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Introduction
Homo oresiensis is a diminutive, primitive hominin
found from Flores, eastern Indonesia (Brown et al., 2004;
Morwood et al., 2004, 2005). The skeletal remains of this
new species are known from the Late Pleistocene strata at
Liang Bua, a limestone cave on the island. Morphology of its
cranium, endocast, mandible, shoulder girdle, pelvis, limb
bones, hand, and foot have been described, analyzed, and
interpreted (Brown et al., 2004; Morwood et al., 2005; Falk
et al., 2005, 2009; Argue et al., 2006, 2009; Larson et al.,
2007, 2009; Tocheri et al., 2007; Gordon et al., 2008; Baab
and McNulty, 2009; Brown and Maeda, 2009; Holliday and
Fransiscus, 2009, 2012; Jungers et al., 2009a, b; Lyras et al.,
2009; Aiello, 2010; Kaifu et al., 2011; van Heteren, 2012;
Baab et al., 2013; Jungers, 2013; Kubo et al., 2013; Orr et
al., 2013; Daegling et al., 2014). However, not all the dental
remains have been described in sufcient detail, and there
even exists controversy as to whether the dental morphology
of H. oresiensis is primitive or modern (Jacob et al., 2006;
Brown and Maeda, 2009).
In this paper, we provide detailed morphological descrip-
tion for all the dental materials of H. oresiensis excavated
during the 2002–2004 eld seasons at Liang Bua (Morwood
and Jungers, 2009). Most of the mandibular teeth of
H. oresiensis have been described by Brown and Maeda
(2009), but we here describe these materials again based on
our own observation of the original specimens and high-
resolution micro-CT scans that were not available to the
previous researchers. We also reassess the previous claims
for primitive and modern aspects of the H. oresiensis teeth.
Materials and Methods
The Liang Bua dental collection: 2002–2004
Bony and dental remains belonging to multiple individu-
als have been excavated and reported from the Pleistocene
levels at Liang Bua (Morwood and Jungers, 2009; Morwood
et al., 2009). In this collection, the cranium (LB1/1) and
mandible (LB1/2) of the individual LB1, as well as the man-
dible (LB6/1) from another individual preserve their denti-
tions. Additionally, there are four isolated teeth as shown in
Table 1 and illustrated in Figure 1, Figure 2, Figure 3, and
AnthropologicAl Science
Vol. 123(2), 129–145, 2015
Descriptions of the dental remains of Homo oresiensis
Yousuke KAifu1,2*, Reiko T. Kono1, Thomas SutiknA3,4, E. Wahyu SAptomo4,3, JAtmiko4,3,
Rokus Due Awe4,3**, Hisao BAbA1
1Department of Anthropology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki 305-0005, Japan
2Department of Biological Sciences, The University of Tokyo, 3-1-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan
3Centre for Archaeological Science, University of Wollongong, Northelds Avenue, Wollongong, NSW 2522, Australia
4The National Research and Development Centre for Archaeology, Jl. Raya Condet Pejaten No 4. Jakarta 12510, Indonesia
Received 19 December 2014; accepted 1 May 2015
Abstract Dental remains of Homo oresiensis excavated during 2002–2004 at Liang Bua, Flores, In-
donesia, consist of one partial maxillary dentition, two nearly complete mandibular dentitions, and four
isolated teeth. We present here morphological descriptions of all these specimens and report aspects of
their dentition, occlusion, and oral health condition. This dental assemblage represents probably ve but
possibly four or six individuals. These different individuals share similar dental characteristics, support-
ing the view that the Liang Bua H. oresiensis assemblage represents a single population. We also re-
assess the previous claims for primitive and modern aspects of the H. oresiensis teeth. The previous
studies reached conicting conclusions: some researchers claim that these teeth are fully modern, where-
as others highlight premolar and other morphologies that suggest their direct evolutionary link with the
African earliest form of Homo or Australopithecus rather than with H. erectus. Neither of these views are
supported. The H. oresiensis teeth exhibit a mosaic of primitive, derived, and unique characters, with
the reported primitive aspects broadly comparable to the morphologies observed in H. erectus sensu lato.
Although a more comprehensive comparative analysis is needed to fully illustrate dental morphological
afnities of this dwarfed hominin species, we nd no grounds for the hypothesis that H. oresiensis
originated from the small-bodied, primitive hominins such as H. habilis sensu lato.
Key words: Homo oresiensis, Flores, Indonesia, dental morphology
* Correspondence to: Yousuke Kaifu, Department of Anthropology,
National Museum of Nature and Science, Tokyo, 4-1-1 Amakubo,
Tsukuba-shi, Ibaraki 305-0005, Japan.
E-mail: kaifu@kahaku.go.jp
** deceased
Published online 18 July 2015
in J-STAGE (www.jstage.jst.go.jp) DOI: 10.1537/ase.150501
Y. KAIFU ET AL.130 AnthropologicAl Science
Figure 4. LB8/2, a right mandibular second premolar, was
recovered by wet-sieving from spit 61 of sector VII
(Morwood and Jungers, 2009). This tooth is unique, show-
ing exceptionally good preservation with transparent texture,
among the bone remains from the Pleistocene levels of
Liang Bua, and is likely a contamination from the upper
stratigraphic level. Therefore, we exclude this specimen
from our H. oresiensis dental sample.
Methods
Tooth crown measurements follow the methods described
in Wood (1991). Both the measured and wear-corrected
Table 1. Inventory and measurements of the H. oresisensis dental remains from Liang Bua
Specimen Individual Tooth Side Sector Spit Wear
Crown diam. Cervical diam.a
Root length
MD MD
corrected BL MD BL
Maxilla
LB15/2 LB15/2 I1L III 51 7 — — ≥6.2 5.2 6.2 12–13
LB1/1 LB1 C1R VII 59 5 8.1 — 8.0 (6.1) 7.8 20.3
LB1/1 LB1 C1L VII 59 6b7.8 — 7.8 6.4 7.7 19.8
LB1/1 LB1 P3R VII 59 3 6.8 7.0 9.7 4.7 9.4 15.9
LB1/1 LB1 P3L VII 59 4 6.9 7.0 8.8 5.0 8.6 14.0
LB1/1 LB1 P4R VII 59 3 6.9 6.9 (8.8)c5.1 (8.1) 16.0
LB1/1 LB1 P4L VII 59 4 7.1 7.1 (8.9)c5.0 8.7 14.9
LB1/1 LB1 M1R VII 59 5 9.3 9.3 11.3 7.4 10.9 12.8, 13.4, 11.9d
LB1/1 LB1 M1L VII 59 5 9.0 9.0 11.5 — — —
LB1/1 LB1 M2R VII 59 4 9.4 9.4 10.9 7.4 10.1 13.7, 13.1, 12.3d
LB1/1 LB1 M2L VII 59 4 9.3 9.3 10.8 7.3 10.3 —
Mandible
LB1/1 LB1 I1L VII 59 6 3.8 — 5.0 3.5 5.0 14.0
LB6/14 LB6/1? I1L XI 53 5 3.7 — 5.0 3.3 5.0 12.2
LB1/2 LB1 I2R VII 59 4 4.7 — 5.7 4.0 5.8 14.3
LB1/2 LB1 I2L VII 59 5.5 4.3 — 5.7 3.8 5.7 —
LB6/1 LB6/1 I2R XI 51 4 4.5 — 5.2 — 5.2 —
LB1/2 LB1 C1R VII 59 4.5 6.7 — 7.4 6.2 7.2 17.2
LB1/2 LB1 C1L VII 59 5.5b6.7 — 7.4 6.0 7.2 16.4
LB6/1 LB6/1 C1R XI 51 4 6.3 — 6.7 — 6.5 —
LB6/1 LB6/1 C1L XI 51 4 6.1 — 6.7 5.2 6.6 —
LB1/2 LB1 P3R VII 59 4 8.6 8.6 8.5 6.4 8.3 12.0
LB1/2 LB1 P3L VII 59 5 8.5 8.5 8.4 6.6 8.5 13.6
LB2/2 LB2/2 P3L IV 43D 4 8.6 8.6 8.7 7.0 8.5 12.4
LB6/1 LB6/1 P3R XI 51 4 8.0 8.2 7.8 — — —
LB6/1 LB6/1 P3L XI 51 4 8.0 8.2 7.8 6.6 7.8 (10.7)
LB6/1 LB6/1 P4R XI 51 4 6.6 6.7 7.8 — — —
LB6/1 LB6/1 P4L XI 51 4 6.1 (6.7) 7.7 5.0 6.7 13.2
LB15/1 LB15/1 P4R III 48 4 7.2 (7.7) 8.5 5.9 7.7 14.3
LB1/2 LB1 M1R VII 59 3.5 9.5 9.6 10.6 8.5 9.1 12.3, 12.5e
LB1/2 LB1 M1L VII 59 6 9.0 (9.6) 10.4 (7.9) 9.1 12.5, 12.4e
LB6/1 LB6/1 M1R XI 51 5b8.8 9.1 10.0 — — —
LB6/1 LB6/1 M1L XI 51 4 9.0 9.3 10.0 — 8.8 —
LB1/2 LB1 M2R VII 59 3.5 9.9 10.1 10.1 8.6 8.8 12.5, 11.8e
LB1/2 LB1 M2L VII 59 4 9.7 10.1 10.0 9.0 9.0 12.5, 11.7e
LB6/1 LB6/1 M2R XI 51 4 9.5 9.8 9.6 — — —
LB6/1 LB6/1 M2L XI 51 3 9.6 9.8 9.5 — — —
LB1/2 LB1 M3R VII 59 4 8.9 8.9 9.5 7.9 8.3 15.8, 14.5e
LB1/2 LB1 M3L VII 59 3 9.8 9.8 9.6 8.5 8.2 15.1, 14.0e
LB6/1 LB6/1 M3R XI 51 3 8.9 9.0 8.8 — — —
LB6/1 LB6/1 M3L XI 51 3 8.9 8.9 8.5 — — —
a Measurements based on CT are in italics.
b Revised from Jungers and Kaifu et al. (2011).
c Estimated original crown diameter allowing for wear.
d Lengths for mesiobuccal, distobuccal, and lingual roots, respectively.
e Lengths for mesial and distal roots, respectively.
DENTAL REMAINS OF HOMO FLORESIENSISVol. 123, 2015 131
values are reported for mesiodistal (MD) crown diameters.
Length of a root was measured on the buccal surface (from
the cervical midpoint to the root tip) except for the lingual
root of the upper molar where the length was taken on the
lingual surface. A single observer (Y.K.) measured the origi-
nal specimens with reference to isolated plaster casts of each
tooth, using a Mitsutoyo digital sliding caliper with pointed
tips. When necessary, the thickness of dental calculus depos-
it was estimated to obtain the original crown diameters.
Cervical diameters were measured either by a sliding caliper
(by Y.K.) or from the CT scan (by R.T.K.) following the
methods of Hillson et al. (2005). Severity of occlusal wear
was scored by the categories (0–8) proposed by Smith
(1984).
High-resolution CT scan was obtained using the microfo-
cal X-ray CT system TX225-ACTIS (Tesco Co.) in April
Figure 1. Maxillary dentition of LB1. Occlusal views (a). Buccal (b) and lingual (c) views of the right dentition. Buccal (d) and lingual (e) views
of the left dentition. Micro-CT scan indicates that right M3 was congenitally absent in this individual. Note that both right and left P4s are rotated
bilaterally. Scale = 10 mm.
Figure 2. Mandibular dentition of LB1. Occlusal views (a). Buccal views of the anterior parts of the right (b) and left (c) dentitions. Lingual
views of the anterior parts of the right (d) and left (e) dentitions. Distal view of the left P3 root (f). Buccal view of the right molars (g). Lingual view
of the left molars (h). (e) and (f) were taken when we disassembled the left mandibular body to correct its reconstruction. Scale = 10 mm.
Y. KAIFU ET AL.132 AnthropologicAl Science
2009 at the University Museum, The University of Tokyo
(Kaifu et al., 2011). Prior to the scan, the system was cali-
brated with known size phantoms. Linear measurement error
in a horizontal plane was less than 0.1% (Kubo et al., 2008).
Original scans were taken at 130 kV and 0.20 or 0.17 mA
with a 0.5 or 1 mm thick copper plate prelter to lessen
beam-hardening effects. Other scanning parameters included
a 512 × 512 matrix, 50 or 80 microns pixel size, and 50 or 80
microns slice thickness and interval. Avizo 8.0 software (FEI
Visualization Sciences Group) was used for segmentation of
the enamel and the dentine to visualize the EDJ surfaces.
Morphological Description
Table 1 reports our measurements of the crowns and
roots. These measurements are in most cases slightly lower
(0.1–0.9 mm, or more in a few cases) than those reported by
Brown et al. (2004) and Brown and Maeda (2009). It should
be noted that there is substantial asymmetry in the degree
and pattern of occlusal wear in LB1. This is because of the
horizontally twisted (rotated) occlusion between the maxil-
lary and mandibular dentitions in this individual, which was
probably caused by posterior positional (deformational)
plagiocephaly (Kaifu et al., 2009, 2010).
Different individuals exhibit similar dental characters to
each other unless noted otherwise. Anatomical terminology
mostly follows Johanson et al. (1982). The following abbre-
viations are used:
MD = mesiodistal(ly)
BL = buccolingual(ly)
LL = labiolingual(ly)
CEJ = cemento–enamel junction
EDJ = enamel–dentine junction
IPF = interproximal facet
Mmr = mesial marginal ridge
Dmr = distal marginal ridge
ASUDAS = Arizona State University Dental Anthropolo-
gy System (Turner et al., 1991)
Incisors
I1: LB15/2 (left: Figure 4)
The specimen is complete except for partial exfoliation of
the cementum from the root surface and a small damage at
the root tip. Extensive occlusal wear has eliminated most of
the crown, leaving a at, slanting occlusal surface that
reaches the CEJ on the distal face but not on the mesial face.
Mesial and distal IPFs are not present. Small patches of den-
tal calculus are attached around the CEJ.
Viewed occlusally, the preserved cervical portion of the
labial enamel surface is nearly straight, suggesting a limited
labial MD convexity of the crown when unworn. In mesial
or distal view, the labial cervical enamel is attened so that
it continues straight to the root surfaces. The remaining
small portion of the lingual enamel suggests that the gingival
eminence was weak. The ASUDAS scores for shoveling and
tuberculum dentale are not available due to the wear.
At the cervical level, the cross-sectional shape of the root
is a MD compressed, rounded triangle with the mesiolingual
face slightly wider and more convex than in the distolingual
face. The root is short (our estimate of the labial length is
12–13 mm), stocky, and retains its thicknesses for 4 mm
above the CEJ before tapering toward the broken tip. This
root length is largely comparable to the mean root length
(distance between the root apex and the center of the cervical
plane) of a recent modern human sample reported by Le
Cabec et al. (2013) (12.9 mm).
I2
This tooth is not represented in the currently available
Liang Bua collection. Although Brown et al. (2004) inferred
Figure 3. Mandibular dentition of LB6/1. Occlusal views (a). Buccal (b) and lingual (c) views of the right dentition. Mesial view of left C1 (d).
Mesial (upper image) and apical (lower image) views of left P3 (e). Mesiolingual view of left P4 (f). (e) and (f) were taken when Y.K. disassembled
the left mandibular body for conservation purposes. Scale = 10 mm.
DENTAL REMAINS OF HOMO FLORESIENSISVol. 123, 2015 133
that the I2 was much smaller than I1 in LB1, this view is not
supported by the partially preserved, moderate-sized right I2
alveolus seen in a CT section (Figure 5a).
I1: LB1 (left: Figure 2), LB6/14 (left: Figure 4)
The former is complete and is in the alveolus of the LB1
mandible. LB6/14 is an isolated tooth that was not described
by Brown and Maeda (2009). It has a slight damage at its
root tip. Its size, morphology, wear, and preservation suggest
that the tooth is from the LB6/1 mandible, but this possibili-
ty cannot be ascertained due to the damage on the latter. In-
cisal wear is extensive in both specimens. The areas of the
CEJ and exposed root surfaces are in part covered by thin
layers of dental calculus.
The preserved basal portions of the crowns do not show
marked lateral aring. Viewed mesially or distally, as in the
I1, the labial cervical enamel is at and continues straight to
the labial root surface. A shallow but distinct vertical furrow
is present on the labial crown face of each specimen. It
reaches and disappears at the CEJ. This may indicate that the
crown had assumed double-shoveling when unworn. On the
lingual face, the gingival eminence is weak and almost ab-
sent.
The root can be examined directly (LB6/14: Figure 4) or
with the CT scan (LB1: Figure 6b). The root is MD com-
pressed and has a vertical furrow on the distal face. Viewed
mesially or distally, the lingual (LB6/14) or both the labial
and lingual (LB1) outlines are markedly convex so that the
maximum LL diameter (5.8–6.0 mm), which measures
around the middle of the root length, considerably exceeds
the LL diameter at the CEJ (5.0 mm). The root lengths,
14 mm (LB1) and 12.2 mm (LB6/14), are largely compara-
ble to the mean values for various modern human samples
(11–13.4 mm: Suwa et al., 2011, Table 3).
I2: LB1 (right and left: Figure 2), LB6/1 (right: Figure 3)
The three specimens are in their alveoli. They are com-
plete except for damages at the root apices of LB1. Wear is
moderate (LB1 right, LB6/1) to extensive (LB1 left). The
left tooth of LB1 exhibits a ‘stepped’ occlusal wear in which
the lingual half is worn more heavily than the labial half. The
CEJs are in part covered by thin layers of calculus.
Basic anatomy is similar to I1, although I2 is larger and its
labial crown surface faces slightly distally. Viewed labially,
the crown is slender MD. On the unworn cervical half of the
labial face, a vertical furrow seen in the I1s does not develop,
but the surface is attened MD. As in the other incisors, the
labial enamel and root surfaces continues straight with little
swelling of the cervical enamel. On the smooth lingual face,
the gingival eminence is weak. Basal parts of the Mmr and
Dmr are barely recognizable near the worn incisal margin of
LB6/1, although the wear prevents ASUDAS scoring for this
trait.
CT scan of LB1 (Figure 6a) indicates that its root form is
generally similar to that of the I1s. The MD compressed root
presents a longitudinal groove on the distal face. The labial
root face shows marked vertical convexity, and LL diameter
reaches its maximum around cervical one-third of the root
length. The root length of the LB1 I2, 14.3 mm, is largely
comparable to the mean values for various modern human
samples (12–14.7 mm: Suwa et al., 2011, Table 3).
Canines
C1: LB1 (right and left: Figure 1)
Both teeth are in their sockets. Each of them is complete
except for a transverse crack in the middle of the root (visi-
ble in the CT scan). Occlusal wear is extensive, leaving only
1–2 mm of the mesial enamels. The right tooth is less worn,
particularly on its labial side, so that its occlusal surface is
strongly sloping lingually. Mesial IPF is lacking. The crown
contacts with the P3 at the distal aspect of its Dmr. An inter-
mittent band of calculus covers the CEJ or root surface,
particularly on the labial side.
The severe wear obscures the labial crown contour, but
a low distal shoulder is evident in the less worn right tooth.
Viewed occlusally, the crown outline is asymmetric with
the convex mesiolabial and attened distolabial aspects. On
the less worn right tooth, a MD broad, rounded gingival
Figure 4. Four isolated teeth of H. oresiensis from Liang Bua: LB
15/2 (left I1), LB 6/14 (left I1), LB 2/2 (left P3), and LB 15/1 (right P4).
From left to right, occlusal (o), mesial (m), labial/buccal (l/b), and dis-
tal (d) views. Scale = 10 mm.
Y. KAIFU ET AL.134 AnthropologicAl Science
eminence dominates the lingual aspect. It does not project
lingually over the cervical line, but smoothly continues to
the prominent basal part of the lingual median ridge. Dmr is
a weak, thin line restricted near the crown’s distal shoulder.
The labial crown face exhibits slight wrinkling.
CT scan indicates that the root is single and robust, with
its labial (right tooth) or lingual (left tooth) face showing
marked vertical convexity. The cross section is MD com-
pressed, oval with the labial section broader than the lingual
(Figure 5a–d). The lengths of the root, 20.3 mm (right) and
19.8 mm (left), are greater than the modern human average
(16.1 mm) and are slightly shorter than the means for Nean-
derthals (22.5 mm) and the Early Pleistocene H. erectus
from Sangiran, Java (21.5–21.6 mm) (Le Cabec et al.,
2013).
C1: LB1 (right and left: Figure 2), LB6/1 (right and left:
Figure 3)
The four specimens from two individuals are complete
and in their alveoli. Approximately half (LB6/1) or slightly
more (LB1) of the crown heights have been lost by wear that
produces largely at occlusal surfaces. Mesial and distal
IPFs are present in the less worn LB6/1 C1s, but mesial IPFs
were probably lost in LB1 by wear.
Figure 6. CT sections of the LB1 mandibular incisor roots. Sagittal
(top) and horizontal (right) sections are shown for right I2 (a) and left I1
(b). The lower left image for each tooth is a surface rendered image of
the labial side, which indicates the levels of the sections (vertical and
transverse lines). Me = mesial, Di = distal, La = labial, Li = lingual.
Note the convex labial and lingual root outlines similar to LB6/14, the
isolated I1 shown in Figure 4.
Figure 5. CT sections of the dental roots of LB1.
The level for each section is indicated in the image
above the relevant section. Upper left (sections a and b):
left maxillary dentition. Upper right (sections c and d):
right maxillary dentition. Lower left (sections e and f):
left mandibular dentition. Lower right (sections g and
h): right mandibular dentition.
DENTAL REMAINS OF HOMO FLORESIENSISVol. 123, 2015 135
The distal crown shoulder is preserved on the relatively
unworn left C1 of LB6/1. Its tip is located 3.5 mm above the
CEJ, and is distinctly lower than the worn mesial shoulder
that measures >5 mm. Apart from this asymmetry, the wear
prevents assessment of labial crown contour. Viewed occlu-
sally, the gently convex buccal crown contour is nearly
symmetric but the lingual contour is asymmetric due to the
distally located gingival eminence. The labial face is charac-
terized by ne, longitudinal wrinkles and a slight thickening
of the cervical enamel (evident in the calculus-free cervical
enamels of LB6/1: Figure 3b). There are no longitudinal
grooves along the mesial and distal aspects of the labial
crown surfaces. The lingual face is dominated by a broad,
prominent median ridge that continues smoothly from the
gingival eminence. Weak lingual Mmr and Dmr are repre-
sented by attened areas beside the median ridge, and the
mesial and distal lineal foveae are no more than shallow de-
pressions.
CT scan (Figure 5e, g) indicates that the root is single and
relatively robust, with vertically convex lingual face and
vertically straight labial face. The root lengths for LB1,
17.2 mm (right) and 16.4 mm (left), are comparable to the
mean values for various modern human samples (13.7–
18.2 mm: Suwa et al., 2011, Table 3).
Premolars
P3: LB1 (right and left: Figure 1)
The specimens are complete except for a crack at the root
of the left tooth, which has dislocated its crown slightly me-
sially and occlusally. The left tooth is worn at with dentine
islands of moderate sizes exposed on the two cusps. On the
less worn right tooth, the lingual cusp (protocone) is worn
at whereas the buccal cusp (paracone) remains high. Mesi-
al and distal IPFs are present, although the left tooth has lost
its contact with the P4 due to the dislocation. A thin layer of
calculus covers the cervical half of the buccal crown surface
of the right tooth. Calculus is present, but less marked on its
lingual face and on the left tooth.
The occlusal outline, partially obscured by the interproxi-
mal wear, is ovoid with MD diameters slightly greater across
the buccal than the lingual crown. The crown is largely sym-
metrical MD, with the lingual cusp tip shifted slightly mesi-
ally. Detailed morphology of accessory occlusal ridges is
unknown due to the wear, but some features are evident. The
longitudinal groove is completely worn away, whereas the
anterior and posterior fovea partially remain unworn. On the
EDJ surface, there is a sharp crest that connects the two
cusps (Figure 7a), indicating the presence of a well-
developed transverse crest when the crown was unworn. The
low Mmr and Dmr are incised by grooves emanating from
the mesial and distal foveae, respectively. On the relatively
vertical and smooth buccal face, the area around the CEJ is
obscured by the calculus deposition, but the presence of a
narrow enamel band along the cervical line is evident on the
left tooth. Mesial and distal buccal grooves are absent al-
though the area of the former is attened in the left tooth.
The lingual face is also smooth and steep.
CT scan (Figure 5a–d) indicates that the root has a buccal
component and a lingual component that are extensively
fused together to form a BL broad, plate-like root complex.
The buccal component is longer than the lingual one. Each
component has its own, single root canal in the right tooth,
whereas the canals are partially connected to each other in
the left tooth. A vertical furrow intervenes between the two
components on the mesial and distal root faces. The furrow
is deeper on the mesial than on the distal faces, and is more
marked on the right tooth.
P4: LB1 (right and left: Figure 1).
The right tooth is complete. In the left tooth, enamel is
chipped off from the distobuccal corner of the crown, and
the root is cracked so that the crown is dislocated slightly
occlusally. Both teeth are rotated ~90° and their original
buccal faces orient mesially in the tooth rows. Occlusal wear
is heavier on the left than on the right tooth. Both teeth con-
tact with their mesial and distal adjacent teeth, although the
mesial side of the left tooth presently lacks the contact due to
the postmortem mesial dislocation of the P3. Calculus depo-
sition is marked on the buccal but much less on the lingual
sides.
The occlusal outline, partially obscured by wear, is ellip-
tical with nearly equal paracone and protocone cusp
breadths. The protocone cusp tip is located slightly mesially
to the crown’s midline (lingually in the dentition because of
the rotation). The worn occlusal enamel surface of the left
tooth is featureless except for a dentine exposure at the par-
acone tip. On the less worn right tooth, the mesial half of the
longitudinal groove has been worn away but its distal half as
well as the anterior and posterior foveae remain intact at
least partially. On the EDJ surface, there is a sharp crest that
connects the two cusps (Figure 7b), indicating that a well-
developed transverse crest was present when the crown was
Figure 7. Occlusal views of the EDJ surfaces of the right P3 (a), P4 (b), M1 (c), and M2 (d).
Y. KAIFU ET AL.136 AnthropologicAl Science
unworn. A short accessary ridge extends from the paracone
cusp tip toward the posterior fovea. The low, incomplete Dmr
remain unworn, whereas the Mmr is obliterated except for
its buccal (mesial in the tooth row) segment. Both the buccal
and lingual faces rise steeply from the CEJ. Mesial and distal
buccal grooves are absent on the less worn right tooth.
CT scan (Figure 5a–d) indicates that the root is largely
similar to those of the P3s. The buccal and lingual compo-
nents of similar lengths are extensively fused to form a BL
broad, plate-like root complex. A longitudinal furrow sepa-
rating the two components is deeper on the mesial face than
on the distal face.
P3: LB1 (right and left: Figure 2), LB2/2 (left: Figure 4),
LB6/1 (right and left: Figure 3)
LB2/2 is an isolated tooth. The other four teeth are in their
sockets. All the ve specimens are complete. Flatly or con-
cavely worn occlusal surfaces are slightly (LB1, LB6/1) to
strongly (LB2/2) beveled distally. Moderate occlusal wear
exposes a large dentine island on the buccal cusp of each
specimen. In the tooth row, the P3s of LB1 are slightly rotat-
ed so that their mesiobuccal surfaces contact with the ca-
nines, whereas the LB6/1 P3s contact with the canines at
their pointed mesial end of the mesial occlusal ridge (thus
these P3s are not rotated, in contrast to the condition in LB1).
Mesial and distal IPFs are also present in the isolated LB2/2,
but the former is located on the mesiobuccal surface, indicat-
ing a slight rotation as in LB1. The buccal and lingual faces
of these teeth are variably covered by dental calculus with
the most severe cases observed in LB1.
Viewed occlusally with the mesial and distal protoconid
(buccal cusp) ridge oriented transversely, the crown is a dis-
torted fan composed of a gently (LB1, 6/1) or moderately
(LB2/2) convex and MD long buccal aspect and distally ta-
pering lingual crown. It is extensive both MD and BL, al-
though the MD diameter for LB1 reported by Brown and
Maeda (2009), 10.4 mm, is an overestimate irrespective of
crown orientation. The prominent buccal cusp supports MD
extensive mesial and distal protoconid ridges (evident from
the preserved enamel and thick lines of exposed dentine).
The lingual cusp (metaconid) is small and is situated distally,
shortly mesial to (LB6/1) the crown’s distolingual corner.
The two cusps are connected by a strong transverse crest that
runs diagonally toward the crown’s distolingual section,
almost in line with the cervical long axis. Mesial to this crest
and lingual to the mesial protoconid ridge, the entire mesio-
lingual segment of the occlusal surface slants and faces me-
siolingually to form an extensive, sloping, generally at but
nely wrinkled enamel surface. This beveled surface ex-
tends inferiorly to the level ~1 mm short of the CEJ. The
short, poorly developed Mmr and the shallow, linear anterior
fovea are restricted to the mesial aspect of this sloping sur-
face. On the distal crown, the talonid is represented by a low,
featureless Dmr that is restricted to the distal crown margin
and delimits a BL oriented, linear posterior fovea. The buc-
cal face slopes lingually and curves moderately (LB1,
LB6/1) or strongly (LB2/2) in a vertical section. There are
no distinct mesial/distal buccal grooves, but the entire buc-
cal face is irregularly wrinkled. A 1 mm wide band of enam-
el develops along the cervical line on the buccal face of LB1
(evident on the calculus-free left side: Figure 2c). The buc-
cal cervical line dips inferiorly so that the crown is deeper on
the buccal than on the lingual aspect.
Based on low-resolution CT scans, Brown and Maeda
(2009) described that LB1 and LB6/1 had two roots ar-
ranged in a ‘MB + D’ pattern (a circular mesiobuccal root
and a larger plate-like distal root) dened by Wood et al.
(1988), and the isolated LB2/2 a Tomes’ root with a deep
mesiolingual cleft resulting from fusion of the original me-
siobuccal and distal components. Our high-resolution CT
scan and direct observation (Figure 2f) of LB1 as well as
direct observations of LB6/1 (Figure 3e) and LB2/2 (Fig-
ure 4) conrm these assessments. The two root components
of LB1 and LB6/1 are closely set to each other, particularly
in their apical portions (Figure 3e (lower), Figure 5f, h). The
cervical outline of the H. oresiensis P3 root is elliptical with
its long axis being almost in line with the transverse crest on
the crown. The root lengths (10.7–13.5 mm: Table 1) are
shorter compared to the mean values for various modern
human samples (12.7–17.0 mm: Suwa et al., 2011, Table 3).
P4: LB6/1 (right and left: Figure 3), LB15/1 (right: Figure 4)
LB15/1 is an isolated tooth that was not described by
Brown and Maeda (2009). The LB6/1 P4 are in their alveoli,
although the mesial root of the left tooth is now visible at the
break of the mandible. The three specimens are complete.
Occlusal wear is moderate, and the less worn LB6/1 right
tooth still retains some cuspal relief. All three teeth have
mesial and distal IPFs that reach the occlusal surfaces. Small
patches of calculus remain on the CEJ and enamel surfaces.
The crown of LB15/1 is larger than LB6/1, and is proba-
bly so compared to LB1 where the space between its left P3
and M1 measures <7 mm. The occlusal contour is asymmet-
ric with slightly protruding mesiobuccal and distolingual
corners. The buccal and lingual cusps are located mesially,
and are connected by a well-developed transverse crest
(evident in LB6/1 and may be so in the worn LB15/1). The
Mmr is incised by a shallow groove emanating from the an-
terior fovea (observable only in the right tooth of LB6/1).
The talonid basin is large, situated at a low level, and sup-
ports a linear distal fovea and a thick Dmr. The Dmr contin-
ues to the crown’s distolingual corner, smoothly exes mesi-
ally to extend toward the lingual cusp tip. Mesial and distal
buccal grooves are absent on the smoothly convex buccal
crown face. The lingual face is unremarkable.
The remaining alveolus for the LB1 left P4 (Figure 5f, e)
and mesial aspect of the LB6/1 left P4 (Figure 3f) indicate
that these two teeth are categorized as Tomes’ root with a
deep mesiolingual cleft. The partial fusion of the LB6/1 root
is not clearly recognized in Figure 13 of Brown and Maeda
(2009) who erroneously reported that this tooth shows
MB + D root pattern. The isolated LB15/1 has a single, ro-
bust root (Figure 4). Its MD diameter is greatest at the cervix
(5.9 mm) and tapers toward the apex, whereas the large BL
diameter remains almost the same in its cervical half (7.7–
7.2 mm). A shallow vertical groove is present on the distal
root face, but the mesial face is generally convex. The root
lengths (13.2–14.3 mm: Table 1) are comparatively shorter
than the mean values for various modern human samples
(12.9–16.5 mm: Suwa et al., 2011, Table 3).
DENTAL REMAINS OF HOMO FLORESIENSISVol. 123, 2015 137
Molars
M1: LB1 (right and left: Figure 1)
The specimens are complete except for cracks at the root
of the left tooth. The lingual half of the occlusal surface is
severely worn, resulted in the exposure of a MD continuous,
concave dentine surface in a form of ‘cupped wear.’ The
buccal half is less worn particularly in the right tooth. BL
extensive mesial and distal IPFs are present. The enamel
surface and CEJ are partially covered by thin calculus de-
posits, particularly on the buccal face of the right tooth.
The occlusal contour is a BL elongated square with a wid-
er mesial segment that results from lingual projection of the
protocone base. The wear has eliminated most of the occlu-
sal ridge and groove structures, but the remaining grooves
indicate that both the right and left teeth had four major
cusps. On the right tooth, much of the buccal transverse oc-
clusal groove remain. On the left tooth, the same groove can
be traced only by an indentation on the buccal occlusal mar-
gin and two tiny, shallow pits near the crown midpoint, the
lingual one of which probably corresponds to the central
fovea. Near the distal occlusal margin and lingual to a tiny
dentine exposure at the metacone apex, a small pit and a
short groove remain. The latter is probably a part of the
groove demarcating the metacone and hypocone. The prom-
inent distolingual corner of the crown suggest strong devel-
opment of the hypocone (ASUDAS grade 4 or 5). The buc-
cal face is obscured by the calculus but the presence of a
0.5 mm wide enamel band is evident along the cervical
enamel. The smooth lingual crown face is marked by a lin-
gual projection of the protocone base, but no expression for
the Carabelli’s trait.
CT scan (Figure 5a–d) indicates that the mesiobuccal,
distobuccal, and lingual roots are divergent from each other.
The lingual face of the lingual faces distolingually (Fig-
ure 1a). The above described lingual swelling of the proto-
cone base is apparently associated with this ‘twisted’ root
orientation.
M2: LB1 (right and left: Figure 1)
The crown on the left side is dislocated slightly buccally
at a large crack on its root. A small crack is also present in
the distobuccal root of the right tooth. Otherwise, the two
specimens are complete. The occlusal surface is worn at
with dentine exposed at two cusps on the right (protocone
and hypocone) and left (protocone and paracone) teeth. Dis-
tal IPF is present only on the left tooth. It measures 2.2 mm
both BL and vertically. A few thin or small patches of calcu-
lus remain mainly on the buccal crown face of the right
tooth.
These are four-cusped teeth. The occlusal outline is a trap-
ezoid with broader mesial section as well as distally extend-
ed hypocone but reduced metacone, which is more marked
in the right tooth. Much of the occlusal enamel structures
have been lost by wear. The buccal transverse occlusal
groove, which runs lingually and mesially, is clearly (right)
or partially (left) recognizable. The central fovea is repre-
sented by a shallow pit near the center of the left tooth, but
no such feature remains on the right tooth. Other occlusal
groove/ridge structures are not evident, although our CT-
based examination of the EDJ topology indicates that tiny
pits on the lingual aspect of the right tooth represent the lin-
gual transverse occlusal groove. A 0.5 mm enamel band is
present along the cervical line of the buccal face. Although
the lingual face shows marked lingual projection mesially as
mentioned above, the entire surface is smooth and Carabel-
li’s feature is absent.
CT scan (LB1 only: Figure 5a–d) indicates that the root
structure is largely similar to the M1s. There are three roots
(mesiobuccal, distobuccal, and lingual) and the lingual root
exhibits distal twisting. However, compared to the M1 con-
dition, these roots are less divergent, the level of split of the
three roots is more apical, and the mesiobuccal and lingual
roots are fused extensively in the left tooth.
M3
Morphology of this tooth can be inferred only from the
left alveolus of LB1 (Figure 5a). Its small, conical form sug-
gests that this tooth had a much smaller crown than M1 and
M2 (Brown et al., 2004).
M1: LB1 (right and left: Figure 2), LB6/1 (right and left:
Figure 3)
These teeth are in their sockets. Small portions of enamel
have been chipped off at the mesiolingual and distobuccal
corners of the left tooth of LB1 either before or after the in-
dividual’s death. Otherwise, the four specimens are com-
plete. Mesial and distal IPFs are present in all of the teeth.
Asymmetric wear pattern in LB1 (less worn lingual cusps of
the right tooth, loss of mesial and distal enamels in the left
tooth, etc.) is primarily because of the unbalanced, rotated
occlusion in this individual (Kaifu et al., 2009). Calculus
depositions are seen around the buccal and lingual CEJs. It
is extremely thick and extensive on the lingual face of the
right tooth of LB1. A lled hole on the buccal face of the
mesial root of the LB6/1 right tooth was made to take sam-
ples for DNA extraction.
The occlusal contour is a MD short, rounded square with
slightly protruding mesiobuccal and distolingual corners.
Much of the enamel surface structures have been worn away,
but the partially remaining occlusal grooves, the arrange-
ments of the exposed dentine patches, as well as the EDJ
surface (Figure 7c, LB1 only) indicate that these are four-
cusped teeth with no development of hypoconulid, Cusp 6,
and Cusp 7. The lingual transverse occlusal groove, parts of
the buccal transverse occlusal groove, and the mesial portion
of the distal longitudinal groove (LB6/1 only) are traceable.
A mesiobuccally oriented, short groove at the central fovea
(evident on the right tooth of LB1 and possibly both teeth of
LB6/1) suggest a substantial contact relationship between
the metaconid and hypoconid. Entoconid is the lingually re-
stricted, smallest cusp. Metaconid is also restricted BL, and
the two buccal cusps dominate on the occlusal surface. At the
buccal termination of the transverse lingual occlusal groove,
the central fovea is shallow with no development of a deep
pit. A sharp, high, and continuous crest connecting the proto-
conid and metaconid on the EDJ surface of LB1 (Figure 7c),
which corresponds to grade 3 of Bailey et al. (2011: Fig. 4),
strongly suggests the presence of a mid-trigonid crest before
the wear (Bailey et al., 2011; Martínez de Pinillos et al.,
2014; the denition of ‘mid-trigonid crest’ follows Martínez
Y. KAIFU ET AL.138 AnthropologicAl Science
de Pinillos et al., 2014). A protostylid is absent in the LB1
right M1, and at least not evident on the moderately worn
other M1s. There is a faint, narrow band (LB1) or swelling
(LB6/1) of enamel along the cervical lines.
CT scan (LB1 only: Figure 5e–h) indicates that M1 has
plate-like mesial and distal roots that are distinctly separated
from each other. The BL broad mesial root is bid apically.
The BL width of the distal root decreases apically. The me-
sial root lengths of LB1 (12.3–12.5 mm: Table 1) are shorter
than the mean values for various modern human samples
(13.2–14.9 mm: Suwa et al., 2011, Table 3).
M2: LB1 (right and left: Figure 2), LB6/1 (right and left:
Figure 3)
The four specimens are complete. The lingual cusps are
variably less worn and the original cuspal reliefs and groove
patterns can be read to some extent. Mesial and distal IPFs
are present, and some amounts of calculus deposition are
seen, particularly on the lingual faces. A lled hole on the
buccal face of the mesial root of the LB1 right M2 was made
to take samples for DNA extraction.
The occlusal outline is a rounded square with slight mesi-
al projection of the protoconid and lingual projection of the
entoconid base. These are four-cusped teeth. Two occlusal
grooves demarcating the entoconid are preserved. Much of
the other occlusal grooves have been worn away, but their
positions can be restricted with reference to the anterior
fovea (LB6/1), distribution of the exposed dentine for the
protoconid and hypoconid (LB1 and LB6/1), the remaining
groove between the metaconid and hypoconid (LB1), and
the topology of EDJ seen in the CT scan (LB1). Cusp ar-
rangement is similar to the M1s: Entoconid is the lingually
restricted, smallest cusp. Metaconid expands slightly more
buccally than the entoconid, but the two buccal cusps are
more expansive BL on the occlusal surface. Metaconid and
hypoconid contact to each other via a short occlusal groove
in LB1, and this seems to be also the case for LB6/1. Cusp 7
assumes a form of a triangular depressed area immediately
distal to the main occlusal ridge of the metaconid in LB1,
whereas this structure is not clearly developed in LB6/1. The
anterior fovea, evident in LB6/1, is a small, triangular notch
that opens mesially. A sharp, high, and continuous crest con-
necting the protoconid and metaconid on the EDJ surface of
LB1 (Figure 7d), which corresponds to grade 3 of Bailey et
al. (2011: Fig. 4), strongly suggests the presence of a mid-
trigonid crest before the wear (Bailey et al., 2011; Martínez
de Pinillos et al., 2014). No deep pit develops at the central
fovea as in the M1s. The distal longitudinal groove disap-
pears before reaching the distal occlusal margin without
forming a distinct posterior fovea. The buccal crown face is
generally smooth except for a narrow, continuous enamel
band stretching along the cervical line over the protoconid
and hypoconid bases. The lingual face is vertically more
convex compared to the M1 condition.
CT scan (LB1 only: Figure 5e–h) indicates that the mesial
and distal root components are fused along their buccal sides
to form a short, cylindrical root complex with a C-shaped
cross section. The mesial root length of LB1 (12.5 mm) is
shorter than the mean values for various modern human
samples (13.1–15.1 mm: Suwa et al., 2011, Table 3).
M3: LB1 (right and left: Figure 2), LB6/1 (right and left:
Figure 3).
The four specimens are complete except for minor cracks
at the root of the LB1 right tooth. The teeth contact with the
M2s, although the LB1 M3s are bilaterally rotated ~30° so
that their mesial faces orient lingually. A lled hole on the
lingual root face of the LB1 left tooth was made to take sam-
ple for DNA extraction.
The occlusal contour is a rounded rectangle with the lin-
gually protruded entoconid (LB6/1) or a pear-shape with the
broader distal crown (LB1). These are four-cusped teeth.
The occlusal groove arrangement is close to a ‘+’ pattern
although metaconid and entoconid slightly contact each oth-
er. The longitudinal groove and the transverse groove are
situated lingually and distally relative to the crown center,
respectively, so that protoconid is the largest cusp. The me-
sial segment of the longitudinal groove incises the Mmr,
whereas its distal segment does not reach the distal occlusal
margin. The anterior fovea is a triangular pit opening mesi-
ally. The central fovea is not very deep, and there is no pos-
terior fovea. Cusp 7 is dened as a triangular depressed area
distal to the main occlusal ridge of the metaconid. A thin
enamel band is expressed along the cervical line on the buc-
cal crown face of LB1, whereas the same face is generally
smooth in LB6/1. The lingual crown face is vertically more
convex than in the anterior molars.
CT scan (LB1 only: Figure 5e–h) indicates that the M3 of
LB1 has a single, pyramidal, long root with a deep buccal
cleft. The mesial root lengths of LB1 (15.8 and 15.1 mm) are
greater than the mean values for various modern human
samples (11.2–14.4 mm: Suwa et al., 2011, Table 3).
Dentition and occlusion
LB1
LB1 had a completed permanent dentition, but had lost all
the maxillary incisors, left M3, right I1, and left P4 postmor-
tem (Brown et al., 2004). Brown and Maeda (2009) reported
that their CT scans suggested the presence of a very small
M3 odontome within the alveolar bone of the LB1 maxilla.
Our micro-CT scan identies no such evidence (the white
materials behind the M2 in Figure 5a–d are probably dirt),
indicating that this tooth was congenitally absent as reported
originally (Brown et al., 2004). Brown and Maeda (2009)
also inferred that the right P4 was originally present in the
jaw, but the right P3 bears no distal IPF and a remnant of the
P4 or its alveolus is not observed in our CT scan (Figure 5g,
h). We suggest that the right P4 was congenitally absent and
a single, BL elongated (4.1 mm wide) mesial IPF on the M1
was a facet for the exfoliated dm2 (Kaifu et al., 2009). The
space between the right P3 and M1 has been reduced to
1.8 mm by mesial migration of the latter.
The maxillary dental arcade shape is described as nearly
square with sharp exion at the canines as seen in H. habilis,
H. ergaster, and Sangiran H. erectus, but is different from
the more rounded arcades seen in Kabwe and Zhoukoudian
13 (Kaifu et al., 2011). The close proximity between the
right I2 and C1 alveoli evident in a CT section (Figure 5a)
does not support the previous inference for a diastema in this
individual (Brown et al., 2004). At variance with the previ-
ous report (Brown and Maeda, 2009), the Curve of Spee is
DENTAL REMAINS OF HOMO FLORESIENSISVol. 123, 2015 139
not strong but weak after the correction of the original left
row (Kaifu et al., 2009). Bilateral rotations are seen in the
P4s (markedly), M3s (moderately), and P3s (slightly).
Kaifu et al. (2009) made a photographic reconstruction of
the centric occlusion of LB1 primarily by matching the oc-
clusal wear facets. The result indicated that the occlusion of
this individual was horizontally twisted so that the left mo-
lars show a Class II relationship (the mandibular teeth are in
a distal relationship with their normal maxillary opponents)
and the right molars a Class III relationship (the mandibular
teeth are in a mesial relationship with their normal maxillary
opponents) (see their Fig. 2). This distorted occlusion is like-
ly a result of the posterior deformational (positional) plagi-
ocephaly, and is not indicative of severe developmental ab-
normality in this individual (Kaifu et al., 2009, 2010).
LB6/1
LB6/1 had a complete set of the permanent mandibular
dentition, but the right and left I1 as well as left I2 were ab-
sent in their alveoli when the specimen was recovered. The
length–breadth ratio of the dental arcade is similar to that of
LB1 before it had been broken (Kaifu et al., 2011). The
Curve of Spee is only slight (Brown and Maeda, 2009). No
remarkable tooth rotation is observed. The occlusal wear
pattern in this specimen indicates occlusion with undistort-
ed, normal Class I relationship.
LB2/2
This individual is represented by a single P3. As men-
tioned above, the position of its mesial IPF suggests a slight
rotation in the tooth row, as in LB1.
Oral health
There is heavy (LB1) or modest (LB6/1) calculus deposi-
tion, and the bone is resorbed in the molar regions of LB1 as
described and illustrated previously (Brown and Maeda,
2009; Jungers and Kaifu, 2011). The other four isolated den-
tal specimens (LB2/2, 6/14, 15/1, and 15/2) also exhibit
some degree of calculus deposition, suggesting its common-
ality in H. oresiensis from the Liang Bua Cave. The heavy
calculus and alveolar resorption in LB1 were probably re-
sulted from the unbalanced, twisted (horizontally rotated)
occlusion in this individual. Dental caries is not observed in
any of the materials described here. No hypoplastic pits or
bands are present on the enamel surfaces. Periapical abscess
cavities are also absent in LB1 (maxilla and mandible) and
LB6/1 (mandible). Previous claims that the oral health con-
dition of LB1 suggests its agricultural subsistence (thus the
individual is from a H. sapiens individual: Henneberg and
Schoeld, 2008) has been rejected by Jungers and Kaifu
(2011).
Discussion
Dental individuals
Probably ve but possibly four or six individuals are rep-
resented in the current dental sample of H. oresiensis.
These are LB1 (skeleton), LB6/1 (mandible) + 6/14 (I1), and
individuals represented by the other three teeth: LB2/2 (P3),
LB15/1 (P4), and LB15/2 (I1). Occlusal wear of the latter
teeth varies from severe (LB15/2), moderate (LB15/1), to
relatively light (LB2/2). At least, the wear state of LB2/2
does not match with either of the other two teeth, and these
three teeth include two or more individuals. In terms of the
P4 crown and root size (Table 1; see above), LB15/1 may
have been the largest individual, followed by LB1 and
LB2/2 whose P3 crown sizes are nearly equal to each other.
LB6/1 is smaller than LB1 in premolar and molar crown
sizes.
Reassessment of the previous evaluations: modern or
primitive?
Previous studies have proposed several potentially taxo-
nomically diagnostic dental characteristics of H. oresiensis.
These are summarized and discussed below.
I2/C diastema
I2/C diastema is frequently observed in Au. afarensis but
is rare in Au. africanus, Paranthropus, and Homo (White et
al., 1981; Kimbel and Delezene, 2009). Brown et al. (2004)
described that, in LB1, “(s)ize, spacing and angulation of the
maxillary incisor alveoli, and absence of a mesial facet on
the canines suggest that incisor I2 was much smaller than I1,
and there may have been a diastema.” This inference is not
supported by our CT scan, which shows close proximity
between the I2 and C1 alveoli (Figure 5a).
Canine size
Brown and Maeda (2009) reported that the mandibular
canine size of LB1 and LB6/1 is ‘small’ like various other
groups of post-1.7 Ma Homo, whereas those of earlier Homo
and Australopithecus were either ‘variable,’ ‘medium,’ or
‘large’ (their Table 3). We here metrically reexamine this
claim. Table 2 compares ‘relative canine size’ among vari-
ous archaic Homo groups and a global sample of modern
humans (H. sapiens). The ‘relative canine size’ is dened as
dimensional proportion of C1 (LL diameter) relative to the
premolar and molar lengths (additive MD diameters for
P3 – M2 or P3 + M1 + M2). The available small fossil sample
indicates that relative canine size tends to be smaller in H.
habilis than in later Homo, contrary to the claim of Brown
and Maeda (2009). The values for the two H. oresiensis
individuals are within the upper range of the variation for
H. habilis, and are also well within the variations exhibited
by the later archaic and recent Homo groups. Therefore, rel-
ative canine size is of limited use to assess the taxonomic
afnities of H. oresiensis.
P4 rotation
The P4s of LB1 are bilaterally rotated parallel to the tooth
row. This was cited as a unique trait (Brown et al., 2004) or
indicative of some developmental abnormality (Hershkovitz
et al., 2007) and/or afnity with local, living ‘pygmy’ groups
of Flores (Jacob et al., 2006). However, tooth rotation is a
relatively commonly observed dental anomaly both in mod-
ern (Jacob et al., 2006; Lukacs et al., 2006) and pre-modern
(e.g. Early Pleistocene Homo from Dmanisi (Rightmire et
al., 2006) and Konso (Suwa et al., 2007)) hominins as well
as other mammals (Natsume et al., 2006), with suggested
etiologies including some genetic mechanism and a lack of
Y. KAIFU ET AL.140 AnthropologicAl Science
space for the normal tooth eruption (Baccetti, 1998;
Natsume et al., 2006). Therefore this trait is not taxonomi-
cally diagnostic and does not necessarily indicate a severe
growth abnormality.
P3 crown
The P3s of H. oresiensis are unique and were a focus of
attention in previous studies (Brown et al., 2004; Brown and
Maeda, 2009). A previous claim that (some of) these are de-
ciduous rst molars (Obendorf et al., 2008) has been effec-
tively rejected by Brown (2012) based on crown and root
morphology as well as the state of wear. Brown and Maeda
(2009) suggested that its MD elongated, asymmetric crown
shape represents a (very) primitive hominin condition,
which changes to a more derived, molarized, bicuspid, and
symmetrical P3 in later australopiths and early members of
Homo. According to these authors, the P3 crown morphology
of H. oresiensis is also similar to ~1.75 Ma Homo from
Dmanisi (Martinón-Torres et al., 2008), but metrically dis-
tinguishable from H. erects (sensu lato) and H. sapiens (their
Figure 12). Jacob et al. (2006) claimed that the “enlarged,
block-like” P3 similar to the condition in LB1 are observed
worldwide in H. sapiens, although no numerical data were
provided to support their view.
Table 2. Dimensional proportion of the mandibular canine and rst premolar relative to the postcanine tooth row (%)a
C1 (LL) P3 (MD)
P3 + P4 + M1 + M2 (MD) P3 + M1 + M2 (MD) P3 + P4 + M1 + M2 (MD) P3 + M1 + M2 (MD)
H. oresiensis
LB1 26.2 30.3
LB6/1 19.7 24.6 24.2 30.1
H. habilis (East Africa, 2.0–1.6 Ma)
Omo75-14 20.2 25.5
KNM-ER 1802 20.1 25.7
KNM-ER 60000 18.6 23.2 20.1 25.0
OH 7 20.9 26.6 19.0 24.2
OH 13 17.0 21.4 20.5 25.7
OH 16 19.8 24.8 20.9 26.1
H. habilis (mean) 19.1 24.0 20.1 25.4
Dmanisi Homo (Georgia, 1.75 Ma)
D211 19.6 24.2 21.0 26.0
D2735 21.5 26.1 22.0 26.7
H. ergaster (East Africa, 1.5–1.0 Ma)
KNM-ER 992 20.4 25.6 21.7 27.1
KNM-WT 15000 21.7 27.4 20.9 26.4
OH 22 22.2 27.7
Early Javanese H. erectus (Sangiran, >1.0 Ma)
Sangiran 22 21.2 26.6 20.6 25.9
European terminal Early Pleistocene Homo (Gran Drina, 0.8 Ma)
ATD H1 23.4 29.0 20.6 25.5
ATD 6-96 20.8 26.0
African early Middle Pleistocene Homo (Tighenif and Baringo, 0.8–0.5 Ma)
Tighenif 1 19.7 24.4
Tighenif 2 19.0 23.7
Tighenif 3 24.7 30.5 21.1 26.0
KNM-BK 8518 21.2 26.5
Chinese Middle Pleistocene H. erectus (Zhoukoudian, 0.75 Ma)
Zhoukoudian B1 24.5 26.3
Zhoukoudian G1 23.3 29.0 20.9 26.0
Zhoukoudian K1 21.1 26.3 22.2 27.7
Post-habilis archaic Homo (mean) 21.9 26.9 21.0 26.1
H. sapiens (mean) 20.7 25.7 19.6 24.3
(N: range) 125: 17.3–24.7 136: 21.6–31.0 167: 17.5–21.9 188: 21.5–27.3
a Measurements for the comparative fossil sample were taken by Y.K. based on high-quality casts produced from the original specimens by Y.K.
or Gen Suwa except for KNM-ER 60000 (Leakey et al., 2012), Dmanisi (Martinón-Torres et al., 2008), KNM-WT 15000 (Brown and Walker,
1993), Gran Drina (Bermúdez de Castro et al., 1999; Carbonell et al., 2005), Tighenif (Bermúdez de Castro et al., 2007), and Zhoukoudian (Weid-
enreich, 1937). The H. sapiens is a global sample from Asia, Oceania, Europe, and Africa, and is based on high-quality casts prepared by Y.K.
DENTAL REMAINS OF HOMO FLORESIENSISVol. 123, 2015 141
In Figure 8, we compare our revised P3 crown diameters
of H. oresiensis with those of a global H. sapiens sample as
well as the Early Pleistocene Homo specimens from East
Africa (H. habilis and H. ergaster), Caucasus (Dmanisi
Homo), and Java (early Javanese H. erectus from the lower
and upper stratigraphic levels of Sangiran). In this chart, the
P3s from three H. oresiensis individuals are situated at the
margin of the large cloud of H. sapiens due to the formers’
relatively larger MD diameters. MD elongated P3 crown
congurations are frequently observed in H. habilis and
Dmanisi Homo, and represent a primitive state for Homo
(Wood and Uytterschaut, 1988; Tobias, 1991; Brown and
Maeda, 2009). However, contrary to Brown and Maeda
(2009), this crown shape is not restricted to H. habilis and
Dmanisi Homo but is also seen in an East Africa specimen
dated to 0.8–1.2 Ma (OH 22: Rightmire, 1980; Antón,
2003). Therefore, H. oresiensis exhibits a primitive, MD
elongated P3 crown shape shared with H. habilis, but such
crown morphology does exist, albeit in small numbers, in
later Homo groups.
P3 of H. oresiensis has a transverse crest that is oriented
distolingually, and, at its end, has a small lingual cusp that is
situated near the crown’s distolingual corner. A similar crest
and cusp arrangement, which contributes to reduce the talo-
nid, is found in D2735 from Dmanisi (Martinón-Torres
et al., 2008) and a few post-habilis African Homo P3s
(KNM-ER 992, KNM-WT 15000 (left), OH 22). Although a
distally oriented transverse crest is a plesiomorphic hominin
condition seen in great apes, Ardipithecus ramidus, and
Australopithecus anamensis (Ward et al., 2001, 2013; Suwa
et al., 2009; Delezene and Kimbel, 2011), the P3s of Au.
afarensis and H. habilis have altered so that the transverse
crest tends to form an acute angle with the mesial protoconid
ridge and the lingual cusp is placed slightly mesial or oppo-
site to the buccal cusp (Suwa, 1990; Suwa et al., 1996;
Delezene and Kimbel, 2011). Thus, the distal location of
lingual cusp in some post-habilis Homo as well as H.
oresiensis P3s is a secondary acquisition of the primitive
pattern that is derived compared to H. habilis. In support of
this view, these Pleistocene Homo P3s also lack other plesio-
morphic features present in early Pliocene hominins such as
an obliquely elongated crown shape (strong mesiobuccal
protrusion of the buccal face) and sharp occlusal crests
(Suwa et al., 1996; Delezene and Kimbel, 2011). Although
the transverse crests of the H. oresiensis P3s have lost their
edges by wear, the unworn portions clearly show that these
crests have thick bases.
The H. oresiensis P3s are, however, unique, showing the
beveled, generally at but wrinkled mesiolingual occlusal
surface as described above. This feature is associated with a
low, mesially restricted Mmr. It is different to the ‘open’
anterior fovea frequently seen in Australopithecus as well as
archaic and modern Homo P3s, where a distinctly elevated
Mmr is deeply incised by a furrow emanating from the pit-
like anterior fovea, and is not associated with ne enamel
wrinkling (e.g., Johanson et al., 1982; Tobias, 1991; Grine
and Franzen, 1994). In Table 2, we compare MD dimension-
al proportions of P3 within the postcanine tooth row, i.e. P3/
(P3 – M2) or P3/(P3 + M1 + M2). This comparison indicates
that the relative P3 lengths tend to be smaller in H. sapiens,
moderately large in the Early–Middle Pleistocene Homo
particularly in post-habilis Homo taxa, and extremely large
in H. oresiensis. The metric data reported by Wolpoff
(1971) also show that P3/M1 size ratio is higher in H. erectus
and Neanderthals than in H. sapiens.
To summarize, H. oresiensis P3 exhibits a primitive
crown shape but is derived from H. habilis in cuspal arrage-
ment, and is unique in its relatively large size and the beve-
led and wrinkled mesiolingual crown.
Mandibular premolar root
Brown and Maeda (2009) emphasized that the bifurcated
or Tomes’ mandibular premolar root forms seen in H.
oresiensis are rare in H. sapiens (Shields, 2005). We agree
that the roots of these (and anterior) teeth of H. oresiensis
are robust and primitive, although we found that the P4 roots
of LB1 and LB6/1 are not bifurcated but should be described
as Tomes’ form with a mesiobuccal cleft (see above).
Brown and Maeda (2009) suggested that such ‘complex’
root forms are more frequently observed in Australopithecus
and East African early Homo than in Sangiran H. erectus
(their Table 3). This is incorrect. Observed frequencies of
double-rooted mandibular premolars do not signicantly
differ between H. habilis (4/13 (P3) and 5/10 (P4)) (Suwa,
1990; Tobias, 1991; Wood, 1991; Leakey et al., 2012) and
the older Sangiran H. erectus (3/6 (P3) and 4/6 (P4)) (Kaifu
et al., 2005b). Single-rooted mandibular premolars do exist
in H. habilis (e.g. KNM-ER 1483, 1501; OH37: Wood,
1991), and non-double-rooted P3s from Sangiran include
Tomes’ root form (S 6a, S 22: Weidenreich, 1945; Kaifu et
al., 2005a). Thus, the available limited information about
Figure 8. Scatter plot of P3 MD and BL crown diameters (mm).
The fossil comparative specimens included are as follows: H. habilis
sensu lato: Omo 29-43, Omo 75-14, KNM-ER 1802, KNM-ER 60000
(Leakey et al., 2012); OH 6, OH 7, OH 13, OH 16, OH 68 (Clarke,
2012); H. ergaster: KNM-ER 992, KNM-ER 1814, KNM-ER 1808,
KNM-WT 15000; Dmanisi Homo: D211, D2735, D2600 (Martinón-
Torres et al., 2008); Sangiran Lower: S 6a, S 9, S 22; Sangiran Upper:
S 7-25. Measurements for the above fossil specimens were taken by
Y.K. based on high-quality plaster casts produced from the original
specimens by Y.K. or Gen Suwa, or from the literature. The H. sapiens
is a global sample from Asia, Oceania, Europe, and Africa, and is based
on high-quality plaster casts prepared by Y.K (n = 208).
Y. KAIFU ET AL.142 AnthropologicAl Science
mandibular premolar root form is not very useful to discuss
the evolutionary origin of H. oresiensis. More detailed
morphometric analyses are needed to investigate detailed
evolutionary changes in root morphology of the Pleistocene
Homo (e.g. Kupczik and Hublin, 2010; Emonet et al., 2012;
Le Cabec et al., 2013).
Molar crown shape
Jacob et al. (2006) listed the following molar traits of LB1
as evidence to link this individual with a modern ‘pygmy
population’ from Flores: (1) a tendency for the longitudinal
ssure to shift away from the buccolingual axis on mandib-
ular molars, (2) rhomboid outlines of upper molars reecting
hypocone reduction, and (3) squared lower molar outlines
related to hypoconulid loss.
The meaning of the rst point is not clear to us. In our
observation, the longitudinal ssures of LB1 and LB6/1 are
remarkable in that their distal segments are shifted extreme-
ly lingually. This trait is more frequently found in the Early
Pleistocene Homo than in H. sapiens, although this observa-
tion needs to be veried metrically in future studies. We will
numerically examine the second and third points elsewhere,
but we here note that it is metacone, not hypocone, that
shows marked reduction on the M1 and M2 of LB1. We con-
rmed that the mandibular molars of the two H. oresiensis
individuals are four-cusped teeth with no hypoconulid. Four-
cusped M1s and M2s have been unknown among H. erectus
assemblages from Indonesia and China (Martinón-Torres et
al., 2007). This morphology is also rare among the Middle–
Late Pleistocene European archaic Homo (Martinón-Torres
et al., 2012). However, Zanolli (2013) recently reported four
four-cusped M2s that may have been derived from the termi-
nal Early Pleistocene Bapang (Kabuh) Formation in the
Sangiran Dome, Central Java (chronology based on Hyodo
et al., 2011). Four-cusped M2s are relatively common in
modern human populations (24%), but four-cusped M1s are
rare (1%) (calculated from the data based on a large global
modern human sample (n = 6790–8638) in Scott and Turner,
1997: Appendix A). Therefore, the condition in H. oresiensis
(both of the existing two individuals have four-cusped M1
and M2) is not a typical observation even for H. sapiens. In
consideration of a report that the loss of hypoconulid is cor-
related with the reduction in mandibular molar size in
H. sapiens (Scott and Turner, 1997), it is possible that
H. oresiensis independently lost the hypoconulid in associ-
ation with the reduction of their mandibular molars.
Molar size sequence
During the course of the Homo evolution, the posterior
molars experienced more marked size reduction than in the
rst molar, resulted in alteration of the molar size sequence
within a dentition, from plesiomorphic ‘M1 < M2 ≥ M3’ to
‘M1 > M2 > M3’ (Wolpoff, 1971; Bermúdez de Castro and
Nicolás, 1995; Kaifu et al., 2005b; Kaifu, 2006). The maxil-
lary and mandibular molar size in H. oresiensis decreases
posteriorly (M1 ≥ M2 > M3). This is a derived character
seen in post-habilis grade Homo (Brown et al., 2004).
Mandibular dental arcade shape
Dental arcade shape, which became wider during hominin
evolution, is a useful character for taxonomic purposes
(Rosas and Bermudez de Castro, 1998; Kaifu et al., 2005b;
Spoor et al., 2015; Villmoare et al., 2015). Brown and Maeda
(2009) suggested that the narrow dental arcades seen in the
LB1 and LB6/1 mandibles are shared with pre-1.7 Ma Homo
and Australopithecus but not present or very uncommon in
Asian H. erectus or later Homo. This view has been dis-
proved by a later metric reinvestigation, which showed that
their arcades are actually wider than seen in H. habilis or
Dmanisi Homo, and are similar to early H. erectus from Java
(Kaifu et al., 2011).
Conclusions
In this paper, we described the dental morphology of
H. oresiensis and corrected some previously reported infor-
mation (e.g. I2 size, I2/C diastema, relative C1 size, P3 crown
dimensions, and P4 root form). The dental assemblage of
Liang Bua H. oresiensis represents probably ve (but
possibly four or six) individuals. They share similar dental
morphologies and represent a single population. We also in-
vestigated primitive and modern aspects of the H. oresiensis
teeth by reassessing the previously reported such characters.
H. oresiensis is primitive compared to H. sapiens in
having a MD elongated P3 crown. Other possibly archaic
features of H. oresiensis mentioned in the present paper
include a prominent canine lingual median ridge, thickening
of the buccal cervical enamel on C1 and P3, a robust anterior
tooth root, a relatively complex and robust premolar root
morphology, and a squarish maxillary dental arcade. H.
oresiensis is derived relative to H. habilis s.l. in having a
distally located P3 lingual cusp, comparatively wider man-
dibular dental arcade, four-cusped mandibular molars, pos-
teriorly decreasing molar size sequence, and a small tooth
size. The previously suggested very primitive features
shared with Australopithecus afarensis, such as a small I2
and presence of I2/C diastema, are actually not evident in the
existing H. oresiensis fossil collection. On the other hand,
the teeth of H. oresiensis are unique, showing an extremely
large relative size and mesiolingually beveled and wrinkled
morphology of the P3 crown.
As in the cases of other skeletal elements (Brown et al.,
2004; Morwood et al., 2005; Larson et al., 2009; Brown and
Maeda, 2009; Jungers et al., 2009b; Kaifu et al., 2011), the
teeth of H. oresiensis exhibit an impressive mosaic of prim-
itive, derived, and unique characters. The derived characters
include four-cusped mandibular molars, a trait that can be
described as modern, like H. sapiens. The primitive features
include those comparable to some of the Early Pleistocene
Homo (P3 crown shape, P3 root morphology, and mandibular
dental arcade). The dental morphology of H. oresiensis has
been controversial: some researchers view that it is fully
modern (Jacob et al., 2006), whereas others point out a few
very primitive features that suggest an evolutionary link
with H. habilis s.l. or Australopithecus (Brown et al., 2004;
Brown and Maeda, 2009). Although more comprehensive
comparative analyses are needed to fully illustrate the dental
morphological afnities of this dwarfed hominin species, the
present study found no grounds for both of these conicting
views.
DENTAL REMAINS OF HOMO FLORESIENSISVol. 123, 2015 143
Acknowledgments
We are grateful to Gen Suwa for the micro-CT scanning
of LB1, and Gen Suwa, Tony Djubiantono, Ian Tattersall,
Ken Mowbray, John de Vos, Philippe Mennecier, Fabrice
Demeter, Nguyen Kim Thuy, and Nguyen Lan Cuong for
access to the specimens in their care. Y.K. thanks the late
Mike Morwood for his invitation and kind support for this
project. This study was supported by grants from the Japan
Society for the Promotion of Science (No. 24247044) and
the National Museum of Nature and Science, Tokyo, to Y.K.
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