Evolutionary changes in the deciduous dentition of near eastern Eastern populations
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Transcript of Evolutionary changes in the deciduous dentition of near eastern Eastern populations
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.