rate, the difference in size between the brain of Neanderthal man and, that of recent man cannot be very considerable.

The conditions in Sinanthropus are quite different. The values obtainable by direct measurements vary between 915 (Skull E) and 1220 cc (Skull L I). However, it is of certainty that 915 cc (Skull E) does not represent the lowest value. For the.fragments of the recently discovered Skull J are so much smaller when compared with the corresponding parts of Skull E that' the cranial capacity of the former could not have been in excess of 850 cc. Sinanthropus, with an average of approximately 1000 cc, therefore, falls considerably below the average of Neanderthal and recent man, though he exceeds the average of the living anthropoids by

more than twice as much. But it must be borne in mind that the maximum value for anthropoids, as represented by the gorilla with a capacity of 623 cc, is not too far removed from the minimum value for Sinanthropus with 850 cc. As to the cranial capacity of Pithecanthropus the newly found skull has a capacity of about 835 cc which exceeds the maximum value for gorilla skulls by only

10 cc.

Bul?etin of the Geological Societ, of China

32

The general increase in brain size is concomitant with a characteristic change in form. The brain of Sinanthro pus in its most primitive form is distinctly flattened like the brain case itself (Fig. 25b.). This flatness gradually becomes less pronounced, that means that the height of the brain increases. The increase is greatest within the area of the vertex and the region immediately behind - obeion region-(Fig. 25c.). Two methods can be used to demonstrate this development, a metric and a graphic one. The former consists of the calculation of the height-length index, that is to say, the entire height of the endocranial cast expressed in percentage of its greatest length. The following table gives the respective averages: According to these figures Sinanthropus ranges between chimpanzee

and gorilla, followed by

Homo

soloensis. and then by the remaining

Neanderthal specimens and finally by recent man. TABLE I

Endocranial LengthHeight Index: 2 Gorilla

61.2

5 Sinanthropus

62.82

average

(61 .3-63.7)

2 Chimpanzee

64.1

2 Homo soloensis

65.5

(63.4-67.5)

5 Neanderthal (65.4-70.6) S Recent nlan*

67.0

g'

75.6

725-807) *including all main races with skull capacities from 935 to 2050 cc and cephalic indices from 68.6 to 90.7.

Phylo genetic development of the hominid brain

33

This development can also be demonstrated by superposing the outlines of the lateral views o the endocranial casts (Fig. 20). Thereby it becomes evident that the braintaking the gorilla and chimpanzee as basis undergoes a continuous increase in' length concomitant with an equal increase in height; but with the beginning of the Neanderthal phase of evolution the vertex-obelion region becäme the most favored part. In representatives of recent man with a very small capacity and a brachycephalic skull the length of the brain may be less than that of Sinanthropus,

but the celation to the height will nevertheless correspond to that which is charactenstic 1er recent man Frontal outlines of endocasts of the region

of the greatest breadth and height (Fig. 21) show the same thing, that is the particularly pronounced increase in the vertex and obelion region. The diagrams exhibit also that the increase in breadth is rather insignificant,

playing a minor role when compared with that of the height (Fig 19). The vertical outlines demonstrate the latter fact still better. All hominids practically coincide in this respect and differ from the anthropoids merely by the size of the brain but not by a special configuration. These statements are of signifcance in more than one way. They

reveal that the general increase in size, especially in length and height, represents an essential factor and that this increase is most pronounced in the vertex and obelion region, while the ratio,het»i'eert length and breadth, regardless of the special types which in principle is the same in all groups of the higher primates varying only within modreate limites, has no phylogenetic significance. According to the endocranial lengthbreadth index as evident in Table II, Neanderthal and recent man are to be placed within the dolichocephalic, mesocephalic and brachycephalic groups, Sinanthropus within the dDlicephalic and mesocephalic groups, Pithecanthropus within the mesocephalic groups, chimpanzee and gorilla within the mesocephalic and brachycephalic groups and orang within the brachyce-

34

Bulletin of the Geological Society of China

TABLE II.

Endocranial Length-Breadth Index Dean Swift (British) Sinanthropus D Australian North Chinese Sinanihropùs I L Le Moustier

New Britain Sinanthropus II L Sinanthropus III L Sinanthropus E Pithecanthropus

NeanderthalDusseldorf Chimpanzee Rhodesia La Chapelle-aux-Saints Gorilla

Chimpanzee I Gorilla I

Orang I Gibraltar Chimpanzee II Gorilla Il Chinese Orang II European (Austrian)

70.7 73.1

73.4 74.2

dolichocephalic

747 74.8 76.1

76.3 76.3 77.2

777 77.8 78.7 78.8 79.0 79.2

80.4 80.5 82.4 83.8 84.9 86.3

mesocephalic

brachy.. cephalic

873 87.3 90.2

phalic group only. Therefore, in contrast to the length-height index, a phylogenetic lire cannot be defined here. Curiously enough, there are anthropologists who consider just this length-breadth index of skull or head

a decisive factor in classifying human races qualitatively in psychical respects.

From the conditions of the endocranial casts, however, it follows

that those interpretatipns are entirely without any scientific foundation, and

Phylogenetic development of the hominid Brain

35

not better substantiated than the speculations of the famous phrenologist Gall more than one hundred years ago.

According to a wide-spread opinion, the frontal lobe should be considered that section which has profited most by the increase of the brain substance during human evolution. Such a viewpoint is erroneous. It was derived from the conclusion that the erect position of the forehead, a striking feature ii recent man, must have been accompanied by a corresponding increase of the underlying brain part (Fig. 3c.) Yet, it was overlooked that the supraorbital ridges óf anthropoids and primitive hominids conceal the actual base of the forehead (Fig. 3 a., b.). However,

when the skull is not viewed from in front but from behind (Fig. 2), it becomes evident immediately that the increase of the brain involves its posterior parts much more than its anterior ones. With respect to the increase of other sections of the brain, there are

no distinct differences, except the temporal lobe and the frontal pole. Davidson Black, and Shellshear and Elliot Smith already pointed this fact out and stated that the temporal lobe of Sinanthropus Skull E is relatively

long and low with a strongly developed lateral eminence and that the entire lobe is separated from the cerebellum by a wide notch. All the endocranial casts of skulls and skull fragmentc recovered subsequently exhibit the same peculiarities, proving thereby that we are dealing with an

appearance typical to Sinanthropus (Figs. 25b.; 27a.). lt is difficult to determine whether the lowness of the temporal lobe must be considered a real under-development or a different configuration of the lobe due to the special width of the temporocerebellar notch. At any rate, it is remark-

able that the temporal lobe of the anthropoids is not underdeveloped in the least in relation to the remaining sections of the pallium whereas the notch is olso rather pronounced (Figs. 25a.; 29).

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Of special phylogenetic interest is the configuration of the frontol Apes (low and high) are characterized by a frontal lobe which becomes narrower gradually and ends in a well marked keel (Fig. 22a., b.). pole.

The conditions in Sinanthropus recall such a primitive feature (Fig. 22c.). In recent man the entire anteror surface of the lobe is broad and flattened, the keel-like prominence being restricted to the bases (Fig. 22g.). Neither

Pithecanthropus (Fig, 22d.) nor Homo soloensis (Fig. 22e.) display this primitive peculiarity of the frontal pole to such an extent as Sinanthropus. The interpretation of the special configuration of the surfaces of the

brain is much more complex than that of the peculiarities of its general shape. First, we must realize that in endocranial casts we are not dealing with the brain itself but with one covered by three membranes, and that therefore the convolutions as well as the fissures in most cases are not re-

cognixable at all or only faintly, indicated (Fig. 29a., b.). In addition, certain prominences and impressions which have nothing to do with cerebral structures may appear on the inner surface of the brain case and become manifest on endocasts. Finally, the variability of Convolutions both in anthropoids and recent man is known to be exceptionally great with oaly the main convolutions showing a certain constancy.

¡ therefore

doubt the possibility of obtaining a better understanding of the.deve][opment

of the brain from a comparative study of impressions and prominences of

the endocast than we already have from other indications. Only one point seems certain, namely that the pattern of the convolutions changes from a relatively simple state to a complicated one, a condition in conformity with the tendency of the brain towards enlarging its surface. For exámpie, the pattern of the frontal lobe, which is remarkably well developed on endocasts of Slnanthropus, is much simpler in Hylobates (Fig. 22a.), chimpanzee (Fig. 22 b.) and SinanthropusFig. 22 c.) and partly also in Plthecanthro pus (Fig. 22d.) than in recent man in whom the

Ph,4o genetic development of the hominid brain

37

convolutions are distinctly more meandering and more subdivided by accessory fissures (Fig 22g.). This is especially evident in the marginal

orbital gyrus which has a surprisingly chimpanzee-like appearance in Sinanthropus.

Considerable significance has at ail times been attached to the so-called 'Affenspalte", the lunate sulcus situated at the boundary between the parietal and occipital lobe (Fig. 29b.). Its persistence in recent man should serve an means for testing the degree of primitiveness. Thus, it was always scarched for on endocranial casts of fossil hominids and be-

lieved to have been actually found in all cases in the form of a shallow furrow in the region of the lambdoid suture. Shelishear and Elliot Smith, for instance, state that the sulcus coincides with this suture in the Sinanthropus endocast of Skull E. Recent investigations carried out by G. Levin have .shown that the lunate sulcus occurs rather frequently in recent

man, regardless of race. On the other hand, I was able to prove that in chimpanzee the brain exhibits a well developed sulcus (Fig. 29b.), whereas

the endocast does not reveal the slightest indication of its existence (Fig. 29a.). All the endocasts of Sinanthropus fail to show any depression within the area of the exact location of the sulcus but instead exhibit a well pronounced depression within the region of the 1anbdoid suture. Whether a specifically developed lunate sulcul really existed in fossil man cannot be concluded from endocasts.

Since the sulcus in question falls into the visual recçptive territory, its alleged strong development in fossil hominids has been brought into close relation with the visual power. This takes me back to the atternpts

mentioned at the beginning of this lecture, namely to deduce special mental faculties from the special development of certain areas of the cerebral surface. Of course, it would be marvelous if we could read off. directly from endocranial surfaces of different stages of hominids their cor-

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responding mental development. What are the real facts?

Davidson

Black deduced the existence of a "cerebral mechanism for the elaboration of

articulate speech' from a strong development of the left orbital operculum in Sinanthropus endocast E, which coincides with Broca's speech center. Shelishear an Elliot Smith accepted his version. This operculum, indeed, is rather pronounced in Sinanthropus (Fig. 30). In Skull Ill of Locus L it is preserved on both sides, but undoubtedly better developed on the right side than on the left. Apart from this fact, however, a strongly develop-

ed orbital operculum is not at all specific to honids, for it occurs in the same degree and with the, saine alternating precedente of the left or the right side in gibbon and anthropoids, particularly in the orang (Fig. 24). Shellshear and Elliot Smith laid special stress upon the formation of

three prominences on the lateral side of the Sinanthropus endocast E. One prominence is found at the posterior end of the Sylvian fissure at the

boundary between parietal and temporal lobes the second has its seat within the region of the posterior end of the second temporal convolution and the third is met in the preoccipital region of the parietal lobe. All these prominences are very distinct on both sides of the Slnanth.rop:zs endo-

casts and more or iess fuse into one single protuberance (Figs. 27a.; 31,). In some cases, they are better developed .on the left side, in others on the right.

Shelishear and Filiot Smith regard the second prominence in con-

nection with the strong development 0f the orbital operculum as an expression of acquisition oí speech, and the third prominence, in connection with the above mentioned one pushing forward toward the temporal pole. an expression of the provision for the attainment of greater skill in move-

ment and particularly in locomotion. I regret to say that I sin not able to follow those authors in their interpretation. There are two reasons, first it is evident that the prominence in question as 'well as the relative recedence of the remaining lateral parts of the hemisphere are first of all in con-

Ph,logenetic development of the hominid brain

39

sequence of the special configuration oi the brain case, a condjtjøn which will be demonstrated subsequently. Second, there is the general question

whether the protrusion of certain areas of the hemisphere really must be considered an expression of a more intensified and differentiated function.

The general tendency of cerebral evolution does not point to the formation of localized prominences in case a special area of the cortex should be developd, but rather to an enlargement of the surface by folding and by internal structural differentiations. The idea inclining to correlate special functional activities with specially developed features rather resembles Galls attempt, who regarded exterior prominences of the brain cases as real seats of psychical qualities, the only difference being that at present these seats have been transferred from the exterior of the skull to the interior side and that Gall's fantastic speculations have been replaced by a somewhat greater rationalism. Let us examine whether there are any indications in support of the assumption of intensification and specialisaflon of psychical activities pro-

duced by changes in the general shape of the brain during its phylogenetic

It - has been proven that the hemispheres increase, in all dimensions and in all sections, particularly within the region between evolution.

bregma and lambda, and that the temporal lobe also becomes higher.

On

the other hand, experiments have shown that a number of areas of the cerebral cortex have a limited but quite specific function and that no essential difference between anthropoids and recent man exist in regard to these localisations. The impulses for movements of the body and its parts

and the sensibility are located within the area around the central gyrus (Fig. 26: 4,6), that is to say, the approximate boundary between frontal and parietal lobe. The acustic center has its real seat in the middle and superior part of the temporal lobe (Fig. 26: 21, 22, 42) and the visual center in the region of the occipital pole (Fig. 26: 17, 18).

All these are-

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as, therefore, are seats of elemental components of the psychical function.

But it follows from these localisations that the middle part of the frontal lobe, the greater part of the parietal lobe, the anterior part of the occipital lobe and the lower part óf the temporal lobe are, so to say, vacant areas

(Fig 26: 5, 7, lO, 19, 20,37-39, 40, 45, 46). These areas, however. are exactly such which have acquired a special enlargement during the course of evolution.

Attempts have been made to attribute certain functional activities to these areas also According to Kleist (Fig. 26) the posterior parietal region is the seat for the.following qualities:4 "activity", writing, memIry for place, calculation, visual, imagery, recognition of numbers and colors; the inferior temporal region is the seat for accustic attention, understanding the meaning of. noises, music and names; the frontal region the seat of sequence in motor activity; thoughts of action, forming sentences, opinions, reasoned activity etc. All these additional

regions combined represent the center proper of association, in other words, they embrace such activities which must be considered to a certain

e,dent specific for human mentality.

lt is, of course, out of my field to express an opinion in how far the assumption of these. localisations is correct or incorrect I am not sufficiently familiar with the question under discussion and therefore unable to state whether the new identifications of

the regions concerned are more sound than the older ones of Gall, who credited the vertex, obelion and occipital regions of the skull, in the order from in front backwards, with the faculty of morality, religion, firmness and pride. The morphological increase of the brain substance, however, is contestable and seemingly also its relation to the areas of associátions, the highest form of mental activity.

Again, the essential principle in the evolution of the hominid brain is the general enlargement of the surface manifested by a general increase *Dr ILS, Lyman very kindly supplied me with the English translation of Kleist's terms.

Phylogenetic deeiopment of the hominid brain

4f

in size and by a concomitant augmentation of the folds which lead to a greatet complication of the pattern of convolutions and fissures. The augmentation of the folds of the cerebral surface takes place without affecting the bony case while the increase in size directly depends upon the expansion of the case. lt has been shown that this expansion does not take place to the same extent in all directions but chiefly in length and height, the vertex region and the region behind it being particularly involved (Fig. 20, 21). Moreover, the temporal lobe becomes not only higher but also undergoes a change in position with the pole turning more downward, so that the wide notch characteristic for the Sinanthropijs phase becomes finally reduced to a relatively narrow deft (Figs. 25a.. b.; 27a., c.) At the same time the angle which the axis of the medulla oblongata forms with the longitudinal axis of the hemisphere amounting to I 150 in chimpanzee

(Fig. 25a.) and to 1100 in Sinanthropes becomes more acute and approximates 800 in recent man (Fig. 25b.). All these transformations are expressions of the fact that the brain originally more flat and straight tends to assume a globular form in that it rolls up. Such a transformation, however, is exactly the characteristic change

undergone by the skull as a whole in the course of evolution leading from the ape stage to that of recent man. Fourteen years ago, previous to the discovery of Sinanthropes, I pointed out that the transformation involving the basal parts of the skull, especially manifested in the change of form and position of the pyramidal portion of the temporal bone, give the impression as it the base of this bone bas become more and more compressed between the sphenoid and the occipital bone and at the same time lifted towards the cavity of the skull. This transformation is also recognizable on the external surface of the skull by the differences in span of three sectors as represented by the outline of the entire skull, the course of the squamosal suture and the enclosure of the mandibular fossa (Fig. 15). In great apes, for instance in the gorilla (a), these three sectors are rather widely spanned

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and become shortened gradually, whereas this span is narrowest in recent

man (c.). With respect to the face, the transformation of the skull causes a shortening of the jaws and their retraction below the forehead. The occiput turns downward, so that the occipital foramen instead of being

directed backward as in anthropoids is finally directed forward in recent man (Fig. 2). The entire brain case becomes higher reaching its greatest expansion behind the vertex.

The increase in height of the temporal lobe,

the change in the direction of its axis and the reduction in width of the temporocerebellar notch appears as a direct consequence of the contraction

of the space occupied by the pyramid.

There is still another change which may also be explained by the process described above. I had called ättention to the fact that the boundary between the parietal and temporal lobe of Sinanthropus projects considerably in lateral direction, thereby producing the three prominences

of Shelishear and Elliot Smith (Fig. 31). This feature corresponds. to the conditions of the brain case of Sinanthropus in which the breadth is greatest in the biauricular diameter, decreasing continuously in upward direction (Fig. 2b.), whereas in recent man the increase in height brings about an increase in breadth also of the superior parts of the parietal bones and hence of the parietal lobes (Fig. 2c.). In consequence thereof the three circums-

cribed prominences will disappear because of the expansion of the entire parietal region of the brain. In Sinanthropusf therefore, we are not dealing so much with a particular development of special areas of the brain surface

but rather with an under development of the remaining parts corresponding to the, more primitive character of the shape of the skulL When viewing the skulls of Sinanthropus and recent man from behind, these differences in appearance of the brain case are easily recognized (Fig. 2).

The form of the brain and the form of the skull are closely related in so far as the brain case proper is concerned. Such a correlation

Phylogenetic development of the hominid brain

43

however, does not apply to the so-called super-structures of the skull, for they are conditioned by the size and robustness of the jaws and by the space required for the masticatory musculature.

The heavy and far pro-

truding supraorbital ridges which dominate the entire brow rigion in gorilla (Fig. la.), chimpanzee and Sinanthropus (Fig. I b.)have a three-

they represent the anterior end of the orbit protruding beyond the brain case proper; second, they serve for the

fold significance; first,

attachment of the foremost portion of the temporal muscle and, third, they

form a heavy buttress supporting the superior part of the face against thc masticatory pressure. These conditions are made apparent by the

topographic relation between the eye-sockets and the frontal end of the brain case (Fig. 18). The development of the ridges, therefore, to a large extent depends upon the development of the jaws and teeth in anthropoids as well as in fossil man. This close correlation was omitted by Bolk when advancing his theory of pedomorphism. Bolk's theory originated from the well-known observation that ridges are almost completely absent in juvenile anthropoids and the forehead correspondingly seems to be in a more

erect position than is the case in adult individuals (cf. Fig. 9a. and la.), thereby giving the juvenile anthropoids a more human-like aspect. He conclúded from this fact that the erect position of the frontal bone represents the genuine and primary condition of both anthropoids and man which should have been lost within the anthropoids but were retained during the entire phylogenetic development of the hominidsi.

Bolk was in

the habit of basing his phylogenetic speculations almost exclusively on observations of ontogenetic character and overlooked more or less completely evidences furnished by palaeontology. Sinanlhropus with his strongly developed supraorbital ridges and his flat and receding forehead (Fig. lb.) shows that the supposed difference between anthropoids and man never existed in reality. But Bolk was wrong also in his presupposition.

First, the frontal part of the brain, as has been proven above, does noatt

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all constitute that portion óf the brain which undergoes a specially favored

development, and, second, a comparison of the frontal lobe of an anthro-

poid child with that of recent man of the same age shows despite the greater similarity of the external appearance of the skulls (Fig. 9) the dif-

ferences of the brains to the same extent as in adults. The similarity of the conditions of the forehead of juvenile anthropoids is feigned by the lack

of super-structures (supra-orbital ridges), thereby making the frontal part of the skull appear higher and more erect, whereas the primarily different conditions of the brain itself remain unaffected. ¡ stated above that a close relation exists between the formation of

the brain and that of the brain case. This suggests to inquire which one represents the primary formation: does the development of the brain phylogenentically precede the special development of the skull or is the reverse the case? in other words, as man became more and more specialized in the development of the brain, was the brain itself the leading motive or have we to search for the final cause outside of the brain?

That size and special form of the brain case are conditioned to an essential degree by the size of the brain can be shown by a sort of natural experiment as is offered by skulls of idiots. Several years ago Mollison observed an idiot who, in early youth had been seriously injured on the head, which consequently arrested the norma! development of the brain. The result was that the bráin case assumed a Neanderthal form: the skull

remained tiot only lw but also developed, strange enough, real supraorbita! ridges, the appearance of which evidently was due to the failure of the brain to expand to completely cover the eye-sockets. That in this

respect we are not. dealing with an exceptional case is taught by the fact that similar conditions may be observed as a very common occurrence in the majority of SkUIISOE idiots.

The skull of the idiot Schilttelndreyer de-

scribed by Carl Vogt, for instance, shows the orbits to be covered by heavy

Phylogenetic dpeoe!oment of 1/ic hominid brain

45

supraorbital ridges. (Fig. 13). These protrude considerably beyond the flat and receding forehead, thereby resembling the general conditions of the great apes (Fig. la.). The same skull, when viewed from above, bears a striking resemblance to that of gorilla or chimpanzee (Fig. 1 1).

In consequence of the extreme smallness of the brain case its greatest breadth falls within the basal parts and the temporal lines approach the midline rather

closely, whereas the jaws generally remain unaltered both in form and size. Such instances prove that the size and form of the brain not only determines the size and form of the brain case but also its topographic and constructive relation to the face. The capacity of Schlttelndreyer's skull

amounts to oniy 365 cc, which is considerably below the average of a gorilla or chimpanzee skull.

The construction of the human foot does not permit any doubt that

hominids were derived from climbing primates and only subsequently acquired the faculty of standing and walking on the ground. Therefore, the question arises whether the special development of the brain is somehow connected with the erect posture.

This question can already be answered satisfactorily by palaeontoegical facts, for it crystallizes in the statement whether limb-bones of fossil hominids approach those of recent man closer than is true of the skull. An

old problem concerning Dubois' Pithecanthropus will thus become vivid again. His brain cap belongs, as proven by von Koenigswald's recent discovery of a second and more complete skull, to a genuine hominid of the same low phylogenetic stage as Sinanthro pus. The femora. however, which were recovered from the Trinil layer and and attributed by Dubois to

the same type of individual as that represented by the skull cap, fail to reveal any real essential differences when compared with the femur of recent man. This peculiarity induced Dubois to give Pilhecanthropus the surname "erectus." Should Dubois' assumption prove true, Pithecanibropus

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would then already have been equipped with a femur like that in recent man. At any rate, there is no longer any doubt that Sinanthropus had already adopted an erect posture from the evidence yielded by the recovery of six femur fragments. Despite certain characteristic differences, these femora in their decisive features closely resemble those of recent man,

and are strikingly different from those of ail the anthropoids (Figs. 45; 47; 51). Sinanthro pus' erect posture can also be deduced from the sediments of his cultural life. For it is evident that he was a hunter and familiar

with the use of fire and tools. That his hands had already been free from their original restriction in serving as organs of locomotion is furthermore

proved by the character of the recently discovered humerus. At any rate the development of the brain and the transformation of the skull of the hominids apparently lagged behind the development of the remaining parts of their body. Fourteen years ago ¡ expressed the opinion that the characteristic rolling up of the skull (Fig. 15) had a close relation to the erect posture and gait on account of its close connection with the sagittal curvatures of the space. in changing from a quadrupedal position to an upright posture, the face with the eyes and mouth turns upward, together with the lifting of the spinal column into a vertical position. in order to bring the face back into a horizontal plane, in which the locomotion occurs, necessitates an extreme flexion of the head and even a bending within the very base of the skull. Such a transformation, indeed, is fulfilled in man (Figs. 48; 52). The base of the skull in evident contrast to the conditions in monkeys and great apes. really shows a considerable "kyphosis" compensating the pronounced lordosis of the neck, both curvatures being specific to man in their special form. Such kyphosis made possible the dorsal expansion of the brain case and thereby produced the required condition for the develop-

ment of the brain in length and height; exactly that which must be considered the essential event in human evolution

Therefore, it seems

Phylogenetic development of the hominid brain

47

justified to regard the erect posture as the decisive factor determining the bominids in their specialisation of development of the brain. This tendency led them through a consecutive orthogenesis to recent man. Many authors, especially W.K. Gregory and Sir A. Keith always considered the erect posture as the preceding and essential factor for the transformation of the hominid skull. The discovery of the limb-bones of Sinanthropus proved the correctness of such a supposition.

May be some are disappointed that I am not able to demonstrate at the hand of morphological facts certain developments of the numerous

mental qualities which adorn present mankind, in contract to his more savage ancestor. But I ani afraid that this will never be realized, since fossil material is not able to supply information as to what took place within the brain substance itself. The only process that we are able to establish is the general increase of the brain and the accommodation of the

enlarged organ within the given conditions of the enclosing brain case. All other conclusions on the mentality must be deduced from cultural remains of fossil man. With regard to this we know that the hominids, even in such a piimitive morphological stage as that represented by Sinanthropus already possessed a rather well developed culture. The average of the skull capacity amounting to 1000 cc qualified them to make

a remarkable cultural achievement and to adopt a real human-like behavior. The increase in cranial capacity by an additional volume of 325 cc, characteristic for recent man, is minimal in relation to the capacity already held in the Sinanthropus stage. There is evidence that Sinanthropus - in striking contrast to the anthropoids - practiced his human qualities in not only hunting for animals but also for members of his own

species whom he killed regardless of age and sex. The acquisition of an additional 325 cc of brain substance in the case of recent man apparent'y

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was not enough to modify in principle that old human habit. Thus, we can only hope for a better application oí the now available brain substance by taking advantage of the possibility oí internal differentiation.

IV.

Did Sinanthropus pekinensis practice cannibalism?

The excavations at Choukoutien which now cover a period of more than ten years have furnished us with many important facts regarding the physical appearance of Sinonthropus pekinensis and his manner of existence. Still there is one mystery to be unraveled, namely: how did the human remains come into the cave and w+iy are substantial skeletal parts missing? Apart from skulls or skull fragments very few other bones have been unearthed hitherto. Of limbs we have some unsignificant fragments of the upper extremity: a collar bone, a small fragment of the lower end of an arm-bone, one complete carpal bone; and only rather recently

fragments of five thigh bones and a humerus were recovered. A small fragment of the first cervical vertebra is the only representative of the spinal column.

Thus, the question arises what has become of the other skeletal parts? There are two alternatives, namely that they were dissolved by natural processes in the course of time or that we (ailed to recognize them when excavating. Generally, both these eventualities may be realized but only in such cases in which the discoveries were restriced to remains of a few individuals. In the case of Sinanthropus, however, the list of material on hand gives an account of the presence of at least 40 individuals, male and female, including 15 children. These individuals were found to be distributed over the entire area of the Choukoutien cave which, according

to accurate plans of the site of excavations, embraces approximately 8000 cubic meters with a height of more than 50 meters and is composed of 15 different sites containing human bones. Many of these individuals are

represented merely by a few isolated teeth. 49

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The complete destruction of all the missing parts of 40 skeletons by dissolution of their organic and anorganic bony substances is rather impro-

bable, all the more so in Choukoutien, where the general conditions for preservation are excellent that, for instance, several skulls of macaques have been recovered which are perfectly intact, including such thin or fragile bones as the zygomatic arch or the inner walls of the eye-sockets. The second alternative that extremity bones may have been overlooked can not be excluded with the same degree of certainty. Previous to my coming to Peiping, I was inclined to accept such a possibility. The methods of excavation at Choukç)utien, however, convinced me immediately that there was no chance whatever of even tiny bones escaping detection. The excavations are carried out systematically with the utmost care. The entire area is divided into regular squares of 4 square meters each and the matrix of each square is gradually removed by a group of spe-

daily trained technicians. When working in sandy layers the earth is scraped with the aid of iron hooks, whereas when working in stony layers the rocks are crushed into small pieces and brought to the laboratory for further preparation. Hence, it would be absolutely impossible to overlook the presence of limb-bones or vertebrae if really present. Rather interesting in this respect is a discovery made by Dr. Pei in the spring of 1 93f. in going through all our material for the purpose of indentifying overlooked fragments of limb-bones, he found among other bones the upper part of a femur with head and trochanters broken off (Fig. 47). This specimen, according to its labelling, had been found in 1929 by B. Bohlin and sub-

sequently stored away because it was considered to be the fragment of an antler. In this case the bone had not been overlooked but rather incorrectly classified.

Thus, all facts lead to the conclusion that only or mainly skulls or skull fragments had been deposited in the cave. There are three even-

Did Sinanthropus pekinensis practice cannibalism?

51

tualities by which these could have become lodged there: the skulls may have been carried by water, by beasts of prey or by man himself. The

first and second alternatives would not be capable of explaining the particular selection of the human remains, for neither water nor animals could make such a choice. Apart from this, there is no indication of water flowing through the cave at any time nor do the finding conditions and the

distribution of the human material suggest that they had once been transported there by the action of water. In case the skulls had been left there by animals, then they would show markings of gnawing which are entirely On the other hand, if all the other bones were consumed by missing. carnivores, why should they save the skulls or fragments thereof? Therefore the only possibility left is that man himself deposited the bones in the cave. Such an assumption presupposes that man was present in the cave. This fact cannot be doubted. For it is manifest by three evidences. The first is the distribution of implements over the entire area within the cave. These implements are composed of stones (flint, quartz, quartzite and other materials) which reveal clear indications of having been worled or used for special purposes. The second evidence being supplied by traces of fire found in numerous places. These traces consist of burnt and split

stones and of burnt and split fragments of animal bones (Fig.

50)

and finally of coal and ashes. The third evidence is that all the material within -the cave is mixed with animal bone fragreients among which deer bones constitute the majority. Most of these fragments are in the form of splinters of long bones as is quite commonly observed in human dwelling places of later palaeolithic time and generally considered an indication that man worked the bones for the purpose of removing the marrow (Fig. 49).

A great number of these animal remains are, as is true of the human material, also restricted to skulls or skull fragments. The same is true of

the macaq.ie material mentioned above of which only skulls and no indication of limb-bones or spine-bones have been found.

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We know from experience of other dwelling places of man in later times that he also used such localities for burial. Such an explanation

could not be applied to Sinanthropus, since a great number of the individuals are only represented by several teeth or fragments of lower jaws or

small fragments of the brain case. Even in those cases in which entire skulls have been found, they cannot be considered to be complete. One may object and consider such a deficiency as being in consequence of the collapse of the cave roof and the falling in of boulders and clods from the outside which may have crushed the originally complete skulls. This, of course, is true. In the case of Skulls 1 and Il of Locus L not only the crushed bones of the brain case were found in situ, but also numerous splintered parts of the facial skeleton which bear markings of secondary destruction. On the other hand, there are cases which lack the slightest indication of such an event having taken place. Hence, it cannot be doubted that the fragmentariness is not a result of a catastrophe which occurred within the cave but rather presents the original state in which the

specimens had been deposited. As further evidence thereof may be quoted not only the frequently found fragments of lower jaws without any indication of the remaining parts or of the skulls belonging thereto, but also the fragments of brain cases which, for instance, may consist of an. isolated temporal bone or a piece of a parietal bone or only of the squarna of an

infantile occipital bone. In the case of the temporal bone designated as Skull H Ill the entire specimen was covered by a thick layer of cristalline matrix, especially thick within the line of breakage. In Locus J, for instance, an isolated but well preserved fragment of a right parietal bone

of an adult, probably male, individual was found composing the mastoid

angle of this bone with the corresponding sutures and not larger than approximately 750 square millimeters. No other human bone which might have belonged to this individual was recovered anywhere near or within

this special site.

The restriction to such isolated and occasionally very

Did Sinanthropus pekinensis practice cannibalism?

53

small fragmeiits definitely excludes the possibility of considering the human material as remnants of a burial site.

The conditions and circumstances under which they came to light are exactly the same as those of the animal bones with which they are mixed over the entire cave region. in the winter of 1938, Dr. W.C. Pci, while sorting specimens lately t.ollected at Choukoutien and subsequently prepared in the laboratory, came across some fragments of thigh-bones which he diagnosed as probably oí human origin. A thorough examination of these fragments proved this diagnosis to be correct. Of these specimens one comprises the entire shaft with the epiphyses missing (Fig. 45). The surface is considerably weathered. The second specimen has already been mentioned above as having first been considered a part of an antler (Fig. 47). Al! the other specimens are fragments of the shaft. Like

in the case of the two first mentioned, the parts missing were apparently broken off before the bones became fossilized and embedded within their present matrix. One of the fragments, the middle part of the shaft (Fig. 51), is considerably blackened on one side, with the dark surface extending in part also over to the other side and in addition to the cross-breakage. Chemical analysis* revealed that the blackening is due to carbonisation of the bone. Another fragment, considerably longer, consists of only the posterior wall of the middle parts of the shaft, with the marrow cavity exposed in its entire length (Fig. 46). One more fragment of a shaft of about the same length shows the same character (Fig. 53). One other fragment is represented by a piece of the upper part of the shaft. lt is not split hut rather weathered, especially along the breakage line. All these specimens show in common that they do not differ in the least from the fragments of long bones of animals found mixed with them *Dr. S.D. Wilson of thc Dipartmeut of Chemistry, Yenching University, very kindly made the analysis for us.

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Bulletin of the Geological Society of China

and under the same conditions (Fig. 49).

This is to say, that they have

been broken or split [or the purpose of extracting the marrow, in part also exposed to fire (Fig. 51). In other words, the human femora were subjected to the same treatment as the bones of animals hunted by man. But it is of special interest that a fragment of a shaft of still a different femur

was found crushed and broken into two parts either by man or by a carnivore bite (Fig. 54). Yet there are still other rather important facts whîcri may serve as

an indication of what had befallen man, woman and children of the Choukoutien population.

The majority of the brain cases at our (hSpOSal

reveal distinct markings of injuries.

Skull E, described by Davidson

Black, shows 1oss of bony substance in various places on the outside of the

two parietal bones, the frontal and the occipital bone which may be considered as having been effected by blows. More definite are the markings on the three skulls of Locus L. in Skull I Locus L there is a deep cut on the left side of the frontal and parietal bone, which runs parallel to the sagittal suture and measures 6 cm within a distance of about 1 cm from the suture (Fig. 7). The surface of this depression is rather smooth but the bone is cracked along the depression and three times across it. Another more round depression with a diameter of about 1.5 cm may be seen just at the posterior end of the first but on the right side immediately bordering the suture and occupying the vertex. This depression is accompanied by a splintering of the bone on the inner side of the brain case. A broad and long crack begins at the depression and runs forward and laterally toward the coronal suture. Skull II shows a shailow and unevenly round depression with a diameter of circa 1.5 cm within the middle of the right parietal

bone and three cracks diverging ¡rom this center. A similar but more sharply bordered depression may be seen on the left parietal bone but the

bone is so broken at this point and partly also lost that it is impossible to

Did Sinarzt!iropus pekinensis practice cannibalism?

55

judge the original extensiveness. A third loss of substance is recognizable on the left side of the bregma, also with three cracks diverging from this

depressed center.

Skull Ill shows a deep cut of approximately 3 cm

length on the right parietal bone leading forward obliquely from the sagittal

suture (Fig. 8). A slight loss of substance may be seen on the same bone within the obeion region. In addition this skull exhibits a distinct scar on the left side of the frontal bone just above the supraorbital ridge. The parietal bone of the skull of Locus J displays a characteristically depressed fracture of 1 cm in diameter and three cracks diverging from the fracture center. Skull D fails to show distinct markings of injury but the entire right side of the frontal bone is missing and the breakage lines are straight and therefore can only be the effects of blows by implements..

The isolated temporal bone of Skull H Ill also exhibits such straight breakage lines and a distinct cut which has its beginning at one of these lines.

In addition, the outer surface of this fragment is covered by several scratches within the anterior region for the attachment of the temple muscle.

There is not the 8hghtest doubt that these depressions were caused by blows on the bone still covered by its soft parts. Some of the depres-

sions have the characteristic appearance of so-called depressed and comminuted fractures which, in the practice of forensic medicine, are considered typical results of heavy blows with more or less pointed implements. The long cuts seem to be the effect of sharply edged tools and the larger and more shallow lesions the results of blows with rounded stones or clubs

lt can be excluded with an appreciable degree of certainty tharthe described depression, cuts or cracks were caused by crushing due to the collapse of the cave roof or by a similar event. For those injuries which may also be observed in the skulls of Sinanfhropus offer an entirely differ-

ent aspect in that the bones in these are neither depressed nor do the breakage lines show regular contours.

Bulletin of the Geological Society of China

56

Thus, the conditions of the Sinanthropus skulls and skull fragments prove that their bearers suffered violent attacks. Such a statement alone,

however, does not explain the presence of isolated skulls within the cave. Thus, we are forced to search for further indications which may throw some light on this mystery. The discoveries of isolated fragments of brain cases without the slightest trace of other parts of the skull or the distribution

of several small pieces of the same skull over a large area, as was true for the skull of Locus J, prove that the skulls had been cut into pieces by man. Moreover, there is further evidence which points to deliberate manipulations of the human bones. For in all five skulls in which the brain case is more or less complete a considerable portion of the base is missing and in particular the anterior part of the occipital foramen and adjacent parts. The

fact that the missing parts are restricted in all instances to this special reg on excludes the possibility that the loss may be an accidental one. The only explanation is that the brain cases were opened from the side of the base, probably for the purpose of reaching their contents, namely the brains. The assumption of deliberate manipulations of the skulls after death

is strongly supported by the state of preservation of the lower jaws and the f emora. Of all the 14 lower jaws recovered hitherto, not a single one is complete. All are broken and represented by more or iess small pieces. As in the case of the fragments of brain cases, these pieces represent the very only remains of jaws which therefore must have been broken before they became embedded within the matrix of the cave. But of particular interest is the fact that the majority of these jaws are broken either exactly

in the midline or immediately next to it. The middle part is preserved only in two instances. A comparison with the respective conditions of the lower jaws within the Neanderthal group of fossil man reveals great differences.

Of the 25 available jaws of this group, only three are broken

I)id Sinanthropus peinensis practice cannibalism?

57

off in the midline and four a short distance from it, whereas in all the other cases the jaws are more or less complete. The percentage of the occu

rerice of complete jaws, therefore, is 48% within the Neanderthal group, whereas it is nil in Sinanthropus. The percentage of the occurrence of broken jaws within or near the middle amounts to 28% in the Neanderthal group and % in Slnanihropus. The breaking within the region of the midline cannot have been caused accidentally due to natural events such as the collapse of the roof of the cave or something similar. For in such cases most of the jaws must either have been hit from one side only or were just lying on one of the sides, a rather remote possibility. The only possible explanation is that the jaws were deliberately broken by man in that one side was fixed and the other pressed against it. I experimented with jaws of recent man and found that in such a case they actually break either within the midline or clase to it. It is of importance to rote that all the above mentioned 28% of the Neanderthal jaws broken in similar manner belong to the Krapina or Ehringsdorf material. But of these sites we have other evidences proving that the bones had been cut into pieces by man. I will return to this question below. As regards the femora their condition also proves that they were broken or split for the extraction of the marrow (Figs. 46, 53).

Taking all these facts into consideration there remains no other choice but to assume that the Slnanthropus population of Choukoutien had

been slain and that subsequently theii heads were severed from the trunk, the brain removed and the limbs dissected. Whether all these manipulations were carried out without oi within the cave is difficult to determine. I am inclined to believe that the work was done outside and the remains thrown into the cave which may have served as a place where the refuse was discarded. But even such an explanation does not de-

finitely answer the question concerning the whereabouts of the remaining

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Bulletin of the Geological Society of China

skeletal parts.

lt seems that the skulls and the limb-bones, especially the

femora, constituted parts of choice and were therefore treated in that special manner. Breuil (Anthropos, Vol. 32, 1932) in his attempt to explain the absence of skeletal parts other than skulls proceeds from ethnological facts of today.

The Australian aboriginal and other primitive people have the custom of exposing corpses on open surfaces or trees for decomposition following which the macerated skulls or lower jaws are carried about as objects of worship. Brown (cited by G. Schmidt, Anthropos, Vol. 32, 1932) describes similar habits of Andaman Islanders.

According

to him, these people open the graves or take the bodies down from trees and bring them back to camp. The skulls and jaw bones are preserved a long time and worn around the neck, either in display in front or behind.

The other parts are also preserved. But these are not considered as precious as the skull and jaw. Although it is certain that every camp houses a number of skulls and jaws, yet it is comparatively infrequent that limbs are found. Howitt reported on the Kurnai, a tribe of South-East

Australia, stating that parents carried the lower jaw of their deceased children as a memento.

It follows from these facts that the Australian and the Anciaman treat the skulls and the jaws with feelings of reverence. But in the case of Sinanthropus the conditions of the human bones certainly do not correspond to such which may have been dictated by considerations of piety. The skulls show marks of heavy blows delivered upon the head while still covered by the scalp, the jaws and femora were forcibly broken into small pieces and the fragments thereof subsequently distributed over the entire area of the cave. This treatment is exactly the saine as that given to the killed game. Breuil made the objection that the absence of cervicai

vertebrae contraditts the idea that heads may have been severed from he trunks. Since then, however, a fragment of a first cervical vertebra was discovered but no other traces of spine bones.

Did Sinathro pus pekinensis practice cannibalism?

59

For these reasons I arrived at the conclusion that the skulls and jaws belong to victims deliberately killed. The deliberate ope ning of the brain case indicates that the brain had been rentoved. Since Sinanthropus already knew the practice oi breaking the long bones so as to obtain the marrow, it seems rather probable that the brain also served him as food. Thus the question arises as to who the killer was.

Boule (L'Anthropologie, Vol. 47, 1937) holds the opinion that the entire physical construction of Sinanthro pus, especially the smallness of his cranial capacity, makes him appear too primitive to be credited with such a high cultural accomplishment like the use of fire or implements or the art of hunting. Boule, therefore, believes that another much more advanced hominid type of fossil man lived contemporaneously with Sinanthropus,

who should be made responsible for the killing of Sinanthropus.

The objection that no bones have been found hitherto justifying the assumption of there having existed an additional human type in Choukoutien is refuted

by Boule on the basis d the fact that we know of numerous undoubted dwelling places of fossil man with no indications of the presence of skeleta]

material. Despite the correctness of this argument, Boule's real starting point cannot withstand a test. The average skull capacity of Sinanthropus

amounts to 1000 cc. That is twice as much as the average for anthropoids of today and only one quarter less than the average for recent mankind. On the other hand, although Skull I of Locus L shows all the characteristics

of the Sinanthropus type, its cranial capacity is 1220 cc.

This volume is

200 cc more than the minimum values obtained for tecent man and approaches rather closely the minimum values estimated for the skulls of the Neanderthal group (Gibraltar 1290 cc). Therefore, the volume of the brain, if it really should be a reliable criterion for the estimation of cultural achievement may be considered perfectly adequate for the cultural accomplishments of Sinanthropus.

Apart from this, Boule's assumption of the

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Bulletin of the GeoLogical Society of China

existence of a second hominid is in certain contradiction to the finding conditins. As mentioned earlier, the excavations now extend to a height of more than 50 m with 15 different sites where human material has been found.

lt is difficult to estimate the time required for the filling of such a large cave. But it is rather improbable that skeletal parts of only one human type should be found if during all this time the alleged two different types of hominids are supposed to have lived side-by-side, one the killer

and proper owner of the cave and the other his prey. Boule suggested that the skeletal parts then known, that is to say, the collar bone, the carpal bone, the atlas and the humerus fragment may have belonged to the killer. This is most unlikely because these specimens apparently belonged, according to their size and robustness, to female and juvenile individuals respectively. in addition, they were found under the same conditions and in the same state as the bones of the alleged victim3.

The same is true of the recently found limb-bones. These also would have to be attributed, in consequence of Boule's idea, to the morphologically advanced

killer whose physical appearance could not have been too different from that of the Neanderthal man. in reality, however, these femora show rather distinct differences and reveal another type well compatible with that represented by the Sinanthropus skulls. I fully realize that it ma prove rather distressing to some sensible people to hear that the most primitive ancestor of recent mankind bad been responsible for such terrible acts as manslaughter of woman and children and cannibalism. But if we should turn to man's later behavior *hich has been recorded of other fossil and prehistoric man, the result would not be any more pleasing. Ten years ago, I was already able to show that the fossil man of Weimar.Ehringsdorf, a representative of the Neanderthal group of the last interglacial period, must have indulged in similar customs.

The skull was broken like those of Sinanthropus, the frontal bone shows

Did .Snanthropus peIinensis practice cannibalism? very characteristic markings of heavy blows (Figs. lO, 12) and the base is missing. The juvenile mandible is broken near the midline and only one

half is preserved as already mentioned above. The remains of the Krapina population, who lived during the same period as the man of eimarEhringsdorf, were so completely broken that it was even impossible

to reconstruct one entire brain case from the numerous bone fragments belonging to some 20 individuals. The recently discovered skull of

Steinheim, who may even be older than the Weimar-Ehringsdorf and Krapina man, also shows markings of manipulations similar to those shown

in the Sinanthropus skulls in that here too the entire surrounding of the occipital foramen is broken off.

The remains of Homo sapiens of later palaeolithic period furnish plain evidences that the individuals had been victims of attacks by man. The most striking example in this regard known hitherto is the case of the "Upper Cave" man of Choukoutien. Here the bones of at least six individuals induding women and children have been unearthed. Three of

the four skulls show tremendous marks of blows proving that these individuals must have been slain not only with pointed weapons but also with heavy clubs (cf. Weidenreich, Chinese Med. J., Vol. 55, 1939). The character of the holes and fractures does not leave room for doubt that the deathly blows had been received during life. In this instance, we have no evidence of cannibalism, the base of the skulls being intact. From the Aeneolithic deposit nf Sha Kuo T'un in North China J.G. Andersson and Davidson Black (Pal. Sin., Ser. D, Vol. I, Fasc. 3, 1925)

recovered the remains of 45 individuals of all ages and both sexes but as Andersson says: "An exceedingly small proportion of the long bones and skulls whatever are preserved in an unbroken condition...." "Many bones were already fractured when they were fresh and probably still had the flesh on," Andersson gathered from this fact that the cave of Sha

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Bulletin of the Geological Society of China

Kuo T'un had either been a dwelling place of people with cannibal habits or a native site, where religious rituals, including the sacrifice of human lives were performed. It is rather strange that evidently late paleolithic people of Central Europe who lived at approximately the same tizne as the population of the Upper Cave of Choukoutien in North China should have suffered the same

in the so-called Of net cave in Bavaria the severed skulls of 33 individuals, mainly children and women, were found buried. Some of fate.

these skulls show large oval holes somewhat pointed at the ends; Mollison infers from this fact that these individuals were killed by heavy blows with

stone axes.

In a very interesting study Wernert (L'Anthropologie, vol. 46, 1936) called attention to the fact that anthropophagy cannot have been performed because of lack of food for there is evidence of the presence of an extraordinary abundance of game in all sites. According to that author

in prehistoric times as well as in historic times up to the present there had always existed an entanglement between the two rites of anthropophagy and head-hunting. Thus, it should have the character of a special rite originating from the idea of increasing the material and spiritual qualities of

an individual or the community taking possession of the corpse of the vanquished. The conditions of all the human bones recovered from the SinonIhropus cave leave no doubt that the individuals concerned were killed and

dismembered. The special treatment undergone by the skulls and the f emora furthermore reveal the removal of their contents, According to similar conditions of later times, we are justified in assuming it was done for the special purpose of consuming the brain and marrow.

Since, before

the discovery of the limb-bones, our material was restricted to skulls only, I

Dtd Sinanthropus peIinensis practice cannIbalism?

63

had concluded that Sincinthropus was a head-hunter not interested in other

parts of the body of his victims. The new finds, however, necessitate a correction in my former assumption but only to a slight extent. For the fact that the upper levels which supplied us with the remains of about 35 individuals failed to shelter other skeletal parts than skulls - with but two exceptionscannot be altered by the recent experience since it refers to the lower levels of the cave. Until the excavations axe completed we shall not be able to draw any more definite conclusion. But at any rate, the evidence of canibalism cannot be eliminated by interpretations.

There is no doubt that the tribes of early man and all their successors have fought and tried to destroy each other. This knowledge is important not only from the standpoint of the history of ethics and religion

but also from that of physical anthropology. For it reveals that from the earliest times there must have existed strong aversions among the various members of the human family. I was able to show that racial differenti

ations began to be manifested already within the Sinanthropus stage, that is much earlier than was previously believed to be the case. The old and recent finds of Pithecanthropus point towards the same direction. The persistence of special morphological characters could be maintained and intensified only by strict isolation in space and time. Mutual aversion may have been a good way to foster such a tendency. The ancient methods of preserving real or supposed specific characters by hunting the competitor and devouring him have disappeared in the meantime. Let us hope that the modern methods applied by the physically moi'e advanced mankind of today in persecuting those people who differ in their physical or mental appearance will also become things of the past in a not too distant future.

V. The classification of fossil horninids and their relations to each other, with special reference to Sinanthropus pekinensis When attempting to classify fossil horninids and to determine the relationship between the individual types, there seems to be no other choice but to define first the morphological characters of the latter and to compare their resemblances and their differences. Neither archaeology nor geology can be of aid, still less act decisively on this question. Comparative anatomy alone is capable of furnishing us with the information required for the recognition of primitive and advanced phases of evolution and for ranging them within phylogenetic lines. Such a procedure merely presupposes that recent man evolved from primitive types in the course of time and that available fossil material indicates the general course oí such a transforma-

tion. We do not believe of venturing too far when stating that such a presupposition has been proved as correct and acknowledged by all present students of natural history, though the opinions on the special condition

and the special course taken by that development may diverge. Such hominid types as resemble the supposed original form most dosely must be considered also the most primitive. The sequence within the line is there-

fore defined by the extent of approach to this form and to that of recent man, respectively. Here we are nainly referring to hominid remains as are represented by skulls. For single mandibles or teeth do not yield a sufficiently reliable basis for the assertion of their systematic position because experience has shown that the variability even in a given type is considerable, and primitive or more advanced features may occasionally occur as individual 64

The classzficaüon of fossil horninids and their relations

65

variations in one and the same group. With this reservation, the most primitive and best known hominid is Sinanthropus pelinnsis. His primitiveness is proved by his dentition which reveals a genuine anthropoid character. For in this respect Sinanthropus approaches the anthropoids

even closer than he does recent man, whereas the differences in the features oi the teeth between Sinanthropus ar.d fossil and recent anthropoids are not greater than those between the single genera of the latter (cf. Lecture Il).

The primitiveness of Sinanthropus can furthermore be deduced from the size and the shape of the brain case. The auricular height in typical types approximates just one half of the length and therewith attains that relation which is characteristic for gorilla and chimpanzee. This ratio in the Neanderthal man is considerably higher. in accordance with the lowness of the skull is the small cranial capacity, being one third less than the average cranial capacity of recent man (cf. Lecture Ill).

As regards special features of brain case and face in Sinanthropus, there are a number which exhibit particular and more primitive characters when compared with the various Neanderthal types. This is especially true for the supraorbital ridges and their relation to the forehead and nasal bridge, for the totus occipitalis and the swellings the parietal and temporal bones connected therewith, foc the position of the foramen magnum, for the configuration of the petrosal bone, for the peculiarities of the interior

wall of the eye-socket, entrance of the aperture piriformis, the anterior surface of the maxila, the course of the zygomatic, arch and its connection with the latter, etc (cf. Lecture I).

With respect to the endocast, the size and special appearance of the temporal lobe, the form and the pattern of the frontal pole and the type of ramification of the middle meningeal artery represent in addition to

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the total flatness and smallness of the brain characteristic differences from the corresponding features within the Neanderthal group The problem as to the real nature of Pithecanthropus, the subject of lengthy discussions. may now be considered as solved. The opinion first

expressed following the recovery of the skull that Pithecanthropus repres-

ents a giant gibon turns out to be erroneous.

Already at the time of the

recovery of the first Sinanthropus skull Davidson Black was able to demon-

strate the close relationship between Sinanthropus and Pilhecanthropus. The discovers' of additional Sinanthropus skulls two of which show a very low ratic between height and length rendered Black's conclusion incon test-

able. The latest discovery of an almost complete brain case of PithecanShropus, the very image of the first as recorded by Dr. von Koenigswald, proves definitely that Pithecanthropus is a genuine hominid of about the same general stage of evolution as Sinanthropus. Such differences as exist, we believe, are merely of racial or regional nature and even if Pltecanthro-

pus should be considered more primitive in one or the other feature, it appears more or less irrelevant as to the mutual position occupied by the two types within the hominid system as a whole. As to the famous femur of Pilhecanthropus the question whether it actually belongs to the same type as the skull cap remains open, despite the four additional specimens found subsequently and attributed by Dubois to the same category. The recently recovered f emoia of Sinanthropus differ considerably from the former and show much more primitive characteristics. This certainly is in striking contrast to the appearence of the skulls.

The newly discovered mandible fragment of Plihecanthropus will in no way alter the statement made above, even if it should belong to the same type as the skulls. The teeth within this mandible exhibit that the molars are primitive in respect to size and proportions, whereas in respect to special features they seem to be more advanced than the molars characteristic for Sinanthropus.

The classification of fossil !iominids and their relations

67

The hominid group commonly called the t'4eanderthal group is everything but uniform when viewed from a comparative morphological standpoint. First of all, Rhodesian Man and Horno soloensis must he scprated from the European types, both of them exhibiting certain remarkable properties abse'it in the European forms. Some of these peculiarities are not only specific to the former and have therefore to be considered as racial

characters but at the same time prove their greater primitiveness. This primitiveness especially involves the entire formation of face, occipital region and temporal lobe in the Rhodesian skull and the lown of the brain case and configuration of the ear region in the Java skulls.

The European Neanderthal stock must again be divided into at least two distinct groups which may tentatively be designated as Ehringsdorf and Spy group. The former is represented by the Ehringsdorf and Steinheirn skulls and probably also by the Krapina man. The principal members of the Spy group include the skulls of Spy, Gibraltar, La Chapel-

le-aux-Saints and Ferrassie. Such a classification, however, does not imply that the various members of each group are absolutely uniform, On the other hand, there is no doubt that the Ehingsdorf group as a whole, when compared with the Spy group and viewed from the purely morphological point of view, represents a more advanced group than the latter. This fact is proved first of all by the much greater J eight of the brain case and the higher bregma angle. The position occupied by the various Palestine skulls cannot be defined with any degree of accuracy until more detailed information is vailable. For the present it seems to us that, notwithstanding certain dif. ferences, the Tabun type with his great cranial capacity (1263 cc) and his high length-height index of 57.3 - according to the results obtained by Sir Arthus Keith and McCown approaches the Ehringsdorf group rather tba the Spy group.

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With respect to fossil hominid remains recovekd from English soil, it seems to be certain that, regardless of their various interpretations, the skull fragments of Putdown - the mandible must be oriitted - represent a recent human type and the same is true of the recently discovered skull fragments of Swanscombe. The morphological dassification of the skulls themselves, therefore, is not too difficult. Kanjera and Kanam of East Africa also have all the essential characteristics of recent man.

When we attempt to range the fossil hominids, according to the degree of their primitiveness, we arrive at the following: TABLE

I.

Morphological sequence of hominid remains General cias-

Sub division

siftct ion Prehomtmds

-

Type

Sinanthropas peklnensls North China pj tjiecanthropus erect us Java Homo soloensis

(Rhodesian group)

Homo primigenlus

Distribution

(Spy croup)

Homo rhodesiensis

N. South Africa

Spy Gibraltar LaChapelie-aux-Saints La Fertassie Le Moustier Mauer mandible

Western Europe

Ehringsdorf

lu

Krapina

-

Putdown (brain case)

(Ehnngsdorf)

Java

(Homo modjokrtensis)

Saccopastore Steinheim Tabún (Mt. Carmel)

Southwestern Eu ... Western Europe Central Europe { Central Europe South Europe Central Europe Palestine

-

Homo sapiens fossills

London skull Swanscombe

England

Skhúl-type(Mt.Carrncl)

Palestine

Kanjera Kanam

} East Africa

The classification of fossil hominids and theIr relations

69

Even if the special morphological position of one or the other type listed herein should not be quite correct, the sequence as a whole is.

This gives rise to the question whether a real genetic or genealogical relationship exists between the main groups in such a way that the respective primitive types may be considered to be the immediate ancestors of the following more advanced types. The zeal to seek for "specialisations" and to proclaim such "specialised" types as "discontinued" sidebranches of human evolution constituted the greatest obstacle in correctly understanding those relations. The literature is full of statements such as this or that fossil hominid cannot be a direct ancestor of any of the following types up to those of recent mankind on account of the special character of certain features There is not a single peculiarity which has not been

taken by some author to represent a unilateral specialisation, beginning with the strong developmeflt of the supraorbital ridges and ending with the conditions of the molar roots. Yet, evidence proving the correctness of such statements is lacking in all the cases. The fact that the greatest care must

be excercised in treating the entire matter is confirmed by the frequently cited "taurodontism." Numerous investigators consider the so-called taurodontism a typical specialisation of Homo primigenius and therefore condemn

the latter to an extinct branch. The only reason for .loing so is the assertion that taurodontism should neither occur in anthropoids nor in recent man. Not one of these presuppositions, however, stand firm against dose

scrutiny. Taurodontisni is characteristic for all fossil hominids recovered hitherto. It occasionally occurs not only in recent man (Eskimo, Bushman, Amerindians and in Palaeoiithic man of the Upper Cave of Choukoutien) but is also found in the chimpanzee and especially in the female orang. Taurodontisin disappears in recent man on account of the reduction in size, robustness and form of the molars. As a matter of fact there is not the slightest justification to exclude a whole group of fossil hominids or certain types thereof from the paternity

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of recent man, at least as far as such an exclusion is made on the basis of morphological facts. As regards Sinanthropus, we believe to b able to prove that he is partly related to a certain group of present Mongols. In support of such an assumption the following facts may be cited: 1) the existence of a torus mandibularis; 2) the occurence of shovel-shaped central and lateral upper incisors; 3) the occurrence of an os epactale (Inca bone): 4) the formation of a very pronounced sagittal crest. All four peculiarities

may be found to exist in 70% to 100% of the Sinanfhropus material available at present, Possibly some of the features of the face point toward the same direction (cf, Lecture 1). At any rate, ail facts known hitherto demonstrate that Stnanthropus must be ranged within the direct line 0f human evolution, a certainty alrea-

dy realized by M. Boule. With the same justification it applies to all other fossil hominids. in the case of Plihecanthropus, for example, the newly discovered skull makes it almost a certainty that Homo soloensjs viewed from the morphological standpoint, is not much more than an enlarged Pithecanthropus. If such close connections between the various fossil hominid types

are evident, then the question arises whether or not th geological conditions support the assumption of direct relations, in other words, does the chronology of the types correspond to the morphology? However, when attempting to adjust the morphological list in Table I according to the sequence of the geological periods, we will meet with great difficulties, in the first place due to the fact that the finding conditions of many of the types are not absolutely secured against the objection of their not really belonging to that geological period to which they were attributed. Such a reser-

vation applies in particular to the types of recent man recovered in England and East Africa. i we disregard the doubtfulness as to the correctness of those geological determinations, we arrive at the following definition:

The classification of fossil hombiids and Their rdaiions

71

TABLE IL

Chrono1oicaI sequence of hominid remains Gacial periods Günz

General and special classification Homo

Homo primigenias II: GünzMindel

Homo primigenius J:

Prehominids: Mmdcl

Tyj

Distribution

I Piltdown 1 Kanam

Western Europe East africa

Mauer

Central Europe

Java

H. modjokerlensls ISinanthropw pekinensis Pithecanthropus erecius

North China

Java

?

Homo sapiens: Mindel -Riss Homo prlmlgenius ¡JI:

Interglacial

Swanscombe

Ehringsdorf group (Steinheirn)

Western Europe Central Europe

Riss

Riss-Wür

Skhül-type Homo prlmlgenius JI!: J Ehringsdorf group t Tablin-type Homo soloensls Homo primigenias I:

W(irin

Homo sapiens:

Homo sapiens:

Homo primigenias II:

Post-Wünn Homo sapiens:

Grimaldi

Palestine Central and Southern Europe Java PalestiEé

Western Europe

Spy gip

Central and

H. sapiens fossills

Europe, Africa. Asia

Western Europe

According to this list Homo sapiens, that is to say, modern man should have existed already in the Lower Pleistocene period of Europe, East Africa acd Asia Minor throughout everyone of the glacial epochs. I do not intend to discuss here all the difficulties which would arise if this supposition were accepted. But there is one point which, to me, seems sufficiently important to be given special emphasis.

Should Homo sapiens

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really be so ancient a type as to have existed already at the beginning of the Ice Age, how are we to exlain his failure in experiencing any changes in his morphological appearance in ail this time, while contemporary prehominids or primigenius types underwent a continuous and characteriatic transformation?

At any rate, some cases at least are sufficiently substantiated to permit the conclusion that morphological and chronological sequences of hominids fail to conform. This fact requires an explanation. The assumption that advanced and primitive types had existed side-by side is insufficient with respect to the extensive duration of those periods. We have no other choice but to abandon the generally accepted conception of there having been only one center of evolution from which from time to time new types thould have been sprung and spread over the whole world, dislodging the older ones. The locality of this supposed center usually is shifted according to the momentary results of excavations.

Once it was in East Africa, then in Central Asia. But as a matter of fact, cer-

tain indications of such an early and exclusive existence of prehominids have not been found as yet in either of the two regions, whereas just the extreme Far East of the Eurasian continent and an island situated in front of its southeastern corner (Java) have each yielded such a type. Moreover, exactly from this most remote locality we have two different primitive

forms - Pitheca'nlhro pus and Homo sol ensis - which, according to their morphological and geological conditions, can be considered as subsequent

stages of the same line and may continue to the Wad jak Man and further to the Australian aboriginal of today, as already suggested by numerous authors. That the SlnantJlro pus branch may have developed in the direction of certain Mongol groups can be demonstrated by anatomical facts. On the other hand, the opinion that Rhodesian Mart is dosely akin to the Negro has just recently also been expressed by Sir Arthur Keith.

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The question concerning the real origin of European hominid remains open. Although the discovery of the Mt. Carmel population contributed greatly

to the solution of this problem, yet Sir Arthur Keith and McCowu consider

the Tabun type as falling within the Ehringsdorf group (Krapina) arid the Skhül type as closely related to the Cro-Magnon form of recent man but nevertheless akin to each other. in this case, the center or one of the centers of the origin of present European mankind must have been located in Palestine. Thus, at least four centers of origination become probable up to the present: one in Asia Minor with relations to the European races, one in East or South Africa with relations to Negroes, one in North China with relations to Northern Mongols and one within the Sunda-Islands with

relations to Australians and Melanesians. On the other hand, fossil evidence proving the existence of only one center is completely lacking. References to migrations of primitive races of present mankind even to the most remote corners oí the world cannot be accepted as convincing proof.

There is no evidence pointing to a real world-wide wandering already in the early stages as represented by the prehominids. If any conclusions may be drawn from later conditions, then we may say that they point exactly to the opposite direction, for the actual existence of races and the surprising tenacity of their characteristics are apparently at variance with the theory of monocentric evolution and the assumption of a relatively late appearaice of racial differentiations. The fact that the latter already existed in prehominids (Pithecanthropus-Sinanthropus) as well as in the Homo primi genius stage (La Chapelle-aux-Saints - Rhodesiaii Man - Homo

soloensis) demonstrate that fundamental regional differentiation must be as

archaic as human evolution itself. In addition, the establishment and fixation of special characters requires - as the history of mankind proves not only extensive periods of time but also strict isolation throughout such a process. Continuous and unlimited mixing, an inevitable consequence of permanent migrations, would have frustrated every step towards fixation.

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I do not deny that certain changes occured in places of habitation in remote times aiso, but I doubt that migrations involved the entire population and extended over whole continents, as presupposed by the monocentric theory. The immigration of the Indians to the American continent cannot

be produced as a counterproof against the latter assertion. Firstly, this wandering took place when man had already reached the stage of his present type, secondly it shows that, on the contrary, the main features of the original anthropological character have been unaltered as a consequence

of strict isolation and that changes began only after descendants from other racial stocks came in touch with the first-corners.

The theory of polycentric evolution makes plausible not only the differentiation of races and the fixation of the radai characters but also the

differences in the time of evolution which must be deduced from the chronological facts. We merely have to admit that human development was not progressing at the same pace everywhere but was accellerated here and retarded there. in such case, it becomes conceivable that within certain regions of the world and at a given time hominids of an advanced type existed simultaneously with more primitive stages in other regions. Such an assumption would serve to explain the contemporaneousness of

types like the Steinheim skull in Europe and Sinanthropus or Pithecanthropus in the Far East during the Middle Pleistocene as weil as the contemporaneousness of Australian natives and European of our time. The conception of a polycentric evolution of man as presented here is not equivalent to a polyphyletic evolution. This term implies an origin from different primate or anthropoid branches gradually transforming into a uniform type by convergence. I believe in a world-wide distribution, at

least of the pre-anthropoids, in the sense of Montandon'8 hologenesis, and as suggested by the actual distribution of the Dryopilhecus types. The general tendency of the development in the direction to recent man must

The classification of fossil horn inids and their relatlon,

75

be the same in principle within all places and times; regional differences concern only minor details which are usually considered racial deviations. But should the European or African Homo sapiens really be as ancient a

type as recorded in Table Il, then that forerunner must belong to a geologically much older period than Sinanthropus himself despite their phylogenetic homogeneousness. On the other hand, the Western Homo

sapiens as such would represent a much older type than the same Horno sapiens of other parts of the woild who reached their destination later. I am fully aware that my view exposed here is of rather hypothetical

nature.

But I failed to see any other way which may lead out of the

difficulties. Should it ever become evident that the remains of Homo sapiens

attributed to the Lower and Middle Pleistocene belonged to much tatet periods, the problem would become simplified but not altered in principle. For other discrepancies existing between the morphological and geological facts would remain and could only be explained by accepting a polycentric evolution of man.

VI. The phylogenetic development of man and the general theories on evolution The .study of the phylogenetic development of hominids as far as it can be deduced from the various fossil remains themselves known hitherto

raises the question which of the general factors may be responsible for the special course taken by man in evolution. So as to relax from the daily routine of describing and comparing, we at times feel the need to devote ourselves to those more general problems. in doing so, it will prevent us from neglecting too badly the general biological laws which govern the creation of new types such as we ourselves represent when compared with the lowest hominid forms.

Edwin L. Goodrich, the well known zoologist of Oxford, in his book on the living organisms in which he attempts to prove once more the correctness of the Darwinian theory, states: "Every theory of evolution must be tested by the results of palaeontology; no condusion can be accepted which is inconsistent with them." Such a reservation is of significance. For, though theories of evolution as first interpreted by Lamarck and Darwin essentially deal with historical matters, they were chiefly deductions inferred from the differing organisations of living organisms.

But their very development was almost completely unknown then, and likewise undetermined whether and in how far the individual phases of evolution actually display the influence of the assumed driving factors of evolution. This implies that tests of accordance between theoretical' presuppositions and objective historical facts were not known at that time. Even today the same omission is often being made. There are geneticists 76

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77

who believe to be able to determine the factors of the phylogenetic evolution

exclusively on the basis of experiments on living organisms.

They fail to

realize that a process which required millions of years to make its appearance can hardly be unravelled by more or less accidental breeding of a few generations and in addition to this under artificial conditions of which no one knows whether they correspond in any way to those actually having been in force in nature.

Hence, as Goodrich admits himself, only

palaeontologists are qualified to definitively answer questions concerning Lamarckism, Darwinism and other theories on evolution. Although palaeontological material at present is rather abundant as a whole, nevertheless really uninterrupted series of special lines are still lacking which may represent the actual sequence of the stages of evolution

and at the same time furnish an opportunity for tracing back the direct causes of those alterations. The horse and the transformation of its foot is one of the best known examples of this kind, despite the gaps which exist

in this case also. But it is not only the question to ascertain the line of evolution but also to prove that all the individual phases really owe their

special form to selection or adaptation. Viewed from this standpoint, one very soon realizes that little advance has so far been made beyond attempts oí rather superficial interpretations.

In this connection the human material serves as an excellent example. With respect to details and variations of the skeleton, man has certainly been better studied than any mammal but, in addition, we are familiar particularly in this case with the statical and dynamic conditions which affect the elements of man's skeletal system. These conditions elucidate the special nature of its architecture much more than is true of other organisms. Is there a real phylogenetic line with distinct differentiations of single phases derived from each other, then we may trace not only that course of evolution but also ascertain whether deviations existed

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and whither they led. The discovery of Sinanthropus pek1nenss made it possible to define the characteristics of a primitive hominid and, in addition,

to trace all his transformations up to recent man with the different types of the Neanderthal group serving as links between the individual forms. in these cases, we are fairly acquainted with the morphological appearance

of the types as we11 as their environment as represented by the fauna, flora, the climatic conditions and the culture. Thus all facts are available which permit us to determine whether radical changes in environmental conditions took place to such an extent as to explain the existing morphological differences in the light of proper adaptations to the new conditions. In addition, we are able to test whether in the course of evolution chance variations really occurred in such a degree as to supply sufficient material at all times from which the fittest could be singled out in

accordance with the requirements of the theory of selection.

Let us begin with the dentition. The tooth structure is considered an excellent example of the effect of adaptation to food habits. Indeed, the teeth seem so completely adapted to the type of food that every palaeontologist takes it upon himself to diagnose the kind of food on the basis of the tooth pattern alone, and to deduce therefrom the general living conditions of the creature in question. lt is supposed thereby that the state

of being adapted indicates at the same time also the manner in which the adaptation was brought about. This implies that the special kind of' food is directly responsible for the special structure of the tooth. For example,

the 'hypsodont" molars of the horse, that is to say, representing high prismatic pillars, should not only be especially adapted to grinding hard grass and grain but also be derived from originally "brachyodont" (low) teeth by a consistent food habit. O. Abel literally states: (in Staemme der Wirbeltiere, 1919, p. 792) that the specialization in question had been acquired by the change from feeding on soft herbs to feeding on hard grass.

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The literature dealing with the teeth of hominids contains a wealth

Each difference evident in the teeth of recent man when compared with those of anthropoids has been explained as being

0f similar statements.

merely due to the omnuvorous character of the human teeth and the frugivorous character of the anthropoid teeth. Such a conception does not include an indirect chemical effect of special ingredients of the food but apparently refers exclusively to a mechanical factor. The actual course of human evolution, however, reveals an entirely

different picture of the change which took place within the dentition. Viewed from the zoological standpoint, the dentition of Sinanthropus is that of an anthropoid. The lower premolars and molars just as an example

are low and long with very stout crowns and roots (Figs. 38a.,c.; 39a., c.; 32a., b.). In addition, they show a distinct cingulum and numerous well pronounced cusps, whereas their entire chewing surface is covered by an abundance of special wrinkles (Fig. 32o, 35a, 36c.). in the course of

human evolution these teeth experienced a characteristic transformation. They became reduced in robustness of both crown and root and especially in length, so that finlly they appear to be higher and narrower than before (Figs. 32c.; 36f.). The cingulum has been lost; the cusps and wrinkles decreased in size and number. Hence, there is a clear tendency to reduce the size of the entire tooth and to substitute its originally complicated pattern by a very simple one. The same process takes place in the incisors and canine, particularly in the latter (Fig. 34b., c.). Can all this really be regarded as consequence of an alteration in food habits? Disregarding the canines, the patterns of the Sinanthro pus

teeth are undoubtedly much more similar to those of anthropoids than to those of recent man (Figs, 35; 38b., d.; 39b., d;). Even his molars are much larger than those of chimpanzee and approximately of the size of those of orang. According to a widely spread belief the descendants of

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Sinarsthropus should have lost their anthropoíd-like dentition by abandoning

the fruit diet to one largely composed of meat. Unfavorable to this theory however, is the evidence that meat had already been induded in the menu

This fact can be proven by his culture: he hunted for game, particularly the deer which oinposs approximately 60%. of the

of Sinanfhrojuis.

fauna recovered from the cave of Choukoutien.

He dissected the animals with the aid of implements and crushed the bones for the purpose of extracting the marrow (Fig. 49). More than that, traces of fire and of burnt

bones (Fig. 50) render it certain that he was already familiar with the preparation of meat too.

As it was shown, the dentition of recent man compared with that of Sinanthropux is muçh weaker and displays a distinct tendency towards simplifying the chewing surface. This condition, however, is exactly in contrast to what may be expected from a meat diet, for the latter certainly requires stronger teeth with a rather uneven chewing surface than a fruit diet - if the character of the dentition really is dependent upon dynamic factors. Objection may be raised that hominids of later times did not eat raw but prepared meat, a habit which apparently required less strengtI. Even in such a case the biting off and the ¿ewing necessitates a greater strength than the chewing of fruit which all the apes peel off carefully first.

On the basis of an unbiased judgement, we may consider the dentition of an anthropoid or that of Sinantliropus as being much better adapted to a

meat diet than is true for that of recent man. in addition, it must be borne in mind that all primates do not feed on fruit exdusively but also on insects, eggs and small birds. The reduction and simplification of the human dentition, therefore1

cannot be the consequence of a mere change in food habits. Another fact may be added. Apes of the Old World show the same general character of dentition as Lemurs and apes of the New World but differ from each

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81

other in a very specific manner, and even within individual groups corsiderable differences exist. The molars of gorilla, orang and chimpanzee exhi-

bit such characteristic peculiarities that they serve as a reliable criterion for the classification of the species itself (Figs. 35; 36). The food habits of all

primates, however, are the same. Nevertheless, New World apes differ korn those of the Old World even in the number of teeth. Their speciality, therefore, cannot be determined by food habits. An additional and most decisive reason for excluding nutrition from being responsible for tooth

patterns is the fact that the patterns become móre or iess completely worn already shortly after the eruption of the tooth, thereby confining the actual

functioning to an extremely short duration (Fig. 32 b., o). This fact alone is incompatible with the idea that the diet is responsible for the special

nature of the pattern and its disappearance in the course of human evolution.

Moreover, the teeth are by no means independent structures. Their tra'isforrnation in size, proportions and pattern is concomittant and in accordance with certain transformations of jaws, face and brain case. The reduction of the roots causes a reduction of the alveolar process of the upper and lower jaws and this again produces the prominence of the chin

(Fig. 43) and the nasal spine and the appearance of "hollow cheeks" (Fig. lb., c.). All these alterations can be swnmarized as the effect of a shortening of the muzzle.

The transformation of the brain case corresponds to such a change. There is a distinct tendency leading from Sinanthropus to recent man to

enlarge the capacity of the brain case at the expense of the face (Fig I). The brain increases in length and height in particular. The frontal bone comes to lie in a more vertical position, likewise the occipital and parietal bones.

The frontal lobe of the brain overlies the orbit completely, causing

in this way the disappearance oí the supraorbital ridges (Fig. 18).

The

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occipital foramen shifts downward and forward with the result that the brain case takes on more and more a globular form (Fig. 15). All the differences in details between the skull of Sinanthropus and that of recent man can be traced back to this essential transformation. Alterations within the dentition and jaws merely represent partial occurrences of this general change and can therefore not be influenced in detail nor in general by such relatively irrelevant factors as slight alterations in food habits.

Just as the direct adaptation must thus be excluded as a possible determinative factor of the special development of the teeth, so also must natural selection. Selection anticipates the presence of a certain number of variations from which the fittest can be singled out. There is no doubt that the number of variations within the Sinanthropus dentition is considerable, involving the size as well as the pattern. All those variations, however, point towaids one direction (Figs. 40, 41), while really specific deviations are completely absent. The trend of the variability, regardless whether it concerns Sinanthropus or the Neanderthal group or recent man always remains unchanged: namely to substitute a simpler pattern for a complicated one. The pendulum oscillates exclusively within this line and no feature outside of it has ever been observed. But even if the variation

in the number of the cusps of the molars or the change in their arrangement, for instance from the Dryopithecus pattern to the so-called "plus"pattern (Fig. 36), should be taken to represent essential variations in the sense of fundamental alterations of the entire type, it is impossible that they are of any significance to the selection, in other words, that extinction or existence of the individual or even the species may depend upon them.

That what is true of the teeth, also holds good for the jaws and the brain case. Within the lower jaw variations fluctuate from coarse types with poorly pronounced relief to such with well modelled surfaces. Besides

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high and narrow jaws, there are iow and broad ones; the ramus in some cases ascends steeply, in other it is more inclined; the mental spine can be present or substituted by a pit, etc. All these variations show the same character, being partly due to differences in sex, partly correspondent to the general direction oí human evolution, and partly fail to reveal any special phylogenetic meaning and may to a certain extent occur in recent man in the same way as in Sinanthropus.

The variations of the brain case of Sinanthropus are of greater interest. They concern the size in particular. Side-by-side with a skull of a capacity of not more than 650 cc there is one with a capacity of 1220 cc. Although it is probable that this variability has to be considered a consequence of sexual differences, it nevertheless follows the general trend

of human evolution.

This view is supported by the fact that the

differences in size are closely correlated with certain differences in the ex-

ternal appearance of the brain case. The supraorbital ridges of the larger skulls are not as distinctly separated from the forehead as those of the smaller ones. The greatest breadth of the larger skulls does not correspond to the biauricular breadth as is true in the small skulls (Fig. 2b.)

but instead has shifted to a higher level.

Real chance vaiiations which fall beyond the character of the type or tend to do so are not found in Sinanthropus or in any of the later fossil hominids known hitherto. Thus, decisive variations are not available which offer a basis for selective action, regardless of the question whether this agent could be effective upon such minor details as specified here.

As a matter of fact, the evolution from the Sinanthropus stage to

that of recent man courses in one and the same direction.

Single

alterations are not brought about by mere local influences but must be considered a consequence of the transformation of the entire organism.

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The evolution of hominids as made evident by the palaeontological facts takes place in the form oí a typical "orthogenesis' in Eimer's sense. lt strives for differentiation with one predominating organ. This organ be-

comes the inducing center of the new organisation, with all the other alterations subordinated to it or representing consequences thereof. Within the ontogenetic development Wilhelm Roux distinguished between "self-

difterentiation" and "dependent differentiation." These terms may also be applied when attempting to define the principles of phylogenetic de-

As to hominids, the brain seems to be the seat of velopment. "sell differentiation," whereas the face and its components represent with regard to the brain "dependent differentiations." The tendency as a whole apparently is directed towards a special differentiation of the brain.

Palaeontological facts lead to an additional result.

There is no

evidence that the phylogenetic evolution of hoininids does progress in leaps.

No indications have been found of the sudden appearance of a new type, or what may be interpreted as a mutation in the sense of de Vries. The single alterations of the teeth are directly derivable from each other and represent a continuous line. The progressive differentiation becomes manifest by the gradual increase in number of individual variations. Many primitive characters may thus be retained even in recent man (atavism). The same is true of the development of the brain case. Real gaps within

the line of evolution do not represent leaps but rather lack of suitable material as has been proven, for instance, by the but recent discovery of the Mt. Carmel pooulation which links the Neanderthal tvoe with modern man.

lt was demonstrated that neither direct adaptation nor chance variation with subsequent selection . can be recognized in individual structures of the organism. Thus, the question arises whether in the course of evolution the influence of those factors may become manifest

The phylo genetic development and general theories

85

within the organism as a whole. Are there external factors which can be interpreted as representing an essential chance of environmental conditions forcing the organism to take a certain course in evolution?

The literature contains numerous such suggestions. We are told that during the glacial age privation and distress stimulated the European Neanderthal man towards acquiring a greater efficiency resulting in the creation of the more advanced type of recent man. But as far as we know now this trans-

formation did not take place in or near the ice covered regions of the world. Moreover, privation and distress are too indefinite to be consider-

ed either as direct or indirect causes.

I even doubt whether general

climatic changes are capable oí causing or driving orthogenetic evolution.

As much as can be concluded from the character of the fauna and flora, the climatic conditions of the Western Hills and the plain of Chihli in North China, the habitation of Sinanfhropus were not too different from the conditions of today. In any case, the climate was certainly unlike that of Java. But Pithecanthropus, a hominid form dosely related to

Sinanthropus and contemporaneous with him, apparently already lived under the tropical conditions of Java. This contemporaneity shows that man within the Sinanthropus-Pithecanlhropus stage of evolution must have

already been adapted to different climates like man of today whose habitation is distributed over polar and equatorial regions.

Heat and cold, humidity and dryness, therefore, can hardly be held responsible for the physical transformation of the hominids.

As to the selection, the majority of investigators agree that this factor in no instance iicapable of producing new types by itself but merely of singling out from different types already present. The problem, how-

ever, is which of the factors actually cause orthogenetic development. The conception that the material for selection is offered by chance variations

must be excluded as far as the hominids are concerned, since the actual

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finds failed to reveal their existence.

or all types known hitherto already

completely fall within the line of evolution with renard to their general character as well as to the details. When speaking of environmental conditions we bear in mind not only the climatic but also other external conditions of existence. I have already mentioned the nutrition and its mechanical effect upon the dentition. Theoretically, it may be possible that the meat diet transformed the anthropoid ancestor of man by the addition of chemically active substances to the humors of the body and thereby stimulated also his brain to enlarge and differentiate. Yet, in what manner such a change in food habit or its

suggested effect on the organism may have brought about orthogenetic development lasting for hundred thousands of years can not easily be understood. Special significance has at all times been attributed to those environmental conditions which are considered responsible for the erect posture of man. This posture requires a skeletal system adapted to particular static and dynamic conditions. Standing and walking in upright posture is, of course, impossible unlessthe foot is constructed accordingly, and the same is true of the lower limbs, the pelvis and last but not least the spine and the skull There is not the slightest doubtall stu(lents of comparative anatomy agree on this pointthat man "in the beginning" cannot have possesed these adaptations but was equipped with a dimbing foot like all the other primates. Some years ago I was able to demonstrate in which way the transformation of this climbing foot may have taken place. in addition, I demonstrated that not only the characteristic sagittal cnrvatures of the human spine but also the so-called basal kyphosis of the skull, an indication of the tendency towards assuming a globular form, must

be consIdered a direct consequence of the upright posture (Figs. 48; 52). In other words, the transformation nf the skull with a special adaptation for expanding the dorsal parts of the brain should be due to changed static

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87

and dynamic conditions as a consequence of the erection of the trunk and the

extreme extension of the lower limbs as is characteristic for upright posture. Hence, such a posture had to precede the development of the brain case and the enlargement of the brain.

Can these conclusions be based upon genuine

palaeoltological finds?

Until very recently limb-bones of Snanthro pus were almost comple-

tely absent in our material. Our collection merely consisted of four small pieces, namely a fragment of an atlas, a collar bone, a small fragment of a humerus and one of the carpal bones.

But within the last year we had the

good fortune to find six fragments of the femur and one large fragment of the

These limb-bones are sufficiently preserved to permit a criterion on their completely human character and their distinct differences from the humerus.

corresponding limb-bones of great apes (Figs. 45; 47; 51). It follows from this fact that there undoubtedly exists a strong contrast between the appearance of the sknll of Slnanthropus and that of the skeletal parts of his body. This phenomenon can only imply that within the phylogenetic evolution the body probably preceded the head, or, in other words, man had already adopted the erect posture while his brain and skull still remained in a more anthropoid-like phase. If such an assumption is correct, then the specific human differentiation mentioned above must again be transferred. Brain or brain case cannot be considered as the organs supplying the decisive impulse3

for further evolution but the erect posture seems to have done so, freeing the hand from locoinotorial f imctions and, at the same time, supplying the brain with the opportunity of gaining sufficient space in which to expand. Now a new question arises: what circumstances forced the ancestor of man to assume an erect posture? Since the ancestor was a climbing primate, it suggests itself that the direct cause may have been the disappearance of forests due to changes in climatic conditions. This, indeed, is a widely spread belief. However, I must say, that I cannot accept such an explanation. Of course,

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discussions on this problem are pure speculations on account of the complete lack of real knowledge as to time and place where the event in question may have occurred. In addition, there are certain suppositions of this theory which

are rather precarious.

The character of the Sinanthropus dentition. indicates

that his forerunner and with him the early hominids must have branched off from the general stem of the anthropoids before the differentiation of the Dryohitpecus line occurred (cf. Lecture II). This suggests the ancestor to Sinanthropus having lived within the Lower Miocene at least. Can it be surmised that during the long period of time required by the transformation

from a climbing foot into one adapted to standing and walking. a hominid was able to survive within an area which not only was lacking in opportunity

for using his foot in the accustomed way but also was barren of shelter and food? It is much more likely that this hominid deliberately or voluntarily assumed the habit of living on the ground because the proportions of his upper and low limbs already in existence then favored such a change.

All the ideas of compulsory influence of the environment upon the development of the hominids are based more or less on the conception that

the evolution evolved within one locality only and was confined to a few In other words, that there should have been one cradle of individuals. mankind fixed geographically and temporally.

Such a presupposition in the first place overlooks that human evolution cannot have been restricted to a certain time.

For this evolution was not an event taking place but once but

rather represents a process which extended over a period of initiions of years.

We are familiar with but the last stages during which the hoininids were distributed over a considerable area of the Old World. It is truc, that as yet continuous series of types restricted to one locality are not available. Fur-

thermore there is a gap in Europe as well as in East Asia between the Neanderthal group and recent man. But the extensive distribution of the

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89

immediate forerunners of present mankind and his early division into various races is proven by every new discovery permitting no other assumption but

that human evolution was neither restricted to one place nor did it progress contemporaneously everywhere. I have been able to prove that Sinanthropus in particular revealed closer relations to certain racial groups of today, especially to some Mongol groups, than to others.

The racial multiformity of mankind is not in the least a relatively late acquisition as many anthropologists believe but a phenomenon forced to accompany the entire evolution (cf. Lecture V). The main races represent geographical varieties of hominids. If so - I fail to find another explanation -

then the tendency to transmute the primitive types into those of recent man must be considered inherent to the form as such. This transmutation, there-

fore, cannot have been caused by accidental and local events which must always differ in place and time. In addition, it seems rather unlike that human evolution evolved at the same rate in every part of the world.

Despite

the scarcity of material, all finds of fossil man point to the fact that the development progressed slower or faster according to the locality. Regardless of his greater primitiveness, Sinanthropus may, therefore, very well have been

contemporaneous with more advanced Neanderthal types of Central Europe, and in consequence thereof, those Mongol groups of recent mankind Which

derived from that Asiatic hominid may constitute later differentiations than those groups of the white which originated from Neanderthal man or other primitive types of Europe or West Asia. We are inclined to consider the primitive races of today as the actual survivors of earlier stages of evolution. The Australian native, for instance, is regarded by many anthropologists as such a stationary palaeolithic man, possibly even identical with the Palacolithic

man of Europe.

But it is also possible that certain races entered into their evolution later than other races and independent from them.

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The fact that despite local and temporal differences the tread of human evolution reveals the same general character at all places and at all times contradicts the assumption of external factors directing the spec:aI course of evolution.

These may determine certain minor differenccs but hardly the

formation of the general character of recent mankind. upon

Changes depending

place and time are incompatible with the nature of orthogenesis.

Environmental conditions and selection can only be of influence by way of elimination of individuals.

According to an opinion frequently expressed neither comparative anatomy nor palaeontology should be qualified to unravel the riddles of evolu-

tion, since they can only deal with phenotype6 but not with genotypes, while the entire evolution should be based exclusively upon alterations of the latter.

The purely physical appearance of the various fossil types representing phenotypes only, would, according to this idea, be rather irrelevant. This conception is partly derived from an erroneous understanding of the transfor¡nation of species. The claim that the ancestor of man was an anthropoid merely implies that man passed through a stage of organisation corresponding to that of the great apes of today, but not that he really represents a transmuted gorilla or chimpanzee.

Therefore, the question is whether he changed his

inherent nature at all by evolving in a special direction which diverged from

that taken by other groups of the same orginal stem. Since phylogenetic development can only be manifest in phenotypes, the genotypical basis of an organism is more or less irrelevant when we trace the course taken by evolution.

For, it even occurs within the human evolution that traits may be specific for the type or for a certain phase thereof tliugh evidently of phenotypic nature. Most of the dependent differentiations may be those phenotypical

phenomena despite their morphological specificness.

Viewed from the com-

parative standpoint this specificness is of much greater importance than their eventual genotypical nature. The behavior of the lateral process of the

The phylogenetic development and general theories

91

tuberositY of the human heel bone is an excellent example of this kind, as I have demonstrated several yeats ago.

given by the brain case.

Another not less instructive example is

Early disturbances in the normal development of

the brain obstructs its growth, and the brain case not only remains small as a

whole but also assumes a very primitive appearance by the formation of supraorbital ridges and the approach of the temporal lines to the midline of the skull (Figs li; I 3). In other worlds, in the case of idiots the skull may become surprisingly similar to that of a primitive hominid like Sinanthro pus

or an anthropoid.

Morphological peculiarities specific to early stages of

human evolution may reappear.

The real significance of this phenomenon is

the fact that those structures which characterize the brain case of recent man even in the present advanced stage are not so inherently fixed as to have been developed quite independently from all the other factors. Hence, they

appear in the form of phenotypical manifestations, whereas the genotypical reaction seems to remain unaltered in principle. 'These facts reveal that the

problem on the phylogenetic evolution can be dealt with without laying decisive weight on the discrimination of the genotypical and phenotypical characters of the individual features.

As regards selection, it was mentioned above that it could never serve as an explanation for the adaptation or the transformation of an organism in Darwin's original sense. Chance variations are not traceable during human

evolution and each essential alteration can be ranged within its direct orthogenetic course. On the other hand, the elimination of individuals is less depend-

ent upon the struggle for better environmental or living conditions than upon

If, for example, the sea transgresses its boundaries or falls dry, the land or sea animals living in these regions will perish, while only those mdivchance.

iduals absent from the catastrophe will survive. In wars not the most gallant

fighters will survive but those who are beyond the range of being hit by

BulletIn of the Geological Society of China

92

bombs or shells. These survivors will have the opportunity of transmitting their own peculiarities to the descendants, regardless of their specific worth or worthlessness. Accidents within evolution do not depend upon the accidental

character of occurring variations but upon thc accidental selection of the survivor.

My conclusion is that internal factors determine not only the special constitution of the organism but also the evolution of its stock. I am not able to explain why constitutions show differing qualities. This question seems to

be connected with the problem of life itself.

For life is not any abstract

conception but is manifest exclusively within constitutions and differentiations.

There exists no organism at present or at any time

rlier without a specific

form and a special adaptation to the corresponding environmental conditions. Neither an organism nor life itself could endure otherwise.

Phylogenetic development like ontogenetic development constitutes nothing else but a phase of life.

In contrast to the ontogenetic development,

however, the phylogenetic development does not refer to the individual but to

the entire group. It does not cover the brief span of individual life but the duration of the whole species represented by countless generations. Like the ontogenetic development which is propelled by intentai impulsesindiffereently known as growththe phylogenetic development takes its course without being influenced decisively by accidental external factors.

Explanation of

Plate I

93

PLATE I.

Figures I -3.

Comparative view of the skulls of (a.) female adult gorilla

No. 324, (b.) a reconstruction of a female Sinanthropus individual and (c.) modern North Chinese male (No. 173). 1/6 The reconstruction of the Sinanthropus skull carried out by the author with the assistance of Mrs. Lucile Swan is based upon real findings of all parts

of the skull, with the exception of the zygomatic arch. The brain case as a whole represents .inanthropus Skull 11 of Locus L (compare Fig. 5 a.). The skulls are orientated in the Frankfort plane. Fig. 1, norma lateralis; Fig. 2, norma occipitalis; Fig. 3 norma f rontalis.

Sinanthropus skull Ill of Locus L. Male individual in 3/4

Figure 4.

profile. 1/3. Note the heavy eyebrow ridges, the receding forehead separated from the ridges by a distinct broad furrow, and the nasal bridge.

Figure 5. Comparative view of the female Sinanthropus Skull II of Locus L (a.), female Pithecanthropus Skull Il recovered by Dr. von Koenigswald in 1937 (b.), and the male Skull No. VI of Homo soloensis (c.). All three skulls reduced to the same length and orientated in Frankfort plane.

Note the similarity in general shape between Pilhecanthroprn (b.) and Homo soloensis (c.) and the difference between Sinanthropus (a.) and Pithecan-

thropn (b.) with regard to the sagittal arch of the brain case: vaulting of the frontal region and depression of the obelion region in Slnanfhroptrs and the reverse conditions in Pllhecanthropus.

Figure 6.

Basal views of the pyramis of the temporal bones of a chimpan-

zee, Sinantlirnpus Skull of Locus E and a recent Chinese.

1/4.

The axis of the pyramis is bent in chimpanzee and Sinanthropus, while n modern man it is straight. In addition, the orientation of the bone as a whole

to the midline of the skull is the same in chimpanzee and Sinanhropas but different in modern man. 94

rariz Weiden reic h: Six Lectures

n Sinanthropus pekinensts

Plate I

Q

N co

u

CV)

Q

In

Explanation of Plate II

95

PLATE TI

of Locus L, showing a long and deep depression on the left side of the bregma region and parallel to the midline, which was apparently caused by a blow while the Male Sinanthropus Skull

Figure 7.

individual was still alive.

I

1/3.

Figure 8. Male Sinanthropus Skull Ill of Locus L, showing a long and narrow depression on the right side of the vertex and approximately parallel to the midline, apparently caused by a blow during life.

113.

Figure 9. Lateral view of the skull of an infantile male chimpanzee No. 389 (a) and an infantile male North Chinese No. 50 (b.). 1/4. Both skulls are of about the saine age in regard to the stage of the dentition: first permanent molar erupting. The forehead of the chimpanzee seems to be more erect and more human-like than in adult individuals.

Figures lO and 12.

Indications of injuries to the skull caused by blows during life in the case of the Weimar-Ehringsdorf skull (Neanderthal type). 2,3. x1 linear depressions caused by a long and sharp implement; x2 straight breakáge lines indicating the use of a similar implement.

Figure I 1.

Vertical view of the skull of a female adult gorilla No. 324

(a.), and that of a modern adult idiot, Schuttelndreyer of Göttingen, (b.).

1/4.

In consequence of a cessation in the development of the brain during fetal life, the skull of the idiot assumed an ape-like form: the skull is much narrower than the face (note the lateral expansion of the zygomatic arch in relation to the breadth of the brain case and the course of the temporal lines approaching the sagittal suture) and the development of veal eyebrow ridges (compare Fig. 13).

96

Franz F'Veidenrek!:Six Leclures on Sinanihroptis pekinensis

Plate 11

Explanation of Plate Ill

97

PLATE 1H

Figure 13.

Skull of an adult male idiot Schittelridreyer of Gdttingen.

114. (cf. explantion of Fig. 12). Note the large size of the face in proportion to the small brain case, the heavy eyebrow ridges and the low and receding forehead.

Figure 14. The torus occipitalis and the formations connected therewith of the female Sinanthropus Skull II of Locus L. 1/2. Abbreviations: cm, crista supramastoidea: corn, crista occipito-mastoidea; li, linea nuchae inferior; is, linea nuchae superior; it, linea temporalis; st; sulcas

supratoralis; t, torus occipitalis; tp. torus anguli parietalis.

Figure 15.

The three skulls of Fig. 1. 1/6. The three lines drawn on each skull represent the curvatures of the

outlines of the brain cases, the courses of the squamosal sutures and the direct

surrounding óf the mandibular fossa. The vertical interrupted line traced through the posterior ends of the mandibular fossae indicates the axis around which the brain cases curve. Note the gradual increase of the curvatures from gorilla (a.) over Sinanthropus (b.) to recent man (c.).

Figure 16. The sagittal crest of the Sinanthropus skull (a.) and that of modern Eskimo, the latter after Filrst and Hansen (b.). 1/6.

Figure 17.

The same as in Fig. 16 observed in modern living individuals: North Chinese (a.); Australian aboriginal after Milducho-Maclday (b.).

Figure 18. The topographical relation between the orbit and the frontal part of the skull cavity in chimpanzee ¿ (a.), Sinanthropus 4. (b.) 1/4. and modern North Chinese c/' (c.) Note the tendency of the brain to overlap the orbit. The skull capacities amount to 475 cc, 1015 cc, 1500 cc.

98

Franz

Weide1

Sir Lectures on Sinanthropus pekinensis

Plate

Is

cm

C.

III

Explanation of

Plate IV

99

PLATE IV

Zones indicating the range o) variation in size of endocasts (brain) within anthropoids (innermost zone bordered by fine lines), Sinantliropus (next zone bordered by heavy full lines), Nean-

Figures ¡ 9-21.

derthal group (the following zone bordered by dotted lines), recent man (outmost zone bordered by interrupted lines).

1 i3.

Fig. 19. Zones in horizontal plane corresponding to the level of the greatest breadth; Fig. 20. Zones 'io sagittal plane; Fig. 21. Zones in frontal plane corresponding to the level of the greatest breadth.

Figure 22.

Frontal lobes of endocasts and brain, respectively, viewed

from in front.

1i4.

a, Sn,phalangus syndact. ¿q; b, chimpanzee ¿q - brain; c, SInanihroxss Skull III Locus L ¿q; d, Pitheconthropus 1 .., e, Homo soloensis ¿q No. V; f, Rhodesian Man ¿';g ,cre ent man (Moravia't-.). Abbreviations; oc, orbital opercilum; 10, Kappers' fissure No. 10.

Figure 23. Ramification of the middle meningeal vessels on endocasts. 14. a, gorilla ¿q; b, Sinanthro pus Skull II Locus L ..; c, recent mau (Moray-

ian.?.); d, Upper Palaeolithic man - Predmost No. X4..

In Sinanthropus like in gorilla the ramus posterior is much longer and thicker than the raums anterior whereas in recent man the reverse is tme. In adition, the ramification is much more abundant in recent roan than in Sinanthropus. c indicates the primitive type of ramification in recent man, d the more advanced one.

Figure 24.

Endocast of a juvenile orang viewed from below.

oo, orbital operculuni.

I (X)

¡/3.

FTCT?Z J4'eidenrec

Plate IV

--Six Lec'v res or Sian!hropus pe'inensTs

20

19

22 a. °

21

g

23

C.

24

Explanation of

Plate V

o'

PLATE V

Figure 25. Comparative view of the shape of endocasts (brains) of an anthropoid (chimpanzee - a.); Sinanthropus (Skull Il Locus L - b.) and recent man (New Britain aboriginal c.). 1/6. The latter has about the same cranial capacity - 93 cc- as Sinanthropw, but the shape of the brain is different.

Figure 26. The special centers of the lateral surface of the human brain after Kleist. Figure 27. The temporal and occipital lobe and the upper part of the cerebellum as manifested in endocats. 114. a, Sinanlhropw Skull H ; b, Neanderthal type (Gibraltar), c, recent man (New Britain aboriginal, cf. Fig. 25. b.). Note the great difference in form and size of the temporal lobe in Sinan thropus on one side and Neanderthal and recent man on the other.

Figure 26.

The special centers of the lateral surface of the human brain

with their supposed significance after Kleist (English translation after Leyman).

Figure 29.

Endocast (a.) and brain (b.) of one and the saine individual

- male chimpanzee - to show that the actual location of the main fissures (Sylvian fissure, sf and lunate sulcus, Is) are obscure on endocasts. Reduced to slightly less than 1/3. Figure 30. Endocast of Sinant hro pus Skull Ill of Locus L from below. 1/4. oo, orbital operculum.

Figure 3 L

Endocast of Sinanthropus Skull Ill of Locus L from above. 1/4.

Note the prominences of the temporal lobe protruding more on the right than on the left side.

102

Franz We!denreich:Stx Lectures on Sinanthropus pel?inenszs

27

26

C.

28

30

31

Plate

V

Explanation of

Plate VI

UX3

PLATE VI

Figure 32.

Lower molar oÍ .Sinanthropus (a. and b.) and recent man

About ;1.

(c.).

Abbreviations: b, buccal view; d, distal view; 1, lingual view; m, mesial vieyv; o, occiusal view.

(a) Upper and (b) lower canine of Sinanthropus4.. About Ill.

Figure 33.

Note the difference in shape and pattern between the two canines. Abbreviations as in Fig. '2.

Upper canine of Sinanthropus p (b.) compared with that of rerent man (c.) and the lower canine of a female fo6sil orang. (a.)

Figure 34.

About I/i. Note the closer resemblance of the Sinanthropas canine to that of orang than to that of recent man. Abbreviations as in Fig. 32.

Figure 35. Comparative view of upper first or second molars of Sinanthropus (a.), Australopithecus Dart (b.), chimpanzee (c.) and gorilla

About I/I. Note the similarity in pattern between Sinonthropw and Austa1opIthecus.

d, distal sides.

Figure 36.

Comparative view of lower first or second molars of gorilla

(a.) orang, (b.), chimpanzee (c.), A ustralopithecu.s (d.), Smanthropus

and recent man (f.). About I/i. Note the likeness in general form and pattern between Sinanthropus and the anthropoids, including Australoplthecus, and che difference from recent man.

Figure 37. Comparative view of the second lower deciduous molar of an Eocene Tarsioid (a.), Sinanthropus (b.), Dryopithecus then. (c.), chimpanzee (d.) and recent American Indian (e.). a,c,d,e after Gregory. Upper Line =buccal view; lower line=ooclusal view.

Note the complete preservation of the trigoitid in the Tarsioid and Sinanthropus and its gradual reduction in anthropoids and recent man.

104

Franz Weidenreic'i: -Six Lectures .n Sinjnthropus pekinensis

Plate VI

N N

o

ri

Explanation of

Plate VII

105

PLATE VII

Figure 38. (rnb)

First lower premolars of Sinantliropus (a. and c.), chimpanzee

and recent man (d.).

About 1/I.

Note that the Sthanlhropas tooth resembles that of the chimpanzee much closer than it does recent man Abbreviations as in Fig. S2.

Figure 39. Second lower preinolars of Sinanlhropus (a. and i:.), chimpanzee (b.) and recent man (d.). About I/I. (cf. remarks in Fig. 38) Figure 40. First lower premolars of chimpanzee. (a.), Slnanthropus (b. and c.) and recent man (d.-f). Photographs. About I / 1. Note the gradual reduction of the talonid and the liligual cusp.

Figure 41. Second lower premolars of chimpanzee (a.), Sinanthrobus (b.-d.) and recent man (e., f.). Photograjhs. About 1/i. Note the gradual reduction of the talonid, a tendency already indicated in Sinanthropus (b.-d.).

Figure 42. Diagrams of the robustness of the crowns of the lower teeth in a female chimpanzee (fine line) and in Sinanthropus (heavy line). The average robustness of each type of tooth can be read off from the scale on the left side. The robustness is expressed by the product of the length and the breadth of the crown. Note that the molars are stronger in Sinanthropus than in chimpanzee, while all of the remaining teeth, especially the incisors, are much weaker.

Figure 43. Mandibles of a female gorilla (a.), a male Sinanthropus G 1 (b.) and recent man (c.).

1/3.

The reconsthiction of the body of the SInanshrqus mandible was made on the basis of the preserved left side.

Note the size, the length of the dental arcade, the chin region and the steepness of the amus.

106

Franz U'etdenrejch:Sx Lectures on Sinanthropus pe4lnensis

Plate VII

Explanation of

Plate VIII

107

PLATE VItJ

Figure 44 Comparative view of the lower canine group of the "heteromorphic" type - female gorilla (a.) and Sivapilhecus sivalensis after Gregory and Heilman (b.) and oi the "homomorphic" type - Sinanthropus (c.).

II I

Tusk-like canine and sectorial type of the first premolar in 1:be heteromorphic group and incisor type of the canine (C) and non-sectorial type of the premolar (P1) in the homomorphic group.

Figure 45. Right femur (IV, Mli) of a female Sinanthropus (a.) and the same femur of a recent European (b.). Upper and lower end of the latter removed. 1/3. Figure 46.

Left femur (VII; MIII) of Sinanthropus, probably male.

1/3.

The bone is split lengthwise as most of the long bones of animals (cf. Fig. 49). Only the posterior wall is preserved.

Figute 47.

Left femur (I; C II) of a male Sinanthrobus. I3.

a., posterior view; b., anterior view.

108

Franz Weidenreich:Six LecEures on Sinanihropus pekinensis

p-

u'

C)

Plate

VIII

Explanation of

Plate IX

109

PLATE IX

Figure 48. Medial sagittal section through the skull and cervical spine of a Mycetes (a.) and the head of recent man (b.). Note the straight line of the skull base in Mycetes and the kyphotic curvature in man.

Figure 49. A heap of long bones of animals as they are usally found in The bones are broken and split lengthwise in order to expose the marrow cavity. Figure 50. Burnt and cracked implement (a.) and animal bones (b. and c.) of Locality I of Choukoutien. Figure 51. Fragment of a left femur (Il; J Il) of a female Sinanthropus. Locality 1 of Choukoutien.

Posterior view.

I Il.

Abbreviation: 1, latera) side.

The anterior and lateral sides of the fragment are blackened by fire; the extension of the blackening on the posterior side is indicated by dotted lines.

Figure 52.

Schemes indicating the curvatures of the middle axis of spine and skull of recent man (a.) and dog (b.). Figure 53. Fragment of a right femur (III; J III) of a male SinanLhropus.

1i2.

Only the medial and a part of the postcridr wall are preserved.

Figure 54. (a.).

Fragment of a left femur (VI; M I) of a male Sinanthropus 1x2.

A portion of the anterior-lateral wall is split off. The splinter (b.) is also preserved. On both pieces corresponling indentationts (a-d.) are evident which may have either been caused by impleinens or by bites of a carnivore.

Figure 55. Endocasts of the skulls of Fig. 9: a, infantile maie chimpanzee; b, infantile male Chinese. 1/6. Note that despite the great similarities in outlines of the skulls, the differences in the form of the endocasts arc not less than in such of adults (cf. Figs. 25 a. and c.). 110

Frar,

Weiden reich: Six Lectures on Sinanthropus pekinensis

Plate

52

53

¡X