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Middle Pleistocene Homo in the Levant Our understanding of the origin, distribution, and evolution of early humans and their close relatives has been greatly refined by recent new information. Adding to this trend, Hershkovitz et al. have uncovered evidence of a previously unknown archaic Homo population, the “Nesher Ramla Homo ” (see the Perspective by Mirazon Lahr). The authors present comprehensive qualitative and quantitative analyses of fossilized remains from a site in Israel dated to 140,000 to 120,000 years ago indicating the presence of a previously unrecognized group of hominins representing the last surviving populations of Middle Pleistocene Homo in Europe, southwest Asia, and Africa. In a companion paper, Zaidner et al. present the radiometric ages, stone tool assemblages, faunal assemblages, and other behavioral and environmental data associated with these fossils. This evidence shows that these hominins had fully mastered technology that until only recently was linked to either Homo sapiens or Neanderthals. Nesher Ramla Homo was an efficient hunter of large and small game, used wood for fuel, cooked or roasted meat, and maintained fires. These findings provide archaeological support for cultural interactions between different human lineages during the Middle Paleolithic, suggesting that admixture between Middle Pleistocene Homo and H. sapiens had already occurred by this time. Science , abh3169 and abh3020, this issue p. 1424 and p. 1429 ; see also abj3077, p. 1395
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PALEOANTHROPOLOGY
A Middle Pleistocene Homo from Nesher Ramla, Israel
Israel Hershkovitz
1,2
*, Hila May
1,2
*, Rachel Sarig
2,3
*, Ariel Pokhojaev
1,2,3
, Dominique Grimaud-Hervé
4
,
Emiliano Bruner
5
, Cinzia Fornai
6,7
, Rolf Quam
8,9,10
, Juan Luis Arsuaga
9,11
, Viktoria A. Krenn
6,7
,
Maria Martinón-Torres
5,12
, José María Bermúdez de Castro
5,12
, Laura Martín-Francés
5,12
,
Viviane Slon
1,2,13
, Lou Albessard-Ball
4,14
, Amélie Vialet
4
, Tim Schüler
15
, Giorgio Manzi
16
,
Antonio Profico
14,16
, Fabio Di Vincenzo
16
, Gerhard W. Weber
7,17
, Yossi Zaidner
18,19
It has long been believed that Neanderthals originated and flourished on the European continent.
However, recent morphological and genetic studies have suggested that they may have received a
genetic contribution from a yet unknown non-European group. Here we report on the recent discovery of
archaic Homo fossils from the site of Nesher Ramla, Israel, which we dated to 140,000 to 120,000 years
ago. Comprehensive qualitative and quantitative analyses of the parietal bones, mandible, and lower
second molar revealed that this Homo group presents a distinctive combination of Neanderthal
and archaic features. We suggest that these specimens represent the late survivors of a Levantine
Middle Pleistocene paleodeme that was most likely involved in the evolution of the Middle Pleistocene
Homo in Europe and East Asia.
Recent dental (1), mandibular (2), genetic
(3,4), and demographic (5) studies have
predicted the existence of an as yet un-
identified African or West Asian Middle
Pleistocene (MP) population that con-
tributed to the evolution of the Neanderthal
clade. This contrasts with the traditional view
that considered the European continent as the
soleplaceoforiginoftheNeanderthalsand
their direct ancestors.
Here we report on the discovery of several
fossils from the recently excavated MP open-air
site of Nesher Ramla (NR), central Israel (Fig. 1),
in association with stone artifacts, and faunal
remains (6).
A nearly complete right parietal bone and
four fragments from the left parietal bone rep-
resent the NR-1 fossil (Fig. 2A and fig. S1). The
NR-2 fossil is an almost complete mandible,
missing only the left ramus, the right condylar
process, and the mandibular angle of the right
ramus (Fig. 3). The lower left second molar
(NR-2 M
2
) and most of the dental roots are still
in place (fig. S2). Both NR-1 and NR-2 were
foundinsituwithinthelowestarchaeological
layer(Fig.1D,UnitVI),togetherwithanimal
bones and flint tools, and most likely represent
thesameindividual(supplementarytextA).
Unit VI is assigned an age of 140 to 120 thou-
sand years (ka) ago, based on the electron
spin resonanceuranium series (ESR-US) dates
of animal teeth recovered in this unit (with a
weighted mean of 125.8 ± 5.9 ka). This age
was corroborated by a series of thermolumi-
nescence (TL) dates of burnt flints from the
archaeological layer immediately above the
fossil (Unit V). This layer yielded a weighted
mean of 127.6 ± 4.0 ka (confirmed by isochron
analysis), which is in agreement with the ESR-
US dates obtained for this unit, ranging be-
tween ~128 and ~120 ka (a weighted mean of
122.3 ± 3.3 ka). This chronological information
is consistent with the previously published
optically stimulated luminescence (OSL) dates
for the entire archaeological sequence [rang-
ing ~170 and ~78 ka; (6)].
The preserved anatomical elements were
thoroughly described and analyzed in compa-
rison to a large number of fossils of different
periods (table S1), using a combination of tradi-
tional approaches based on linear and angular
measurements, as well as three-dimensional
(3D) landmark-based geometric morphomet-
ric (GM) methods (supplementary text C to E).
The overall morphology of the NR-1 parietal
bones (supplementary text C and tables S2
and S3) is indicative of an archaic, low cranial
vault, which is typical of MP Homo specimens
and is substantially different from early and
recent H. sapiens,whichinsteadmanifesta
curved parietal bone with a pronounced emi-
nence (7,8).
Further support for the rather archaic mor-
phology of the NR Homo comes from the angle
formed by the coronal and sagittal sutures (c/s
angle), 91.1° in the NR-1 specimen. This angle
increased during the evolution of Pleistocene
Homo (fig. S3): H. erectus and African MP Homo
exhibit a mean angle of 92.1° ± 2.1°; the angle
opens to 94.9° ± 3.4° in European MP Homo/
Neanderthals and reaches 99.4° ± 4.2° in early
and recent H. sapiens.Thec/sangleissignif-
icantly different between these three groups
(H=22.5,p<0.001).Thec/sangleforNR-1is
similar to that of archaic Homo,particularly
African MP Homo (91.1° ± 1.1°), and falls out-
side the range of variation of H. sapiens.
The NR-1 parietal bone is considerably thick,
mainly in the parietal eminence area (figs. S4
and S5). Regarding this aspect, the NR-1 parietal
is similar to that of European MP Homo spec-
imens (e.g., Petralona, Atapuerca SH) (fig. S4).
It is generally thicker than the parietal of
Neanderthals (e.g., Amud 1, Guattari, and La
Chapelle-aux-Saints) and most early H. sapiens
(except for Laetoli H18 and Omo 2), and it is
much thicker than that of recent H. sapiens.
The 3D GM analysis, used to assess NR-1
shape variation with respect to a comparative
sample of Pleistocene and recent Homo (sup-
plementary text, Fig. 2C, fig. S1, and table S1),
confirms the archaic morphology of NR-1. The
first three principal components (PCs) explain
74.5% of the total shape variance. The first
PC (34.9%) differentiates early and recent
H. sapiens from all other groups, including
Asian H. erectus, European and African MP
Homo, and Neanderthals, owing to their marked
curvature along both the sagittal and the cor-
onal planes (Fig. 2C). The second PC (21.3%)
is not taxonomically informative (fig. S1). The
third PC (18.3%) separates Asian H. erectus
and African MP Homo from Neanderthals and
European MP Homo (Fig. 2C), based on the
relative development of the parietal eminence
and its relative antero-posterior position. The
European MP group is characterized by an
antero-posteriorly and supero-inferiorly flatter
parietal bone (Fig. 2C). NR-1 is distinct from
H. sapiens; it is at an intermediate position be-
tween the Neanderthal and MP Homo clusters
(Fig. 2C). An unrooted phylogenetic analysis,
based on the mean shape of each Homo group,
placed NR-1 close to the origin of the branch
RESEARCH
Hershkovitz et al., Science 372, 14241428 (2021) 25 June 2021 1of5
1
Department of Anatomy and Anthropology, Sackler Faculty of Medicine, Tel Aviv University, Tel Aviv, Israel.
2
The Shmunis Family Anthropology Institute, the Dan David Center for Human
Evolution and Biohistory Research, Sackler Faculty of Medicine, Tel Aviv University, Tel Aviv, Israel.
3
Department of Oral Biology, the Goldschleger School of Dental Medicine, Sackler Faculty of
Medicine, Tel Aviv University, Tel Aviv, Israel.
4
UMR7194, HNHP, Département Homme et Environnement, Muséum national dHistoire naturelle, CNRS, UPVD, Paris, France.
5
CENIEH (National
Research Center on Human Evolution), Burgos, Spain.
6
Institute of Evolutionary Medicine, University of Zurich, Zurich, Switzerland.
7
Department of Evolutionary Anthropology, University of
Vienna, Vienna, Austria.
8
Department of Anthropology, Binghamton University (SUNY), Binghamton, NY, USA.
9
Centro UCM-ISCIII de Evolución y Comportamiento Humanos, Madrid, Spain.
10
Division of Anthropology, American Museum of Natural History, New York, NY, USA.
11
Departamento de Geodináica, Estratigrafía y Paleontología, Facultad de Ciencias Geológicas, Universidad
Complutense de Madrid, Ciudad Universitaria s/n, 28040, Madrid, Spain.
12
Department of Anthropology, University College London, London, UK.
13
Department of Human Molecular Genetics
and Biochemistry, Sackler Faculty of Medicine, Tel Aviv University, Tel Aviv, Israel.
14
PalaeoHub, Department of Archaeology, University of York, York, UK.
15
Thuringian State Office for the
Preservation of Historical Monuments and Archaeology Weimar, Germany.
16
Department of Environmental Biology, Sapienza University of Rome, Roma, Italy.
17
Core Facility for Micro-Computed
Tomography, University of Vienna, Vienna, Austria.
18
Institute of Archaeology, The Hebrew University of Jerusalem, Jerusalem, Israel.
19
Zinman Institute of Archaeology, University of Haifa, Haifa, Mount
Carmel, Israel.
*Corresponding author. Email: anatom2@tauex.tau.ac.il (I.H.); mayhila@tauex.tau.ac.il (H.M.); sarigrac@tauex.tau.ac.il (R.S.) These authors contributed equally to this work. Present address: Natural
History Museum, University of Florence, Florence, Italy.
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leading to African MP Homo, close to the split
from the H. erectus branch and to European MP
Homo and Neanderthals (includi ng Atapuerca
SH), and far from early and recent H. sapiens
(Fig. 2B).
With regard to the configuration of the
endoparietal surface (fig. S5), NR-1 is polygo-
nal, i.e., the surface is clearly oriented accord-
ing to three distinct planes (fig. S5). Instead,
Neanderthals and H. sapiens manifest an
arched endoparietal surface. The flatness of
the superior parietal lobule, seen in the NR-1
virtual endocast (fig. S6), is one of the most
characteristic features of MP Homo (9,10). Other
important characteristics of the NR-1 endocast
and that are also typical of MP Homo are the
very low position of the maximum endocra-
nial width at the superior part of the first tem-
poral convolution (fig. S6), the very short parie tal
lobe (fig. S7), and the differing lengths of the
maximal endocranial width and intraparietal
width as well as their posterior position on the
parietal bone (table S3). These features can
sometimes also be seen in Neanderthals (table
S3) (9,10). Conversely, recent H. sapiens speci-
mens exhibit subequal maximal endocranial and
intraparietal widths, which are located much
higher and more anteriorly than in NR-1 (9).
Thevascularpatternofthemiddlemenin-
geal vessels in NR-1 is simple. Only a few, short
ramifications are visible and anastomoses are
absent,asisthecaseinotherMPHomo and
Neanderthals (figs. S9 and S10) (11). The pos-
terior branch of the middle meningeal vessel in
NR-1 is as developed as the anterior one, a pattern
persistent among MP Homo. Both Neanderthals
(e.g., La Quina H5 and La Chapelle-aux-Saints)
and recent H. sapiens show a dominance of the
anterior branch; the latter also possesses com-
plex vascular endocranial imprints (fig. S10).
The NR-2 specimen is a robust mandible
(Fig. 3); the corpus is medio-laterally wide, and
the cortical bone is thick (Fig. 3, fig. S11). Its
most pronounced feature is the short ramus
relative to the body height, with a sturdy, low,
and wide coronoid process (Fig. 3). This spe-
cimen displays several archaic features (e.g., no
trigonum mentale or incurvatio mandibulae, a
wide incisura submentalis, a developed pla-
num alveolare, a strongly developed planum
triangulare, and a mandibular corpus that
presents fairly parallel alveolar and basal mar-
gins) commonly seen in MP Homo (12,13)
(supplementary text D and table S4A).
We combined taxonomically relevant man-
dibular features into a hierarchical cluster-
ing analysis (fig. S12). Modern and Pleistocene
humans form the two main clusters: NR-2 is
placed on a side branch of the latter, together
with MP Homo from Atapuerca SH, Tighenif 3,
Arago XIII, and one Neanderthal (fig. S12).
Thediscretetraitsunderscorethemosaicna-
ture of the NR mandible, showing archaic mor-
phology together with some Neanderthal traits.
The metric dimensions of the NR-2 mandi-
bular body are presented in fig. S13. The sym-
physeal area is considerably thick (16.6 mm),
close to the values of European MP Homo man-
dibles (16.9 ± 2.1 mm), and moderately tall
(33.7 mm), close to the Neanderthal mean
(34.0 ± 4.6 mm). The body (measured between
the first and the second molar) is thick (17.7 mm),
within the range of European MP Homo (18.1 ±
3.1 mm), yet taller (32.7 mm) than that of
European MP Homo (30.2 ± 1.6 mm) and
Neanderthals (29.9 ± 3.3 mm), close to the val-
ues of early H. sapiens (33.0 ± 4.0 mm).
The results of the 3D GM analysis (fig. S14
and tables S5A and S5B) for the NR-2 man-
dible are illustrated in Fig. 3C. The first two
principal components explain 47.5% of the
total variance. Variation along PC1 (37.9%) is
driven by changes in the length of the man-
dibular body, the shape of the ramus (shorter
and broader among archaic Homo), and the
expression of the mental area. Variation along
PC2 (9.6%) reflects changes in the body height
(mainly in the mental region), and the transi-
tion from a bodys parallel alveolar and basal
marginstoonesthatconvergeposteriorly.In
the PC1-PC2 plot, early and recent H. sapiens
separate from the other Homo specimens,
whereas European MP Homo and Atapuerca
SH are distinguished from Neanderthals (in-
cluding the Levantine Amud 1) and Asian
H. erectus. NR-2 falls between Neanderthals
and the European MP Homo specimens (in-
cluding Atapuerca SH), far outside the range
of the variation of H. sapiens. The phyloge-
netic analysis, based on the mandibular mean
shape of each hominin group, placed NR-2
on a separate branch (together with Tabun
C1), close to the split between MP European
fossils and Neanderthals, and far from
H. erectus,AfricanMPHomo,andH. sapiens
(Fig. 3B). This result, based on metrics alone,
largely echoes the results of the cluster analysis
based on discrete traits and confirms that NR-2
belongs to an archaic group with Neanderthal
affinities.
Hershkovitz et al., Science 372, 14241428 (2021) 25 June 2021 2of5
Fig. 1. The Nesher Ramla site and its stratigraphy. (A) Map of the eastern Mediterranean region showing
the location of the site. (B) View of the excavation from the east. (C) Section through the general
archaeological sequence placed within the local geology (the vertical scale on the left shows meters above
sea level). (D) Magnification of Units IV to VII. A red star denotes the location of the NR-1 (parietal bone) 8 m
below the surface in Unit VI. NR-2 (mandible) was found in the same unit, 2 m to the north.
RESEARCH |REPORT
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The lower second molar (NR-2 M
2
)iscom-
plete and shows some occlusal wear causing a
slight exposure of the dentine horns (Fig. 4A
and supplementary text E). The occlusal sur-
face of the NR-2 M
2
reveals four well-developed
cusps and a hypoconulid. The presence of
five main cusps is typical for most (70%) of
the Atapuerca SH Homo (14)andNeander-
thals (15). The NR-2 M
2
has a clear continuous
mid-trigonid crest and a discontinuous distal
trigonid crest on the dentine surface, corre-
sponding to grade 3 of Bailey et al.(16)(fig.
S15). A mid-trigonid crest is present in more
than 90% of Neanderthals and MP Homo from
Atapuerca SH (14,15). A grade 3 expression of
the mid-trigonid crest, as in the NR-2 M
2
,is
present in nearly 60% of the Neanderthal spe-
cimens, but it is absent in H. sapiens (16). The
Qesem Cave M
2
specimen (QC-J15) (17)shows
a similar pattern of a continuous mid-trigo nid
crest and a discontinuous distal trigonid crest
(fig. S15). The Ehringsdorf G specimen presents
only a mid-trigonid crest (but no distal crest)
(fig. S15), whereas the Mauer specimen does
not manifest a mid-trigonid crest at all.
The NR-2 M
2
has a single, pyramidal root bi-
furcating at the apical fourth of the root (Fig.
4, C and D). The large pulp cavity extends to
the middle of the root and branches out into
shortrootcanalsthatextendintotheapices,a
configuration of the roots known as tauro-
dontism (Fig. 4). This pyramidal root, with a
taurodontic pulp cavity, is frequent in Nean-
derthals (18). In modern humans, the second
lower molars possess separate mesial and
distal roots with some variation in the canals.
The root of the NR-2 M
2
(Fig. 4 and fig. S15) is
relatively long (16.4 mm), falling toward the
higher end of the range of the variation of both
Upper Paleolithic H. sapiens (11.3 to 16.8 mm)
and Neanderthals (14.3 to 16.5 mm).
The 3D GM analysis for the dentinal crown
shape (landmark configuration combining the
information from the enamel-dentin junction
or EDJ, and that from the cemento-enamel
junction or CEJ: fig. S16 for the measurement
template, table S6 for the landmark defini-
tions, and fig. S17 for the PC1-PC2 plot and the
PC1-PC3 plot) showed that the NR-2 M
2
falls at
the upper distant margin of the Neanderthal
range, close to the Krapina specimens and
Ehringsdorf G.
Shape variation along PC1 (30.6% of the
total variance) is driven by the relative height
of the crown and by the bucco-lingual expan-
sion of the EDJ relative to the dentine outline.
Like the M
2
of Neanderthals and H. sapiens,
NR-2 M
2
exhibits a relatively high crown and
a bucco-lingually expanded EDJ. Along PC2
(14.7%), the NR-2 M
2
plots toward the most
extreme range of the distribution, opposite
to the H. sapiens,AtapuercaSH,andAfrican
MP specimens. The associated shape is char-
acterized by the expansion of the distal as-
pectofthedentinecrown,afeaturethatNR-2
M
2
shares with some Neanderthal specimens
(Krapina and El Sidrón) and the European MP
Homo Ehringsdorf G (supplementary text E).
Differently from the parietal and mandible,
the unrooted phylogenetic treesconstruction,
based on the combined CEJ-EDJ data (Fig.
4B), resulted in a clear affiliation with Nean-
derthals, whereas Qesem QC-J15 associated
with Atapuerca SH. Concerning crown size,
NR2 M
2
is outside the modern human range
(fig. S18).
The cumulative evidence from the three
analyzed anatomical elements (parietal bone,
mandible, and M
2
) reveal a unique combi-
nation of archaic and Neanderthal features,
supporting the existence of a local, Levantine
population at the final MP. The results of the
Hershkovitz et al., Science 372, 14241428 (2021) 25 June 2021 3of5
Fig. 2. The NR-1 parietal bones and their analyses. (A) Virtual reconstruction of the middle portion of
the calvarium including both parietals. The almost complete right parietal was mirrored-imaged and the left
parietal fragments (represented in various colors) were superimposed, showing a very good overlap with
the contralateral part. (B) An unrooted phylogenetic tree constructed on the mean shape of each group
using the neighbor-joining method. NR-1 is positioned on the line leading to African MP Homo, close to the
splitting point from H. erectus and not far from the European MP Homo and Neanderthals. (C) PCA
plot in shape space for the parietal bone. PC1 separates early and recent H. sapiens from the rest of the
Homo sample. European MP Homo and Neanderthals (including the Levantine Amud 1) overlap and are
distinguished from Asian H. erectus and African MP Homo along PC3; NR-1 is intermediate between these
clusters and is close to Petralona and some late Asian H. erectus. Extreme shapes along the PCs are
shown from the anterior and medial views.
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quadratic discriminant analyses (QDAs) (table
S7) reinforce this observation, showing that an
affiliation of the NR fossils with early and
recent H. sapiens is highly unlikely, but that it
is impossible to establish whether NR fossils
are more likely to be classified as MP Homo,
Neanderthal, or H. erectus (the latter for the
parietal only). Consequently, the discriminant
function plot (fig. S1) shows that the NR-1
parietal falls between the H. erectus/African
MP Homo group and the European MP Homo/
Neanderthals, with a similar likelihood of be-
longing to either cluster (H. erectus =0.41,MP
Homo =0.34,Neanderthal=0.25,basedonthe
first three PCs).
The earliest that Neanderthal features in
Levantine fossils have been discernible in the
MP was around 400 ka ago at Qesem Cave
(19), the earliest modern humans were present
in the Levant around 180 ka ago (20), and
unequivocal Neanderthals did not appear in
theMiddleEastbefore~70kaago.NRbridges
a gap in this record, by displaying a highly
heterogeneous, yet archaic morphology. The
parietal documents a rather archaic shape of
the braincase; the mandible is similar to that
of MP Homo; the molar is quite Neanderthal-
like, similar to Ehringsdorf G.
Arsuaga et al.(21) advocated an earlier evo-
lutionary development of the masticatory
apparatus, compared with the braincase in
Neanderthals. Similarly, the Jebel Irhoud fos-
sils from North Africa possess a more primi-
tive neurocranium but a more H. sapienslike
face and dentition (22). Archaic populations
carrying Neanderthal-like features were also
present across much of the Eurasian continent
during the MP, revealed by the Chinese find-
ings of Maba, Xujiayao, and Xuchang (2327).
The existence of MP Asian populations de-
viating markedly from the H. erectus paradigm
has been repeatedly proposed, for instance,
for the Tongzi teeth (28)ortheSambungmacan
3cranium(29); the latter (together with
Ngandong 6 and 7) shows strong morpholog-
ical affinities with the NR-1 parietals.
The NR fossils could represent late-surviving
examples (140 to 120 ka) of a distinctive South-
west Asian MP Homo group, predating Levan-
tine Neanderthals from Amud, Kebara, and
Ein Qashish (70 to 50 ka). On the basis of their
mosaic morphology showing a different de-
gree of Neanderthal features, other MP Levan-
tine fossils, whose taxonomic affinities have
long been debated, from the sites of Qesem
Cave (19), Zuttiyeh Cave (30), and probably
Tabun Cave (31), might also be attributed to
this group (supplementary text F). Adopting
the cautious approach advocated by Mayr
(32), we suggest addressing this Levantine
MP paleodeme as the Nesher Ramla Homo.
Its presence from ~420 to 120 ka ago in a
geographically restricted area may have al-
lowed for repeated interbreeding with modern
Hershkovitz et al., Science 372, 14241428 (2021) 25 June 2021 4of5
Fig. 3. The NR-2 mandible and its analysis. (A) Different views of the reconstructed mandible. (B)Unrooted
phylogenetic tree constructed using the mean shape of each Homo group using the neighbor-joining method.
NR-2 and Tabun C1 have a common ancestor that is close to the split between Neanderthals and European
MP Homo and far from early and recent H. sapiens, as well as from H. erectus and African MP Homo.
(C) PCA plot for the mandible in shape space. The combination of PC1 and PC2 separates early and recent
H. sapiens from the other Homo specimens and distinguishes European MP Homo and Atapuerca from
Neanderthals (including the Levantine Amud 1) and Asian H. erectus. NR-2 plots between Neanderthals and
European MP Homo. Extreme shapes along PC1 and PC2 are shown from a lateral view.
RESEARCH |REPORT
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human populations such as the people from
Misliya Cave (20), a notion also supported by
their shared technological tradition [(6); sup-
plementary text F]. This scenario is compatible
with evidence of an early (200 to 400 ka ago)
gene flow between modern humans and
Neanderthals (3,4) and helps explain the vari-
able expression of the dental and skeletal
features of later Levantine fossils from the
Skhul and Qafzeh populations, a phenomenon
noted by anthropologists since the 1930s (31,33).
Moreover, a recent study of the Atapuerca SH
and Arago dental remains (1)suggestedthe
existence of more than one Homo lineage in
MP Europe [see also (34)] and hypothesized
the contribution of Levantine Homo groups
carrying Neanderthal-like traits to European
Homo lineages. The NR Homo,carryingNean-
derthal-like traits, could thus represent the
sourcepopulation postulated in the demog-
raphic sources and sinksmodel (5), accord-
ing to which Western Europe was repopulated
through a series of successive migrations.
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ACKNO WLED GME NTS
M. Prévost made the drawings in Fig. 1.P. Hervé made the drawingsin
fig. S10; A. Ehrenreich photographed the parietal bone in figs.S1D
and S9A. E. Santos performed the virtual reconstruction of the SH
mandibles. F. L. Bookstein supplied the R script for the quadratic
discriminant analysis. J. J. Hublin, Department of Human Evolution,
Max Pla nck Ins titute fo r Evoluti onary Ant hropol ogy, Leipzi g, Germany,
supplied some of the specimens for the study. Microcomputed
tomography scans ofthe NR fossils were done by S. Ellenbogenat the
Shmunis Family Anthropology Institute, Dan David Center for Human
Evolution and Biohistory Research, Tel Aviv University. Funding: This
work was funded by grants from the Dan David Foundation; the
Shmunis Family Anthropology Institute; the Leakey Foundation; the
CareArchaeological Foundation;the LabEx Sciences Archéologiques de
Bordeaux (LaScArBx ANR-10-LABX-52); the Dirección General de
Investigación of theMinisterio de Ciencia, Innovación y Universidades,
grant nos. PGC2018-093925-B-C31 and C33 (MCI/AEI/FEDER, UE);
and the Israel Science Foundation (1936/18, 1773/15). C.F. and V.A.K.
were financially supported by the Swiss National Science Foundation
(grant nos. 31003A_156299/1 and 31003A_176319). V.S. acknowledges
funding from the Alon Fellowship. Author contributions: I.H., H.M., R.S.,
G.W.W., and Y.Z. conceived the project, analyzed the data, and wrote
the manuscript. C.F., V.A.K., and Ar.P. performed the investigation
of the 3D datasets, from segmentation to data collection and
morphometric analysis. C.F. and G.W.W. performed the QDA
analysis. D.G.H., L.A.-B., and E.B. performed the endocast analysis.
R.Q., J.-L.A., C.F., and V.S. helped interpret the work and supervised
writing the manuscript. Y.Z. provided data on the lithic industry,
site formation, environmental conditions, and subsistence. M.M.-T.,
J.-M.BdC., L.M.-F., A.V., T.S., G.M., A.Po., and F.D.V. provided crucial
data on their fossils and thoroughly discussed the manuscript.
All authors drafted the manuscript. Ar.P. prepared the figures.
Competing interests: The authors declare no competing interests.
Data and materials availability: Data related to the new fossils
are available from the Shmunis Family Anthropology Institute website
(https://sfai.tau.ac.il/virtual_fossils_archive). Formal applications to
access the fossils should follow the regulations listed at https://en-
med.tau.ac.il/dan_david_center.
SUPPLEMENTARY MATERIALS
science.sciencemag.org/content/372/6549/1424/suppl/DC1
Materials and Methods
Supplementary Text
Figs. S1 to S18
Tables S1 to S7
References (35139)
1 March 2021; accepted 28 April 2021
10.1126/science.abh3169
Hershkovitz et al., Science 372, 14241428 (2021) 25 June 2021 5of5
Fig. 4. The lower left second molar (NR2 M
2
) and its analysis. (A) From the left, the NR2 M
2
in an
occlusal view, with and without an enamel cap, and in mesial, buccal, distal, and lingual views. The mesial and
distal interproximal wear facets are visible in the buccal and lingual views. (B) An unrooted phylogenetic
tree construction, based on the mean shape of each hominin group using the neighbor-joining method. NR2
M
2
is close to Neanderthals and far from H. sapiens.(C) Root canals (pulp cavities) in the mesial, buccal,
distal, and lingual views. The pyramidal roots and the taurodontic pulp cavity extend into the apex before
branching out into short root canals. (D) The dental roots (in brown) and their canals (in red) are presented
in glassimages of the mandible from a lateral view, revealing the presence of taurodontism.
RESEARCH |REPORT
on June 24, 2021 http://science.sciencemag.org/Downloaded from
from Nesher Ramla, IsraelHomoA Middle Pleistocene
Weber and Yossi Zaidner
Viviane Slon, Lou Albessard-Ball, Amélie Vialet, Tim Schüler, Giorgio Manzi, Antonio Profico, Fabio Di Vincenzo, Gerhard W.
Quam, Juan Luis Arsuaga, Viktoria A. Krenn, Maria Martinón-Torres, José María Bermúdez de Castro, Laura Martín-Francés,
Israel Hershkovitz, Hila May, Rachel Sarig, Ariel Pokhojaev, Dominique Grimaud-Hervé, Emiliano Bruner, Cinzia Fornai, Rolf
DOI: 10.1126/science.abh3169
(6549), 1424-1428.372Science
, abh3169 and abh3020, this issue p. 1424 and p. 1429; see also abj3077, p. 1395Science
time. had already occurred by thisH. sapiens and HomoPaleolithic, suggesting that admixture between Middle Pleistocene
findings provide archaeological support for cultural interactions between different human lineages during the Middle
efficient hunter of large and small game, used wood for fuel, cooked or roasted meat, and maintained fires. These
was anHomo or Neanderthals. Nesher Ramla Homo sapienstechnology that until only recently was linked to either
environmental data associated with these fossils. This evidence shows that these hominins had fully mastered
present the radiometric ages, stone tool assemblages, faunal assemblages, and other behavioral andet al.Zaidner in Europe, southwest Asia, and Africa. In a companion paper,Homolast surviving populations of Middle Pleistocene
140,000 to 120,000 years ago indicating the presence of a previously unrecognized group of hominins representing the
authors present comprehensive qualitative and quantitative analyses of fossilized remains from a site in Israel dated to
'' (see the Perspective by Mirazon Lahr). TheHomo population, the ''Nesher Ramla Homopreviously unknown archaic have uncovered evidence of aet al.greatly refined by recent new information. Adding to this trend, Hershkovitz
Our understanding of the origin, distribution, and evolution of early humans and their close relatives has been
in the LevantHomoMiddle Pleistocene
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... If we consider the dispersal path as a whole, then the paradox of the seemingly ruptured biogeographic region of ancient hominin, which includes three disjunct areas: (1) the East African Rift Belt, (2) the carbonate platform of the Mesozoic Terrane Belt, and (3) the East Anatolian-Caucasian zone of the Alpine orogen. This topological paradox is well illustrated by numerous paleo-archaeological, anthropological, paleontological, paleomagnetic, radiometric and landscape-ecosystem research methods [1,3,7,[12][13][14][15][16][17][18][19][20][21][22]. However, from the standpoint of regional geology and the evolution of terrestrial processes, it has not been systematically analyzed due to the fact that the data of global geodynamic mapping and zonation of this complex region became the subject of generalization only recently [23][24][25][26][27][28][29]. ...
... Human origins on the Earth and the ways of his dispersal are, in fact, among the most important problems in the field of natural sciences [1,3,7,15,19,21]. The discoveries of the past decades have allowed for a significant breakthrough in the field of ancient ecosystems, where the way of life of ancient hominins, artifacts of that time, and elements of biotic and abiotic habitats were studied. ...
... At the end of the Late Akchagylian regression period (about 1.1-0.8 Ma ago), there was the development by Man [21] of southern Eurasia and northern Africa [3,52]. ...
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... Critically, the eco-demographic niche-construction engine differentially affected the limiting conditions and possibilities for biocultural evolution across strong ecological productivity clines in southwestern Asia. Thus, small groups with divergent recent ancestry-alternatively from Subsaharan Africa, South Asia, or western Eurasia, as indicated by anatomical variability in Middle and Late Pleistocene fossil finds in the Levant (Freidline et al., 2012;Hershkovitz et al., 2021Hershkovitz et al., , 2018Hershkovitz et al., , 2015-would have sporadically occupied arid and semi-arid areas, exploiting highly dispersed wetland patches, along with limited seasonal resources across large territories. According to the demographic modelling reviewed above, tightly knit groups of ca. 150 would usually have persisted for centuries or millennia before stochastic fluctuations in demographic rates would have contributed to local extinctions. ...
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... Similarities are also observed between Crvena Stijena and certain sites in the Levant. For instance, Nesher Ramla in Israel-dated to MIS 5 and renowned for hominin fossils (Hershkovitz et al., 2021)-exhibits nearly all the technological features characteristic of Crvena Stijena: preferential and centripetal Levallois technology, COF (i.e., Kombewa cores and flakes), TFP, déjeté and convergent sidescrapers, and artifact thinning (Prévost and Zaidner, 2020). ...
... Therefore, rather than tracking favorable territories to the south, Neanderthals most likely declined during the harsh glacial phases, and only after an increase in average temperatures, new groups from glacial refuge zones repopulated the northern areas (Hublin and Roebroeks, 2009;Roebroeks et al., 2011). Some scholars broaden the regional extension of the glacial refugia proposing that the Levant was the Neanderthal population's source area, with recurrent recolonizations stemming externally from the continent (Dennell et al., 2011;Bermúdez de Castro and Martinón-Torres, 2013;Hershkovitz et al., 2021;Mihailović et al., 2022). From an archaeological perspective, these models' applications pose challenges due to the absence of prolonged cultural site comparisons within same regions. ...
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Chapter
The Shanidar Neandertal sample consists of two infants, Shanidar 7 and 9, three young adults, Shanidar 2, 6, and 8, and four older adults, Shanidar 1, 3, 4, and 5. It is difficult to tell exactly how old each of the adults was at death, but it is possible to rank the Shanidar partial skeletons in terms of age. The best arrangement would be in ascending order: 7 and 9; 2; 6 and 8; 1 & and 4; 3 and 5. The age of Shanidar 8 is highly uncertain so its position in the ranking could be considerably different. Among the Shanidar adults, three of the individuals—Shanidar 1, 3, and 4—appear, on the basis of pelvic evidence, to be male. Two other individuals—Shanidar 2 and 5 are probably male, largely on the basis of the large size of their skulls. Only two of the adults—Shanidar 6 and 8—appear to be female. It may be assumed that there is not necessarily a systematic bias in the sexing of the Shanidar Neandertals. The Shanidar fossil sample preserves males and females, infants, young adults, and old adults. However, it is clearly dominated, in numbers and degree of preservation, by elderly males. Even though there are other sites that have yielded the remains of adult Neandertals of similarly advanced ages at death, Shanidar is the only site that has provided the remains of several elderly male Neandertals. This should be kept in mind with respect to the evaluation of their morphologies as males tend to be larger and more robust than females among the Neandertals and many of the abnormalities present on the Shanidar specimens are undoubtedly correlated with the advanced ages of these individuals.
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In 1972 and 1983, four hominin teeth were recovered from Yanhui Cave, Tongzi, southern China and assigned to later Homo erectus or archaic Homo sapiens. The teeth can be dated to approximately 172,000-240,000 years before present. Here, in addition to the standard morphological comparisons, we reevaluate the morphology of the hominin teeth using geometric morphometric analyses and micro-computed tomography (micro-CT) scans. The Tongzi teeth were primarily compared to hominins from the same chronological period (late Middle Pleistocene) and/or the same geographic area (East Asia), although a wide range of hominins were included in the comparative sample. This study reveals that the Tongzi teeth do not fit the morphological pattern of classic H. erectus, and expands our understanding of the morphological diversity of the Asian Middle Pleistocene hominins. Overall, our results point to the existence of more than one paleodeme in East Asia during this period: one that can be taxonomically classified as H. erectus sensu stricto (represented by fossils such as Zhoukoudian, Hexian, and Yiyuan) and a second that is characterized by the expression of derived traits more commonly found in later Homo (such as the crown symmetry, lingual reduction, and simplified EDJ surface of the third premolar). More fossil and genetic findings will help assess the taxonomy of the "non-erectus" Asian Middle Pleistocene populations, like the Tongzi hominins.
Article
The Sima de los Huesos (SH) endocranial sample includes 16 complete or partial endocasts corresponding to European Middle Pleistocene hominins. Different anatomical and molecular studies have demonstrated that these hominins are phylogenetically related to Neanderthals, thus making them the earliest unquestionable representatives of the Neanderthal lineage. The description of endocranial variation in this population is fundamental to shedding light on the evolution of the Neanderthal brain. In this contribution, we analyze and describe endocranial variation in this sample, including aspects related to brain size (endocranial volume and encephalization) and brain organization (through qualitative descriptions and quantitative analyses). Our results indicate that the SH hominins show a transitional state between a primitive hominin endocranial configuration (which is found in Homo erectus and non-SH Middle Pleistocene Homo) and the derived configurations found in Neanderthals and modern humans, without a clear anticipation of classic Neanderthal endocranial traits. In comparison with other cranial and postcranial traits that show a fully Neanderthal or clear pre-Neanderthal condition in the SH collection, endocranial variation in these hominins is surprisingly primitive and shows no Neanderthal affinity. These results and the comparison with other cranial traits confirm that Neanderthals evolved in a mosaic fashion. Traits related to mastication (dental, facial and mandibular anatomy) led the Neanderthalization process, whereas neurocranial anatomy must have acquired a fully Neanderthal condition considerably later.
Book
Cambridge Core - Quantitative Biology, Biostatistics and Mathematical Modeling - A Course in Morphometrics for Biologists - by Fred L. Bookstein
Article
Paleoneurology deals with the study of brain anatomy in fossil species, as inferred from the morphology of their endocranial features. When compared with other living and extinct hominids, Homo sapiens is characterized by larger parietal bones and, according to the paleoneurological evidence, also by larger parietal lobes. The dorsal elements of the posterior parietal cortex (superior parietal lobules, precuneus, and intraparietal sulcus) may be involved in these morphological changes. This parietal expansion was also associated with an increase in the corresponding vascular networks, and possibly with increased heat loads. Only H. sapiens has a specific early ontogenetic stage in which brain form achieves such globular appearance. In adult modern humans, the precuneus displays remarkable variation, being largely responsible for the longitudinal parietal size. The precuneus is also much more expanded in modern humans than in chimpanzees. Parietal expansion is not influenced by brain size in fossil hominids or living primates. Therefore, our larger parietal cortex must be interpreted as a derived feature. Spatial models suggest that the dorsal and anterior areas of the precuneus might be involved in these derived morphological variations. These areas are crucial for visuospatial integration, and are sensitive to both genetic and environmental influences. This article reviews almost 20 years of my collaborations on human parietal lobe evolution, integrating functional craniology, paleoneurology, and evolutionary neuroanatomy.
Article
The variability observed in the growing Middle Pleistocene hominin fossil record of Europe continues to trigger much debate on taxonomic issues and the biological processes that gave rise to Neanderthals. Here we present a metric and morphological comparative study of the dental samples recovered from the sites of Arago (southeast France) and Sima de los Huesos (SH) in the Sierra de Atapuerca (northern Spain). These sites are key to providing answers to these questions since they have yielded the largest hominin samples so far recovered for this time period. Despite the geographical proximity of the two sites and the contemporaneity of their hominin assemblages, we have observed remarkable metric and morphological differences between the teeth at Arago and SH. Whereas the SH teeth present an almost morphological identity with European Neanderthals, the Arago teeth exhibit a combination of plesiomorphic as well as some Neanderthal-derived features. In addition, the Arago crown dimensions are remarkably larger than those from SH, the differences being statistically significant for most variables. We hypothesize that during the Middle Pleistocene the European continent was settled at different points in time by hominin groups coming from Southwest Asia, probably from a common mother population evolving in this latter region. These first settlers can be identified by their more plesiomorphic morphology, whereas the most recent settlers are closer in appearance to Neanderthals. In addition, genetic processes such as isolation, genetic drift, directional adaptation or hybridization would have given rise to the puzzle we observe in the current fossil record.