Patricia Smith

Department of Anatomy and Embryology, Hebrew University Hadassah Schools of Medicine and Dental Medicine, Jerusalem, Israel

Received April 1977 and accepted December 1978

Keywords : evolution, deciduous dentition, Near East

Evolutionary Changes in the Deciduous Dentition of Near Eastern Populations*

Deciduous teeth from Middle and Upper Pleistocene sites in Europe and Israel were examined. In addition, data on a fairly complete series of teeth from the Epipaleolithic down to recent times was used to obtain better evaluation of temporal trends in tooth size and morphology. Some reduction in tooth size was found between Riss and Wfirm I - I I specimens, but the rate of reduction remained slow throughout the Wfirm, then accelerated in the early post-Pleistocene period. Little change in tooth size has taken place over the past 6000 years. Morphological differences as opposed to metric differences were most pronounced between the Neanderthals and later groups.

1. I n t r o d u c t i o n

The h u m a n deciduous dent i t ion is usual ly considered to be more conservat ive than the p e r m a n e n t dent i t ion in r ega rd to morpho logy , size and s tabi l i ty o f tooth n u m b e r a n d to resemble homologous teeth in the A n t h r o p o i d e a more t han do the p e r m a n e n t teeth. This conclusion is ma in ly based on compar isons m a d e be tween the tee th o f recent m a n a n d those of A n t h r o p o i d e a a n d appl ies especial ly to compar isons based on first m o l a r morpho logy (Gregory, 1922 ; R e m a n e , 1927; Jorgensen , 1956; Braban t , 1965; Koenigs- wald , 1967). However , wi th the except ion of Braban t ' s s tudy (1965), l i t t le a t t en t ion has been given to the discussion of long- te rm trends in the evolut ion of the dec iduous dent i t ion , a l though de ta i l ed s tudies have been publ i shed of the dec iduous den t i t ion of Aus t ra lop i th ic ines (Robinson, 1956), Homo erectus (Weidenre ich , 1937), Neande r tha l s (~enyiirek, 1959; T h o m a , 1963; Legoux, 1970; Smi th & Arensburg , 1977) and recent popula t ions (Jorgensen, 1956; H a n i h a r a , 1963, 1967), a n d these p rov ide a basis for eva lua t ing evolu t ionary trends in the dec iduous dent i t ion of recent man .

T h e purpose of this repor t is to descr ibe a n d define the ra te a n d p a t t e r n of changes found in the deciduous dent i t ion of homin ids from Near Eastern sites, over the past 40,000 years. I have c o m p a r e d this d a t a wi th tha t ava i lab le for E u r o p e a n sites over the same per iod, and fur ther examined dec iduous teeth from A r a g o and K r a p i n a in o rder to ob ta in a be t te r long te rm perspect ive of changes found.

2. M a t e r i a l s and M e t h o d

The dec iduous teeth of fossil specimens f rom European a n d Near Eas tern sites l isted in Tab les 1 a n d 3 were examined .* T e e t h showing slight to m o d e r a t e a t t r i t ion were

* Supported in part by grants in aid from the Wenner Gren Founda- tion, and the Joint Research Fund of Hadassah and the Hebrew University-Hadassah School of Dental Medicine, founded by the Alpha Omega Fraternity, and the Hadassah Medical Organization.

* Except where otherwise indicated, the specimens listed were examined by me during 1973-75. For permission to examine the specimens in their care I would like to thank L. Y. Rachmani, Rockefeller Museum, Jerusalem; I. Crnolatac, Norodne Museum, Zagreb; R. Gessain, Mus6e de l'Homme, Paris; J. Piveteau, J. Lehmann, and B. Vander- meerch, Facult6 des Sciences, Universit6 de Paris, Paris; L. Balout, Institut de Pal6ontologie Humaine, Paris; H. de Lumley and M. A. de Lumley, Facult6 des Sciences, Universit6 de Marseilles, Marseilles and T. Molleson, Natural History Museum, London.

Journal of Human Evolution (1978) 7, 401-408 0047-2848/78/050401 +09 $02.00/0 �9 1978 Academic Press Inc. (London) Limited

402 P. SMITH

measured using vernier calipers with ground Points, permitting an accuracy of measure- ment of 0.05 mm. Measurements taken were (1) maximum diameter of the tooth mesiodistally (MD) taken in a line parallel to the long axis of the tooth and (2) the maximum diameter of the tooth buccolingually (BL) taken at right angles to the first measurement.

Morphological observations were made of shovelling, lingual tubercle configuration, incisal edge reduction, molar cusp number and pattern, degree of buccal convexity and presence of accessory cusps and styles as defined by Remane (I 921), Jorgensen (1956) and Hanihara (1963). Specimens were grouped by period and region and mean values of tooth dimensions calculated for each group.

3. R e s u l t s

Odontometrics

In the Near Eastern sample all teeth showed some reduction from BP 40,000 to the present (Tables 1,2). The rate of reduction was, however, minimal during the Upper Paleolithic, so that there was very little difference in tooth size between Middle Paleolithic and Epipaleolithic specimens. Most of the reduction in size occurred between the Epipaleo- lithic and Chalcolithic periods, with teeth reaching modern dimensions in this period: thus little change in deciduous tooth size has occurred over the past 6000 years.

The incisors and especially the maxillary incisors show most reduction over the period studied (20% difference in mesiodistal dimensions between Middle Paleolithic and Chalcolithic specimens). The canines and first molars show approximately 8-10% differ- ence, while the upper second molar does not seem to have altered significantly in length (MD) at all over the past 40,000 years.

The European sample shows similar trends for the same interval of time (Tables 3,4). Teeth from the earlier Middle Paleolithic site of Krapina are the same size as those from the earlier, Lower Paleolithic site of Arago, and some 20% larger than those of the other Middle Paleolithic specimens. The sample sizes are small, but indicate that deciduous teeth were decreasing in size throughout the Middle and Upper Pleistocene, with the rate of molar reduction slowing down towards the end of this period, while incisor reduction has continued to the recent past. However the range of tooth size shows consider- able overlap between the different periods. Two of the Middle Paleolithic specimens (Subalyuk and Shanidar) have exceptionally small anterior teeth (Thoma, 1963; ~enyfirek, 1959) while some teeth, and especially molars, from specimens in more recent periods, reach Middle Paleolithic dimensions.

Morphology

The size differences present between the fossil and recent teeth are associated with differ- ences in the shape of the teeth. The fossil teeth are relatively thicker buccolingually so that they appear more robust and rounded (Plate I). All surfaces are markedly convex and taper inwards towards the occlusal surface. A buccal cingulum is frequently present on the buccal surfaces of the canines and molars (Plates 2 and 3). The incisors and canines are typically shovel shaped with a large lingual tubercle or marked lingual swelling; the incisal edge of the lateral incisor is rounded, with reduction of the distal lobe (Remane's type C) and the canines typically possess a slender distolingual cuspule. The upper first molars are usually four cusped with the occlusal surface forming an oblique angle to the main curve of the dental arch. The fourth cusp, the hypocone is very small and in the

Plate 1. Deciduous dentition of the Neanderthal infant from Kebara.

Plate 2. Lingual aspect of Neanderthal maxillary deciduous canine (Kebara).

Plate 3. Occlusal aspect of Neanderthal deciduous maxillary first molar (Kebara). 402

Tab

le

1 M

esio

dls

tal

mea

sure

men

ts

of d

ecid

uou

s te

eth

in

Nea

r E

aste

rn

grou

ps

Gro

up

N

o.

di 1

di

s d

c d

m 1

X

Max

. M

in.

No.

X

M

ax.

Min

. N

o.

X

Max

. M

in.

No.

X

M

ax.

Min

. N

o.

dlT

l ~

X

Max

. M

in.

(a)

Max

illa

Wfi

rm I

-II

8 7.

9 8-

1-7"

1 6

6.3

6.9

-5.7

6

7.5

8.5

-7.0

9

7"9

8.5

-7.4

8

9.4

Ep

ipal

eoli

thic

2

7.6

7.8

-7.4

2

5-7

6.1

-5.6

3

7.4

7-6-

7.3

8 7.

7 8

.6-7

.5

10

9.5

Neo

lith

ic

5 6.

9 7-

2-6-

7 4

5-3

5"6-

4-9

7 7"

3 7-

1-6.

3 7

7"2

8.5

-6.9

8

9.3

Ch

alco

lith

ic

14

6.6

7.3-

6.1

10

5.4

5.8-

5.1

17

7'0

7.5

-6.5

8

6.9

7.2-

6"6

11

9.0

M.

Bro

nze

II

..

..

..

..

.

17

7.1

7.9

-6.5

17

8"

8 Ir

on

I

19

6.5

7.0

-5.9

8

4-7

4.9

-4.6

23

7'

0 7

.5-6

.2

33

7"2

8.1

-6.6

34

9.

0 R

om

an

.

..

..

..

..

9

7.0

7.6

-6,6

7

8.9

Rec

ent

11

6.3

6.7

-5.8

13

5"

1

5'6

-4'6

9

6"7

7-5-

6.1

19

7'2

8.8

-6,2

i

I 9

'2

(b)

Man

dib

le

Wti

rm I

-II

5 4.

6 5-

1-4.

2 6

5.1

5'5

-4'6

7

6'7

7

.4-5

-4

7 8.

9 9

'2-8

-7

8 10

.6

Ep

ipal

eoli

thic

3

4.6

4-7

-4.5

3

5"2

5.5

-4.9

7

6"6

6-8

-5.8

15

8.

6 9.

5-7"

5 17

10

.4

Neo

lith

ic

6 4,

5 4

.8-4

.2

6 4.

9 5

.2-4

.9

5 6'

3 6

.4-6

-0

7 8.

6 8

.8-8

,0

8 10

.5

Ch

alco

lith

ic

4 4.

0 4

'2-3

-7

9 4.

9 5

.5-4

.6

3 6'

1 6.

2-5"

6 12

7.

9 8"

3-7.

5 13

9.

9 M

. B

ron

ze I

I .

..

..

..

..

17

8"

2 8"

6-7"

3 19

10

'0

Iro

n I

19

4

'3

4"7-

3-7

25

5-0

5"5

-4'9

23

5"

9 6-

7-5"

5 33

8"

1 8"

3-7"

4 38

10

'0

Ro

ma

n

..

..

..

..

.

12

8.1

8.5

-7.4

11

9

'9

Rec

ent

7 4.

0 4

.0-3

.8

4 4.

6 4

.8-4

.4

8 5.

8 6.

1-5.

1 12

8.

1 9

.0-7

.5

12

9.8

10.0

-8.9

10

-7-8

.1

10.5

-9.1

9

.6-8

.3

10.3

-7.9

10

.1-7

-4

9.3

-8.4

10

.5-8

.2

11.3

-10.

0 1

0.7

- 8"

8 1

0'9

-10

'0

10

.6-

9"0

10"5

- 9

'0

10

.3-

9'0

1

0.2

- 9-

2 1

0.4

- 8"

0

Spec

imen

s exa

min

ed

1.

Mid

dle

Pal

eoli

thic

K

ebar

a, A

mu

d,

Tab

un

, S

kh

ul,

Qu

afze

h,

Sh

anid

ar

2.

Ep

ipal

eoli

thie

H

ayo

nim

, E

1 W

ad,

Keb

ara,

Ey

nan

, N

ahal

O

ren

3.

N

eoli

thlc

Je

rich

o,

Ab

u G

ho

sh

4.

Ch

alco

lith

lc

Azo

r 5.

B

ron

ze A

ge

Taa

nac

h

6.

Iro

n A

ge

Jeru

sale

m

7.

Ro

ma

n

Jeru

sale

m

8.

By

zan

tin

e K

hir

bet

Sh

ema

9.

Rec

ent

Bee

r S

heb

a (B

edou

in)

Th

e m

easu

rem

ents

for

Sh

anid

ar a

nd

Am

ud

wer

e ta

ken

fro

m ~

enyf

irek

(1

959)

an

d S

uzu

ki

& T

akal

(19

70).

Ta

ble

2

Bu

cco

lln

gu

al

mea

sure

men

ts

of

dec

idu

ou

s te

eth

in

N

ear

Ea

ster

n

gro

up

s

Gro

up

N

o.

di 1

di

2

dc

dm

1

dm2

X

Max

. M

ia.

No

. X

M

ax.

Min

. N

o.

X

Max

. M

in.

No

. X

M

ax.

Min

. N

o.

X

Max

. M

in.

Wii

rm I

-11

8

5"8

Ep

ipal

eoli

thic

2

5.8

Neo

lith

ic

5 5-

2 C

hal

coli

thlc

14

5"

1 M

. B

ron

ze I

I --

--

Ir

on

I

19

5"2

Ro

ma

n

--

--

Rec

ent

11

5.1

6.2

-5.8

6

5.7

6.0

-5.2

6

.0-5

-7

3 5.

7 5

-9-5

.5

5-4

-3.7

5

4.8

5.2

-4.4

5

.64

.7

10

4.8

5.2

-4.0

5.3

-4.5

8

4.5

4.6

-4.2

5.4

-4.8

13

4.

7 5

.3-4

.2

Wa

rm I

-II

5 4-

6 4-

8-4"

2 6

5.0

5-1-

4"8

Ep

ipal

eoli

thic

3

4.1

4.1

-3-8

3

4.8

4.9

-4.6

N

eoli

thic

6

3"9

4.3

-3 '7

6

4"5

5.0

-4.0

C

hal

coli

thic

4

3.8

4-0

-3.6

9

4.3

4.8

-4.0

M

. B

ron

ze I

I .

..

..

.

Iro

n I

18

4.

1 4"

7-3"

7 25

4

'7

5"2-

4'2

Ro

ma

n

..

..

..

R

ecen

t 7

3.7

4-0-

3"2

4 4.

3 4

.4-4

.1

(a)

Max

illa

5 7.

1 7

.5-6

.6

8 9"

2 1

0.2

-8.7

9

10.4

10

.8-9

"8

4 7.

4 7

.5-6

-9

8 9.

5 10

.6-9

.0

9 10

.5

11.0

-9-7

9

6.5

7.0-

5"7

7 9"

0 9

.5-8

.0

9 10

'2

11.4

-9"6

17

6"

2 6

.9-5

.6

8 8"

5 9

'0-7

"9

11

10.0

10

.5-9

"5

--

--

--

14

8.6

9.6

-7.3

10

9.

9 10

.4-9

.0

23

7.0

7.3

-6'8

33

8.

6 9

.5-6

-6

34

10"0

11

.0-9

.0

--

--

--

9 8"

3 9

.3-7

.6

7 9"

7 11

"1-8

"3

9 6.

5 6

.9-5

.3

19

8.4

9-1

-6.2

11

9-

5 10

.1-8

:0

(b)

Man

dib

le

5 6.

0 6

.5-5

.7

7 7.

4 7-

6-6"

9 8

9.4

10-0

-9"2

5

6"3

5.9-

5"8

15

7.4

8.6

-6.5

17

9.

3 10

.5-8

.1

5 5.

9 6

.2-5

.5

7 7"

2 7.

5-6"

9 8

9"2

9.6

-8.6

3

5.5

5.7

-5.4

12

6.

7 7

.3-6

.4

13

8.5

9.1

-7-9

--

--

--

17

7"

0 7.

8-6"

1 19

8"

8 10

.5-8

-3

23

5"5

6.1-

5"1

33

7-1

8"0

-6'5

38

8-

7 9

"5-7

'9

8 5"

5 6"

1-5"

2 11

6"

9 7"

5-6-

0 11

8"

5 9

.3-7

'7

8 5"

5 6"

1-5"

2 12

6"

8 7"

7-6"

3 12

8"

6 9-

3-8"

0

Spec

imen

s exa

min

ed

1.

Mid

dle

Pal

eoli

thic

K

ebar

a, A

mu

d,

Ta

bu

rn,

Sk

hu

l, Q

uaf

zeh

, S

han

idar

2.

E

pip

aleo

lith

ic

Hay

on

im,

E1

Wad

, K

ebar

a, E

yn

an,

Nah

al

Ore

n

3.

Neo

lith

ic

Jeri

cho

, A

bu

Gh

osh

4.

C

hal

coli

thic

A

zor

5.

Bro

nze

Ag

e T

aa

na

ch

6.

Ir

on

Ag

e Je

rusa

lem

7.

R

om

an

Je

rusa

lem

8.

B

yza

nti

ne

Kh

lrb

et S

hem

a 9.

R

ecen

t B

eer

Sh

eba

(Bed

ou

in)

Th

e m

easu

rem

ents

for

Sh

anid

ar a

nd

Am

ud

wer

e ta

ken

fro

m S

eny

fire

k

(195

9)

and

Su

zuk

i &

Tak

ai

(197

0).

Tab

le

3 M

eslo

dls

tal

mea

sure

men

ts

of

dee

ldu

ous

teet

h

(Eu

rop

e)

dP

d

i 2

dc

dm

1

dm

2

No

. X

S.

D.

No

. X

S.

D.

No

. X

S.

D.

No

. X

S.

D.

No

. X

S.

D.

Ara

go

K

rap

ina

W

arm

I-

II

Up

pe

r P

aleo

lith

ic

Mes

oli

thic

<

N

eo

lith

ic

Ro

ma

n

Re

ce

nt

Ara

go

K

rap

ina

r

Wfi

rm

I-II

U

pp

er

Pal

eoli

thic

Z

M

eso

lith

ic

Ne

oli

thic

R

om

an

R

ec

en

t

1 8

.40

.

..

.

1 8

.80

--

4

8.3

5

--

1 7

.06

--

5

7.6

6

--

3 7

.10

0

.5

2 6

.0

0.3

5

7-0

4

0.6

3

7.6

0

.3

3 5

.6

0.2

2

7.4

0

.3

1 7

.50

--

2

5.3

5

0.5

5

7.1

8

0.2

5

2

.66

6

--

41

5

.42

--

9

2

6.8

0

--

15

6.2

5

--

22

5

.17

--

3

5

6.8

7

--

61

6.1

0

--

60

4

-90

--

6

3

6.6

7

--

--

--

--

1 5

.70

--

1

7.5

0

--

--

--

--

1 5

.80

.

..

.

5 4

.64

--

6

5.0

5

0.3

7

6.7

4

0.3

1

4.0

--

1

4.8

0

--

--

6.8

0

--

4 4

.25

0

.4

6 5

.15

0

.3

8 6

.46

0

.6

35

4

.07

--

4

2

4.9

1

--

62

5

.80

--

18

4

.01

--

17

4

-63

--

3

6

5.9

4

--

60

4

.10

--

6

2

4.4

0

--

71

5.5

0

--

2 9

.05

.

..

.

4 8

.25

--

9

10

.10

--

4

8.1

3

1 "2

4

9.3

3

0.2

10

7

.3

0.5

5

9.6

0

0.8

7

7.3

3

0.3

7

9.2

9

0.3

10

7

.47

--

6

6

9.0

1

--

31

7.1

7

--

46

9

.16

--

7

3

6.7

7

--

87

8

.97

--

1 1

0.2

0

--

2 1

0.7

5

--

2 9

.60

--

7

10

.88

--

7

8.8

9

0.2

8

10

.58

0

.5

--

9.0

3

--

--

10

-90

--

9

8.4

0

.7

9 1

0.2

9

0.5

7

9

8.5

3

--

10

3

10

.17

--

4

5

7-7

9

--

50

9

.88

--

9

0

7,7

8

--

88

9

.57

--

1.

Lo

we

r P

ale

oli

thic

2

. M

idd

le

Pa

leo

lith

ic

3.

Up

pe

r P

aleo

lith

ic

4.

Me

soli

thlc

--M

ug

e

5.

Ne

oli

thic

6

. Ir

on

Ag

e

Ro

ma

n

Re

ce

nt

Ara

go

(a

) K

rap

ina

(b

) P

ec

h

de

l'A

ze

, G

ibra

lte

r,

Ch

ate

au

ne

uf,

C

om

be

G

ren

al,

En

gis

, A

rch

i, S

ub

aly

uk

F

on

tec

he

va

de

, Is

turi

tz,

La

Ch

aise

, L

e P

ey

rat,

L

e R

ois

, B

run

iqu

el.

Th

e

me

asu

rem

en

ts

for

Arc

hi,

S

ub

aly

uk

, a

nd

N

eo

lith

ic

to

Re

ce

nt

Eu

rop

ea

n

sam

ple

s a

re

ba

sed

o

n

Asc

en

zi

&

Se

gre

(1

97

1),

T

ho

ma

(1

96

3),

an

d B

rab

an

t (1

97

4).

T

he

m

ea

sure

me

nts

fo

r M

ug

e

we

re m

ad

e

av

ail

ab

le

by

D

. F

rey

er.

A

ll

oth

er

spe

cim

en

s w

ere

me

asu

red

b

y t

he

a

uth

or.

Tab

le

4 B

ucc

olin

gual

m

easu

rem

ents

of

d

ecid

uou

s te

eth

(E

uro

pe)

di 1

d

i 2

dc

No

. X

S.

D.

No

. X

S.

D.

No

. X

S,

D.

dm

1

dm

2

No

. X

s.

D.

No

. X

S.

D.

Ara

go

K

rap

ina

Wii

rm

I-II

U

pp

er

Pal

eoli

thic

M

eso

lith

ic

N

Neo

lith

ic

Ro

man

R

ecen

t

Ara

go

K

rap

ina

Wfi

rm

I-II

2;

U

pp

er

Pal

eoli

thic

<

Mes

oli

thic

N

N

eoli

thic

R

om

an

Rec

ent

1 6"

00

--

1 --

--

--

7"

40

--

4 6"

75

--

2 6

'10

--

5

6"48

--

3

5"67

0-

4 3

5"10

0"

1 5

6"46

0"

7 3

5-3

0-3

2 5"

1 0"

3 2

6"7

0

0'2

1

5'3

0

--

41

4"9

0

0"1

5 6

'26

0"

3 52

5

"24

--

22

4

"99

--

92

6

'27

--

15

5"

09

--

60

4

-86

--

35

6

'05

--

61

4

"98

--

--

4

"49

--

6

3

6"0

4

--

..

..

5

"00

--

--

6

-80

--

.

..

.

4.8

0

..

..

5

4"5

6

0.3

6 4

.96

O

.1

5 6

-04

O

" 1

..

..

..

1

6.2

0

--

4 3

.98

0-

1 6

4.5

7

0.2

8 5

.78

0

.4

35

3.9

8

--

42

4-4

6

--

68

5

-86

--

18

3

.75

--

17

4

-27

--

3

6

5.3

6

--

60

3

.90

--

62

4

.00

--

71

5

.10

--

--

10"0

1 4

8"95

5

9"13

9

8"7

7 8"

57

70

8"91

31

8"

56

73

8"5

8

2 8

"10

7

7"39

3

7"7O

9

7"20

79

7"

13

45

7"

05

90

6"

91

0"3

0-6

0"4

--

10.0

--

9 10

-54

--

4 10

.3

0.7

10

10

.5

0.5

7

10.2

1 0-

4 6

6

10-1

8 --

4

6

9.91

--

8

7

9.7

7

--

--

2 9

-55

--

--

7

9.6

0

--

0"1

8 9.

41

0-3

0.4

6

9.1

7

0"2

0'3

9

9.21

0

.3

--

103

9.1

6

--

--

50

8

.88

--

--

8

8

8.7

0

--

1.

Lo

wer

P

aleo

lith

ic

2.

Mid

dle

P

aleo

lith

ic

3.

Up

per

P

aleo

lith

ic

4.

Mes

oli

thic

--M

ug

e 5.

N

eoli

thic

6.

Ir

on

A

ge

Ro

man

R

ecen

t

Ara

go

(a

) K

rap

ina

(b)

Pec

h

de

l'A

ze,

Gib

ralt

er,

Ch

atea

un

euf,

C

om

be

Gre

nal

, E

ng

is,

Arc

hi,

S

ub

aly

uk

F

on

tech

evad

e,

Istu

ritz

, L

a C

hai

se,

Le

Pey

rat,

L

e R

ois

, B

run

lqu

el.

Th

e m

easu

rem

ents

fo

r A

rch

i,

Su

bal

yu

k,

and

N

eoli

thic

to

R

ecen

t E

uro

pea

n

sam

ple

s ar

e b

ased

o

n

Asc

enzi

&

S

egre

(1

97

1),

T

ho

ma

(19

63

), a

nd

B

rab

ant

(19

74

).

Th

e m

easu

rem

ents

fo

r M

ug

e w

ere

mad

e av

aila

ble

b

y

D.

Fre

yer

. A

ll

oth

er

spec

imen

s w

ere

mea

sure

d

by

th

e au

tho

r.

E V O L U T I O N OF T H E DECIDUOUS D E N T I T I O N 407

more modern specimens such as Skhul and Quafzeh it is usually missing, while the paracone and metaeone are fused at least partially, so that the tooth is bicuspid.

The lower first molars are four or five cusped with the trigonid and talonid portions fairly equal and the occlusal surface rectangular. Both upper and lower second molars are somewhat elongated mesiodistally with well defined anterior fovea and a high frequency of accessory cuspules; Carabelli 's cusp is c o m m o n . . , in upper molars as are sixth and seventh cusps in the lowers.

In the recent specimens, teeth are more slender and less convex, with a marked slope present only on the buccal surface of the first molars, where it is associated with a tuber- culum molare. The frequency and intensity of expression of shovelling and accessory cuspules is low and shows considerable regional variability. Upper first molars are usually bicuspid and lower first molars may have 3-5 cusps, the mesiolingual cusp is often high and the occlusal surface triangular rather than rectangular.

4. D i s c u s s i o n

The data presented here suggests that there was considerable reduction in tooth size between the Anteneanderthals and Neanderthals followed by a period of little change in Upper Paleolithic Homo sapiens. Reduction in deciduous tooth size then accelerated between the Epipaleolithic and the Chalcolithic periods in the Near East and since then tooth size has remained fairly stable.

The morphological differences found are those associated with reduction, such as de- creased frequency of shovelling and accessory cuspules. The changes found in the incisors, canines and second molars are similar to those found in the permanent dentition.

The first molar differs from other deciduous teeth in that it resembles no tooth in the permanent dentition. The bicuspid form of the upper molar typical of recent populations was recognized by Jorgensen (1956) as resulting from reduction of a more fully molarized, four cusped ancestral form. However in his study of the lower first molar, he preferred to consider the reduced triangular form as the ancestral type. This, as outlined earlier, was based on comparison of this tooth with the homologous tooth in Anthropoidea. Neither he nor Koenigswald (1967), were prepared to include the Australopithicinae, with their heavily molarized first molars, in the hominid lineage: indeed, for Koenigswald this feature provided additional grounds for excluding them from the Hominidae.

The results of this study do not support their conclusions. Both upper and lower first molars in the Lower and Middle Paleolithic specimens examined were more fully molarized than those of recent populations and it is probable that the tooth was even more heavily molarized in the larger toothed earlier hominids. In Middle Paleolithic hominids, reduction of this tooth was well under way, and in more advanced specimens from Skhul and Quafzeh the tooth was essentially of modern form, although large in size. The changes found in the first deciduous molar are reminiscent of those found in the lower second permanent molar, in which cusp number changed from five to four cusps in Upper Paleolithic Homo sapiens, but was not invariably associated with a reduction in size (Brabant, 1969).

5. C o n c l u s i o n s

Comparison of evolutionary trends in the deciduous dentition of hominids in Europe with those in Israel show considerable similarities, both regarding tooth size and morphology in early specimens, and the timing and rate of change in both areas.

408 P. SMITH

A reduction in tooth size was noted between Lower and Middle Paleolithic specimens but the rate of reduction appears to have accelerated between the Epipaleolithic and Chalcolithic periods with teeth reaching modern dimensions at this time. Throughout, anterior teeth show more reduction than the posterior teeth, the second deciduous molar showing little change in size. The morphology of the anterior teeth and first molar has changed most and may be of value as diagnostic criteria for differentiating between fossil hominids and Homo sapiens.

R e f e r e n c e s

Ascenzi, A. & Segre, A. G. (197I). A new Neandertal child mandible from an Upper Pleistocene site in Southern Italy. Nature 233, 280-283.

Brabant, H. (1965). Observations sur l '6volution de la denture temporaire humaine en Europe Occident- ale. Bull. Group. Int. Reeh. So. Stomat. 8~ 235-302.

Brabant, H. (1969). La denture humaine au Paldolithique superieure d 'Europe C.N.R.S. In (G. Camps & G. Oliver, Eds) L'Homme de Cro-Magnon Paris: C.N.R.S.

Brabant, H. (1974). Etude odontologique des restes humaines decouverte dans la n6cropole Gallo- Romaine et merovingienne de Diene (Meuse) en France. Bull. Group Int. Rech. Se. Stomat. 6, 239-261.

Gregory, W. K. (1922). The Origin and Evolution of the Human Dentition. Baltimore: Maryland- Baltimore Press.

Hanihara, K. (1963). Crown characters of the deciduous dentition of the Japanese American hybrids. In (D. R. Brothwell, Ed.), DentalAnthropology, pp. 105-124. New York: Pergamon Press.

Hanihara, K. (1967). Racial characteristics in the dentition. Journal of Dental Research 46, 923-926. Jflrgensen, K. D. (1956). The deciduous dentition. Acta odontologia scandinavica 14, Supplement 20. Koenigswald, G. H. R. von (1967). Evolutionary trends in the deciduous molars of the Hominidea.

Journal of Dental Research 46, 779-786. Legoux, P. (1970). Etude odontologique de l 'enfant Ndandertalien du Pech-de l'Az6. Archive de

l'Institut de Palgontologie humaine 33, 53-87. Remane, A. (1921). Beitr~ige zur Morphogie des Anthropoidengebisses. Wiegmann Archly f l i t Natur-

geschiohte 87, 1-179. Quoted inJorgensen (1956), pp. 177-178. Remane, A. (1927). Studlen uber die Phylogenie des Menschichen Eckzahns. Quoted in Jorgensen

(1956), p. 94. Robinson, J, T. (1956). The dentition of the Australopithicinae. Memorandum 9, Transvaal Museum,

Pretoria. Senyfirek, M. (1959). A study of the deciduous teeth of the fossidal Shan~da Infant. A comparative study

of the milk teeth of fossil men. Faculty of Languages, History and Geography, University of Ankara, No. 128.

Suzuki, H. & Takai, R. (Eds) (1970). The Amud Man and his cave site. Tokyo : University of Tokyo. Smith, P. & Arensburg, A. 1977. New discoveries in Mt. Carmel: The Mousterlan Infant from Kebara

In (Eretz-Israel, vol XI I I ) Israel: Exploration Society, Jerusalem pp. 64-76. Thoma, A. (1963). The dentition of the Subalyuk Neanderthal child. Z. Morph. Anthrop. 54, 127-150. Weidenreich, F. (1937). The dentition of Sianthropus Pekinensis. Palaeontologica Siniea N.S.D. no. 1.

Whole Series no. 101.