International Journal of Osteoarchaeology

Int. J. Osteoarchaeol. (2015)

Published online in Wiley Online Library
(wileyonlinelibrary.com) DOI: 10.1002/oa.2451

The Retromolar Space: A Morphological

Curiosity Observed Amongst the
Protohistoric Arikara and Mandan
C. DE LA COVA1,2*
1
Department of Anthropology, University of South Carolina, Columbia, SC 29208, USA
2
African American Studies Program, University of South Carolina, Columbia, SC 29208, USA

ABSTRACT The retromolar space (RMS), defined in palaeoanthropology as a space posterior to the third molar, between
the distal edge of the tooth and the anterior margin of the ascending ramus when the mandible is held in
lateral view, has been described as an autapomorphic trait unique to Neanderthals despite its presence in
anatomically modern humans. This study examined RMS prevalence in a sample of protohistoric Arikara
and Mandan Amerindians to determine what craniofacial morphology is correlated with the RMS. It was
hypothesised that the feature would be present in the Amerindians studied and associated with a long cranial
length, a large nasal height, midfacial prognathism, a broad mandible, and dental wear. The results indicated
that RMSs were present in the Arikara and Mandan and significantly correlated with cranial length, cranial
breadth, nasal height, bizygomatic breadth, basion-nasion length, basion-nasiospinale, mandible length,
gonial angle, bigonial breadth, and dental wear. Thus, RMSs are associated with a dolichocephalic skull,
wide cranial and facial breadth, a prognathic face, large nose and a corresponding wide and long mandible
with a reduced gonial angle. This suggests that the RMS is the result of these features merging together in the
craniofacial complex and should not be considered a Neanderthal autapomorphy. Copyright © 2015 John
Wiley & Sons, Ltd.

Key words: Arikara; craniofacial morphology; Mandan; retromolar space

Introduction to aspect of the last erupted molar. In contrast, the

This study examines the retromolar molar space (RMS)
in a sample of protohistoric Arikara and Mandan
Amerindians dating from 1679 to 1832. The definition
of this feature varies in regard to discipline. Otolaryn-
gologists define the RMS as the part of the buccal
cavity that bilaterally lies between the two dental
arcades and behind the roots of the last upper and lower
erupted molars (Barbosa, 1959; Malhotra, 2005).
Dentistry and orthodontic scholars also define the
feature in regard to the last erupted molar. Some refer
to it as the ‘lower third molar space’ or the space
between ‘the distal surface of the second molar crown
and the anterior border of the mandibular ramus’ (Zelić
& Nedeljković, 2013:924). The space is not restricted to
the third molar, but comprises the area from the
ascending ramus of the mandible to the most posterior
* Correspondence to: Carlina de la Cova, Department of Anthropology,
University of South Carolina, Gambrell Hall 440A 817 Henderson Street
Columbia, SC 29208, USA.
e-mail: delacova@mailbox.sc.edu
RMS is defined by palaeoanthropologists as the space
posterior to the mandibular third molar, ‘or the
presence of a clear separation between the distal
margin of the third molars and the anterior margin
of the ramus,’ when the mandible is held in lateral
view (Franciscus & Trinkaus, 1995:577). Thus, by
palaeoanthropological standards, this feature is a
non-metric trait that is only considered present when
the skull is viewed in norma lateralis and comprises a
visible space from the anterior margin of the ascending
ramus to the most distal edge of only the third molar,
as illustrated in Figure 1.
Anthropological research on the RMS has mostly
focused on Palaeolithic hominins as some scholars con-
sider the feature a Neanderthal autapomorphy, despite
its occurrence in anatomical modern humans (AMHs)
from the Upper Palaeolithic and Neolithic periods
(Frayer, 1992; Franciscus & Trinkaus, 1995; Nara
et al., 1998; Trinkaus, 1987, 2003, 2007). Carleton
Coon (1962) was the first to define the RMS, indicat-
ing that it was distinctive to Neanderthals and resulted
from increased midfacial prognathism. This allowed for

Copyright © 2015 John Wiley & Sons, Ltd. Received 17 November 2013
Revised 28 February 2015
Accepted 20 March 2015

Figure 1. Illustration of the retromolar space as defined in
palaeoanthropology (after Aiello & Dean, 1990).
the expansion of the internal nasal cavity surface area,
thus providing an adaptation to frigid temperatures by
warming incoming cold air.
Palaeoanthropologists and skeletal biologists have
continued in the same vein as Coon, defining the
RMS as singular to Neanderthal cranial anatomy. Vari-
ous studies have attempted to provide a morphological
explanation of why the feature exists in Neanderthals
(Rak, 1986; Trinkaus, 1987; Frayer, 1992; Spencer &
Demes, 1993; Franciscus & Trinkaus, 1995). A more
anteriorly placed dentition and retracted zygomatics
(Howells, 1975), a reduction in mandibular ramus
breadth and shortening of the dental arcade (Trinkaus,
1983, 1987), facial prognathism and forward migration
of the mandibular tooth row (Frayer, 1992), an adapta-
tion to heavy loading on the anterior dentition,
anterior migration of the postcanine dentition and mas-
seter and temporalis muscles, and shortening of the
dental arcade (Spencer & Demes, 1993) have all been
attributed to the creation of an RMS. However, the
trait is not exclusive to Neanderthals and is found
in AMHs, including Palaeolithic individuals from
Predmostí, Qafzeh, Skhul and Vogelherd, and Neo-
lithic Jomon populations (Frayer, 1992; Franciscus &
Trinkaus, 1995; Nara et al., 1998; Trinkaus, 1987,
2003, 2007b). Despite its presence in these prehistoric
groups, few studies have examined RMS prevalence in
recent populations. This research does so by determin-
ing if RMSs exist in a sample of protohistoric Arikara
and Mandan, and if so, what anatomical features in
the craniofacial complex of these Amerindians correlate
with this feature.
Most anthropological research focused on the RMS
has defined and scored it based on a non-metric assess-
ment of its presence or absence in norma lateralis. Few
studies in the discipline have attempted to metrically
define or establish what measurement size or metric
Copyright © 2015 John Wiley & Sons, Ltd.
C. de la Cova
parameters qualify as an RMS. This study differs from
previous works as it will not only record retromolar
space presence (RMSP), but it will also metrically ex-
amine the length of the feature to determine if there
is a relationship between RMS size and other
craniometric traits. This will allow for a better under-
standing of the anatomical relationship between the
RMS and the dimensions of the skull. The Arikara
and Mandan were selected for this study because of
their craniofacial morphology, which echoes traits ob-
served in Neanderthals, including a long skull, tall face
and large mandible. Based on these shared features, it is
hypothesised that RMS presence will be correlated
with cranial length, facial prognathism, nasal height,
mandibular length and breadth, and dental wear in
the Amerindians studied.
The Arikara and Mandan
The Arikara and Mandan Amerindians inhabited the
American Great Plains and resided in small villages
along the Missouri River from South Dakota to North
Dakota, from the 15th to 19th centuries. The Arikara
are Caddoan speaking and closely related to the
Pawnee (Mariotti, 1995:1). They migrated up the
Missouri River during the 17th century and experi-
enced European contact in the 18th century (Mariotti,
1995). The Arikara settled at the previously Mandan-
occupied site of Greenshield, North Dakota (32OL17),
the main focus of this paper, at the end of the 18th cen-
tury, between 1785 and 1790 (Mariotti, 1995:7). The
Mandan were a Siouan-speaking tribe that also lived in
western North Dakota. By the time the Arikara arrived
at Greenshield, the Mandan had moved north, following
the Missouri River (Mariotti, 1995:7–8). Both were semi-
sedentary horticulturalists that relied heavily on buffalo
during the winter, leaving their villages and camping in
skin lodges whilst in pursuit of these animals (Denig,
1961; Trimble, 1994). Although buffalo was their main
staple, deer, elk, pronghorns and bears were also hunted
for consumption and trade (Robertson, 2001:159).
Tribes were agrarian during the spring and summer
months, living in dome-shaped lodges that held 10 to
30 members of an extended family (Robertson,
2001:160). Towns included 12 to 150 or more concen-
trated lodges that were laid out randomly and so close
together that inhabitants had to walk in single-file lines
in order to pass each structure (Denig, 1961). Corn,
squash, beans, pumpkins, and sunflowers were produced
on small patches of land, with planting occurring from
mid-April to the beginning of May, depending on
the weather (Denig, 1961:44; Robertson, 2001:159).
Int. J. Osteoarchaeol. (2015)
The Retromolar Space: A Morphological Curiosity
Crops were stored away and cached until the following
spring, when buffalo were not amply available (Denig,
1961). The diet was supplemented with wild Indian
potatoes, chokecherries, prairie turnips, artichokes,
grapes, plums, currants, and prickly pear cactus fruit
(Robertson, 2001:159). The Missouri River also pro-
vided numerous resources, including fish, mollusks, and
terrapins.
French contact in the Upper Missouri River Valley
introduced the Arikara and Mandan to the fur trade.
They primarily sold and bartered buffalo robes with
Europeans and Americans at Fort Clark Trading Post
in North Dakota and the American Fur Company
based in St. Louis (Denig, 1961; Robertson, 2001). In
addition to hides, Arikara-grown corn and squash were
exchanged for numerous items, including combs,
beads, knives, hoes, ammunition, paints, and tobacco
(Denig, 1961:46).
European contact also resulted in the rapid spread of
infectious diseases and viruses. Smallpox, bubonic
plague, malaria, and yellow fever plagued the Missouri
Valley from the end of the 18th century until
the beginning of the 19th century (Mariotti, 1995;
Robertson, 2001). During this period, the Mandan,
Arikara and Hidatsa tribes experienced a devastating
succession of measles, smallpox, and cholera epidemics
that greatly reduced their populations. Between 1780
and 1796, a series of smallpox epidemics spread
amongst the Arikara. The years of 1780 to 1782 were
particularly lethal for the tribe, whose numbers de-
clined from 15 000 to 1500 (Isenberg, 2001). Popula-
tion losses and cultural upheaval resulted in the
formation of a coalition with their former enemies,
the Mandan, for survival (Mariotti, 1995; Robertson,
2001). This union resulted in gene flow between the
groups, which has been observed by skeletal biologists
(Jantz, 1973, 1977; Key & Jantz, 1981; Owsley et al.,
1981; Key, 1983; McKeown, 2000). Early Arikara sam-
ples have shorter cranial vaults and faces. As time
passes, Arikara crania become morphologically similar
to the Mandan, exhibiting an increase in cranial length
and facial height (Key & Jantz, 1981; Owsley et al.,
1981; McKeown, 2000).
Between 1837 and 1838, the infamous Great Plains
smallpox epidemic, brought to the Missouri River
Valley by infected passengers on the American Fur
Company’s steamer, St. Peter, swept through the surviv-
ing Arikara and Mandan groups (Robertson, 2001).
Half of the remaining Arikara and 90% of the Mandan
population perished (Mariotti, 1995). Today, their de-
scendants live on the Ford Berthold Reservation in
North Dakota as part of the federally recognised
Mandan, Hidatsa and Arikara Nation.
Copyright © 2015 John Wiley & Sons, Ltd.
Materials and methods
A skeletal sample of protohistoric Arikara and Mandan
Amerindians (n = 34) collected by A. Bowers from the
Greenshield and Larson protohistoric village sites and
curated at Indiana University, Bloomington, were
examined for the presence of RMSs. Greenshield
(320L17), located in west-central North Dakota on
the west bank of the Missouri River, dates from 1785
to 1830 (Mariotti, 1995). It was originally occupied
by the Mandan and later abandoned, before the
Arikara settled there in the late 18th and early 19th
centuries. The Larson site (39WW2) was a protohis-
toric Arikara village and cemetery that was occupied
from 1679 to 1733 and located south of Mobridge in
Walworth County, South Dakota, on the left bank of
the Missouri River.
All adults with complete, non-fragmented skulls
and fully erupted mandibular third molars present
were studied. Mandibles were examined in norma
lateralis for a visible RMS between the distal margin
of the third molar and the anterior aspect of the as-
cending ramus, as described by Franciscus & Trinkaus
(1995) and illustrated in Figure 1. RMSP was bilater-
ally recorded and scored as present or absent. An ad-
ditional ‘RMSP overall’ variable was created, which
combined the findings from the left and right sides
of the mandible so that RMSP could be accurately re-
ported for each individual, especially those with a
unilateral RMS. Retromolar space length (RMSL)
was measured with a GPM sliding calliper from the
posterior distal side of the third molar to the buccal
portion of the ascending ramus along the internal al-
veolar ridge, as shown in Figures 2 and 4(a). If the gap
between these anatomical landmarks was measureable,
then the RMS was scored as present. However, if the
space was too small for calliper measurement, the
RMS was recorded as absent, but a dummy variable
of 0.1 cm was entered so that statistical analyses could
be performed to determine if there was a relationship
between RMSP and RMSL and other craniometric
features.
Cranial and mandibular measurements, listed in
Table 1, were taken on all complete skulls. Measure-
ments of the length of the lingual and buccal/labial sides
of the mandibular tooth row were also recorded
[Figures 3 and 4(b)]. Buccal/labial mandibular tooth
row length was documented using a white cord and
measuring from the left oblique line to the right oblique
line [Figures 3(a) and 4(b)]. Lingual tooth row length
was also taken in a similar manner along the lingual sur-
face of the mandible utilising a white cord and measur-
ing from the extramolar sulcus, at the left endocoronoid
Int. J. Osteoarchaeol. (2015)
 C. de la Cova

Spearman’s rho correlation analyses to determine what

craniofacial metrics were correlated with RMSP and
RMSL.
Since the completion of this research, the collection
has been repatriated.

Figure 2. Retromolar space length was measured from the posterior
distal side of the third molar to the buccal portion of the ascending ra-
mus along the internal alveolar ridge of the mandible. (a) The measure-
ment in lateral view. (b) The anatomical landmarks that comprised the
measurement (modified from Buikstra & Ubelaker, 1994).
ridge, to the right endocoronoid ridge [Figure 3(b)]. For
both measurements, the white cords were marked based
on where the described left and right anatomical land-
marks aligned on each mandible. The marks on the
cords were then measured with sliding callipers. Dental
wear, which has been linked to RMS presence (Nara
et al., 1998), was also recorded as present or absent.
All measurements were analysed using one-tailed
Results
Of the 32 skulls available for study, only 10 complete
crania maintained their third molars and could be ex-
amined (Table 2). Five additional mandibles were in-
cluded in regard to RMS features and mandibular
measurements (Table 2). Eleven (73.3%) of the 15
Arikara analysed had non-metric and measurable RMSs
(Table 3; Figure 5). The average RMSL when present
was 0.8545 cm. All RMSLs are reported in Table 3.
Statistical results (Table 4) indicated that RMSP was
highly correlated with cranial length (left: r = 0.739,
p = 0.011), cranial breadth (left: r = 0.696, p = 0.019),
nasal height (left: r = 0.617, p = 0.038; right: r = 0.731,
p = 0.013), bizygomatic breadth (left: r = 0.713,
p = 0.023; right: r = 0.630, p = 0.047), basion-nasion
length (left: r = 0.606, p = 0.042), basion-nasospinale
(overall: r = 0.688, p = 0.014), mandible length (left:
r = 0.486, p = 0.039), gonial angle (left: r = 0.649,
p = 0.006), and bigonial breadth (left: r = 0.579,
p = 0.019).
RMSL was also significantly correlated with cranial
length (left: r = 0.819, p = 0.003), glabella-inion (left:
r = 0.662, p = 0.026), cranial breadth (left: r = 0.679
p = 0.022; right: r = 0.587, p = 0.048), nasal height
(right: r = 0.678, p = 0.022; maximum: r = 0.553,
p = 0.049), bizygomatic breadth (left: r = 0.630,
p = 0.047), basion-nasion (left: r = 0.672, p = 0.024),
basion-nasospinale (maximum: r = 0.612, p = 0.030),
gonial angle (left: r = 0.522, p = 0.028), and bigonial
breath (left: r = 0.633, p = 0.010; maximum: r = 0.508,
p = 0.032). Analyses of the relationship between dental

Table 1. Measurements

Cranial Measurements Instrument Mandibular measurements Instrument

G-OP (cranial length) Spreading Mandible length Mandibulometer

G-I (glabella-inion) Spreading Gonial angle Mandibulometer
EU-EU (cranial breadth) Spreading Ascending ramus height Mandibulometer
N-NSP (nasion-nasospinale) Sliding Bicondylar breadth Sliding Calliper
N-PR (nasion-prosthion) Sliding Bigonial breadth Sliding Calliper
N-GN (nasion-gnathion) Sliding Retromolar space length Sliding Calliper
BZ-BZ (bizygomatic-breadth) Spreading Lingual tooth row from left endocoronoid Cord/sliding calliper
B-N (basion-nasion) Spreading Ridge to right endocoronoid ridge
B-NSP (basion-nasospinale) Spreading Buccal/labial tooth row from left Cord/sliding calliper
B-PR (basion-prosthion) Spreading Oblique line to right oblique line

Copyright © 2015 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. (2015)
The Retromolar Space: A Morphological Curiosity

Table 2. Sample composition

Complete skulls Site Mandibles only Site

32–3 (Ar1–3) Greenshield 32–24 Greenshield

32–4 (Ar1–4) Greenshield 34–1 (Ar 3–1) Greenshield
32–6 (Ar1–6) Greenshield 35–3 (MND 1–3) Larson
32–7 (Ar1–7) Greenshield 35–5 (MND 1–5) Larson
32–16 (Ar1–16) Greenshield 37–1 Unknown
32–17 (Ar1–17) Greenshield
32–19 (Ar1–19) Greenshield
32–21 (Ar1–21) Greenshield
oB–9 Over Museum
978 Over Museum

wear and RMSP revealed no significant associations

(Table 5). However, there was a positive correlation be-
tween RMSL and dental wear (left: r = 0.528, p = 0.026).

Figure 3. Measurements of the buccal/labial (a) and lingual (b) mandib-
ular tooth row lengths (modified from Buikstra & Ubelaker, 1994).
Discussion
The findings of this study contribute to current anthro-
pological debates related to the RMS and shed light on
the possible origins of this morphological curiosity. Re-
sults indicated that 73.3% of the Arikara and Mandan
studied had RMSs, or a visible gap between the ascend-
ing ramus of the mandible and the most distal aspect of
the third molar, that was measureable (Figure 5). RMSP

Figure 4. Measurements of retromolar space length (a) and buccal/labial mandibular tooth row length (b) demonstrated on a subject. This figure is
available in colour online at wileyonlinelibrary.com/journal/oa

Copyright © 2015 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. (2015)
 C. de la Cova

Table 3. Retromolar space presence (RMSP) and length (RMSL)

RMSP left side RMSL left side (cm) RMSP right side RMSL right side (cm) RMSP overall Max RMSL (cm)

oB–9 Unobservable N/A Present 0.85 Present 0.85

32–3 Absent 0.40 Unobservable N/A Absent 0.40
32–4 Absent 0.40 Absent 0.20 Absent 0.40
32–6 Present 1.00 Present 1.00 Present 1.00
32–7 Present 0.50 Present 0.70 Present 0.50
32–16 Absent 0.20 Present 0.60 Present 0.60
32–17 Present 1.00 Present 0.80 Present 1.00
32–19 Absent 0.35 Present 0.60 Present 0.60
32–21 Absent 0.10 Absent 0.25 Absent 0.20
32–24 Present 0.80 Present 0.80 Present 0.80
34–1 Present 1.00 Present 1.00 Present 1.00
35–3 Present 1.00 Unobservable N/A Present 1.00
35–5 Absent 0.40 Absent 0.10 Absent 0.40
37–1 Present 0.90 Present 0.95 Present 0.95
978 Present 0.90 Present 0.70 Present 0.90
Percentage of individuals with RMSs
Total present 11 (73.3%)
Total absent 4 (26.7%)
Average size of space
Present 0.8545
Not Present 0.3625
Left 0.6393
Right 0.6577
Overall 0.7233

and RMSL were negatively correlated with gonial angle
and positively correlated with cranial length, cranial
breadth, nasal height (nasion-nasospinale), facial
breadth (bizygomatic breadth), midfacial prognathism
(basion-nasion and basion-prosthion), mandible length,
and mandible breadth (bigonial breadth). These find-
ings supported the research hypothesis by demonstrat-
ing that RMSs amongst the Arikara and Mandan are
strongly correlated with a dolichocephalic skull; wide
cranial breadth; a large nasal height; a wide, prognathic
face; and a square, long, broad and projecting
mandible.
Whilst these findings may be unique to the Amerin-
dians examined, or the result of a small sample size,
previous studies on numerous Arikara and Mandan re-
mains have noted their tall facial heights and long
skulls, which become more defined as gene flow
increased between the two groups (Key & Jantz,
1981; Owsley et al., 1981; McKeown, 2000). The
craniometric features observed in this study are also
found in Neanderthals, although Neanderthals display
the extreme of these traits, including a low and long
cranium, large nose, long face, a large and broad man-
dible, midfacial prognathism, and an RMS. Given that
much research has been performed on Neanderthal fa-
cial morphology and the RMS, it is worth discussing
what causes an RMS in Neanderthals and whether
any of these elements apply to the Amerindians
examined.
Many have argued that Neanderthal craniofacial
morphology, including the RMS, resulted from cli-
matic and behavioural adaptations. Behavioural argu-
ments have sought to explain anterior dental wear in
Neanderthals, asserting that the hominin’s face devel-
oped as a response to paramasticatory use of the teeth
to resist biomechanical stresses associated with re-
peated heavy loading (Brace, 1962; Smith, 1983;
Trinkaus, 1987; Rak, 1986; Spencer & Demes, 1993).
In the same vein, some researchers believe the Nean-
derthal face was selected for bigger teeth to resist high
levels of dental wear (O’Connor et al., 2005). The pres-
ence of large, robust incisors, taurodontism, and high
levels of attrition in the anterior Neanderthal dentition
support this. However, this has been contradicted by
studies that suggest Neanderthals were not capable of
producing great anterior bite forces and probably did
not generate anterior dental loads larger than AMHs
(Antón, 1996; O’Connor et al., 2005).
The most common assertion made about the prog-
nathic Neanderthal face is that it evolved as environ-
mental adaptation to a dry, glacial climate. Its large
internal nasal margin and expanded sinuses may have
provided additional surface area for nasal mucosa and
cilla to warm and humidify incoming air before it
reached the lungs (Laitman et al., 1996; Schwartz &
Tattersall, 1996). Forward shifting of the face and
expansion of the sinuses would have resulted in a
more forward-placed tooth row and, thus, an RMS.

Copyright © 2015 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. (2015)
The Retromolar Space: A Morphological Curiosity
Figure 5. Retromolar spaces measured and observed in the Amerindians studied. This figure is available in colour online at wileyonlinelibrary.com/journal/oa
However, Rae et al. (2011) argue that Neanderthal si-
nuses were not larger than AMHs and possessed a
reduction in sinus volume associated with cold adapta-
tion. Nathan Holton and colleagues (2011:625) dis-
agree with this, pointing out that Rae et al. (2011)
ignored ‘the more climatically relevant aspect of the
naso-facial skeleton, the internal nasal passages, which
are the primary site of heat and moisture exchange with
respired air’. Even modern humans adapted to cold cli-
mates have taller, narrower, and longer nasal cavities
that allow for increased mucosal surface area and effi-
cient moisture and heat exchange by warming and
moisturising cold, dry air (Franciscus & Long, 1991;
Yokley, 2009; Holton et al., 2011). If this is true for
modern humans, then it may be true for Neanderthals.
Facial prognathism may also be related to cranial ca-
pacity and cranial length, as basion-nasion length is in-
fluenced by endocranial volume (Trinkaus, 2003).
Further analyses of the Amerindians examined in this
study supported this and revealed that basion-nasion
length was highly correlated with cranial length
(r = 0.792, p = 0.003). Like Amerindians and AMHs,
Neanderthals were dolicocephalic, had big brains,
with an average cranial capacity of 1400 to 1600 ccs,
and an increased basion-nasion length. However,
Copyright © 2015 John Wiley & Sons, Ltd.
when compared with most Pleistocene non-Neanderthal
hominins, Neanderthal faces are not longer or more
prognathic, but average and reduced in size (Trinkaus,
2003). It is AMHs that are more derived in
numerous ways from previous Homo species, with their
uniquely human autapomorphies including their small,
orthognathnic faces, gracile mandibles, and projecting
mental eminences (Trinkaus, 2003).
Studies of Neanderthal and AMH facial ontogeny
support this, illustrating that both hominins had
different patterns of craniofacial and mandibular devel-
opment that resulted in their dissimilar facial morphol-
ogies (Bastir et al., 2007). Mandibular growth in AMHs
includes ‘supero-inferior expansion of the mandible and
a retraction of the dental arcade’ (Bastir et al.,
2007:1129). This leads to increased ramus height, a
narrower posterior mandible and the formation of a
prominent chin. Neanderthals, in contrast, experienced
forward growth of the mandibular corpus and dental ar-
cade, which resulted in a more anteriorly placed alveo-
lar process and an increase in height of the anterior
dental arcade and mandibular body (Bastir et al.,
2007). These ontogenetic shifts in Neanderthals re-
sulted in shorter and broader mandibular rami, and
more than likely, the creation of the RMS. In other
Int. J. Osteoarchaeol. (2015)
 C. de la Cova

0.508 (0.032)* 8
words, the RMS serves no functional or evolutionary

0.553 (0.049)*

0.612 (0.030)*
0.455 (0.093)
0.276 (0.220)
0.472 (0.084)

0.129 (0.361)
0.208 (0.282)
0.515 (0.078)
0.454 (0.094)

0.385 (0.136)
0.229 (0.206)
0.400 (0.070)
0.382 (0.089)
0.202 (0.275)

0.364 (0.091)
0.355 (0.097)
Maximum

importance and is the result, or side effect, of other

r (sig.)
RMSL

factors related to craniofacial growth, development,

and anatomy.
This is further iterated by the fact that the RMS is
not unique to Neanderthals. Its presence in the Arikara
and Mandan examined in this study is also the result of
other anatomical characteristics and cultural behaviours
0.587 (0.048)*
0.678 (0.022)*
0.388 (0.151)
0.249 (0.259)

0.249 (0.259)
0.235 (0.271)
0.539 (0.084)
0.409 (0.137)
0.517 (0.077)
0.288 (0.227)
0.312 (0.150)
0.474 (0.051)
0.067 (0.417)
0.194 (0.296)
0.452 (0.060)
0.260 (0.195)
0.331 (0.135)
Table 4. Spearman’s correlation analyses of retromolar space presence (RMSP) and length (RMSL) with cranial and mandibular measurements

RMSL right

related to the craniofacial complex. Studies on Arikara

r (sig.)
side

cranial ontogeny have indicated that the Arikara

underwent temporal changes that included a reduction
in frontal bone length, an increase in maxillary height,
a decrease in orbit size, expansion of the zygomatics,
and increased alveolar prognathism (Strand &
0.630 (0.047)*
0.672 (0.024)*

0.522 (0.028)*

0.633 (0.010)*
O’Higgins, 2003). Craniometric studies of Arikara from
0.819* (0.003)
0.662* (0.026)
0.679* (0.022)
0.556 (0.060)
0.300 (0.217)
0.429 (0.125)

0.477 (0.097)
0.250 (0.258)
0.432 (0.061)

0.600 (0.015)
0.462 (0.090)

0.416 (0.070)
0.307 (0.143)
RMSL left

the 16th to 19th centuries support this (Jantz, 1970,

r (sig.)
side

1972, 1973, 1977; Key & Jantz, 1981; Owsley et al.,

1981; Key, 1983; McKeown, 2000). The Arikara orig-
inally had shorter cranial lengths, cranial heights, and
facial heights (Owsley et al., 1981; McKeown, 2000).
As time passed, their skulls became longer and more
0.688 (0.014)*
0.191 (0.298)
0.076 (0.417)
0.305 (0.196)
0.501 (0.070)
0.114 (0.377)
0.190 (0.300)
0.413 (0.135)
0.267 (0.228)

0.424 (0.111)
0.018 (0.475)
0.210 (0.226)
0.059 (0.420)
0.130 (0.352)
0.216 (0.229)
0.175 (0.266)
0.350 (0.100)

prognathic, frontal bone height reduced, and facial

Overall

height increased (Owsley et al., 1981; Key, 1983;

r (sig.)
RMSP

McKeown, 2000). Whilst reduction in vault height

and lengthening of facial height has been associated
with temporality, it is also related to northward move-
ment along the Missouri River (Key, 1983). However,
it seems unlikely that these shifts were related to cli-
0.731 (0.013)*

0.630 (0.047)*
0.364 (0.168)
0.260 (0.250)
0.468 (0.102)

0.416 (0.133)
0.414 (0.134)

0.312 (0.207)
0.574 (0.053)
0.211 (0.292)
0.073 (0.406)
0.196 (0.261)
0.197 (0.270)
0.000 (0.500)
0.269 (0.187)
0.122 (0.345)
0.245 (0.210)

matic change as they occurred over a short time span

r (sig.)
RMSP
Right

of 200 years (McKeown, 2000). Historical, archaeolog-

ical, and bioanthropological evidence indicates that in-
creasing gene flow through time with the Mandan,
who had dolicocephalic skulls, long faces, and alveolar
prognathism, resulted in the shifts in Arikara craniofa-
cial morphology (Owsley et al., 1981; Key & Jantz,
0.649 (0.006)**
0.739 (0.011)*

0.713 (0.023)*
0.606 (0.042)*

0.486 (0.039)*

0.579 (0.019)*
0.696* (0.019)
0.617* (0.038)
0.522 (0.075)

0.217 (0.287)
0.433 (0.122)

0.522 (0.075)
0.524 (0.074)

0.416 (0.079)
0.314 (0.188)

0.252 (0.192)
0.251 (0.193)

1981; Key, 1983; McKeown, 2000).

r (sig.)
RMSP

These craniofacial changes would have caused ana-

Left

tomical shifts in regard to mandibular placement. An

extended cranial length, wide cranial breadth, and prog-
nathic Arikara face resulted in an increase in mandibular
length and gonial breadth, with forward shifting of the
mandible so that it could efficiently articulate with the
17.820
17.640
13.020
5.400

11.720
13.944
10.135
9.770

8.390
23.333

12.109
10.154
13.615
14.353
7.160

9.800

6.650
x

cranium. From a functional perspective, the associated

mandibular dentition had to shift to a more anterior
**Significant at the 0.01 level.
*Significant at the 0.05 level.

position so that it could come into occlusion with the

Ascending ramus height

maxillary teeth (Frayer, 1992; Wolpoff, 1996). Thus,

Buccal/labial tooth row
Bizygomatic breadth

the RMS becomes a side effect or end result of other

Nasion-nasospinale

Basion-nasospinale

Bicondylar breadth

morphological changes in the skull.

Bigonial breadth
Lingual tooth row
Nasion-prosthion

Mandible length
Basion-prosthion
Cranial breadth

Nasion-gnathion

Basion-nasion
Cranial length

The Amerindians analysed also had heavy dental at-

Glabella-inion

Gonial angle

trition (Ungar et al., 1997), which was significantly cor-

related with RMSL (r = 0.528, p = 0.003). This indicates
that tooth wear plays a role in RMS formation and
length. Chronic dental attrition results in wear on the
Copyright © 2015 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. (2015)
The Retromolar Space: A Morphological Curiosity
Table 5. Spearman’s correlation analysis of dental wear and retromolar space presence (RMSP) and length (RMSL)
Left RMSP Right RMSP Overall RMSP
Dental wear 0.417 (0.069) 0.225 (0.230) 0.123 (0.331)
*Significant at the 0.05 level.
occlusal and interproximal surfaces of a tooth, includ-
ing the mesial and distal sides of the crown. Over time,
this causes a reduction in tooth size. As tooth crowns
become smaller, the dentition will slowly start to mi-
grate anteriorly, which results in the creation or length-
ening of an RMS (Nara et al., 1998). This has been
observed in prehistoric Japanese Jomon populations
and found to be directly related to age and wear
(Nara et al., 1998).
Thus, in regard to craniometrics, both Neanderthals
and the Arikara and Mandan examined in this study
share similar cranial morphological traits that result in
the formation of an RMS. Both are dolicocephalic,
have wide skulls, long facial and nasal heights, and
prognathic faces with robust, broad, long and square
mandibles that have anteriorly placed tooth rows,
which are the result of facial prognathism, increased
mandibular length, and chronic dental wear. The RMS
results from the merging of all these features in the skull.
This means that it cannot be derived, is unique, or an
autapomorphy of one hominin species. Based on the
results of this study, it is merely a morphological end
result, or curiosity with no functional purpose, of other
cranial traits asserting themselves.
Conclusion
This study examined RMS prevalence in a sample of
protohistoric Arikara and Mandan Amerindians, dating
from 1679 to 1832. Results indicated that RMSs were
present in 73.3% of the sample. This supports asser-
tions made by other researchers that the RMS should
not be considered an autapomorphic Neanderthal trait
based on its presence amongst other groups of AMHs
(Frayer, 1992; Franciscus & Trinkaus, 1995; Trinkaus,
2003, 2007; Rosas & Bastir, 2004; Bastir et al., 2007).
Spearman’s rho analyses of cranial measurements
revealed that RMSP and RMSPL were correlated
with a long cranial length, wide cranial breadth, tall
nasal height, increased basion-nasion length, wide
bizygomatic breadth, mandible length, reduced gonial
angle, a broad bigonial breadth, and dental wear. The
findings of this study suggest that the RMS is merely
a result of these important craniometric features. The
long cranial length, wide cranial breath, and increased
Copyright © 2015 John Wiley & Sons, Ltd.
Left RMSL Right RMSL Maximum RMSL
0.528* (0.026) 0.415 (0.079) 0.399 (0.070)
basion-nasion length observed on the Arikara resulted
in a prognathic face that caused the forward migration
of the maxillary dentition and dental arcade. A longer
and wider skull meant the mandible had to extend in
length and bigonial breadth. The mandibular dentition
would have also migrated anteriorly to occlude with
the maxillary teeth and articulate with the cranium.
Furthermore, dental wear would have contributed to
increasing the size of the RMS. When all of the
aforementioned anatomical features are considered, the
morphological curiosity of the RMS appears to be the
end result of numerous cranial traits that can be
found in both historically recent AMH and Neanderthal
skulls.
Acknowledgements
The author would like to thank Kevin Hunt for his sup-
port and suggestions, which helped immensely with
this project. Much appreciation is given to Della Cook
for advice on this manuscript and access to skeletal
collections at Indiana University. The author is also
very grateful to of this manuscript, who offered
thoughtful insights and comments. Any errors are the
fault of the author.
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