THE ANATOMICAL RECORD (NEW ANAT.

) 269:148 –156, 2002

ARTICLE

A Closer Look at Neanderthal Postcanine Dental

Morphology: The Mandibular Dentition
SHARA E. BAILEY*

Neanderthals are known to exhibit unique incisor morphology as well as enlarged pulp chambers in postcanine teeth
(taurodontism). Recent studies suggest that their overall dental pattern (i.e., in morphologic trait frequencies) is also
unique. However, what this means in a phylogenetic sense is not known. Although exploring the polarity of dental
morphologic characters is essential to understanding the phylogenetic implications of unique patterns of variation,
few have undertaken this task. This study moves beyond standard scoring methods, which are based on modern
humans, to include several postcanine traits that have not been considered previously. In addition, Homo erectus is
used as an outgroup to Neanderthals and modern humans to explore the polarity of these traits. The findings of this
study suggest that Neanderthals are not only unique in their pattern of dental trait frequencies (as found in previous
studies) but that they present several dental autapomorphies, as well. These include a high frequency of the
mid-trigonid crest in lower molars and unique morphology of the lower premolars. Interestingly, these characters are
not observed in the Mauer mandible, which some have claimed to be a member of a chronospecies that is a unique
ancestor to Neanderthals. Anat Rec (New Anat) 269:148 –156, 2002. © 2002 Wiley-Liss, Inc.

KEY WORDS: comparative anatomy; dental morphology; human evolution; hominoid postcanine teeth; Neanderthals; Homo
sapiens; Homo heidelbergensis; Homo erectus

INTRODUCTION
Neanderthals are a group of extinct
humans that inhabited Europe and
Western Asia between approximately
200,000 and 30,000 years ago. They
are also the best-known and most ex-
tensively studied extinct human
group. Despite the totality of morpho-
logic and cultural evidence in support
of the theory that Neanderthals are a
distinct group of archaic humans
(Tattersall and Schwartz, 2000), pa-
leoanthropologists continue to debate
the biological specializations that
help define their clade. This debate
Ms. Bailey is a paleoanthropologist
completing her doctorate at Arizona
State University. She specializes in den-
tal anthropology and is in the process of
developing new standards for scoring
variation in fossil hominids. Currently
her interest lies in using dental morphol-
ogy to investigate evolutionary patterns
in the fossil record and to address is-
sues regarding the origin of modern hu-
mans.
*Correspondence to: Shara E. Bailey,
American Museum of Natural History,
Division of Anthropology, Central Park
West at 79th Street, New York, NY
10024-5192. Fax: 212-769-5334; E-mail:
shara.bailey@asu.edu
DOI 10.1002/ar.10073
Published online in Wiley InterScience
(www.interscience.wiley.com).
concerns the origins of modern hu-
mans and the nature and degree to
which Neanderthals participated in
our evolution (Box 1).
Traditionally, researchers have fo-
cused on cranial (and to a lesser ex-
tent postcranial) features to address
the question of evolutionary relation-
ships among Neanderthals and mod-
ern humans. However, this kind of re-
search has been confounded by a lack
of agreement regarding how morpho-
logic characters should be inter-
preted, that is, whether they are prim-
itive or derived (in a cladistic sense).
This disagreement may be due, at
least in part, to the nature of working
with skeletal features that can be af-
fected more by a variety of external
influences such as diet and behavior.
Teeth are good anatomical candidates
for making phylogenetic inferences
for several reasons, not the least of
which is because they are the most
abundantly preserved elements in the
fossil record. In addition, teeth con-
tain a large array of morphologic in-
formation that is controlled to a
greater extent by genes than are skel-
etal features. Once formed, crown
morphology does not change, except
through tooth wear. Dental morpho-
logic studies have shown that dif-
ferent geographic populations— e.g.,
Asia, Europe, Africa, and India—
can be characterized by regional den-
tal morphologic patterns (Hanihara,
1969; Mayhall et al., 1982; Turner,
1990a; Irish, 1994; Hawkey, 1998).
Studies also show that variation in
dental morphologic characters can be
successfully used to work out biologi-
cal affinities among human popula-
tions (Sofaer et al., 1986; Hanihara,
1989; Irish and Turner, 1990; Turner,
1990b Lipschultz, 1997; Bailey et al.,
1998; Hawkey, 1998). Because teeth
preserve extremely well over long pe-
riods of time, the same approach can
be easily applied to both present and
past populations.
NEANDERTHAL DENTAL
MORPHOLOGY
Neanderthals are known to have had
large anterior teeth marked by strong
shoveling, marked labial convexity,
and prominent lingual tubercles, as
well as postcanine teeth with enlarged
pulp chambers (taurodontism) (Fig-
ure 1). Beyond this, it is generally as-
sumed that Neanderthal postcanine
tooth morphology is just like that in
modern humans (e.g., Smith, 1976).

© 2002 Wiley-Liss, Inc.

ARTICLE THE ANATOMICAL RECORD (NEW ANAT.) 149

BOX 1. THE MODERN HUMAN ORIGINS DEBATE

For the past two decades Neanderthals have been in-
terpreted in terms of one of two competing models for
modern human origins (see Figure). Supporters of the Re-
cent African Origin model (a) posit that Neanderthals were
replaced by emigrating African modern humans and con-
tributed little or nothing to the modern human gene pool
(e.g., Stringer, 1992; Stringer and Bräuer, 1994). Support-
ers of Multiregional Evolution (b) believe Neanderthals may
have played a significant role in modern human origins
(even if they stress that the role of Neanderthals is not
central to their model) (Frayer et al., 1993; Wolpoff et al.,
1984). Where proponents of these models disagree most is
in deciding whether or not Neanderthals were too different
from modern humans to have played a significant role in
their evolution. Intermediate models have been proposed
but prove difficult to test with the fossil record. Note:
dotted lines indicate gene flow between ancient popula-
tions.

Figure. Two competing models for modern human origins: Recent African origin model (a) and multiregional evolution (b).
(Modified from Stringer and Gamble, 1993).

However, recent studies indicate that
this may not be the case.
The Arizona State University Dental
Anthropology System, or ASUDAS, is
a standardized system for recording
variation in the tooth crowns and
roots of modern humans (Box 2). Sev-
eral studies within the past 5 years
have shown that the system can be
applied effectively to fossil hominids
as well (Stringer et al., 1997; Irish,
1998; Tyrell and Chamberlain, 1998;
Bailey and Turner, 1999; Bailey,
2000b; Coppa et al., 2001). Results
from these studies suggest that the
differences between Neanderthals and
modern humans go beyond those ob-
served on the incisors and postcanine
roots (at least in trait frequencies).
For example, when the frequencies of
10 morphologic traits in Neanderthals
were compared with ranges observed
in large groups of geographically di-
verse contemporary modern humans,
the frequencies for five traits fell
within the modern human range,
whereas the frequencies for the other
five fell outside the modern human
range (Figure 2) (Bailey, 2000a).
Based on all the dental evidence that
has been analyzed thus far, the Nean-
derthal dental pattern can be de-
scribed in terms of low and high fre-
quency ASUDAS traits (Table 1).
It is now clear that relative to mod-
ern humans, Neanderthals exhibit a
unique dental morphologic pattern.
Teeth contain a large
array of morphologic
information that is
controlled to a greater
extent by genes than
are skeletal features.
However, to date, no study has sys-
tematically explored what this
uniqueness means in an evolutionary
sense. We do not know, for example,
whether or not the Neanderthal dental
pattern is a primitive one or even
whether or not there are yet-to-be-de-
scribed dental morphologic charac-
ters that are uniquely derived in (au-
tapomorphic for) Neanderthals. This
is likely due to the fact that ASUDAS is
based on modern human variation
and was developed for the purpose of
deciphering population relationships,
not evolutionary ones. Therefore,
when applied to fossil hominids, this
system is biased in that only those
characters in fossil hominids that are
present and variable in modern hu-
mans are evaluated. Characters that
are present but invariable or absent in
modern humans are not included in
the system, as they would not be use-
ful for the purpose for which it was
designed.
On the basis of descriptive studies
of Mid to Late Pleistocene hominid
teeth (Patte, 1959; Genet-Varcin, 1962,
1966, 1972; Smith, 1976; Trinkaus,
1978; Tillier, 1979, 1991; Tillier et al.,
1989), there is reason to believe that
the Neanderthal dentition differs
from that of modern humans in more
than trait frequencies and includes
dental traits that are not currently
part of the ASUDAS. Unfortunately,
characterization and quantification of
150 THE ANATOMICAL RECORD (NEW ANAT.)
Figure 1. Marked shoveling (1), labial convexity (2), and lingual tubercles (3) on maxillary
incisors (a) and taurodontism (arrow) of mandibular molars (b) have been the primary focus
of dental morphologic research on Neanderthals. (Photo (a) by E. Trinkaus.)
these traits and their variation have
generally been lacking.
One of the goals of my research is to
correct this problem, in part, by devel-
oping a supplemental scoring system
for fossil hominids. The purpose of
this particular study is to describe and
define the variation in some mandib-
ular postcanine tooth crown traits
that are not part of the ASUDAS and
to draw attention to the fact that these
variations (and others) appear to have
phylogenetic significance. Of primary
concern is determining whether sig-
nificant differences exist between Ne-
anderthals and modern humans. The
results presented here are part of an
ongoing study of Neanderthal postca-
nine dentition. In the interest of
space, only the mandibular dentition
is discussed; the maxillary dentition
will be discussed in a subsequent
study.
THE STUDY
The Neanderthals used in this study
include specimens from both Europe
and Western Asia. Previous studies
have shown that, although some geo-
graphic variation exists between these
samples, they always form a group
separate from modern humans
(Bailey and Turner, 1999; Bailey,
2000a). The modern human sample
includes early modern humans, Upper
Paleolithic modern humans, and con-
temporary modern humans from six
geographic regions (see Table 2). To
ascertain the polarity of these traits, a
small outgroup sample (Homo erec-
tus) is included in the analyses. Fi-
nally, comparisons are made with a
ARTICLE
and Turner, 1988). This strategy re-
sults in smaller sample sizes but
avoids biasing the results toward indi-
viduals with both antimeres present.
For the most part, specimens were in-
cluded if they exhibited only minimal
to moderate wear (fissure pattern vis-
ible and cusp tips only moderately
worn). However, for some traits (e.g.,
P4 transverse crest, see below), assess-
ment of trait’s presence could be as-
certained even when the tooth was
quite worn.
The premolar traits used in the
analysis include lingual cusp number,
transverse crest, lingual crown con-
tour, and metaconid position (Figure
small sample of teeth from Arago and 3). Of these, only lingual cusp number
Mauer, which are European Middle is part of the ASUDAS. The other
Pleistocene hominids sometimes at- traits were chosen because of their ap-
tributed to the species Homo heidel- parent low variation of expression in
bergensis. The samples in which the modern human populations (Kraus
postcanine teeth could be adequately and Furr, 1953; Ludwig, 1957; Bigger-
scored are presented in Table 2. The staff, 1969) and because previous re-
individual count method used in this search suggested to me that they may
study involves scoring both right and have evolutionary (phylogenetic) sig-
left sides of the dentition but only by nificance. No reference plaques have
using the antimere with the highest yet been developed to represent ex-
trait expression in the analysis (Scott pression of these new traits.
BOX 2. THE ARIZONA STATE UNIVERSITY DENTAL
ANTHROPOLOGY SYSTEM
The Arizona State University Dental Anthropology System (ASUDAS) is
becoming the worldwide standard for morphologic study. It currently consists
of more than 28 morphologic traits of proven utility for population compari-
sons. Traits are scored by using a combination of plaques (see figure) and
written descriptions of the variation (Turner et al., 1991). Although many more
traits are present on teeth than are included in the ASUDAS (see Morris, 1965),
those that are used were chosen because they are relatively easy to score,
remain even after moderate wear, and have proven useful in characterizing
intra- and interpopulation variability and relationships.
Figure. A sample scoring plaque from the ASUDAS (canine distal acessory ridge).
ARTICLE THE ANATOMICAL RECORD (NEW ANAT.) 151

Figure 2. Bar graphs illustrating how frequencies of Arizona State University Dental Anthropology System traits found in Neanderthals
compare with those observed in recent human populations. Of the 10 traits for which comparisons were possible, 5 fall within the observed
ranges of variation in recent humans (a) and 5 fall outside the observed ranges of variation in recent humans (b). Groups exhibiting high
frequencies of traits are shown in black and those exhibiting low frequencies are shown in grey. Note that the Neanderthal dental pattern
does not correspond to that of any living modern humans. SS, sub-Saharan; W, western, N, north. All modern human data are taken from
Scott and Turner (1997).

The mid-trigonid crest (MTC) is the
only mandibular molar trait reported
in this study. The MTC is an enamel
bridge that joins the protoconid and
metaconid (Figure 4). This trait was
chosen because of its apparent low
frequency in modern humans (Wu
and Turner, 1993) and high frequency
in Neanderthals (Zubov, 1992). It is
currently part of the ASUDAS, but the
scoring procedure used here differs
from the one described in Turner et al.
(1991). Table 3 presents the trait de-
scriptions, scoring procedures, and
the presence/absence breakpoints used
in this analysis.
NEANDERTHAL POSTCANINE
DENTAL MORPHOLOGY
After examining 26 Neanderthals P4s, a
distinct configuration emerged— one
that is marked by a complex occlusal
surface topography and an asymmetri-
cal lingual contour. The tooth’s com-
plex topography results from a combi-
nation of a strong and continuous
transverse crest, a high and well-devel-
oped metaconid, and extra fissures,
ridges, and lingual cusps (Figure 5a).
The modern human sample con-
sisted of 125 P4s. In contrast to a typ-
ical Neanderthal P4, modern human
P4s, more often than not, exhibit a
greatly simplified occlusal morphol-
ogy. Although multiple lingual cusps
are not uncommon in modern popu-
lations, P4s of both contemporary and
fossil modern humans typically lack a
well-developed continuous transverse
crest, as well as other accessory ridges
and fissures. Moreover, modern hu-
man P4s almost invariably exhibit a
symmetrical lingual crown contour.
One of the most notable features of
the modern human P4s is the occur-
rence of a metaconid that is so re-
duced that it forms only a lingual shelf
or is lacking altogether (Figure 5d).
This condition was never observed in
the Neanderthal sample.
Figure 6 illustrates how the traits
mentioned above are distributed
among Neanderthals and modern
human populations. As suspected,
TABLE 1. High and low frequency dental morphological
traits* found in Neanderthals
High frequency traits
Incisor shoveling
Incisor labial convexity
Incisor lingual tubercles
Cusp 5 M1
Y-groove pattern M2
Canine mesial ridge
Carabelli’s cusp M1
Presence of M3
*Based on the Arizona State University Dental Anthropology System (Turner et al.,
1991).
Neanderthals exhibit the highest fre-
quencies for multiple lingual cusps
(93%), transverse crest (88%), and
asymmetrical lingual crown contour
(96%). Although multiple lingual
cusps are also found in moderate to
high frequencies in modern groups
(50 – 80%), the other two traits occur
in much lower frequencies (0 – 6% in
recent groups and 18% in the Upper
Paleolithic group). The presence of a
transverse crest is more frequent in
the early modern human sample
(50%) than it is in other modern hu-
man samples; however, a sample
size of two individuals requires cau-
tious interpretation. Frequencies for
metaconid placement are presented
in Figure 6 to illustrate that a mesi-
Low frequency traits
Incisor double shoveling
Four cusped M2
Three cusped M2
M1 enamel extension
152 THE ANATOMICAL RECORD (NEW ANAT.) ARTICLE

TABLE 2. List of samples and teeth used in this study

Neanderthals
Europe
P4 and M1or2 Pontnewydd, Hortus, Petit Puymoyen, Le
Moustier, Krapina
P4 only Monsempron
M1or2 only Montmaurin, Ehringsdorf, La Quina, La
Ferrassie, Gibraltar, Grotta Brueii
Western Asia
P4 only Kebara, Amud, Shanidar
M1or2 only Tabun
Early modern humans
P4 and M1or2 only Qafzeh, Skhul
Upper Paleolithic modern
humans
P4 and M1or2 Le Rois, St. Germaine-la-Rivière, Dolni
Figure 3. Premolar crown characters re-
Vestonice, Gough’s Cave, Oberkassel ferred to in the text: transverse crest (a),
Brno metaconid (b), lingual contour (c), and lin-
P4 only Fourneau du Diable gual cusps (d). The symbol indicates orien-
M1or2 only Fontéchevade, Isturits, Vachons, Abeilles, tation of the tooth: B, buccal or cheek side
La Ferrassie, Mieslingtal, Fontana of the tooth; L, lingual or tongue side; M,
Ranuccio, Madeleine, Greze mesial or toward the front of the mouth; and
Contemporary modern humans D, distal or toward the back of the mouth.
Africa North Africa
Europe British Neolithic, Hungary, Germany, Austria, anderthals show the highest frequen-
Yugoslavia, Bulgaria, Greece, Crete cies for this trait (90 –100%). Notable
North Asia Japan, China, Korea as well is that, in modern humans, the
West Asia Tel Hesi, Jericho MTC is even less frequently observed
India India
in M2 and M3, but in Neanderthals the
Australasia Australia, New Guinea
frequencies are relatively constant in
Homo erectus
Asia: all three molars.
P4 and M1or2 Zhoukoudian, Sangiran To determine whether these charac-
Africa: ters are primitive retentions or derived
P4 and M1or2 KNM-ER, KNM-WT, Olduvai, Tighenif features, high definition casts of 7 P4s, 9
M1or2 only Sidi Abderrahman M1s, and 7 M2s from East African and
Homo heidelbergensis Asian Homo erectus were examined.
P4 and M1or2 Arago, Mauer Superficially, Homo erectus P4s resem-
ble Neanderthal P4s in size and mor-
phologic complexity. However, they
ally placed metaconid (quite com- observation is not remarkable in and of
lack the unique Neanderthal morphol-
mon in modern humans) is not itself, as occlusally complex molars typ-
necessarily associated with an asym- ify most fossil hominids and are found
metrical tooth crown (rare in mod- with some frequency among modern
ern humans). humans as well. The most remarkable
Further analysis reveals that, al- feature observed on Neanderthal man-
though traits that contribute to the dibular molars is its wide and deep an-
unique Neanderthal P4 morphology terior fovea bordered mesially by a low
(such as a continuous transverse mesial margin and distally by a contin-
crest, a large metaconid, and an asym- uous MTC (Figure 8a). In contrast,
metrical lingual crown contour) may modern human mandibular molars
be present in modern populations, the typically lack a bridge between these
combination of all three traits in an cusps. Although the crests of C1 and C2
individual is not. In fact, only 2.4% of may be well developed, they generally
the modern humans exhibited even do not join to form a continuous ridge
two of the traits in combination. In (see Figure 8). In this case, the cusps are
marked contrast, 35% of Neander- separated by the sagittal sulcus, which
thals exhibited two traits and 59% ex- either terminates in the anterior fovea
hibited all three traits in combination (b) or runs the length of the tooth (c).
(Figure 7). Most often the crests are not very well
Figure 4. Molar crown characters referred
As is true of the P4s, Neanderthal developed.
to in the text: protoconid (a); metaconid
mandibular molars also tend to possess The frequencies for the MTC are (b); anterior fovea (c); mid-trigonid crest
extra occlusal fissures and crests. This given in Figure 9. It is clear that Ne- (d). Symbol orientations as in Figure 3.
ARTICLE THE ANATOMICAL RECORD (NEW ANAT.) 153

TABLE 3. Scoring procedure and presence/absence breakpoints for traits used in this study

Trait Definition and scoring procedure Presence/absent breakpoint

Transverse crest (P4)
Metaconid position (P4)
Lingual crown contour (P4)
Mid-trigonid crest (M123)
Presence of a crest or ridge connecting the Presence ⫽ Grades 2–3
buccal and lingual cusps
0 ⫽ absent; 1 ⫽ weak expression or
interrupted crest; 2 ⫽ moderate
expression; 3 ⫽ marked expression
Position of the metaconid relative to the Presence ⫽ mesial
position of the protoconid apex.
Scored as mesial, central or distal.
Shape of the lingual contour of the tooth Presence ⫽ asymmetrical
when viewed occlusally.
Scored a symmetrical or asymmetrical
An enamel ridge that connects the Presence ⫽ Grade 2
protoconid (Cusp 1) and metaconid
(Cusp 2).
0 ⫽ absent; 1 ⫽ crests are well developed
but do not form a bridge; 2 ⫽ crests form
a complete bridge

ogy described above. Instead, they ex- the sample size is small (3 P4; 3 M1; 5 characters, some general observations
hibit a symmetrical (almost square) lin- M2; and 2 M3) and this investigation is can be made. All P4s resemble those of
gual contour and the transverse crest is based on only a few morphologic Homo erectus and Neanderthals in
either completely lacking or is divided
by the longitudinal fissure (Figure 5b).
In this way, they resemble the modern
human P4 more than the Neanderthal
P4. The mandibular molars of Homo
erectus are morphologically complex,
and the crests of C1 and C2 are often
well developed. However, the MTC so
frequent in Neanderthals is rare or ab-
sent in this sample (Figure 8b): only 1 of
the 14 teeth examined exhibited a
bridge-like crest.
Table 4 summarizes the characters
and character states discussed above.
Although some variation in each group
exists, certain character states can be
said to typify a particular group. A com-
parison of these states among the sam-
ple groups indicates that (assuming
Homo erectus presents the primitive
conditions) the Neanderthal dental pat-
tern is the result of derived dental mor-
phology rather than primitive reten-
tions. It also indicates that, relative to
the derived conditions observed in Ne-
anderthals, modern humans retain the
primitive condition for several of these
morphologic features.
The samples Arago and Mauer were
also included in this study to investi-
gate whether dental morphology
would support the claim that they, as Figure 5. A comparison of mandibular second premolars. The Neanderthal P4 (a) presents
members of the species Homo heidel- an asymmetrical lingual contour, marked transverse crest, a mesially placed metaconid
bergensis, should be considered a and extra lingual cusps. The Homo erectus (b) and modern human P4s (c,d) all have
symmetrical lingual contours and lack a prominent transverse crest. The marked metaconid
chronospecies that gave rise exclu-
reduction and shelf-like morphology seen in d is common in recent modern human P4s. but
sively to Neanderthals (Rosas and not found in Neanderthals or in Homo erectus. a, Krapina; b, Zhoukoudian; c, Dolni
Bermúdez de Castro, 1998). Although Vestonice; d, recent Australian. Symbol orientation as in Figure 3.
154 THE ANATOMICAL RECORD (NEW ANAT.) ARTICLE

exhibits a strong MTC, as do two of

the four M2s (Figure 11b), whereas
the single scorable M3 does not ex-
hibit an MTC. In sum, certain teeth in
the Arago sample possess traits that
occur in uniquely high frequencies in
Neanderthals. This evidence could be
used as tentative support for the hy-
pothesis that they are members of a
group that gave rise exclusively to Ne-
anderthals (Rosas and Bermúdez de
Castro, 1998). It would be more diffi-
cult to make this case based on the
Mauer mandible alone, as it does not
show any particular affinity to Nean-
Figure 6. A comparison of P4 trait frequencies among samples. Neanderthals show the derthals based on its tooth crown
highest frequencies of multiple lingual cusps (MLC), presence of a well-developed contin- morphology. It is important to keep in
uous transverse crest (TRCRST), and lingual crown asymmetry (ASYM mesial). Lingual con- mind, however, that the dental char-
tour asymmetry does not appear to be directly related to the position of the metaconid.
acteristics discussed here comprise
METPOS, mesial; UP, Upper Paleolithic; NE, North East; W., West.
only a small portion of the entire set of
dental traits available for analysis. My
on-going research will take into con-
sideration all dental traits in order to
better test the dental affinity of the
Middle Pleistocene hominids.

CONCLUSIONS
Neanderthal premolars and molars
have received less attention than their
incisors owing to the assumption that
Neanderthal postcanine dental mor-
phology is much like our own. Recent
research, however, has shown this not
to be the case, as Neanderthals exhibit
a unique pattern of dental morpho-
logic trait frequencies (Stringer et al.,
1997; Irish, 1998; Bailey, 2000a;
Coppa et al., 2001). The results of this
particular study corroborate earlier
ones by providing evidence for some
Figure 7. The frequency at which a transverse crest, a well-developed metaconid, and newly described dental morphologic
lingual crown asymmetry occurs in combination in modern humans and Neanderthals. See traits that are rare-to-absent in mod-
text for explanation.
ern humans. In particular, the mandib-
their size and morphologic complex- ular P4s present a unique combination
ity. The Mauer and Arago 13 P4s are
quite similar to each other and overall
resemble Homo erectus P4s: both ex-
hibit symmetrical lingual contours
and lack a continuous transverse crest
(Figure 10a,b). The Arago 28 P4, how-
ever, exhibits a mixed morphology: it
possesses a mesially placed meta-
conid together with a strong trans-
verse crest but a symmetrical lingual
contour (Figure 10c). As was the case
for the P4, the Mauer molars also re-
semble the Homo erectus condition:
completely lacking a MTC (Figure
Figure 9. A comparison of mid-trigonid crest frequencies among sample groups. Neander-
11a). The Arago molars are variable thals show the highest frequencies for all three molars (M1–3). UP, Upper Paleolithic; NE,
(as were the P4s). One of the two M1s North East; W., West.
ARTICLE THE ANATOMICAL RECORD (NEW ANAT.) 155

of dental characters that was neither

observed in modern humans nor in the
Homo erectus outgroup. This finding
suggests that the characters are not
primitive retentions but are instead Ne-
anderthal dental autapomorphies.
The preponderance of evidence
thus far suggests that, relative to mod-
ern humans and regardless of geolog-
ical age or geography, Neanderthals
are uniquely derived in their dental
Figure 8. A comparison of mandibular molars. Arrows point to the mid-trigonid crest (a) or morphology. Not surprisingly, this
the sagittal sulcus (b,c). A, Grotta Brueil, Neanderthal [flipped]; b, KNM-WT 15000, Homo finding also holds true when Neander-
erectus; c, Abeille, Upper Paleolithic Homo sapiens. Symbol orientation as in Figure 3.
thals are compared with a hypotheti-
cal “dental ancestor” of modern hu-
TABLE 4. Characters and character state distribution mans (Stringer et al., 1997). However,
that Neanderthals exhibit a unique
Character H. erectus Neanderthal H. sapiens dental morphologic pattern may not
be particularly surprising, given that
P4 Lingual crown outline Symmetrical Asymmetrical Symmetrical
P4 Transverse crest Absent/divided Present Absent distinct dental patterns are found in
Frequency of a well- Frequent Frequent Infrequent contemporary modern humans of dif-
developed P4 ferent geographic origins (Hanihara,
metaconid
Mid-trigonid crest (M1) Absent/divided Continuous Absent/divided
Mid-trigonid crest (M2) Absent Continuous Absent
The preponderance of
evidence thus far
suggests that, relative to
modern humans,
regardless of geologic
age or geography,
Neanderthals are
uniquely derived in their
dental morphology.
Figure 10. A comparison of Homo heidelbergensis P4s. All resemble Homo erectus in their
symmetrical lingual crown contour and complex occlusal topography. The continuous
transverse crest present in Arago 28 (c) is a typical Neanderthal character. It is absent
1969; Mayhall et al., 1982; Turner,
in both Mauer (a) and Arago 13 (b). See text for explanation. Symbol orientation as in 1983; Irish, 1994; Hawkey, 1998).
Figure 3. What is interesting is that modern hu-
man dental patterns appear to have
great antiquity. For example, contem-
porary Europeans link closely with
Upper Paleolithic modern humans,
and contemporary Africans link
closely with Late Pleistocene Africans
(Bailey, 2000a). Yet, the Neanderthal
dental pattern has no modern repre-
sentatives and, instead, seems to dis-
appear with its possessors. In addition,
incipient Neanderthal dental traits can,
however, be found among earlier
archaic Homo sapiens (specifically
Arago—this study), suggesting that the
Neanderthal dental pattern may have
been evolving separately from modern
Figure 11. A comparison of the Mauer M1 (a) and an M2 from Arago (b: flipped). Both exhibit
a complex tooth crown, but Mauer lacks a mid-trigonid crest (a typical Neanderthal charac-
humans for a long time. Moreover, nei-
ter), and in Arago, it is present. See text for explanation. Symbol orientation as in Figure 3. ther the unique Neanderthal dental pat-
156 THE ANATOMICAL RECORD (NEW ANAT.)
tern (in trait frequencies) nor the mor-
phology described herein has been
found in Upper Paleolithic modern hu-
mans, as may be expected if the two
groups were interbreeding for a sub-
stantial length of time.
ACKNOWLEDGMENTS
Thanks to Bill Kimbel, Don Johanson,
and Ken Mowbray and the anony-
mous reviewer for their helpful com-
ments on the manuscript and to all
the curators who provided access to
the many fossils included in this
study. This research was made possi-
ble by funding from the LSB Leakey
Foundation and National Science
Foundation (Award No. BCS-0002481).
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