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Journal of Human Evolution 61 (2011) 186e196

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Journal of Human Evolution

journal homepage: www.elsevier.com/locate/jhevol

A human mandible (BH-1) from the Pleistocene deposits of Mala Balanica cave
(Si

cevo Gorge, Nis, Serbia)
Mirjana Roksandic a, *, Dusan Mihailovi
c b, Norbert Mercier c, Vesna Dimitrijevi

c b, Mike W. Morley d,
Zoran Rako c e, Bojana Mihailovi

cevi
c f, Pierre Guibert c, Jeff Babb g
a
Department of Anthropology, University of Winnipeg, 515 Portage Avenue, Winnipeg, MB, R3B 2E9 Canada
b 
Department of Archaeology, Faculty of Philosophy, Belgrade University, Cika Ljubina 18-20, 11000 Belgrade, Serbia
c
Centre de Recherche en Physique Appliquée à l’Archéologie, Université de Bordeaux Maison de l’Archéologie 3607 Pessac cedex, France
d
Human Origins and Palaeo-Environments (HOPE) Group, Department of Anthropology and Geography, Oxford Brookes University, Gipsy Lane, Oxford OX3 0BP, UK
e
Center for Digital Radiography, Faculty of Dentistry, Belgrade University, Rankeova 6, 11000 Belgrade, Serbia
f
National Museum, Trg Republike 1a, 1100 Belgrade, Serbia
g
Department of Mathematics and Statistics, University of Winnipeg, 515 Portage Avenue, Winnipeg, MB, R3B 2E9 Canada

a r t i c l e i n f o a b s t r a c t

Article history: Neandertals and their immediate predecessors are commonly considered to be the only humans inhabiting
Received 23 October 2009 Europe in the Middle and early Late Pleistocene. Most Middle Pleistocene western European specimens
Accepted 21 February 2011 show evidence of a developing Neandertal morphology, supporting the notion that these traits evolved at
the extreme West of the continent due, at least partially, to the isolation produced by glacial events. The
Keywords: recent discovery of a mandible, BH-1, from Mala Balanica (Serbia), with primitive character states
Neandertal
comparable with Early Pleistocene mandibular specimens, is associated with a minimum radiometric date
Archaic Homo sapiens
of 113 þ 72  43 ka. Given the fragmented nature of the hemi-mandible and the fact that primitive
European hominin dispersals
Central Balkans
character states preclude assignment to a species, the taxonomic status of the specimen is best described as
an archaic Homo sp. The combination of primitive traits and a possible Late Pleistocene date suggests that
a more primitive morphology, one that does not show Neandertal traits, could have persisted in the region.
Different hominin morphologies could have survived and coexisted in the Balkans, the “hotspot of
biodiversity.” This first hominin specimen to come from a secure stratigraphic context in the Central
Balkans indicates a potentially important role for the region in understanding human evolution in Europe
that will only be resolved with more concentrated research efforts in the area.
Ó 2011 Elsevier Ltd. All rights reserved.

Introduction
The settlement of Europe has a long history, with hominins
present at the gates of the continent in Dmanisi (Georgia) as early as
1.7 million years ago (Gabunia et al., 2000). The recent recognition of
several new species of Homo in Europe, including H. georgicus
(Gabunia et al., 2002) in Dmanisi, Georgia, H. antecessor (Bermúdez
de Castro et al., 1997) in Atapuerca, Spain, and H. cepranensis from
Ceprano in Italy (Mallegni et al., 2003), as well as H. heidelbergensis
(Schoetensack, 1908), suggests considerable diversity in the Euro-
pean hominin record. While western European specimens from the
* Corresponding author.
E-mail addresses: m.roksandic@uwinnipeg.ca (M. Roksandic), dmihailo@f.bg.ac.rs
(D. Mihailovi
c), norbert.mercier@u-bordeaux3.fr (N. Mercier), vesnadim@beotel.rs
(V. Dimitrijevi
c), mmorley@brookes.ac.uk (M.W. Morley), mailzrakoc@yahoo.com
(Z. Rakocevi
c), salitrena@gmail.com (B. Mihailovi

c), pierre.guibert@u-bordeaux3.fr
(P. Guibert), j.babb@uwinnipeg.ca (J. Babb).
Middle and Late Pleistocene show derived Neandertal traits, this is
not necessarily true for the East and the South of the continent. The
existence of areas of refuge in the Mediterranean peninsulas could
have played an important role in maintaining the variability of
hominins in Europe, by a combination of migratory pulses and in situ
evolution. With a revised date, the Ceprano skull from the Apennine
Peninsula could be regarded as one example where primitive
morphology persisted into the Middle Pleistocene (Muttoni et al.,
2009). As a result, the peopling of Europe in the Pleistocene may
have been very complex and the current fossil record may represent
any number of different genetic processes including drift, founder
effect, directional adaptation and hybridization. Reconstructing
human evolution in the regional European context is therefore
inseparable from understanding the migrations of early hominins
into the area and their later movements into and out of Europe. The
Balkan Peninsula, with its river valleys used as migratory routes for
animals and humans in prehistoric as well as historic times, repre-
sents the most likely corridor from Asia Minor into Europe and is no

0047-2484/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jhevol.2011.03.003
 Author's personal copy
M. Roksandic et al. / Journal of Human Evolution 61 (2011) 186e196
Figure 1. The context of the find: (a) Map of Southeast Europe indicating sites with fossil human remains: 1 Krapina, 2 Vindija, 3 Crvena stijena, 4 Balanica, 5 Bacho Kiro, 6
Petralona, 7 Theopetra, 8 Lakonis, 9 Apidima. Small letters indicate newly discovered Paleolithic sites in Serbia (aeh) and Monte Negro (iek) without hominin fossils. (b) The
location of Balanica cave complex in the Si
cevo Gorge.
doubt of extreme importance in understanding Paleolithic migra-
tions (Mihailovi
c, 2009; Dennell et al., 2010). However, the region
has a very limited Paleolithic research history and very few fossil
finds (for a recent overview of Paleolithic research in Greece see
Harvati et al., 2009). New systematic prospection of several regions
in Serbia and excavations of newly identified Paleolithic sites
(Mihailovi
c, 2008) (Fig. 1a), show that the dearth of fossil evidence
from southeast Europe should be attributed to the lack of systematic
research in the area.
A recently discovered hominin fossil from Mala Balanica1 cave is
the first hominin specimen in the Central Balkans recovered from
controlled excavations with firm stratigraphic context. Here, we
present a detailed description of the partial human hemi-mandible
from Mala Balanica (BH-1) in its stratigraphic context and discuss
its potential significance. The preservation of the specimen (see
below) precludes a more complete analysis of its species status,
while its primitive morphology raises questions about the role that
the Balkan Peninsula, the least researched glacial refugium in
Europe, played in hominin evolution on the continent.
Geoarchaeological and cultural context
Mala Balanica (N43 20.2110, E22 05.1150 ) is an elongate karstic
chamber, 8 m wide, 25 m deep and 2.8 m high, measured from the
present-day surface in the interior of the cave. The site is situated at
an elevation of 332 m above sea level, approximately 100 m above
the Nisava River, with the cave mouth facing SSW across the valley
(Fig. 1b). It is separated by a distance of 7 m from the entrance to the
larger Velika Balanica cave with which it forms the Balanica Cave
Complex. The cave complex is situated on the southern slopes of
the Svrljig Mountain chain, formed of Cretaceous and Jurassic
limestone in the area where the Rodop Mountains meet the
younger chains. The Si
cevo Gorge is cut through by the Nisava River,
which provides an important communication between two
1
pronounced Bal
anitza.
187
adjoining river valleys. Situated at the exit from the gorge, the caves
are both easily accessible and secluded, as is often the case with
Middle Paleolithic localities (Mellars, 1996). Today the catchment
area is characterized by a moderate continental climate with
deciduous, oak-dominated forests. The site is situated in an active
karst setting (Woodward and Goldberg, 2001) and is subject to
groundwater infiltration through cracks and fissures in the host
bedrock. The present development of the cave system appears to be
driven primarily by solution through infiltration and colluvial slo-
pewash processes.
In 2005, a test excavation of a small 2 m2 area near the western
wall of the cave identified a number of quartz artifacts and Quina
scrapers immediately below the present-day surface, and revealed
a clandestine pit in the southwest corner of the cave. During the
following four seasons, the excavation of artefact-bearing layers
(averaging 30 cm in depth) was extended to a 12 m2 surface (Fig. 2),
and the backfill from the clandestine excavations was cleared. The
latter allowed us to expose a profile over 2 m below site datum,
which was subsequently used to gather geoarchaeological and
palaeontological information (Table 1; Fig. 3).
In very broad terms, the stratigraphy exposed in the studied
profile comprised well-bedded, horizontal to sub-horizontal layers
of limestone gravel contained within a silt- and sand-dominated
matrix, with major variations in the ratio of coarse (gravel) to fine
(silt-dominated) components. The profile has been bulk sampled
for detailed geoarchaeological analysis, which will provide impor-
tant paleoenvironmental context to the archaeological material
recovered from this area of the site, including the BH-1 mandible.
This work is currently ongoing and the results will be presented
elsewhere. For the purposes of the current paper, the stratigraphic
succession will be described in terms of field observations made
during detailed sediment logging as part of both the archaeological
and geoarchaeological fieldwork.
The exposed sequence has been divided into three facies with
the aim of describing groups of sedimentary units with similar
depositional (and post-depositional) histories. These facies are
differentiated chiefly by colour, composition and the inclusion of
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188 M. Roksandic et al. / Journal of Human Evolution 61 (2011) 186e196

Figure 2. Surface contours of the cave with the excavation grid. Inset shows the details of the excavation area where the specimen was found.

anthropogenic material (e.g. lithics). Facies 1 is a thin (c. 15 cm),
sub-horizontal layer of silt and limestone clasts, which represents
Holocene and sub-recent deposition at the top of the sequence.
Facies 2 (a e h) is characterized by clast-supported, coarse angular
to sub-angular gravels held within a matrix of reddened silts and
sands. Facies 3 (a e c) is typified by matrix-supported, sub-angular
to sub-rounded gravel in a matrix of brown silts and sands. At the
base of the sequence, unit 3c consisted of fine-grained silts and
clays, some of which were well-cemented, with a paucity of lime-
stone clasts. Facies 2 and 3 represent Pleistocene accumulation.
At the macroscopic level, the sequence is essentially minero-
genic in nature, with no major organic component within the
sedimentary matrix except for occasional faunal material.
Unequivocal evidence of human activity was noted only in Facies 2
(aec), which could be the result of the very limited excavation area
in which the profile was observed, as well as the limited depth of
the excavation in the remainder of the squares. Animal bones are
present throughout the sequence, although they are less common
in Facies 2de2h. Though the physical characteristics of the litho-
logic units are relatively uniform, a number of preliminary obser-
vations concerning the sedimentary sequence can be made. From
the base of the sequence upwards there is a general trend towards
coarser sedimentary components, with the frequency of medium to
large limestone clasts increasing towards the base of Facies 2. Unit
3c, at the base of the sequence, comprised well-cemented, fine-
grained silts and clays, suggestive of a wetter climate, possibly with

Table 1
Features of the sedimentary layers at Mala Balanica.

Layer Summary characteristics of lithologic facies and units Archaeological material Faunaa
1 Friable clay silt with mod-freq, sub-ang Recent and sub-recent Ovis aries
to sub-r, fine-medium limestone gravel. material, charcoal
2a Mod compact silt with mod-freq, Present only near the Canis lupus, Felis silvestris Schreber,
ang to sub-ang, fine-med limestone gravel. Western wall of the cave, Cervus elaphus, Capra ibex.
Middle Paleolithic artefacts
2b Friable silt with freq med-large, ang to sub-ang, limestone clasts. Throughout the cave, Middle Castor fibre, Lepus sp., Canis lupus,
Clast frequency increases towards base. Paleolithic artefacts, charcoal Ursus sp., Panthera pardus,
Felis silvestris.
2c Clast-supported, poorly sorted, ang to sub-ang, Present sporadically, Middle Apodemus sp., Castor fibre,
fine-med limestone gravel in a silt matrix Paleolithic artefacts, charcoal Ochotona pussila, Canis lupus,
2d Mod compact, slightly sandy silt with mod-freq, sub-ang to sub-r, Very few artefacts in the Vulpes vulpes, Martes martes,
fine limestone gravel e coarser towards base. topmost segment Mustelidae indet., Ursus arctos,
of the layer, no charcoal Ursus sp., Crocuta spelaea,
2e/2f Mod compact sandy silt with occ-mod, No artefacts, no charcoal Felis silvestris, Cervus elaphus,
fine-med, sub-ang to sub-r limestone gravel. Dama dama, Capreolus capreolus,
2g Matrix-supported, mod-sorted, ang to sub-ang, fine limestone gravel in a No artefacts, no charcoal Rupicapra rupicapra, Capra ibex.
sandy silt matrix. Appears clay enriched and more compact towards base.
Freq degraded limestone fragments and flecks
2h Matrix-supported, mod-sorted, ang to sub-ang, fine-med limestone No artefacts, no charcoal
gravel in a silt matrix. Clast size increases towards base
3a Matrix-supported, poorly sorted, sub-ang to sub-r, med-coarse No artefacts, no charcoal Ursus arctos, Ursus sp., Equus sp.,
limestone gravel in a calcareous sandy silt matrix. Freq CaCO3 inclusions. Capreolus capreolus.
Mod-freq large bone frags
3b Matrix-supported (occ clast-supported towards upper contact), No artefacts, no charcoal Canis sp., Ursus sp., Crocuta spelaea,
mod to poorly sorted, ang to sub-r, fine to medium (but occ coarse) Layer in which the BH-1 Cervus elaphus, Dama dama, Capra ibex.
limestone gravel. Darker and less calcareous than layer 3a mandible was found
3b0 Very well carbonate-cemented silt with occ fine gravel. Also contains No artefacts, no charcoal Canis sp., Ursus sp., Crocuta spelaea,
nodular calcium carbonate inclusions which Cervus elaphus, Dama dama, Capra ibex.
rest on upper contact of underlying layer
3c Dark, compact silty clay with occ poorly sorted, fine-med limestone No artefacts, no charcoal Carnivora indet., Cervus elaphus

Abbreviations are as follows: occ (occasional); mod (moderate); freq (frequent); ang (angular); r (rounded).
a
Additionally, the following taxa were identified from mixed contexts: Meles meles (Linnaeus), Bos/Bison.
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M. Roksandic et al. / Journal of Human Evolution 61 (2011) 186e196
Figure 3. The profile excavated along the northern wall of the clandestine pit. Sedimentary units are described in Table 2. Layers 2ae2c produced Mousterian artifacts, layer 3B
produced the hominin mandible, BH-1.
water pooling at the site. Unit 3b appeared clay- and fine silt-
enriched towards its base, again indicating a wetter climate
allowing fine-grained allogenic sediments to be washed into the
site. The limestone clasts observed in Facies 3 were rounder than
the angular to sub-angular clasts of Facies 2, implying partial
dissolution as a consequence of the throughput of percolating
groundwaters. With the coarse angular gravel of Facies 2 typifying
a cooler climate where cryoclastic processes promoted the gener-
ation of eboulis-like autogenic sediments (though issues of equi-
finality should be borne in mind, see Bailey and Woodward, 1997),
preliminary observations advocate a warm e cooling phase repre-
sented in the Facies 3e2 up-profile sequence. This proposal would
broadly concur with the faunal observations of warm-adapted
species recovered from unit 3b. A marked variation was also
observed in the unit-to-unit variability in carbonate content. This
parameter can be used as a paleoenvironmental proxy in identi-
fying warmer, wetter climatic episodes (Woodward, 1997;
Woodward and Goldberg, 2001; Schuldenrein, 2001; Straus et al.,
2001) and this may help refine the paleoenvironmental recon-
struction at a later stage. At present it is unknown whether these
carbonates are primary (e.g., aeolian carbonate silt) or secondary
(i.e., reprecipitated).
Stratigraphic context of the find
The mandibular specimen, BH-1, originated from layer 3b,
characterized by the presence of larger animal bones, fewer and
less angular limestone clasts, and no other evidence of anthropo-
genic activity. BH-1 was situated 1.5 m below the lowest recorded
artifacts. The layer in which the specimen was found was excavated
on a very restricted surface below the clandestine pit. The bottom of
189
the clandestine pit was reached at the top of layer 3b, in squares E,
F, G 19/20; consequently, the pit did not disturb the underlying
sediments. Controlled excavation in squares D18a, b and E18d failed
to reach bedrock and the stratigraphic sequence continues below
the current level. The specimen was found in square D18, layer 3b,
three arbitrary 5 cm spits below the base of the clandestine pit. The
mandible was removed together with some adjoining sediment.
Other animal bones and nodular calcium carbonate were found and
recorded in situ in the same layer. No other fragments of bone were
found in the faunal material from this and the surrounding exca-
vation loci. The faunal remains showed the same adhered sediment
and colouration as observed on the mandible. While the area
excavated is small and a detailed site formation reconstruction is
currently beyond our reach, there is no indication that the strati-
graphic sequence at this particular part of the cave was disturbed
by geological, anthropogenic or biogenic processes. Fragmentation
of faunal material, evidence of gnawing by carnivores and rodents,
and the general uniform slope of the sediments towards the
western wall throughout the excavated area, indicate that accu-
mulation could have originated towards the middle of the cave,
gradually fanning out towards the sides of the cave. Evidence of
water logging on the mandible cannot be readily interpreted; it
could be due to a relatively long exposure prior to burial, or to the
water pooling in layer 3c suggested by geoarchaeological obser-
vations, or a number of other factors.
Fauna from the Mala Balanica cave
Animal bones are present throughout the cave sequence. The
bones and teeth of large mammals are well preserved, but highly
fragmented. Many are damaged by carnivore and rodent gnawing.
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190 M. Roksandic et al. / Journal of Human Evolution 61 (2011) 186e196
Some fragments have polished surfaces and randomly distributed
striations that are probably due to trampling of the bones and bone
fragments, and their friction with the sediment particles. Anthro-
pogenic modifications are visible from layers 2a to 2d as traces of
fire damage and cut- and chop-marks on animal bones. Given the
small area from which the faunal material has been excavated
below these strata, it is hard to ascertain whether the lack of
anthropogenic modifications in the lower strata is genuine.
The identified taxa are listed in Table 1. Because of the limited
volume of excavations in the lower levels and the low yield of
faunal remains, it is not yet possible to conclude whether there are
significant differences in faunal composition throughout the
sequence. In layers 2ae2c, the most numerous remains are of ibex,
represented by cranial and postcranial elements of several adult
and juvenile animals. Preliminary morphometric analysis points to
a large-sized ibex, which is commonly found in Upper Pleistocene
cave deposits in Serbia (Dimitrijevi
c, 1997). In terms of the pre-
vailing environmental conditions, the most important taxa are two
cervid species, roe deer (Capreolus capreolus) and fallow deer
(Dama dama). Both are present in layers 3 and 2c - 2h and disappear
in layers 2a and 2b. Both species were more common in warm than
in cold stages of the Pleistocene, and preferred a wooded habitat.
Fallow deer were completely absent in Last Glacial deposits in
Serbia (OIS 2e4), while roe deer were absent from most of the sites
where Last Glacial age was ascertained (Dimitrijevi
c, 1997). The
fallow deer finds from layers 3 and 2 correspond in size to Last
Interglacial Dama dama, the species considered characteristic of the
Upper Pleistocene interglacials (Gibbard, 2003). The appearance of
murids (Apodemus sp.) suggests interglacial affinities for the lower
portion of the sequence, as cave deposits of the Last Glacial in
Serbia are dominated by steppe species of arvicolids and cricetids
(Dimitrijevi
c, 1997). The presence of the typical steppe species
Ochotona pusilla in the upper portion of the sequence indicates
changes towards a more open landscape.
A possible interglacial origin for layers 3ce3a would also be
supported by the sedimentology, as the fine-grained sediments of
3c immediately below the layers from which the mandible was
found, suggest a warmer, wetter climatic regime with a cooling
trend towards upper levels of the sequence. Calcium carbonate
levels appear to increase from 3b to 3a, which could indicate
a reduction in the throughput of percolating groundwater, also
suggested by the increase in angularity of the gravel noticeable in
levels 3a up to 2g.
Materials and methods
The mandibular preservation and morphology are described
following the terminology of Rosas (1995). The CT scan was per-
formed at the Centre for Radiological Diagnostics (University of
Belgrade) using a Siemens multi-slice Somatom sensation 16
scanner to obtain sections (interslice distance 0.75 mm) in axial,
sagittal and frontal planes, and were used subsequently to create
a 3-d reconstruction.
The radiometric date of the mandible was obtained by the
U-series method and, in particular, the disequilibrium between the
uranium and thorium radioelements (Table 2). We applied a non-
destructive approach by detecting the gamma rays emitted from
the specimen itself by the decay of radioelements of the U-series
chain. The mandible was placed directly on a broad energy
Germanium detector (thickness ¼ 30 mm, diameter ¼ 76 mm) for
almost two weeks. As gamma rays have a long freepath through
matter, they can be detected with a low background Germanium
detector; the analysis is consequently non-destructive. The lead
screen surrounding the detector and the mandible allowed us to
detect the following gamma ray emissions: 234Th (63.3 and
Table 2
Activities of 234Th, 234U, 234mPa, 235U, 230Th and 214Bi were determined in analyzing
the gamma ray lines whose energies are given in the text.
U-238 (p.p.m.) U-234/Th-232 Th-230/U-234 Age (ka)
2.86  0.04 80 0.649  0.170 113 þ 72  43
The radioisotopic contents of the natural clay used for making the cast were also
determined by gamma spectrometry in using the GSN (granite) standard and cor-
rected for taking account of the difference in densities between the clay and the GSN
standard.
92.8 keV), 234U (53.2 keV), 234mPa (1001.0 keV), 235U (185.8 keV),
230
Th (67.7 keV), 214Pb (53.2, 295.1 and 351.9 keV), 214Bi (609.3 and
1120.3 keV) and 210Pb (46.5 keV). In addition, we recorded the
gamma rays coming from radioelements of the Th-series chain
(228Ac, 212Bi, 212Pb, 208Tl). To obtain a reference standard, we
submitted a cast of the specimen made of a natural clay rich in
radioelements of the U-series chain. This clay is used as a reference
standard because its radioisotopic contents are well known. Since
the cast had the same shape and size as the mandible, the gamma
spectrum recorded during the measurement of the mandible could
be compared with the gamma spectrum recorded for the cast. As
the radioisotopic contents of the cast are known, by comparison of
these two spectra, it is possible to deduce the radioisotopic
contents of the mandible, and consequently its age (see Table 2).
The age estimate of the mandible was calculated assuming an early
uptake model for the uranium, i.e., one in which this radioelement
penetrated the fossil in a relatively short time in comparison to the
burial duration. Since this assumption cannot be confirmed inde-
pendently, the radiological age should be understood as the
minimum date for the specimen. The date obtained was compared
to faunal and geoarchaeological data to help estimate the age of the
mandibular specimen.
Statistical procedures
Cluster analysis was run for morphological traits of BH-1 and 26
specimens reported in Mounier et al. (2009) that had all of the 16
variables observable in BH-1: variables IeL and NeU and variables
OO, PP, QQ and UU (as defined in Mounier et al., 2009: Table 5, Fig. 1
and Table 11, Fig. 1, respectively). An agglomerative hierarchical
clustering procedure was applied using the average linkage crite-
rion, which measures the similarity between any two clusters as
the mean of the similarities between the objects in one cluster and
the objects in the other cluster. The distance measure was
Euclidean distance, applied to non-standardized data. To evaluate
the extent to which the dendrogram reflects the relationships in
the original distance matrix, we calculated the coefficient of
cophenetic correlation, which represents the linear correlation
between entries of the original distance matrix and corresponding
Table 3
Hominin fossils used in cluster analysis with assigned groupings. From
Mounier et al. (2009).
Early Pleistocene (EP)
Homo erectus s.l.: D211, D2600, Sangiran1b (S1b), KNM-ER 992
Homo antecessor: ATD6-96
Middle Pleistocene (MP)
Homo erectus: (s.l.) Tighenif 1 (T1), Tighenif 2 (T2), Tighenif 3 (T3),
Sinanthropus H1 (SH1)
Homo heidelbergensis or Pre-Neandertals: Mauer, AT-888, AT-950,
Arago II (ARII), Arago XIII (AR XIII), Montmaurin, Ehringsdorf F (Eh F)
Upper Pleistocene: Neandertal (NT)
(NT): Krapina J, Krapina G, Spy 1, Shanidar I, Regourdou, Bañolas,
La Ferrassie 1, La Quina H5, Amud 1, Zafarraya
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M. Roksandic et al. / Journal of Human Evolution 61 (2011) 186e196 191

Table 4
Data used in the cluster analysis.

# G Specimen I J K L N O P Q R S T U OO PP QQ UU
1 UU BH-1 3 1 1 2 2 1 1 2 1 2 2 3 2 2 2 1
2 EP D211 3 2 1 1 2 1 2 2 1 2 3 2 1 2 2 1
3 EP D2600 1 2 1 1 3 2 2 2 2 2 2 2 2 1 2 1
4 EP Sangiran1b 3 2 1 2 2 2 1 2 1 2 2 2 2 2 2 1
5 EP KNM-ER992 3 1 1 3 2 2 1 3 1 2 2 2 1 1 2 1
6 EP ATD6-96 3 2 1 2 2 1 1 2 1 2 2 2 2 1 2 1
7 MP Tighenif 1 3 2 1 1 2 1 2 3 1 2 1 2 1 1 2 1
8 MP Tighenif 2 3 1 1 2 2 1 2 2 2 2 2 2 2 1 2 1
9 MP Tighenif 3 3 2 2 2 3 2 2 3 2 2 1 3 1 1 2 1
10 MP Mauer 2 2 2 2 3 2 2 3 3 2 1 3 1 1 2 1
11 MP AT-888 3 1 3 1 2 1 2 2 3 3 1 2 3 1 2 1
12 MP AT-950 3 2 2 1 3 2 2 3 3 3 1 2 1 1 2 2
13 MP Arago II 3 1 3 1 2 2 1 3 3 2 1 3 1 2 2 2
14 MP Arago XIII 3 2 2 1 2 1 2 3 2 2 1 2 1 1 2 2
15 MP Montmaurin 3 2 2 2 2 2 1 2 2 2 1 2 2 1 2 2
16 MP Sinanthropus SH1 3 2 2 2 3 2 2 2 2 3 2 1 2 2 2 1
17 MP Ehringsdorf F 3 2 3 2 2 1 2 1 3 3 1 1 1 2 2 2
18 NT Krapina J 3 2 3 1 3 2 2 2 3 3 1 1 3 1 1 1
19 NT Krapina G 3 2 3 2 3 2 2 1 3 3 1 1 3 3 2 2
20 NT Spy 1 2 1 3 2 1 1 1 2 3 3 1 2 3 2 1 2
21 NT Shanidar I 2 1 2 1 3 2 2 2 3 3 1 2 3 1 1 2
22 NT Regourdou 3 2 3 1 2 1 2 1 3 3 1 2 3 1 1 1
23 NT Ban olas 3 1 2 3 3 2 2 3 2 3 2 1 2 1 1 2
24 NT La Ferrassie 1 3 2 3 2 2 2 1 2 3 3 1 1 3 3 1 2
25 NT La Quina H5 3 1 2 1 2 1 2 1 3 3 1 2 3 3 1 2
26 NT Amud1 3 1 3 1 2 1 2 1 3 3 1 2 3 1 1 1
27 NT Zafarraya 3 2 3 1 3 2 2 1 2 3 1 1 3 2 1 1

G (Grouping): UU (Unknown), EP (Early Pleistocene), MP (Middle Pleistocene), NT (Neandertal). Data for 27 specimens on 16 variables representing 12 selected morphological
characteristics of the lateral face of the corpus (I through L, and N through U) and four morphological characteristics (OO, PP, QQ and UU) of the medial face of the mandibular
corpus described by Mounier et al. (2009).

entries of the cophenetic matrix of merge distances between
taxonomic units (Sneath and Sokal, 1973). The specimens that were
available for statistical analysis are listed in Table 3 and 4.
Results
Radiometric dating of the specimen
Despite the outlined limitations of the dating method, the
mandible can be attributed a minimum age of 113 þ 72  43 ka.
This large uncertainty has two main origins: the counting statistics
of the 230Th (67.7 keV) peak remains low despite the two week
duration of the experiment, and the 234U (53.2 keV) peak is largely
dominated (in a ratio of about 1:10) by 214Bi gamma rays emitted at
almost the same energy. This age estimate, however, must be
considered as a minimum age since there is no independent eval-
uation for the validity of the assumed early uptake model. Contrary
to this assumption, the activities of the post-230Th radioelements
(214Pb, 214Bi, 210Pb), which are about three times lower than the
230
Th activity, suggest that the 226Ra and/or its daughter, 222Rn,
were mobile during at least the last 10 ky. Since there is no
complementary information for older periods, we need to take into
account the possibility that part of the uranium was also mobile
during the burial time.
Lithic industry (Fig. 4)
Only layers 2ae2d show unequivocal evidence of human activity
in the form of stone tools, charcoal fragments and cut-marks on
animal bones. Lithic analysis of the material from both caves of the
Balanica complex identified the presence of different facies of the
Mousterian. The industry of Mala Balanica is limited, with only 102
artefacts, of which 93% were excavated from layers 2ae2c, with an
overall depth of 20 cm. Artefacts made of quartz predominate
(81.4%), the Levallois technique is completely absent, and retouched
artefacts represent 26.7% of the assemblage. Scrapers, some with
typical Quina retouch, are the most common tool type (62.1%), while
denticulates represent 24.1% of the lithic sample. Though limited in
scope, the industry of Mala Balanica is consistent with Charentian-
type Mousterian (Fig. 4). This facies is ascertained on a much larger
scale in the adjoining Velika Balanica cave (Mihailovi
c, 2008, 2009).
Charentian-type Mousterian, present in both Balanica caves, has not
been previously confirmed south of the Sava and the Danube Rivers,
although it is known in adjoining areas, both in Central Europe
(Betalov Spodmol, Erd, Krapina) and in Anatolia at Karain Cave. This
facies cannot be associated with a particular chronological period;
the Proto-Charentian at Karain (complex BeE) was tentatively dated
to 300e330 ka (Koz1owski, 2002), Krapina was dated to 130 ka (Rink
et al., 1995; Simek and Smith, 1997), while Betalov Spodmol and Erd
could be as late as 40 ka (Gábori-Csánk, 1968; Osole, 1991). The
Charentian industry found at the site could represent an early phase
in the development of the Middle Paleolithic in the Central Balkans,
a regional facies, or a particular type of site use by human pop-
ulations in the area.
Geoarchaeological, archaeological, and faunal data seem to
indicate an Early Upper Pleistocene age for the entire sequence. The
lower portion of the sequence (layers 3ce3a) is consistent with
a wooded habitat and likely represents a relatively mild and wet
period compatible with OIS5 (e, c, a). In the upper portion of the
sequence, we observe a cooling trend and the presence of steppe
species, indicating changes towards a more open landscape. This
could be related to environmental changes at the beginning of the
Last Glacial, i.e., OIS 4. This would be in accordance with the
radiometric date of 113 þ 72 -43 years for the lower layers.
However, until the ESR and U-series dates are available, we cannot
exclude the possibility that the find could be older.
The BH-1 specimen
The BH-1 specimen is a 67 mm long left hemi-mandible,
preserved from the posterior margin of the canine alveolus to the
mesial aspect of the ascending ramus with all three molars present
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192 M. Roksandic et al. / Journal of Human Evolution 61 (2011) 186e196
Figure 4. A sample of tools from Mala Balanica layers 2a-2c: Transverse (a,b) and
lateral scrapers (c,d) made of quartz (a) and flint (bed).
in their sockets. The lower half of the mesiolingual root of the M3 is
missing and the remaining roots are exposed due to the destruction
of the adjacent endomandibular lamina. The mesial portion of the
mandible shows an old breakage filled with sediment, whereas all
of the breaks on the distal end are fresh. The alveoli of the P3 and P4
are complete and are for the most part filled with sediment, while
the posterior portion seems to have been subject to water infil-
tration resulting in substantial fragility that, together with the
damage sustained during excavation, could explain its poor
Figure 5. The BH-1 specimen: (a) exomandibular view, (b) endomandibular view, (c) occlusal view, (d) dentition: from left to right M1, M2 and M3. Mesial (m) is up, distal (d) is
down, buccal (b) is left, and lingual (l) is right. Scale in centimetres.
preservation. The posterior portion of the mandible was recon-
structed by refitting the fragments with small amounts of Paraloid
B72 dissolved in acetone. One positively identified small fragment
could not be refitted because it had no direct contact with the rest
of the bone.
Age and sex
Complete eruption and closure of the root apex of the M3 indi-
cate an adult individual, while minimal wear on the M3 and slight
to moderate wear on the M1 and M2 each suggest a relatively young
adult. Given the mandible’s unclear taxonomic position, it is
impossible to determine the sex of the individual.
Exomandibular aspect (Fig. 5a)
The BH-1 mandible is lacking the symphyseal region and the
presence/absence of the mental eminence could not be ascertained.
In the lateral view, the basal and alveolar margins are almost
parallel. The mandible shows only very faint relief in the exo-
mandibular view. The superior marginal torus is poorly defined. It
is represented by a slight change in the orientation of the lamina to
the axes of the horizontal branch above and below the mental
foramen. It transitions smoothly into the lateral prominence,
located at the level of M1/M2, equidistant from the alveolar and
basal margins. The ascent of the oblique line begins just above the
posterior marginal tubercle, 18.5 mm below the alveolar border at
the level of M1/M2 vertically, and the mental foramen horizontally.
While the position is the same as in a comparative sample of
modern humans, the ascent is not as steep. The lateral prominence
is more anterior than in Neandertal and pre-Neandertal samples,
where it is commonly located under the M3 (Rosas, 2001). The
fragment of the exomandibular lamina of the vertical branch, which
could not be refitted but could be identified on the basis of the
visible interior structure of the mandibular canal, shows very slight
relief at the masseteric fossa, with no pronounced rugosities.
The reconstructed root of the vertical branch does not indicate the
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M. Roksandic et al. / Journal of Human Evolution 61 (2011) 186e196
presence of a retromolar space. However, it does not obscure the M3
in lateral view. This is most likely due to a wide extramolar sulcus,
as the vertical branch overlaps with the edge of the M3 in the
occlusal view.
The mental foramen is situated below the P4 alveolus, 17.0 mm
from the alveolar margin and 16.4 mm from the basal margin. It is
oval in shape, set obliquely to the sagittal and transverse planes,
and measures 5.2 mm by 2.9 mm. In contrast, the mental foramen is
located under the M1 in up to 80% of Neandertal specimens and 54%
of the Middle Pleistocene samples from Sima de los Huesos (Rosas,
2001). This position is often interpreted to be a reflection of the
development of marked midfacial prognathism (Quam and Smith,
1998). The more anterior position of the mental foramen in this
specimen and the possible absence of a retromolar space, both
plesiomorphic traits observable in Homo erectus, indicate that the
mandible lacks autapomorphies of Neandertals and their Middle
Pleistocene precursors. Similarly plesiomorphic is the equidistant
position of the mental foramen in relation to the alveolar and the
basal margins. The anterior marginal tubercle could not be
observed in this specimen, as the basal margin is missing mesially
from below the mental foramen.
Endomandibular aspect (Fig. 5b)
The internal lamina is preserved from the canine alveolus to the
M2/M3 septum in the alveolar portion, and from below the P3/M1
septum to below the mesial edge of the M3 alveolus in the basilar
border. There is visible flaking and root etching all over the endo-
mandibular surface. The mesiolingual root of the M3, the lingual
lamina of the M3 alveolus, and the adjacent area above the basilar
edge are missing. A trowel mark can be observed on the M3 and the
interior lamina.
While the bone is robust, the relief of the internal surface is not
marked. The alveolar border shows thickening on the lingual side
from P3 to M2 (and possibly beyond), forming a mandibular torus just
below the alveolar process. The width of the alveolar process
decreases mesially. The width of the subalveolar plane increases
towards the middle portion of the mandible, forming a shelf-like area
(planum alveolare) that extends from below the P3 towards the canines
and the symphysis, best observed in the occlusal view (Fig. 5c). The
mesial endomandibular increase in width is observable towards the
upper third of the mandibular height below the alveolar border. Since
the lower portion of the mandible is broken in the canine area, we
have combined overlapping CT images at the canine alveolus and at
the mental foramen showing thickening at the level of the canine and
an oblique rather than subvertical alveolar planum (Fig. 6).
The subalveolar plane (sublingual fossa) is flat rather than
concave. The profile of the submandibular fossa is moderately
concave. The expression of the mylohyoid line is moderate, and is
Figure 6. Cross-section of the mandible showing the alveolar planum. The figure is
based on two superimposed CT cross sections: one at the level of the canine and the
other at the level of the mental foramen. The photos were inverted and rendered in
grey-scale in Photoshop CS2.
193
noted as a change in orientation between the subalveolar plane and
the submandibular fossa rather than a sharply delineated line. The
level at which it begins cannot be ascertained as the lower portion
of the endomandibular face is destroyed in that area. However, it
seems to extend towards the P3. Its ascent is not steep and it is still
present at the level of the mesial alveolar margin of the M3 beyond
which it can no longer be observed due to the breakage. The line
rises from 27.6 mm below the mesial edge of the M1 alveolus to
12.0 mm from the distal edge of the M2 alveolus.
Occlusal view (Fig. 5c)
The mandible is very thick in the buccolingual dimension. The
mesial thickening is endomandibular while the distal thickening is
exomandibular with respect to the dental arcade. The occlusal view
shows that the mandibular torus decreases in width from the M3 to
P3, while the shelf-like thickening of the alveolar plane increases in
width from the M1 towards the symphysis, and the ridge formed
descends towards the mid-symphyseal region. This latter structure is
responsible for the endomandibular thickening of the bone towards
the symphysis. The retromolar trigone could not be observed due to
breakage. The extramolar sulcus is very wide, accentuated by a low
and non-steep oblique line. The substantial width of the extramolar
sulcus is further accentuated by a pronounced curvature of the dental
arcade towards the sagittal plane.
The mandible measures 34.2 mm in height at the mandibular
foramen, recedes slightly towards the M3 where it measures
31.2 mm, and is most consistent with a “parallel” character state for
alveolar margin orientation. The width of the mandible varies from
19.1 mm at the canine alveolus, becoming more restricted towards
the mandibular foramen (17.8 mm) and M1 (17.5 mm) and
increasing towards the M2 (18.4 mm) and the M3 (23. 8 mm).
Dentition (Fig. 5d and Fig.7)
Only the three left molars are present in the BH-1 specimen. The
teeth are well preserved except for the mesiolingual root of the M3,
which shows recent breakage. The occlusal outline is sub-rectangular
and elongated mesiodistally. The teeth are not occlusally complex as
they show no extra fissures or crests. All three teeth have the five
main cusps (protoconid, metaconid, hypoconid, entoconid and
hypoconulid). M1 amd M2 have a “Y” groove pattern, while M3 shows
a “þ” pattern. The hypoconulid is large and buccally aligned. There is
an easily observed, wedge-shaped cusp 7 (tuberculum intermedium)
(Scott and Turner, 1997), separated by clear fissures in all three
molars. A protostylid is not present on any of the teeth. The mesial
marginal ridge shows a proper ridge in M1 with no anterior fovea.
This feature is continuous and depressed (very low) in M2 and
accompanied by an anterior fovea that is relatively poorly defined. It
is represented by a wide depression rather than a deep triangular
depression, as described by Scott and Turner (1997). The mesial
marginal ridge shows a tubercle on the M3 and a possible but not
clear anterior fovea (Hillson, 1996). A well developed anterior fovea is
common in Neandertals (87% according to Bailey, 2002) and variable
in modern humans with a 83% frequency in a sample of modern
Croatians (Gauta et al., 2010) and a frequency of 92% in a sample of
modern Javanese (Artaria, n.d.). The M2 and M3 present a distal
trigonid crest that can be assessed by a short transverse fissure,
slightly oblique to the buccolingual fissure. The M2 and M3 show the
same number of cusps as the M1, with less advanced degree of wear
on M2 and no wear on M3. None of the teeth show the continuous
midtrigonid crest e considered to be indicator of Neandertal affinity
as it occurs in 96% of Neandertals (Bailey, 2002). The presence of cusp
7 is non-diagnostic. While not unknown in Neandertals (18.8%), it is
much more common both in Homo erectus (40%) and is extremely
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194 M. Roksandic et al. / Journal of Human Evolution 61 (2011) 186e196

should be considered as a plesiomorphic trait in the Homo lineage

(Ackerman et al., 1973). Notable is the substantial distance between
root apices and the mandibular canal, which could be explained by
a reduction in tooth size coupled with the large size of the mandible.

Discussion

While the shape of the dental arcade and molar morphology

place BH-1 in the genus Homo, its fragmentary nature and plesio-
morphic character of its traits preclude a more precise taxonomic
designation for this specimen (Carcraft, 1989). We assign it at
present to Homo sp. The presence of the alveolar planum and the
overall robusticity of the specimen indicate a non-modern
morphology and we can narrow the designation to an archaic Homo
sp. We compared available measurements and character states of
the mandibular body with specimens from the Early, Middle and
Upper Pleistocene (Table 3) published in two recent articles that
both claim to be able to differentiate among species on the basis of
a set of metric (Lague et al., 2008) and morphological traits
(Mounier et al., 2009) of the mandible. While the comparative
samples in these two articles are not ideally suited to examine the
position of BH-1 on the hominin spectrum, they provide a good first
Figure 7. CT scan of the mandible in: (a) mesiodistal (med) cross-section, (b) bucco- view of the affinities of the specimen. Raw data on mandibular
lingual (bel) slice at the root of M1 showing substantial subcortical thickening and height, width and robusticity (Table 5) were compared with data
a mesotaurodontic M1, (c) horizontal slice showing four diverticles of the M1 root, (d)
from Carbonell et al. (2005). A principal components analysis of
Three superimposed CT sections showing the morphology of the P4 alveolar socket
with possible root bifurcation. Scale in centimetres. metric data from Lague et al. (2008) (measurements were kindly
provided by Nicole Collard) did not provide any further insight. The
scores for the BH-1 specimen show that the specimen lies at the
variable in modern human populations (3e61%) (Bailey, 2002), with very margin of the spread for Neandertals and anatomically
highest frequencies recorded in Africa (Scott and Turner, 1997). The modern humans (AMH), in the area where they overlap, and very
expression of the distal trigonid crest is highly variable (Scott and close to the H. erectus range. On the basis of this, the specimen
Turner, 1997) and according to Martinón-Torres et al. (2008) often cannot be excluded from any of the three groups.
underscored. It is, however, expressed in higher frequencies in When compared with the table provided by Carbonell et al.
Dmanisi and Sangiran populations (Martinón-Torres et al., 2008). (2005), the height of the mandible falls within the range of Nean-
While the M1 and M2 have the same buccolingual width (10.9 mm) dertals and H. heidelbergensis. The corpus thickness, however,
and mesiodistal length (11.5 mm), the M3 is longer mesiodistally exceeds the limits of the Neandertal range and falls towards the
(12.1 mm) and narrower buccolingually (10.5 mm). upper limit of the H. heidelbergensis values. Corpus thickness, rather
than corpus height seems to show a general trend of decrease with
Radiological features (Fig. 7) time in the human lineage. Mesiodistal and buccolingual diameters of
the teeth fall within the range of variation for European Neandertal,
The CT scan of the mandible reveals a very dense subcortical Krapina, and Atapuerca samples (Wolpoff, 1979; Bermúdez de Castro
bone of almost homogenous structure (1600-1800H, corresponding et al., 2001).
to massa compacta). The thickness varies in both buccolingual, and Cluster analysis (Fig. 8) shows clear Early, Middle and Late
mesiodistal directions; the area adjacent to the lingual cortical Pleistocene clusters. The cophenetic correlation coefficient
lamina is almost twice the width of the buccal one (Fig. 7b) and the obtained equals 0.78, which is very close to taxonomic validity,
thickness increases from the molars towards the symphyseal region usually considered as 0.8 (Rohlf and Fisher, 1968; Sneath and Sokal,
(Fig. 7c). Trabeculation of the spongious bone is visible only in the 1973). BH-1 clusters with the Early Pleistocene sample, including
central part of the mandibular body superior to the mandibular specimens from Sangiran (1b), Atapuerca (ATD6-96), Dmanisi
canal; the trabeculae are coarse with prominent marrow spaces. (D211) and Koobi Fora (KNM-ER 992), as well as a Middle Pleisto-
There is no visible thickening of the lamina dura, nor the walls of cene specimen from Tighenif (2) and slightly later in the sequence
the mandibular canal.
The CT scan of the P4 alveolus shows a clear separation towards Table 5
the distal portion of the root socket, however, only the apex of Comparison of mandibular thickness, height and robustness of BH-1 with different
the root is clearly bifurcated. This could indicate a possible (or species of Homo as reported by Carbonell et al. (2005: Table 2).
partial) root bifurcation, or alternatively, a groove on the distal side Species Thickness, mm (n) Height, mm (n) Robustness (n)
of the root (Fig. 7d). Since the root itself is missing, it is not possible Homo habilis 19.7  2.3 (5) 29.3  2.2 (4) 64.5  5.3 (4)
to ascertain the presence or absence of two diverticles in the P4. The H. ergaster 19.8  1.4 (8) 31.2  2.8 (8) 63.8  5.0 (8)
M2 and M3 are mesotaurodontic (Mena, 1971). The M1 has two H. erectus 19.8  2.6 (4) 36.3  1.1 (4) 54.7  7.4 (4)
mesial and two distal diverticles (Fig. 7c). All three aspects of the root H. mauritanicus 18.0  1.1 (4) 35.8  1.5 (4) 50.2  1.8 (4)
H. pekinensis 16.5  1.8 (7) 28.6  3.3 (7) 58.0  5.6 (7)
morphology are primitive. While taurodontism has often been cited
H. antecessor 16.4 (2) 27.6 (2) 62.6 (2)
as a Neandertal trait, its varying frequencies in many different H. heidelbergensis 16.4  1.7 (16) 30.8  3.4 (16) 53.4  5.1 (16)
modern human populations (Terezhalmy et al., 2001), as well as its H. neanderthalensis 15.3  1.7 (21) 32.1  3.3 (21) 47.9  5.1 (21)
appearance in the Atapuerca sample (Bermúdez de Castro et al., BH-1 17.5 (1) 33.1 (1) 52.8 (1)
1997, 1999) and Homo erectus (Conroy, 1997) indicate that it
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M. Roksandic et al. / Journal of Human Evolution 61 (2011) 186e196
Figure 8. Cluster analysis on mandibular body character states using Average Linkage, Euclidean Distance. UU: unknown age, EP: Early Pleistocene, MP Middle Pleistocene, NT:
Neandertals. The coefficient of cophenetic correlation ¼ 0.78.
with Sinanthropus H1 and Neandertal Bañolas specimen. This result
is consistent with the absence of observable derived Neandertal
traits in BH-1 and the presence of the alveolar planum, four diver-
ticles in M1 and a possible bifurcation of the P4, all three of which
are primitive traits inconsistent with Neandertals and AMH speci-
mens. When considering raw data, BH-1 shares 13/16 traits with
Sangiran (1b) and Atapuerca (ATD6-96), 12/16 with Tighenif (2),
and 10/16 with Dmanisi (211) and Koobi Fora KNM-ER 992. While
the Bañolas mandible clusters relatively closely, it shares only four
character states with the BH-1 mandible.
Comparative analysis of the specimen indicates a number of
primitive traits: the alveolar planum, possible bifurcation of the P4,
four diverticles in M1 and an anteriorly positioned single mandibular
foramen that may signal the absence of midfacial prognathism. Given
the associated Upper Pleistocene date, it is possible that this specimen
is a primitive Neandertal. However, the complete lack of derived
Neandertal traits and clustering of BH-1 with the Early Pleistocene
specimens requires us to examine other plausible explanations.
While the paucity of remains from the area precludes any further
discussion of the meaning of this partial specimen, the particularities
of the Balkan Peninsula as a refugium for temperate deciduous forests
and associated biota (Eastwood, 2004; Tzedakis, 2004) and a “hotspot
of biodiversity” (Lesbarreres, 2008) warrant further research. Of the
three southern European peninsulas that were the source of most
repopulation of Europe for both animals and plants, the Balkan
195
Peninsula emerges as the source of all of the species of the East and
many of the West of northern Europe (Hewitt, 2000, 2004; Tzedakis,
2004). It shows substantial faunal heterogeneity (Griffiths et al.,
2004) and high levels of endemism (Zima, 2004). While the role that
refugial areas of the Iberian and Apennine Peninsulas played in human
evolution during glaciations in the Northern hemisphere has been well
established in recent research (Finlayson, 2004; Finlayson et al., 2008),
the importance of Southeastern Europe in providing glacial refugium
and maintaining gene flow between continents, while proposed for
a number of animal species (Griffiths et al., 2004), is rarely discussed
with regards to human evolution. High levels of biodiversity in the
Balkans (Hewitt, 2004) and the refugial character of its flora
(Eastwood, 2004; Tzedakis, 2004), coupled with a lack of isolating
mechanisms during Pleistocene glaciations, could have played an
important role in preserving variability, maintaining relic species, and
allowing gene flow throughout the Pleistocene for all biota, including
hominins. This lack of isolating mechanisms in the Balkans could have
resulted in the preservation of the primitive morphology seen in BH-1,
or the group of archaic humans it belonged to.
Conclusion
The recent discovery of a mandible BH-1, from Mala Balanica
(Serbia), with primitive character states comparable with the
Early Pleistocene mandibular specimens, is associated with
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196 M. Roksandic et al. / Journal of Human Evolution 61 (2011) 186e196
a minimum date of 113 þ 72  43 ka. The BH-1 specimen, despite
geographic proximity and possible contemporaneity with the
Krapina Neandertals, does not share any observable derived
Neandertal traits. BH-1 clearly represents an individual from an
archaic Homo sp., possibly non-Neandertal population, with
a suite of features falling at the primitive end of the range of
variation for the genus Homo.
Acknowledgements
The research was supported by a grant from the offices of the VP
Academic and VP Research at the University of Winnipeg, an NSERC
grant (371077-2010) to MR, and a grant from the Ministry of
Culture and the Ministry of Science and Technological Develop-
ment of the Republic of Serbia (177023) to DM. MR wishes to thank
Mary Silcox, Fred Smith and Katerina Harvati for comments on the
first draft, Nicole Collard for mandibular measurements, William
Sanders and Zoran Pavlovi
c for help with conservation, Rob Hoppa
and his team for a 3-D printout of the specimen and anonymous
reviewers for their valuable comments.
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