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611
The Human Cranium from the Pes¸tera
Cioclovina Uscata˘, Romania
Context, Age, Taphonomy, Morphology, and
Paleopathology
Andrei Soficaru,Catalin Petrea,Adrian Dobos¸, and
Erik Trinkaus
Institutul de Antropologie “Fr. J. Rainel,” Eroii Sanitari 8,
P.O. Box 35-13, 76241 Bucures¸ti, Romania / Institutul de
Speologie “Emil Racovit¸a,” str. Calea 13 Septembrie nr. 13,
50711 Bucures¸ti, Romania / Institutul de Arheologie “Vasile
Paˆrvan,” Henri Coanda˘ 8, Sector 1, 010667 Bucures¸ti, Ro-
mania / Department of Anthropology, Campus Box 1114,
Washington University, St. Louis, MO 63130, U.S.A.
(trinkaus@artsci.wustl.edu). 15 I 07
Reanalysis and direct dating of an early modern human neu-
rocranium from the Pes¸tera Cioclovina Uscata˘ (Cioclovina
1), in combination with excavation and reanalysis of the re-
maining deposits in the cave, establish Cioclovina 1 as one
of a small number of European early modern humanssecurely
dated prior to ca. 28,000
14
C BP (ca. 32,500 cal BP). The
original stratigraphic context and archeological association of
Cioclovina 1 are unknown (and probably unknowable), but
sedimentological analysis and dating of cave bear remains
suggests substantial Late Pleistocene geological reworking of
deposits within the cave, which probably altered its context
prior to the mining operations which unearthed the neuro-
cranium. The otherwise excellent (if incomplete) preservation
of Cioclovina 1 raises questions as to what, if any, human
behaviors resulted in its burial within the Pes¸tera Cioclovina
Uscata˘. Lesions are limited to the minor exocranial traumatic
changes common among Late Pleistocene humans. Morpho-
logically, Cioclovina 1 presents a suite of distinctive, derived
modern human neurocranial features, associated with aspects
of the superior nuchal morphology best known for European
Neandertals, a mosaic pattern increasingly in evidence among
early modern humans in Europe.
Our understanding of the spread of early modern humans
into Europe during oxygen isotope stage (OIS) 3 has been
revised in the past decade as the application of direct radi-
ocarbon dating to human remains purportedly from the crit-
ical time period has removed several samples from the Pleis-
tocene (Hahno¨fersand, Velika Pec´ ina, Vogelherd), assigned
other remains to later phases of the Late Pleistocene (e.g.,
Cro-Magnon, La Rochette, Zlaty´ku˚n), and provided secure
䉷2007 by The Wenner-Gren Foundation for Anthropological Research.
All rights reserved. 0011-3204/2007/4804-0008$10.00
early ages for additional specimens (e.g., Mladecˇ, Pes¸tera cu
Oase, Pes¸tera Muierii) (Trinkaus et al. 2003; Wild et al. 2005;
Trinkaus 2005; Soficaru, Dobos¸, and Trinkaus 2006). One of
the important specimens is a largely complete human neu-
rocranium from the Pes¸tera Cioclovina Uscata˘ (Cioclovina
Dry Cave), Bos¸orod, Hunedoara County, Romania (45⬚35⬘
N, 23⬚07⬘E). Traditionally assigned to the Aurignacian (Rai-
ner and Simionescu 1942; Necrasov and Cristescu 1965), the
Cioclovina human neurocranium (fig. 1) has variably been
included in considerations of early modern humans in
Europe.
Context
The Pes¸tera Cioclovina Uscata˘ is a 2,002-m-long fossil karstic
cave gallery (fig. 1). It is the superior part of a 7.8-km-long
karstic system, the Ponorici Cioclovina, and served during
the Pleistocene principally as a hibernation cave for Ursus
spelaeus. It has been known and variably described since the
1880s, and karstic geological and vertebrate paleontological
work has been conducted intermittently (Breban et al. 2003;
see also Caˆrciumaru 1999; Pa˘unescu 2001). Sometime before
1911 a human cranium was purported to have been found
within the cave associated with U. spelaeus crania; the human
specimen was never located, but the report prompted M.
Roska to undertake archeological excavations in the cave in
1911 and 1921 (Roska 1923, 1930a). Roska and subsequent
excavators of the deposits unearthed samples of techno-
typologically Mousterian and Aurignacian lithics. However,
the excavations consisted of a series of widely dispersed
soundings, the stratigraphic relationships of the lithic remains
are uncertain, and most of the pieces were assigned to one
of these technotypological complexes on the basis of individ-
ual piece attributes rather than assemblage composition (Pa˘u-
nescu 2001). The principal remains within the cave are those
of U. spelaeus, with rare remains of U. arctos,Canis lupus,
Panthera spelaea, and Capra ibex (Ra˘dulescu and Samson
1959; Pa˘unescu 2001). It therefore has the faunal profile of
a cave bear hibernation site that was occasionally used as a
den by brown bear and wolf. As with a number of cave sites
in the Carpathians (Pa˘unescu 1980, 1989, 2001; Caˆrciumaru
and Dobrescu 1997), the presence of human activity at the
Pes¸tera Cioclovina was probably minor.
In 1940–41 the cave was exploited for phosphates by the
Societatea Nitrogen, and in the process, as reported by Sim-
ionescu (1942), a partial human cranium was recognized by
workmen and placed with guano, three lithic artifacts, and
animal bones in a mining cart, where the team’s engineer
identified it as of interest. It was subsequently claimed to have
come from a sealed layer of Sector 1 of the exploitation, which
also yielded the three artifacts along with crania and vertebrae
of U. spelaeus (Rainer and Simionescu 1942). There is no
documentation of the original positions of these remains, and
there is no factual basis for using the artifacts or any specific
612 Current Anthropology Volume 48, Number 4, August 2007
Figure 1. Plan of the primary paleontological and archeological portions
of the Pes¸tera Cioclovina Uscata˘, showing possible but unconfirmed lo-
cation of the original position of the Cioclovina 1 neurocranium, loca-
tions of the excavations by Roska in 1911 and 1921, and locations of the
recent excavation by C. Petrea (1) and the source of the Ursus spelaeus
radiocarbon samples (2). The gray area indicates the presence of U.
spelaeus remains. Plan modified from one by C. S. Protheus Hunedoara.
U. spelaeus remains to date or provide a technotypological
context for the cranium.
An association with U. spelaeus provides little chronological
control other than a Late Pleistocene age. The three artifacts
(a blade with denticulate retouch, a blade with fine retouch,
and a subtriangular flake) are commonly referred to the Au-
rignacian but are undiagnostic and too few to permit a tech-
notypological attribution. There are a number of other arti-
facts from the cave, and similarly their exact proveniences are
not known, but some 20 have been consistently attributed to
the Aurignacian (Roska 1923, 1930b; Breuil 1925; Pa˘unescu
2001; Dobrescu 2004). It is also unclear from which of the
chambers of the cave the human fossil derived. One can only
infer that it must have been found in an area where there
was a significant guano layer. Roska’s (1923) description of
the stratigraphic sequence in Sala Mare contains no mention
of such a deposit, so presumably the cranium came from a
different portion of the cave. For these reasons, even though
it has traditionally been attributed to the early Upper Paleo-
lithic (Rainer and Simionescu 1942; Necrasov 1971), others
(e.g., Churchill and Smith 2000) have pointed to the need for
a secure archeological context and/or a direct radiometric age
prior to the integration of the specimen into the European
early modern human sample.
Age
In an effort to assess the ages of the cave’s deposits and the
important remains, geological work was undertaken by one
of us (CP) in 2004 on one of its few remaining intact sedi-
mentological profiles. This profile is a 4-m-high alternation
of siltic-clay beds (matrix support) with ruditic beds (grain
support, intraspeleic limestone clasts), indicating alternating
low- and high-energy hydraulic transport of sediments in the
cave as a result of climate change. Comparison of preliminary
magnetic susceptibility data with the reference section devel-
oped for cave sediments from southern Europe (Ellwood et
al. 1998, 2001) suggests that its upper portion contains sed-
iments deposited during the last glacial maximum (OIS 2).
A fibula of U. spelaeus from the upper part of the profile
(50–60 cm below datum) was radiocarbon-dated to ca.
31,500–32,000
14
C BP (table 1). Its d
13
C and C : N values are
well within acceptable ranges (see below), and its percent
collagen yield is relatively high for Late Pleistocene bone,
providing confidence in its age. A second U. spelaeus specimen
from 65–80 cm below datum yielded a date of ca.
40,000–50,000
14
C BP; its collagen yield is lower, but both the
yield and the associated d
13
C and C : N values are within
acceptable ranges. The substantial difference between the
613
Table 1. Radiocarbon Dating Results for Cioclovina 1 and Ursus spelaeus from 2004 Excavations at the
Pes¸tera Cioclovina Uscata˘
Specimen
Age
(
14
C yrs. BP)
Age
(cal yrs. BP)
a
d
13
CC:N
Sample
Weight
(mg)
Collagen
Weight
b
Burn
weight
c
(mg) %C
d
%Nmg %
Cioclovina 1 temporal (LuA-5229) 29,000 Ⳳ700 33,726 Ⳳ1,132 – – 536 – – – – –
Cioclovina 1 occipital (OxA-15527) 28,510 Ⳳ170 33,212 Ⳳ693 ⫺19.6 3.4 680 40.4 5.9 6.3 44.4 15.9
Ursus spelaeus fibula (OxA-15528) 31,700 Ⳳ190 36,667 Ⳳ261 ⫺20.4 3.3 1,400 173.0 12.4 6.0 43.6 15.0
U. spelaeus metapodial (OxA-16328) 40,550 Ⳳ600 44,237 Ⳳ710 ⫺20.3 3.2 680 25.6 3.8 5.3 42.4 15.3
Sources: Olariu et al. (2005), A. Olariu (personal communication), and this study.
a
Based on CalPal (http://www.calpal.de) version 1.4 (May 2006).
b
Ultrafiltered gelatin yield.
c
Gelatin combusted for graphitization.
d
On combustion.
probable OIS 2 age of the deposits and the ages of the bear
bones indicates transport of the bones after the animals’
deaths. Consequently, even if we could identify the original
location and stratigraphic context of the human remains, it
is likely that they were in a secondary position within the
cave.
Two separate efforts have been made to date the Cioclovina
1 specimen directly. In 2001 a sample of the external right
mastoid process was taken by A. Olariu and E. Alexandrescu
and provided a date of
14
C BP (LuA-5229) (Pa˘-29,000 Ⳳ700
nescu 2005; Olariu et al. 2005) (table 1). This date was suf-
ficient to confirm the Late Pleistocene age of Cioclovina 1,
but the absence of associated chemistry and the large error
value limited its utility. It was therefore decided to redate the
specimen using ultrafiltration pretreatment (Higham et al.
2006; Higham, Jacobi, and Bronk Ramsey 2006) rather than
the standard HCI extraction. A small piece of the posterior
left foramen magnum border (lateral of opisthion and du-
plicating the preserved right side) that had become detached
in storage was taken with the permission of T. Neagu and
submitted to the Oxford Radiocarbon Accelerator Unit. The
median value of the OxA-15527 date of
14
CBP28,510 Ⳳ170
is slightly more recent than LuA-5229, but, given the large
standard deviation on LuA-5229, the two dates are statistically
in agreement (table 1). Moreover, the d
13
C value for OxA-
15527 falls in the middle of the acceptable range for bone,
and its C : N ratio is within the acceptable range of 2.9 to 3.5
for well-preserved bone (Bronk Ramsey et al. 2004). The per-
cent collagen weight is moderate for bone, but the collagen
and nitrogen burnweights are acceptable.
The Cioclovina 1 neurocranium is therefore firmly dated
to OIS 3. As such, it joins the Pes¸tera cu Oase (Trinkaus et
al. 2003, 2005a, 2006; Rougier et al. 2007), Pes¸tera Muierii
(Soficaru, Dobos¸, and Trinkaus 2006), and Mladecˇ (Wild et
al. 2005) human remains as one of a small number of di-
agnostic and directly dated early modern human samples
from Europe prior to ca. 28,000
14
C BP. Although it is similar
in age to the later stages of the Aurignacian in southeastern
Europe (Chirica, Borziac, and Chetraru 1996; Dobrescu
2004), it may also overlap the beginning of the Gravettian in
the same region (Chirica 1989; Caˆrciumaru 1999).
Taphonomy
It is not clear how the Cioclovina 1 neurocranium came to
be contained within the principally ursid context of the Pes¸-
tera Cioclovina Uscata˘ . Considering its discovery during min-
ing operations purportedly from a sealed level with cave-bear
remains, it is unusual in its preservation (figs. 2 and 3). The
external vault shows little or no abrasion, and there is no
carnivore damage to the bone. The postmortem damage to
the cranium consists of loss of the facial skeleton and zygo-
matic arches, erosion of the tips of the mastoid processes,
and loss of portions of the basicranium. With respect to the
last, the posterior orbital roofs are absent, the medial petrous
processes are incomplete, and the basioccipital is lacking. Yet,
greater wings of the sphenoid bones are present, and the
sphenoid body is largely present with a portion of the vomer.
The fragile superior ethmoid bone with the cribiform plate
remains, along with most of the posterior margin of the fo-
ramen magnum. Moreover, both frontozygomatic sutures are
preserved without abrasion, and the superior 10 mm of the
nasal bones remain along the intact nasofrontal and fronto-
maxillary sutures.
There is a series of cracks through the middle of the right
parietal bone, centered around a depressed section of bone
close to the sagittal suture and radiating onto adjacent bones.
This damage has been interpreted (Ullrich 1979, 2005) as the
result of perimortem violence. However, there is no mention
of the damage in the original publication of Rainer and Sim-
ionescu (1942), even though they mentioned the other exo-
cranial lesions (see below), and Rainer was an experienced
anatomist, pathologist, and skeletal biologist. Moreover, the
original photographs show no evidence of the damage.
It remains uncertain how complete the remains of Cio-
clovina 1 were when they were deposited in the cave, whether
614 Current Anthropology Volume 48, Number 4, August 2007
Figure 2. The Cioclovina 1 neurocranium in norma lateralis left. Scale in
centimeters.
the individual succumbed within or near the cave, and
whether it was initially in the superficial deposits of the cave
or buried. Interestingly, although intentional burials are now
unknown from the European Aurignacian (Henry-Gambier
2002), there are three karstic caves in Europe (Mladecˇ, Pes¸tera
cu Oase, and Pes¸tera Muierii) in which directly dated human
remains from this time period have been found on or near
the surface (Gheorghiu and Haas 1954; Svoboda 2000; Zilha˜o
et al. n.d.). As is suggested by Svoboda (2000) for Mladecˇ, it
is possible that these represent intentional disposals of human
remains, subsequently displaced within the caves by geological
processes. Could the pattern of preservation of the Cioclovina
1 neurocranium be the result of such a disposal of remains?
Unfortunately, the circumstances of discovery of the Mladecˇ,
Muierii, and Cioclovina remains, the clear intrakarstic hy-
draulic transport of the Oase remains, and the probable hy-
draulic displacement of Cioclovina 1 make it impossible to
test this hypothesis.
Morphology
The Cioclovina 1 neurocranium has been described from the
beginning as that of an early modern human, with an age-
at-death probably in the fourth decade based on extensive
fusion but not obliteration of the cranial vault sutures. Rainer
and Simionescu (1942) suggested that it might be female, an
attribution often repeated (Necrasov and Cristescu 1965; Ni-
cola˘escu-Plops¸or 1968) but also questioned (Smith 1984;
Churchill and Smith 2000). The prominent superciliary arches
and large (if damaged) mastoid processes of Cioclovina 1,
especially with respect to presumed females such as Mladecˇ
1 and 2 and Muierii 1 (Wolpoff, Frayer, and Jelı´nek 2006;
Soficaru, Dobos¸, and Trinkaus 2006), suggest that a diagnosis
as female may not be accurate, although the specimen is less
robust than presumed males such as Mladecˇ 5 and 6 (Frayer
et al. 2006). Similarly, given the large estimation errors for
age assessment from sutural fusion (Meindl and Lovejoy
1985), its age-at-death is probably best considered as mature
but not geriatric.
Osteometrics and some morphological details have been
provided by Rainer and Simionescu (1942) and further data
will be published elsewhere. However, a few salient morpho-
logical characteristics can be mentioned here.
The supraorbital region (figs. 3 and 4) presents prominent
superciliary arches which extend laterally to slightly lateral of
the supraorbital foramina, with a slight depression above gla-
bella. There is no evidence of a sulcus separating them from
the lateral trigone, but each side has a distinct concavity above
the lateral orbital margin between the orbital margin, the
temporal crest, and the lateral superciliary arch. Nasion is
anteriorly prominent relative to the frontozygomatic sutures.
615
Figure 3. The Cioclovina 1 neurocranium in approximate norma frontalis
(left) and norma occipitalis (right). Scale in centimeters.
Figure 4. Supraorbital (above) and occipital (below) details of the Cio-
clovina 1 neurocranium.
616 Current Anthropology Volume 48, Number 4, August 2007
Cioclovina 1 therefore lacks a full supraorbital torus and has
the pattern of superciliary arch development seen in Mladecˇ
5 and male Gravettian specimens (e.g., Cro-Magnon 3, Dolnı´
Veˇstonice 16, Pavlov 1, Prˇedmostı´ 3, Sunghir 1). Although
approached by Mladecˇ 1 in supraorbital development, it has
greater superciliary arch and glabellar projection than Mladecˇ
2, Muierii 1, and the (albeit adolescent) Oase 2 cranium.
The neurocranial vault is relatively high and rounded, with
strongly and evenly curved frontal, parietal, and occipital sag-
ittal contours (fig. 2). There is a modest change of curvature
near lambda with an associated posterior protrusion of the
superior occipital squamous; this is the configuration referred
to by Smith (1984) as a hemi-bun. However, it is smaller than
the full occipital buns (indicated by a strong change in cur-
vature near lambda and/or a distinct protrusion of the squa-
mous occipital) evident in Mladecˇ 3 and 5 and Muierii 1
(Frayer et al. 2006; Minugh-Purvis, Viola, and Teschler-Nicola
2006; Soficaru, Dobos¸, and Trinkaus 2006), as well as Cro-
Magnon 3 (Broca 1868). The nuchal plane has semispinalis
capitis fossae which meet at the midline, with a clear midline
crest along most of the nuchal plane. The juxtamastoid em-
inences are on the occipital bone and project little beyond
the surface of the nuchal plane.
The temporal bones (fig. 2) resemble those of recenthuman
males, with laterally and (originally) inferiorly prominent
mastoid processes, high and rounded squamous sutures with
posteroinferiorly sloping parietomastoid sutures, a vertically
oriented auditory porous, and the zygomatic root above the
porous. The laterally rugose mastoid processes lack anterior
tubercles and have straight anterior and posterior margins.
There are prominent supramastoid crests, and the tempo-
rozygomatic sulcus slopes markedly anteroinferiorly. The
postglenoid processes are barely perceptible, and the glenoid
fossae are deep, with vertical anterior surfaces.
These features combine to give Cioclovina 1 a coronal out-
line in norma occipitalis (fig. 3) that is largely rounded across
the sagittal suture, has rounded angles at the prominent pa-
rietal bosses, continues vertically with minor irregularities
across the squamous sutures to the supramastoid regions, and
ends with the prominent mastoid processes. This coronal pro-
file contrasts with both the ovoid forme en bombe of most
Neandertals and the more pentagonal forme en maison of
many modern humans (see Hublin 1983); it is also found on
Muierii 1, Oase 2, and possibly Mladecˇ 2, whereas at least
Mladecˇ 5 has the forme en maison pattern.
In the context of this morphologically modern human neu-
rocramial pattern, the region of the superior nuchal line pres-
ents an unusual pattern for a modern human (figs. 3 and 4).
There is no external occipital protuberance, eventhough there
is a small, nonprojecting triangular area below the nuchal
lines resulting from the confluence of the superior nuchal
lines and the midline crest between the semispinalis capitis
insertions. There is a modest but clear rounded nuchal torus
that extends primarily across the middle half of the occipital
bone, fading out laterally near the lateral extents of the semi-
spinalis capitis fossae. The nuchal line is moderately promi-
nent across the full occipital bone from asterion to asterion,
and it lies to a large extent on the nuchal torus instead of
inferior to it. Above the midline nuchal torus there is a shallow
but distinct, irregularly rugose, transversely oval depression
ca. 34 mm wide and ca. 14.5 mm high—a suprainiac fossa
morphologically similar to and contrasting only in depth with
those found universally on Neandertal occipital bones. It is
distinct from the supranuchal fossa (Sla´dek 2000) found on
Mladecˇ 5 and a number of Gravettian specimens (Frayer et
al. 2006), which is a triangular depression bounded inferiorly
by elevated nuchal lines curving inferiorly toward a prominent
external occipital protuberance. It also contrasts with the
smoother iniac regions of Mladecˇ 1 and 3, Muierii 1, and
Oase 2 (Wolpoff, Frayer, and Jelı´nek 2006; Minugh-Purvis,
Viola, and Teschler-Nicola 2006; Rougier et al. 2007).
The combination of three of these features—absence of a
distinct occipital protuberance, a modest nuchal torus limited
to the medial half of the occipital bone, and a transversely
oval suprainiac fossa—is principally found on Neandertal cra-
nia, absent from Middle Paleolithic modern humans, and
rarely found on other contemporaneous or older human cra-
nia (Hublin 1978; Santa Luca 1978; Trinkaus 2004). Cio-
clovina 1 contrasts with Neandertals in this anatomical region
principally in that the superior nuchal line is relatively prom-
inent and traverses the nuchal torus. However,if one separates
out the two complexes (suprainiac fossa, nuchal torus, and
external occipital protuberance, on the one hand, and the size
and position of the superior nachal lines, on the other), it is
apparent that Cioclovina 1 shares the former with the Nean-
dertals and the latter with early modern humans. Distin-
guishing the Cioclovia 1 suprainiac fossa complex from the
Neandertal range of variation would be possible only if one
applied different criteria to Neandertal versus modern human
crania.
Paleopathology
The Cioclovina 1 cranium provides evidence of antemortem
trauma in the form of two shallow depressions on the external
frontal squamous portion. One of them is ca. 12 mm in
diameter centered 52 mm above nasion just to the right of
the midline, and the other is ca. 9 mm in diameter 52 mm
above right superior orbital margin and 7 mm medial of the
temporal line; both were noted by Rainer and Simionescu
(1942). They are limited to the external table, healed long
before death, and similar to lesions common on Late Pleis-
tocene human crania (Berger and Trinkaus 1995; Trinkaus et
al. 2005b; Frayer et al. 2006). It is unclear whether these minor
injuries were the result of interpersonal violence (see Walker
1997), hunting (see Berger and Trinkaus 1995), or incidental
accidents. Independent of their origin, they would not have
affected the individual once healed.
Discussion
Given the fortuitous discovery of the human cranium, the
617
nature of previous archeological and paleontological work in
the cave, and the dearth of undisturbed sediments remaining,
we will probably never know how the human remains came
to be preserved in the cave. However, several aspects have
been clarified by recent work.
The human cranium is securely dated to between 28,000
and 29,000
14
C BP. This brackets it between the Mladecˇ and
Muierii remains and the Cro-Magnon sample. It is possible
the Cioclovina 1 postdates the primary hibernation period
for the bears, but it remains untestable whether human or
geological processes were responsible for the introduction of
the human remains into the karstic system.
The specimen is of an early modern human adult of un-
certain sex. The overall morphological pattern of the specimen
is similar to that of earlier Upper Paleolithic modern humans
from Europe, but its superior nuchal region has distinctly
Neandertal-like aspects. This combination of principally
modern human features with archaic morphological aspects
is becoming increasingly evident among early modern hu-
mans across Eurasia. The human remains from Pes¸tera cu
Oase (Trinkaus et al. 2003; Rougier et al. 2007), Pes¸tera Mui-
erii (Soficaru, Dobos¸, and Trinkaus 2006), Mladecˇ (Frayer et
al. 2006; Wolpoff, Frayer, and Jelı´nek 2006; Minugh-Purvis,
Viola, and Teschler-Nicola 2006), and Les Rois (Vallois 1958)
present a variable morphological mosaic that is difficult to
explain as a simple OIS 3 expansion of modern human pop-
ulations derived solely from those of Middle Paleolithic East
Africa and extreme southwestern Asia. Although overwhelm-
ingly presenting derived modern human features (Trinkaus
2006), these remains (now joined by Cioclovina 1) present a
variable sprinkling of traits that are either distinctly Nean-
dertal-like, generally archaic Homo features that had been lost
in the Middle Paleolithic modern human sample, or unusual
in either potentially ancestral sample (potentially descendant
groups and more recent Paleolithic specimens are irrelevant).
This mosaic is most parsimoniously explained as the result
of a modest level of admixture with Neandertals as modern
humans dispersed across Europe. Given the reproductive
compatibility of many closely related species (Jolly 2001; Hol-
liday 2006) and the culturally mediated nature of mate choice
in humans, such admixture should be neither rare nor un-
expected. Moreover, current analyses of recent (Templeton
2005; Evans et al. 2006; Garrigan and Hammer 2006) and
ancient (Serre et al. 2004; Zilha˜o 2006; Noonan et al. 2006)
human DNA support and/or permit such a level of admixture
in the Late Pleistocene. The morphological patterning of these
European early modern human fossils is beginning to doc-
ument such populational interactions.
Acknowledgments
The 2004 excavation in the Pes¸tera Cioclovina was undertaken
by CP through the auspices of the Institutul de Speologie
“Emil Racovit¸a,” Academia Romaˆna˘. The reanalysis and direct
dating of Cioclovina 1 was made possible by T. Neagu and
funded by the Wenner-Gren Foundation (grant 7290) and the
National Science Foundation (BCS-0509072). A. Olariu pro-
vided the available limited data on the original dating of the
Cioclovina 1 specimen. To all of them we are grateful.
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