*Supported by the Russian Science Foundation (Project
A.V. Zubova, T.A. Chikisheva, and M.V. Shunkov
Institute of Archaeology and Ethnography, Siberian Branch, Russian Academy of Sciences,
Pr. Akademika Lavrentieva 17, Novosibirsk, 630090, Russia
E-mail: firstname.lastname@example.org; email@example.com; firstname.lastname@example.org
The Morphology of Permanent Molars from the Paleolithic Layers
of Denisova Cave*
The article describes the morphology of two permanent molars from the Pleistocene layers of Denisova Cave, the
Altai Mountains. Denisova 4 is an upper left third or second molar, and Denisova 8 is an upper left third molar. Both
specimens were examined using the extended trait battery. The results indicate a high informative potential for dental
traits in the analysis of group variation within the genus Homo. They support the view that Denisovans, or H. altaiensis,
were a distinct group of hominins, differing from both H. sapiens and H. neanderthalensis not only genetically but
morphologically as well. The distinctive dental features of the Denisovans include extremely large dimensions, and
afﬁ nities with Homo erectus of Sangiran and the Middle Pleistocene hominins of China, such as Xujiayao. On the basis
of the morphological analysis of Denisovan upper molars, it is proposed that the unidentiﬁ ed part of the Denisovan
genome may stem from Homo erectus. Dentally, Homo altaiensis is a very conservative taxon.
Keywords: Denisovans, Homo altaiensis, Homo erectus, Homo neanderthalensis, Middle Paleolithic, Upper
Paleolithic, dental anthropology.
ANTHROPOLOGY AND PALEOGENETICS
Archaeology, Ethnology & Anthropology of Eurasia 45/1 (2017) 121–134 Email: Eurasia@archaeology.nsc.ru
© 2017 Siberian Branch of the Russian Academy of Sciences
© 2017 Institute of Archaeology and Ethnography of the Siberian Branch of the Russian Academy of Sciences
© 2017 A.V. Zubova, T.A. Chikisheva, M.V. Shunkov
The archaeological site of Denisova Cave is located in the
south of the West Siberian region, in the Altai Mountains,
close to the Russian Federation’s border with China,
Mongolia, and Kazakhstan.
The remains of Pleistocene hominins from the
Denisova Cave have been studied for more than 30
years. First dental remains were excavated here in 1984:
a deciduous molar named Denisova 2 was found in the
stratigraphic layer 21.1 (Shpakova, Derevianko, 2000).
In the same year, one more tooth was found in layer 12.
It was initially attributed as an upper permanent incisor
(Turner, 1990; Shpakova, Derevianko, 2000); but
later, a comparative study demonstrated that the tooth
actually belonged to an animal from the Bovidae family
(Viola et al., 2011: 209). In 2000, a well-preserved
upper permanent molar was recovered from layer 11.1
(Denisova 4); and in 2010, fragments of the crown of an
upper permanent molar (Denisova 8) were found in the
bottom of stratigraphic layer 11.4, in the contact zone
with layer 12.
According to geochronological data, the Denisova
4 and Denisova 8 permanent molars are dated to 50–40
ka BP; however the layer containing the Denisova 4 is
younger than the layer where Denisova 8 was found
(Sawyer et al., 2015).
Sequencing of mtDNA and the nuclear genome
from the distal phalanx of the carpal minimus of a
6–7 year old girl (which was found in layer 11.2
(Denisova 3)) and the Denisova 4 molar has shown that
both specimens represent a formerly unknown hominin
species. The species was named “Denisovan” (Reich
A.V. Zubova, T.A. Chikisheva, and M.V. Shunkov / Archaeology, Ethnology and Anthropology of Eurasia 45/1 (2017) 121–134
et al., 2010), or Homo altaiensis (Derevianko, 2011)
by the authors of the study. It was the ﬁ rst case in the
history of physical anthropology when a new taxon was
described by the results of genetic analysis, rather than
by a morphological study.
The analysis of the mtDNA has shown that the
hominin lineages ancestral to Denisovans, Neanderthals,
and Homo sapiens diverged about a million years ago
(Krause et al., 2010). The study of the nuclear DNA
revealed a much later date for the separation of the
line leading to H. sapiens from the hominin population
ancestral to H. neanderthalensis и H. altaiensis, about
800 ka BP (Meyer et al., 2012). The divergence between
Denisovan and Neanderthal lineages was initially thought
to have occurred 640 ka BP (Reich et al., 2010); but
accordin g to the results of recent studies, it happened only
some 430 ka BP (Meyer et al., 2016).
Denisovans have occupied the Altai Mountains for a
long period of time. Judging by the rate of accumulation
of mutations in the mtDNA of present-day humans, it
can be hypothesized that the Denisova 2 and Denisova 8
specimens are some 65 thousand years more ancient
than the Denisova 3 and Denisova 4 samples (Slon et al.,
2015). It was found that Denisovan genome contained
alleles associated with a dark skin-color, chestnut hair,
and brown eyes (Meyer et al., 2012). According to the
results of analysis of the mtDNA of all individuals found
at Denisova Cave, the level of the intragroup genetic
diversity in the Denisovan population was much lower
than in both Neanderthals and modern humans (Meyer
et al., 2012; Slon et al., 2015).
The place of origin of the Denisovans, as well as
their position in the hominin taxonomy, have been
hotly debated. On the basis of archaeological data,
A.P. Derevianko put forward a hypothesis that the
Denisovans are a subspecies of polymorphic H. sapiens,
contemporaneous with other subspecies, e.g. H. sapiens
neanderthalensis, H. sapiens africaniensis, and
H. sapiens orientalensis (Derevianko, 2011). According
to this hypothesis, the subspecies originated from local
Asian populations of H. erectus (Ibid.). However, recent
paleogenetic data on the Pleistocene hominins from Sima
de los Huesos in Spain suggest that the mitochondrial
genomes of this population and those of Denisovans were
similar (Meyer et al., 2016). This result questions an Asian
origin for the most ancient components in the genome of
This study presents the results of an extended
morphological analysis of the Denisova 4 and Denisova 8
permanent molars. The results describe peculiarities of
H. altaiensis dentition, and infer relationships of the
Denisovan population with other taxa: H. erectus s.l.,
H. heidelbergensis s.l., H. neanderthalensis, and
H. sapiens s.l.
The dental samples from Denisova Cave were previously
described using standard ASUDAS protocol and the
protocol of Neanderthal apomorphies, during the
paleogenetic study of the Denisovan hominins (Krause
et al., 2010; Sawyer et al., 2015).
We carried out an examination of the Denisova 4
and Denisova 8 upper molars employing a substantially
extended set of traits. The set was generally based on
the ASUDAS protocol (Turner, Nichol, Scott, 1991;
Scott, Turner, 1997) and the conventional set of traits
used in Russian dental anthropology (Zubov protocol),
which includes a comprehensive description of the
pattern of crown grooves (Zubov, 1968, 1974, 2006;
Zubov, Khaldeyeva, 1989, 1993). The protocol of the
Neanderthal complex markers (Bailey, 2002; Bailey,
Skinner, Hublin, 2011) was also employed in our study,
as well as a protocol previously developed for accounting
of plesiomorphic traits in modern human populations
Each of the protocols has some advantages, and thus
the combination of these protocols can help to extract as
much information as possible from the dental samples.
The ASUDAS protocol employs the most diversified
scales for ﬁ xing dental traits that are particularly precise
in terms of describing the dental patterns found in modern
populations. The main advantage of the traits put forward
by S. Bailey is their increased sensitivity to the presence
of Neanderthal genes in a population. The Zubov protocol
permits not only study of the details of the macrorelief
of the crown, but also the pattern of its grooves, which
represent an independent and hierarchically organized set
of traits: odontogliphics.
In total, 60 dental traits were scored. In the molars
studied, the following traits were observed: reduction of
the hypocone and metacone, rhomboid shape of the upper
molars, metaconulus, mesiostylid, enamel extension,
Carabelli cusp, distal and mesial accessory cusps of the
upper molars, epicrista, plagiocrista (crista oblique),
entocrista, posterior fovea, and odontoglyphic traits.
Hypocone reduction. The trait describes the dynamic
of the hypocone’s size relative to the protocone and
metacone. The ASUDAS and Zubov protocol scales
have some differences: the former includes 6 grades
(0 – absence of the cusp, 5 – maximal development of the
cusp) (Turner, Nichol, Scott, 1991: 18), while the latter
has only 4 grades (4, 4–, 3+, 3), where grade 4 stands for
the maximal development of the cusp, and grade 3 for the
absence of the cusp (Zubov, 1968: 152).
Metacone reduction. This trait describes the dynamic
of size of the metacone relative to the paracone. In
ASUDAS, the trait is scored in the same way as the
hypocone reduction (Turner, Nichol, Scott, 1991: 18),
A.V. Zubova, T.A. Chikisheva, and M.V. Shunkov / Archaeology, Ethnology and Anthropology of Eurasia 45/1 (2017) 121–134 123
while the Zubov protocol employs a 5-level scale (1 to 5),
where grade 1 means the absence of reduction, and
grade 5 its maximal development (Zubov, 1968: 160).
Rhomboid shape of the upper molars. This trait
describes upper permanent molars showing a non-
reduced hypocone, strongly developed in the disto-
lingual direction, and a reduced metacone, which forms
an oblique disto-vestibular corner of the crown (Bailey,
Metaconulus. A small cusp in the depth of the enamel
of the axial ridge of the metacone, in its central part
(Zubov, Khaldeyeva, 1993: 68).
Carabelli cusp. A styloid cusp in the mesio-lingual
part of the crown. Grades 0 to 7 in ASUDAS: 0 – absence,
7 – maximal development (Turner, Nichol, Scott, 1991:
19). Zubov protocol employs 5 grades from 0 to 5 (1968:
157). The grades of the two protocols correspond as
Distal accessory cusp of the upper molars. A styloid
cusp at the distal marginal ridge, at the interface between
the metacone and hypocone. A scale, in which grade 1
correspond to the absence of the cusp and grade 6 to
maximal development, is employed in both protocols
(Zubov, 2006: 56; Turner, Nichol, Scott, 1991: 19).
Mesial accessory cusp of the upper molars. A cusp
at the mesial margin of the crown. It is delimited by an
additional groove that falls into the ﬁ ssure separating the
paracone and protocone. Phylogenetically, it differs from
the cusps formed by the distal segments of the paracone
and protocone (Zubov, Khaldeyeva, 1993: 67; Scott,
Turner, 1997: 45). A scale for evaluating the degree of
development of this trait is absent.
Epicrista. A ridge connecting protocone and metacone.
The trait is present when the ﬁ ssure dividing the cusps is
Plagiocrista. A ridge connecting the metacone and
protocone. This study employs a 4-level scale for this trait
(Zubova, 2013: 114).
Entocrista. A marginal ridge connecting the protocone
and hypocone (Zubov, Khaldeyeva, 1989: 62). It is almost
never found in modern humans.
Posterior fovea. An elongated depression in the distal
part of the upper and lower molars, which can vary in
length. In modern humans, it is typically located at the
occlusal surface of the metacone. Mesially, it is delimited
by the distal segment of the metacone, and distally by
the distal marginal ridge (Zubov, 2006: 61). In ancient
specimens, it can reach the surface of the hypocone as
well. In such cases, it is delimited by the same elements
as in the metacone. Scales for evaluating the degree of
development of this trait are absent.
Odontoglyphic elements. Like other dental traits,
these elements are not equal in terms of taxonomic value.
Combinations of the elements reﬂ ect the evolutionary
status of the population (Zubov, 1974). The nomenclature
of the odontoglyphic elements has been repeatedly
changing during its development. As a result, different
researchers employ rather different nomenclatures for
these elements (Zubov, 1974, 2006; Zubov, Khaldeyeva,
1989; Hillson, 1996). In our study, we follow the
methodology described in the last publications of
A.A. Zubov (Zubov, Khaldeyeva, 1989; Zubov, 2006). In
the next section, the classiﬁ cation and description of the
odontoglyphic traits are outlined.
We employ three types of the grooves of the occlusal
surface of the crown. The ﬁ rst and most ancient type
comprises the intertubercular ﬁ ssures of the ﬁ rst order.
They separate major cusps of the crown, and are
designated as I–IV for the upper molars, and I–VI for
the lower molars. Fissure I separates the metacone and
the paracone; ﬁ ssure II separates the paracone and the
protocone; fissure III separates the metacone and the
protocone; and ﬁ ssure IV separates the hypocone from
both the metacone and the protocone.
Tubercular grooves of the second order delimit
the axial ridges of each cusp, dividing a cusp into
three segments. In some studies, these segments are
designated as mesial, central, and distal (Carlsen,
1987; Bailey, Skinner, Hublin, 2011). Intersegmental
grooves are tagged as 1 and 2 plus the two (or three for
the lower) ﬁ rst characters of the name of the cusp (e.g.
“pa” stands for the paracone, “prd” for the protoconid)
In the metacone and paracone, these grooves separate
the axial segment from mesial one (groove 1), and the
axial segment from the distal one (groove 2). The grooves
are designated as 1me, 2me, 1pa, and 2pa, respectively.
In the protocone and hypocone, groove 1 separates the
axial and distal segments (1pr, 1hy), groove 2 the axial
and mesial segments (2pr, 2hy).
Grooves of the third order are divided into two
categories. The ﬁ rst includes accessory grooves dividing
the mesial and distal segments of each cusp into two parts
in the sagittal direction, and lying parallel to grooves 1
and 2. These are designated as 1’ and 2’, respectively. The
second category includes accessory grooves of the axial
ridges of the major cusps, designated as 3 and 4 (Zubov,
Khaldeyeva, 1989; Zubov, 2006).
In the metacone, 1’me falls into ﬁ ssure I, and divides
the mesial segment in the vestibular direction, while 2’me
falls into ﬁ ssure III parallel to the posterior fovea and
divides the distal segment in vestibular direction. In the
A.V. Zubova, T.A. Chikisheva, and M.V. Shunkov / Archaeology, Ethnology and Anthropology of Eurasia 45/1 (2017) 121–134
paracone, 1’pa falls into ﬁ ssure I and divides the distal
segment in vestibular direction, 2’pa falls into ﬁ ssure II
and divides the mesial segment in vestibular direction,
3pa falls into ﬁ ssure II and divides the axial segment in
vestibular direction, and 4pa divides the axial segment
in medio-distal direction. In the protocone, 1’pr falls
into ﬁ ssure IV and divides the distal segment in lingual
direction, 2’pr falls into ﬁ ssure II and divides the mesial
segment in lingual direction, 2’’pr duplicates 2’pr in the
mesial segment and falls into ﬁ ssure II or 2’pr, 3pr falls
into ﬁ ssure II and the central fovea and divides the axial
segment in lingual direction, 4pr bisects the axial segment
in transverse direction. In the hypocone, 1’hy falls into
ﬁ ssure IV or the posterior fovea and divides the distal
segment in lingual direction, 2’hy falls into ﬁ ssure IV or
the basin of the talon and divides the mesial segment in
The total number of tubercular grooves and their
directions differ in various species of the genus Homo.
The grooves of the third order are the most variable. Many
of them are reduced, and very rare in modern humans.
An increased irregularity of enamel is typical of early
Homo, and teeth of these hominins can exhibit additional
grooves of the third order, which are absent in later species
of Homo. The positions of the points of contact between
tubercular grooves and the inter-cusp ﬁ ssures can also
vary. Furthermore, the tendency to ridge formation is
more prominent in extinct representatives of the genus
Homo than in modern humans.
The traits studied can be divided into three groups. The
ﬁ rst group includes evolutionary stable and taxonomically
neutral variables related to the basal teeth morphology in
the hominin lineage and its evolutionary continuity (e.g.
four-cusped upper molars, ﬁ ve-cusped lower molars or the
number of intertubercular grooves of the crown). Traits of
this group are present in all Homo species.
The second group is composed of so-called generalized
archaic markers. These are plesiomorphic traits displaying
a negative temporal dynamic. In other words, they
are found with the highest frequency in early Homo,
and become increasingly rare in later Homo species.
The second group includes such traits as derivatives
of the cingulum; ridge-forming structures of molars;
M1<M3<M2 and М1<M2<M3 patterns; phylogenetic
diastems; and the sub-squared shape of the crown of the
upper and lower molars lacking prominent angles, etc.
(Khaldeyeva, Kharlamova, Zubov, 2010; Zubova, 2013;
Gomez-Robles et al., 2007).
The third group includes traits that are evolutionarily
progressive among members of the genus Homo, and
is composed of two blocks of traits. One of the blocks
includes traits describing reduction of the dentition:
hypocone and metacone reduction, decrease in the size of
distal teeth in a row, small size or absence of the styloid
cusps in the distal part of the lower molars, axial position
of the hypoconulid, simpliﬁ ed odontoglyphic pattern,
etc. The frequency of these traits steadily increases with
time in the hominin lineage. Another block includes
apomorphic patterns of various taxa.
When describing the upper molars from Denisova
Cave, most attention was paid to the traits of the second
and third groups, namely plesiomorphic traits and the
markers of Neanderthal and modern dental patterns.
The frequency traits were then compared with the
patterns typical of H. erectus s.l., H. heidelbergensis s.l.,
H. neanderthalensis, and H. sapiens s.l.
Previously published raw data, as well as high-
deﬁ nition images and morphological descriptions of ﬁ nds,
were used as reference data (see Table).
Our sample comprises two permanent molars. The
Denisova 4 molar from the lithological layer 11.1 was
previously described by B. Viola as an upper left third
molar (Viola et al., 2011). But the pattern of attritional
facets provides some evidence that the tooth might also
be a second molar, if the individual lacked the third molar.
In the study cited, the patterns of enamel macrorelief
and dental metrics were published. The authors pointed
out a similarity in dental metrics between the Denisovan
specimen and early Homo (and even australopithecine)
samples, but not Neanderthal dentition.
The Denisova 8 molar also belongs to the left side of
the upper jaw. A.P. Buzhilova (2014) determined it to be
an upper second or third permanent molar, while Viola
pointed that this was most probably the third upper left
molar (Sawyer et al., 2015). This tooth is worse preserved
than the Denisova 4 molar: the roots were completely
lost, the crown was reconstructed from fragments, and
the mesial part of the crown at the interface between the
paracone and protocone was destroyed. The macrorelief
of the cusps of the trigon was almost completely worn
off, as well as most intertubercular ﬁ ssures. But there is
almost no attrition at the hypocone and the distal part of
the metacone; just one small contact-facet can be seen on
the top of the main ridge of the hypocone, in its mesial
part. On the interproximal surface of the distal part there
is no contact-facet.
Left upper permanent second or third molar (М2/3) from
the layer 11.1, Denisova 4 (Fig. 1). This belonged to a
young adult male (Slon et al., 2015).
The tooth is very massive, with a long neck. The roots
are strongly divergent in the mesial and distal norms,
the crown exhibits rounded corners. The bucco-lingual
A.V. Zubova, T.A. Chikisheva, and M.V. Shunkov / Archaeology, Ethnology and Anthropology of Eurasia 45/1 (2017) 121–134 125
Materials used for comparison
Region Taxonomic status of
the ﬁ nd*Site Specimen No. Source
12 3 4 5
Africa Anatomically modern
Fish Hoek – Schwartz, Tattersall, 2003
Homo helmei Florisbad – Ibid.
Homo habilis Hadar A.L. 666 "
Homo heidelbergensis Kabwe – "
Homo erectus Koobi Fora KNM-ER 1813, KNM-ER 3733 "
Homo ergaster Nariokotome KNM-WT 15000 Khaldeyeva, Zubov,
Homo habilis Olduvai Gorge OH 6, OH 13, OH 16, OH 24 Schwartz, Tattersall, 2003,
Homo erectus Konso KGA 4-14, KGA 11-350 Suwa et al., 2007
Early Homo sapiens Dar es-Soltan DS II – H9, DS II – NN, DS II – H5,
DS II – H10
Hublin et al., 2012
" Smugglers’ Cave Ctb H7, Ctb Ib 19, Ctb T4, Ctb T3b Ibid.
West Asia Early Homo sapiens Jebel Qafzeh Qafzeh 4, Qafzeh 5, Qafzeh 6,
Qafzeh 9, Qafzeh 11
Schwartz, Tattersall, 2003,
Homo neanderthalensis Skhul Skhul IV, Skhul V Ibid.
" Tabun Tabun I, Tabun T I, Tabun T II "
" Kebara KNM 24, KNM 21 Tillier et al., 2003
Early Homo sapiens Qesem – Hershkovitz et al., 2011
Archaic Homo sapiens Jinniushan – Schwartz, Tattersall, 2003,
Homo sapiens Liujiang – Ibid.
Homo erectus Sangiran Sangiran 4, 7, 17, 27, NG 91- G10
No1, NG 0802.1, NG 0802.3,
NG 92.3, Njg 2005.05,
Bpg 2001.04, PDS0712, NG0802
Ibid.; Zanolli, 2013; Kaifu
et al., 2007; Zaim et al.,
Homo sapiens Wadjak Wajak 1, Wajak 2 Schwartz, Tattersall, 2003,
Homo erectus Zhoukoudian,
ZKD 169.25, ZKD PA 327, ZKD
Homo sapiens Zhoukoudian,
PA 101, PA 102, PA 103 Ibid.; Turner, Manabe,
Early Homo Xujiayao PA 1480, PA 1481, PA 1500 Xing et al., 2015
Homo sapiens Daoxian DX 1, 4, 5, 6, 8, 12, 14, 16, 17, 20,
21, 24, 28, 31,33, 35, 36, 39, 41, 47
Liu et al., 2015
Homo erectus Liang Bua LB1 Kaifu et al., 2015
North Asia Homo sapiens Malta 1, 2 Zubov, Gokhman, 2003;
Zubova, Chikisheva, 2015
Homo neanderthalensis Chagyrskaya 10, 51.1, 57 Unpublished data of
Homo neanderthalensis Obi-Rakhmat – Glantz et al., 2008;
unpublished data of
" Teshik-Tash – Unpublished data of
A.V. Zubova, T.A. Chikisheva, and M.V. Shunkov / Archaeology, Ethnology and Anthropology of Eurasia 45/1 (2017) 121–134
12 3 4 5
Europe Homo sapiens Abri Pataud Pataud 1 Schwartz, Tattersall, 2003,
Homo heidelbergensis Arago Arago 21, 14, 31 Ibid.
Homo antecessor Atapuerca: Gran
ATD 6-69 "
Homo heidelbergensis /
de los Huesos
AT-16, AT, 3177, AT-138, AT-406,
AT-139, AT-26, AT-959, AT-20,
AT-2076, AT-812, AT-944, AT-196,
AT-2071, AT-4317, AT-3424,
AT-587, AT-46, AT-4326, AT-960,
AT-824, AT-2179, AT-407, AT-4319,
AT-4336, AT-12, AT-2175, AT-815,
AT-821, AT-15, AT-170, AT-602,
AT-816, AT-274, AT-3181, AT-171,
AT-826, AT-601, AT-945, AT-1471,
AT-2393, AT-3183, AT-194,
AT-5082, AT-2150, AT-140
Martinón-Torres et al., 2012
Homo erectus Dmanisi D 2882, D 2700 Schwartz, Tattersall, 2003;
Martinón-Torres et al.,
Homo sapiens Engis Engis 2 Schwartz, Tattersall, 2003,
" Grimaldi Barma grande 2 Ibid.
" Isturitz Ist 71 "
Homo neanderthalensis Krapina 45, 46, 47, 48, D 119, D 120, D 170,
D 180, D 136, D 164, D 178, D 188
Radovčić et al., 1988
" La Quina H 5, H 18 Schwartz, Tattersall, 2003,
" Le Moustier – Ibid.
Homo sapiens Mladeč 1, 2 "
Homo neanderthalensis Pech-de-l’Azé – "
" Saccopastore 2 "
" Sakajia – "
" Scladina – "
" Spy 1, 2 "
" Subalyuk 2 "
Homo sapiens Akhshtyrskaya – Unpublished data of
" Rozhok-1 – Same
" Caldeirao 1 Trinkaus, Bailey, Zilhao,
" Sungir 2, 3 Zubov, 2000
" Kostenki 14, 15, 17, 18 Khaldeyeva, 2006;
unpublished data of
" Visogliano 6 Abbazzi et al., 2000
" Galeria da
– Trinkaus et al., 2011
Early Homo Peştera cu Oase 2 Trinkaus, 2010
*Since the taxonomic status of some ﬁ nds is debatable, information in this column is given according to the opinion of the
authors referenced in this article.
A.V. Zubova, T.A. Chikisheva, and M.V. Shunkov / Archaeology, Ethnology and Anthropology of Eurasia 45/1 (2017) 121–134 127
diameter of the metacone is small relative to the paracone,
while the mesio-buccal corner demonstrates only
moderate obliquity. As a result, the crown is narrowed
in its distal portion. But since the hypocone is not biased
lingually, the crown does not display the rhomboid shape
typical of the Neanderthal upper molars (Fig. 2). The
apexes of the major cusps are inclined towards the center
of the crown. The intertubercular ﬁ ssures are very deep.
Fissures I and III are visible only on the occlusal surface,
while ﬁ ssures II and IV expand to the vertical surfaces
of the tooth: ﬁ ssure IV, which divides the hypocone and
protocone, reaches the lingual surface, and ﬁ ssure II,
which separates the protocone and paracone, is present
on the mesial surface (Fig. 3).
The paracone is divided into three segments by deep
grooves, which dissect the margin of the crown and
continue in the upper portion of the buccal surface. The
mesial ridge of the paracone is wider than its distal ridge,
which disappears in the middle of the axial ridge. The
groove delimiting the mesial ridge joins the intertubercular
ﬁ ssure, which separates the metacone and paracone. The
paracone exhibits deep grooves 1ра and 2ра, the ﬁ rst of
which falls into ﬁ ssure I, and the second into the central
fovea. The terminal segments of the grooves lie on the
border of the occlusal plane, and change their direction
towards the apex of the axial ridge (Fig. 3). So the above-
mentioned grooves delimit elements of the marginal ridge
of the vestibular surface (eocrista). The latter are found
in the central branch of the ridge, and are not present in
modern humans. At the interface between the paracone
and metacone, the ridge is intercepted by ﬁ ssure I and a
short parallel groove 1’pa.
The metacone is not reduced. Its mesio-distal diameter
is not less than that of the paracone. On the surface of
the metacone, an axial, a mesial, and a distal segment
can be distinguished. The axial segment is very massive,
and its distal portion is divided into separate fragments
by accessory transversal grooves. The distal and mesial
segments of the metacone are clearly visible only at the
vestibular margin of the crown.
The plagiocrista is almost completely interrupted by
the central groove; only a thin enamel bridge in the distal
portion, lying parallel to ﬁ ssure IV, is left.
The terminal segments of the ﬁ rst and second grooves
of the metacone continue on the vestibular surface. They
form separate apexes of the mesial and distal portions
of the cusp. Groove 1me falls into ﬁ ssure I just below
1pa, and 2me merges with the basin of the talon. Two
accessory enamel ridges branch out of the axial ridge of
the metacone in the distal direction. These are oriented
towards the accessory cusps of the distal margin of
the crown, but do not reach it, being intercepted by
the basin of the talon. The ridges are separated by an
accessory groove of the third order. The groove delimits
elements of the metaconulus in the axial ridge, and does
not match precisely with conventional elements of the
odontogliphic pattern of the molars of modern humans
(Zubov, 2006; Zubov, Khaldeyeva, 1989). Following
the nomenclature of other cusps, it is referred to as 4me.
In the vicinity of the central fovea, the axial segment
of the metacone is divided in the axial plane by one
more accessory groove of the third order (3me), which
originates approximately in the middle third of the ridge
and falls into ﬁ ssure III.
The hypocone is large, sub-oval in shape, and
elongated in the vestibular-lingual direction. A massive
central ridge is prominent on its surface. The mesial
segment of the cusp is very thin, and the groove that
delimits it is almost merged with ﬁ ssure IV. The distal
segment is somewhat better pronounced. On the occlusal
surface, the hypocone is separated from the metacone
and protocone by a wide and elongated basin of the
talon, which merges with the elements of the posterior
fovea at the metacone. In its distal portion, the hypocone
is separated from the metacone by a rounded accessory
cusp in the marginal ridge (С5 grade 1 ASUDAS). Both
maj or grooves of the hypocone (1hy and 2hy) fall into
the basin of the talon. These are not particularly long, but
rather deep. They are duplicated in the mesial and distal
segments by parallel grooves of the third order, which
are very rarely found in modern humans. Similarly to
corresponding grooves of other cusps, they can be referred
to as 1’hy and 2’hy. An element of the marginal ridge is
prominent in the apex of the cusp, as is the case in the
metacone and paracone.
The protocone is massive: it is the largest cusp of the
crown. Such a large size is related to an increase in size
of its basal portion owing to the Carabelli cusp, which
contacts with the mesio-lingual groove separating the
hypocone and protocone. The Carabelli cusp occupies the
whole base of the protocone in its cervical portion (Fig. 1).
In the middle third of the height of this cusp, there are
four apexes formed by cingular ridges (Fig. 2). Thus,
the genetic potential of the upper molar growth is more
fully realized in this specimen than in modern human
teeth. Multiple apexes in such cases are a manifestation
of rudimentary derivatives of the cingulum, which are
referred to by P. Hershkovitz (1971) as entostyles.
Finding a correspondence between the morphological
pattern described above on the one hand, and the grades
of standard dental protocols on the other, is a complicated
task, since none of the protocols has grades to describe
a cusp with multiple apexes. In ASUDAS, the pattern, in
which the Carabelli cusp contacts with the intertubercular
groove, is referred to as grade 5; in Zubov protocol it is
The protocone, like other major cusps, is segmented
into three portions. The central segment is the largest,
followed by the mesial, and the distal segment is the
smallest. The second groove of the protocone (2pr)
merges with 2’pr and forms an isolated triradius. One
more groove of the third order, 2’’pr, falls into 2’pr in
its terminal segment. The 1pr groove is reduced, and
its distal segment is outlined by 1’pr. The latter appears
as a fovea isolated from ﬁ ssures III and IV by a narrow
enamel bridge. The bridge connects the distal segment of
the protocone with the distal portion of the axial ridge of
the metacone, thus forming a continuous element of the
plagiocrista (Fig. 3).
The protocone and paracone are separated by an
accessory mesial cusp, which is formed by accessory
grooves 2’ра and 2’pr both falling into fissure II. In
modern humans, this cusp is usually round in shape, and is
formed by the terminal triradius of ﬁ ssure II strictly at the
mesial marginal ridge. In our case, the cusp appears as a
segment lying parallel to ﬁ ssure II and almost reaching the
central fovea. The triradius is biased towards the central
third of the intertubercular fissure, which is thereby
substantially shortened. “Sprigs” of the triradius are
similar to the major tubercular grooves. The epicrista is
interrupted. The anterior fovea is absent, as is the enamel
extension on the vestibular side of the tooth.
Initially, the tooth had three roots: lingual, mesio-
buccal, and disto-buccal (Fig. 4). The lingual root, the
longest and the most massive, was destroyed during
the paleogenetic investigation. It is oval in section,
ﬂ attened in the bucco-lingual direction, and branches off
the buccal roots at a very large angle. The lingual root
separates from the distal root at a level of 3.6 mm from
the cementoenamel junction, and from the mesial root at
the level of 4.1 mm.
The roots of the buccal side separate only in its lower
third, 8.2 mm from the cementoenamel junction. Above
this level they are connected by a cemental lamina. The
mesial root is curved in the middle third. It is ﬂ attened
in the mesio-distal direction, and its vestibular portion is
more massive than the lingual one. The distal root is the
least massive and almost ﬂ at in section.
Left upper permanent third molar (М3) from the
base of the layer 11.4 at the interface with the layer 12,
Denisova 8 (Fig. 5). This is an upper left molar of an adult
male (Slon et al., 2015), who was slightly older than the
Denisova 4 individual. After restoration of the crown, it
became possible to assess its contours. It is of oval shape,
without an obliquity in the mesio-vestibular portion, but
with an expansion in the vestibular portion as compared
to the lingual portion (Fig. 6).
The protocone looks fairly massive despite postmortem
destruction. The axial ridge of the cusp is very wide, and,
together with the axial ridge of the mesial portion of
the metacone, it forms a wide ridge, plagiocrista. It was
impossible to determine reliably if the plagiocrista was
continuous or discrete. Groove 1pr is worn off, while
groove 2pr was probably deeper than 1pr. This conclusion
is based on the traceability of 2pr despite substantial
attrition of the mesial portion of the crown. It segmented
the marginal ridge of the protocone and extended to the
vestibular surface of the crown (Fig. 7).
The metacone is very massive, and divided into two
parts; it is much larger than the paracone and hypocone.
One part is composed of the axial portion of the cusp with
a reduced mesial segment, the second part comprises the
distal segment and distal marginal ridge. Groove 1me is
worn off, while 2me, whi ch separates the two portions of
the cusp, is of substantial length and depth and is similar
to the intertubercular fissures. In the disto-vestibular
Fig. 4. Denisova 4: root system.
Fig. 1. Denisova 4: disto-
Fig. 2. Denisova 4: occlusal
Fig. 3. Denisova 4: odontoglyphic pattern.
A.V. Zubova, T.A. Chikisheva, and M.V. Shunkov / Archaeology, Ethnology and Anthropology of Eurasia 45/1 (2017) 121–134 129
portion of the crown, 2me divides the marginal ridge,
and extends well into the external surface of the tooth
wall, reaching its lower third. In the central portion, 2me
falls into ﬁ ssure III. The axial ridge of the metacone is
as massive in Denisova 8 as in Denisova 4. From the
distal side, it is divided by accessory grooves 4me and
4’me, which delineate elements of the metaconulus in its
structure (Fig. 7).
A massive ridge is prominent in the cusp that is formed
by the distal part of the metacone. This ridge contacts the
axial ridge of the hypocone, and forms a structure that is
parallel to the plagiogrista but is interrupted by the sagittal
groove dividing the metacone and hypocone. The ridge is
divided into two parts by an accessory groove falling into
2me on one side, and into the posterior fovea on the other
side. The fovea delimits the ridge distally, and separates it
from the distal marginal ridge. On the vestibular surface of
the metacone, at the interface with the paracone, there is a
well-deﬁ ned mesiostyle. The enamel extension is absent
on the vestibular side of the tooth.
The paracone is substantially smaller than the
metacone. The mesio-distal diameter of the former
visually corresponds to the mesial portion of the latter.
Owing to strong attrition and postmortem destruction,
it is difficult to describe the paracone. We can only
hypothesize the presence of an accessory groove 4ра,
which dissected the axial ridge parallel to ﬁ ssure II.
The hypocone is substantially reduced, and it
does not form the disto-lingual corner of the crown.
The cusp is smaller than the metacone and protocone
and not as prominent as in Denisova 4. According to
A. Dahlberg’s scale, which is used in th e Zubov protocol,
the hypocone can be assigned to grade 4, and according
to ASUDAS, to grade 5 (Turner, Nichol, Scott, 1991).
The cusp is segmental in shape, with the apex lying at
the intersection between ﬁ ssures III and IV, and it is
strongly morphologically differentiated. Three ridges
can be observed on its surface: mesial, central, and
distal marginal. The distal segment of the major portion
Fig. 7. Denisova 8: odontoglyphic pattern.Fig. 5. Denisova 8 molar before the
Fig. 6. Denisova 8 molar
after the restoration.
of the cusp is reduced. The mesial ridge is very narrow
and short, and appears as a thin enamel crest branching
out of the margin of the lingual surface in the vestibular
direction. It is separated from the axial ridge by the 2hy
groove, which merges with intertubercular ﬁ ssure IV
near the lingual margin of the crown (Fig. 7). The axial
ridge of the hypocone is outstandingly massive. It lies
in the vestibular-lingual direction, parallel to the crista
oblique (plagiocrista). It is delimited in distal portion
by 1hy, which is almost merged with the posterior fovea
separating the axial ridge of the hypocone from its distal
marginal ridge. The fovea is very narrow, and appears
as a deep groove interrupted by an accessory distal
cusp (grades 3–4 ASUDAS). The ﬁ ssure separating the
hypocone and protocone terminates on the surface of
the marginal ridge, instead of extending to the lingual
surface as is usually found. A thin continuous bridge
between the hypocone and protocone remains at the
intersection between the occlusal and lingual surfaces
of the tooth. It can be interpreted as a rudiment of
the archaic ridge called “entocrista”, which is found
in primate dentition, but in modern humans is only
observed in the deciduous second molar buds (Zubov,
The root system of the tooth is destroyed; but the
morphology of the remaining fragments points towards
the presence of three roots: lingual, distal, and mesial.
On the taxonomic status of the dental pattern
of the Denisovan upper molars
The permanent molars from Denisova Cave have a
number of common traits, which suggests that both
specimens belong to the same taxon. The most speciﬁ c
features of their dental pattern are: large size and rounded
corners of the crowns; absence of notable reduction of
the distal teeth in the row of molars; exceptionally strong
development of the grooves of the ﬁ rst and second orders,
which usually extend to the walls of the crown or form
accessory cusps; and presence of accessory grooves of
the third order, which are not found in modern humans.
In both teeth, there are cingular structures represented by
numerous entostyles in the Carabelli complex, accessory
marginal cusps, and mesiostyle of the upper molars.
This series of features also includes a pronounced trend
towards formation of ridges on the occlusal surfaces of
major cusps. This trend is evidenced in the formation
of a wide and complex plagiocrista; a segmentation of
the metacone accompanied by the emergence of a ridge
parallel to the plagiocrista; a segmentation of major cusps;
and in the persistence of the entocrista and elements of
According to the results of the comparative analysis,
the combination of traits observed in Denisovan
molars does not match the dental patterns of any
European hominin taxa. Most traits typical of Denisovan
pattern are highly archaic markers. Despite this, the
Denisovan dental complex lacks Neanderthal features
(Khaldeyeva, Kharlamova, Zubov, 2010; Bailey, 2002)
that might suggest an afﬁ nity of the population studied to
H. heidelbergensis. The latter taxon is characterized by
the increased frequency of archaic markers, accompanied
by the presence of Neanderthal traits (Martinón-Torres
et al., 2012). Markers of H. sapiens lineage, such as
strong reduction of the hypocone of the second upper
molar and reduction of the grooves of the third order, are
not present in the Denisovan dental pattern. A substantial
reduction of the hypocone of the upper third molar is the
only relatively advanced evolutionary feature observed in
Denisovan teeth. At the moment, there are not enough data
to discuss apomorphic features of this species.
The morphological features of the molars from
Denisova Cave are generally consistent with the pattern
typical of the Archantropus evolutionary stage. They are
mostly similar to the dental complexes found in H. erectus
of Southeast Asia; in particular, in Sangiran specimens.
Sangiran complexes are extremely variable, but there
are megadontic specimens among them, comparable in
size to Denisovan teeth (Lovejoy, 1970: Tab. 2; Orban-
Segebarth, Procureur, 1983: Tab. 2; Tyler, 2001). Almost
the full set of archaic markers typical of Denisovans
is observed in the Sangiran specimens. First, the
frequency of cingular derivatives (ectostyle and ectostylid
extensions on the vestibular and lingual surfaces of the
upper and lower molars and accessory cusps) is increased
in the Sangiran specimens: NG 8503 (Kaifu, Aziz, Baba,
2005), Sangiran 5, 6, 7, 9 (Schwartz, Tattersall, 2003:
Vol. II), Sangiran 8 (Kaifu, Aziz, Baba, 2005), Sangiran
22, 27, 33 (Ibid.), and Bpg 2001.4 (Zaim et al., 2011). The
trend towards the formation of ridges is in many cases as
pronounced in Sangiran hominins as it is in Denisovans.
However, it is unclear from published data if the entocrista
and eocrista persisted in the specimens from Java, and
if they displayed some archaic odontoglyphic variants.
For instance, a posterior fovea appearing as a T-shaped
ﬁ ssure was observed in specimen Bpg 2001.4 (Ibid.).
Trapezoid contours with rounded corners, similar to those
of Denisovans, were described in the upper molars of
Sangiran 4, 27, and Bpg 2001.04 specimens (Schwartz,
Tattersall, 2003: Vol. II; Zaim et al., 2011).
Another location of dental specimens morphologically
similar to the Denisovan teeth is Xujiayao, a ﬁ nal Middle–
early Upper Pleistocene site in northern China (Xing
et al., 2015). The upper permanent molars from Xujiayao
display the following set of features: very large size, sub-
square shape of the crown, accessory mesial and distal
marginal cusps, complex shape of the Carabelli cusp,
differentiated odontoglyphic pattern with extremely
pronounced major tubercular grooves, and a tendency
towards fragmentation of the metacone and hypocone
(Ibid.: Fig. 2). An extremely strong divergence between
vestibular and lingual roots, similar to Denisova 4, was
observed in the РА 1481 and РА 1500 upper molars (Ibid.:
Fig. 3). The denta l specimens from Xujiayao display
a mosaic morphology, which is substantially different
from the samples of early modern humans from China,
but retain an archaic component that makes them similar
t o the dentition of Middle Pleistocene hominins from
East Asia: Sangiran, Zhoukoudian, Longtandong, and
Chaoxian. From Neanderthal complex markers, only non-
speciﬁ c traits, broadly found in the samples of the Middle
Pleistocene hominins, were observed in the Xujiayao teeth
The taxon omic status of the Xujiayao samples relative
to H. sapiens and H. neanderthalensis has not yet been
determined. But the strong similarity in morphology of
the upper molars from Xujiayao and Denisova Cave might
suggest that the two populations could have belonged to
the same taxon, exhibiting a long persistence of erectoid
traits. If future research conﬁ rms this similarity, this will
become a strong argument to support the hypothesis that
Denisovans were widespread in East Asia (Reich et al.,
2010; Derevianko, 2011). It is of note, though, that the
complex of archaic features common to H. erectus from
Sangiran and Denisovans is substantially reduced, or
absent, in other Chinese ﬁ nds (Turner, Manabe, Hawkey,
2000; Wu, Poirier, 1995; Schwartz, Tattersall, 2003:
Vol. II; Xing, Zhou, Liu, 2009; Liu et al., 2010).
The skeletal remains of H. ﬂ oresiensis (Brown et al.,
2004), a species that emerged as a result of long
island isolation, confirm the possibility of long-term
conservation of erectoid morphology. The dental pattern
typical of Denisovans and Xujiayao hominins suggests the
presence of one more locus of evolutionary conservation
in East Asia.
The simil arity between dental complexes of the Upper
Paleolithic population from Altai, the Middle Pleistocene
hominins from China, and the Lower Paleolithic
A.V. Zubova, T.A. Chikisheva, and M.V. Shunkov / Archaeology, Ethnology and Anthropology of Eurasia 45/1 (2017) 121–134 131
population from Southeast Asia does not contradict the
results of paleogenetic studies. The estimated time of
divergence between the ancestors of Denisovans and the
common ancestor of H. sapiens and H. neanderthalensis
coincides with the latest dates obtained for H. erectus
ﬁ nds from Java (Pope, Cronin, 1984). In Altai, the Karama
site has a similar age (Bolikhovskaya, Derevianko,
Shunkov, 2006). Thus, a migration becomes a feasible
explanation for the similarity between the dental
patterns of Denisovans and those of H. erectus from
Java. Importantly, the complex of archaic morphological
features is more pronounced in later Javanese H. erectus
than in earlier specimens (Kaifu et al., 2005).
The great est genetic impact of Denisovans is found in
modern populations from Southeast Asia and Melanesia
(Reich et al., 2010), Papua-New Guinea, Polynesia, and
Fiji (Reich et al., 2011). The results of these studies
have shown that admixture of Denisovan and basal
modern human genomes could have occurred in neither
the northwest nor the west of the Asian continent. The
admixture between these two species most probably took
place in Southeast Asia (Ibid.: 523). The paleogenetic data
also suggest that Denisovan genes were widespread in this
region before the advent of modern humans.
Our results have brought us to the following major
conclusions. First, the conservation of archaic components
without any replacement by more progressive features
was the main evolutionary trend in the emergence of
H. altaiensis. The most prominent feature of dental
morphology of this species is the set of erectoid traits
found in both molars studied. The set is fully present
in both Denisovan individuals, despite the high level of
genetic divergence between them. Genetic diversity was
generally very low in the Denisovan population (Meyer
et al., 2012; Slon et al., 2015), which sharply contrasts
it to the maximally broad adaptive radiation and genetic
diversity typical of modern humans. In this respect,
Denisovans were more similar to Neanderthals who, as
compared to modern humans, were a more specialized
species with a lower level of genetic diversity (Reich
et al., 2010: 1055).
Second, the peculiar morphology similar to the
Denisovan molars is found only in Asian hominins,
but not in any European specimens. Thus, the origin
of H. altaiensis is most probably related to Asian
H. erectus, which is supported by archaeological data
Judging by the prevalence of erectoid features
in the morphology of the Denisovan molars, we can
hyp othesize that the part of De nisovan genome related
to an unknown hominin species (Krause et al., 2010)
might have belonged to H. erectus. This is just a very
tentative suggestion, since most of the archaic traits in
the Sangiran hominins dentition are not apomorphic,
but rather inherited from more ancient species of the
genus, H. habilis and H. rudolfensis. However, from the
point of view of dental morphology data, the presence of
some genetic heritage of Asian H. erectus in Denisovans
appears fairly well-based.
The results of the present study confirm the high
importance of dental traits in detecting interspecific
differences in the genus Homo. The results have also
conﬁ rmed the presence in Altai of a speciﬁ c hominin
population referred to as H. altaiensis, and different from
H. sapiens and H. neanderthalensis not only genetically
but morphologically as well. Peculiar features of dental
morphology of this population are megadontia; and a
long-term conservation of the dental markers typical of
the Middle Pleistocene hominins of Northern China, and
of H. erectus from Sangiran.
Thus, H. altaiensis exhibits a very conservative mode
of morphological evolution.
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morphology has conﬁ rmed the equal validity of genetic
and morphological criteria for differentiating hominin
species, which has been a matter of hot debate in
paleoanthropology. Moreover, this morphological
analysis has enabled us to put forward a well-based
hypothesis according to which the unidentiﬁ ed portion of
the Denisovan genome belongs to H. erectus s.l. Thus, our
results emphasize the fact that classical dental studies still
retain an independent value not lessened by the advent of
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