Gene 585 (2016) 9–12

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Gene

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

Letter to the Editor

The Out of Africa hypothesis and the ancestry of recent humans:

Cherchez la femme (et l'homme)
Úlfur Árnason
Department of Brain Surgery, Faculty of Medicine, University of Lund, Sweden

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

Article history: The Out of Africa hypothesis (OOAH) has been a mainstay in the discussion of human evolution since its presen-
Received 29 December 2015 tation in the 1980's. However, recent advances in palaeontology and molecular genetics have made it possible to
Received in revised form 3 March 2016 examine the hypothesis in a manner that was inconceivable at the time of its proposal. The palaeontological prog-
Accepted 14 March 2016
ress relates to early Homo finds in the Caucasus, Denisova finds in the Altai Mountains and Neanderthal finds in a
Available online 17 March 2016
wide range of localities from the Altai Mountains, the Caucasus, the Levant, Asia Minor, southern and Central
Keywords:
Europe and the Iberian Peninsula. The Eurasian location of these finds and recognition of the principle of Last
Human evolution common ancestor (LCA) lend no support to OOAH. The same conclusion is drawn from genomic findings,
Out of Africa hypothesis which (a) have revealed the presence of Denisovan and Neanderthal nuclear DNA, primarily in the genomes of
Neanderthals recent Eurasians and (b) have shown genomic introgression from early modern humans into Neanderthals in
Denisovans the Altai Mountains. Similarly, archaeological finds in Sulawesi and the discovery of ≈ 100,000 years old
Palaeontology human teeth in southern China constitute strong independent challenges to OOAH. The genomic and
Palaeogenomics palaeogenomic results and the new palaeontological and archaeological discoveries suggest (a) that the ances-
Biogeography
tors of modern humans had their origin in a Eurasian (largely Asian) biogeographic region which may also
LCA, last common ancestor
have extended into NE Africa, and (b) that the founders of basal African lineages became separated, geographi-
Askur/Embla hypothesis
cally and genetically, in the westernmost part of this region and spread from there to different parts of the
African continent.
© 2016 Elsevier B.V. All rights reserved.

1. Introduction
“… They came across two logs and created people out of them. …
The man was called Ask (Icelandic: Askur), the woman Embla, and
from them were produced the mankind …” (Sturluson, ≈ 1220).
OOAH posits a dispersal of the ancestors of recent humans from
Africa. Cann et al. (1987) provided the essential molecular basis of the
hypothesis in a study that included a phylogenetic tree based on restric-
tion maps of human mitochondrial DNA molecules (mtDNAs). The basal
split of the tree was between a branch that contained exclusively
African individuals and a branch that split between an African branch
and a branch that consisted of mtDNAs of different geographic origins.
The interpretation of the finding was that the basal split among the an-
cestors of modern humans had taken place in Africa between a branch
Abbreviations: A/C-60, the molecular calibration point implying the divergence
between ruminant artiodactyls and cetaceans (whales) set at 60 million years before
present; Hs, H. sapiens, the ancestor of Hss and Hsn; Hss, H. sapiens sapiens, modern
humans; Hsn, H. sapiens neanderthalensis, Neanderthals + Denisovans; Hsnd, d for
Denisovans; Hsnn, n for Neanderthals; LCA, last common ancestor; mtDNA, mitochondrial
deoxyribonucleic acid; MYBP, million years before present; OOAH, Out of Africa hypothesis;
YBP, years before present.
E-mail address: ulfur.arnason@gmail.com.
that had remained intact in Africa and another African branch from
which other human lineages had departed at different times to other
parts of the world.
OOAH has become acknowledged in a great number of molecular and
non-molecular studies of human evolution. The hypothesis has also been
taken as rebutting the so-called multiregional hypothesis on the origin
and evolution of modern humans advocated by M. H. Wolpoff and co‐
workers (e.g. Wolpoff et al. 2000). There is, however, a fundamental
and hitherto disregarded circumstance connected to OOAH, namely the
African placement superimposed on the root of the tree. That position
may have appeared reasonable at the time of the initial studies due to
the paucity of the non-African hominid palaeontological record. Even so it
should have been apparent that the very implementation of this root had
the automatic effect that the direction of any early human transfer could
never be into Africa, only out of or within that continent. O'Regan et al.
(2011) and Turner and O'Regan (2015) examined the palaeontological
record related to the dispersal of large mammals between Africa and
Eurasia during Pliocene–Pleistocene. Although the studies were inconclu-
sive with respect to OOAH they showed that mammalian dispersal
between Africa and Asia was by no means a one-way route.
The phylogenetic relationship of recent humans, Neanderthals and
Denisovans as recovered in studies of complete mtDNAs is shown
schematically in Fig. 1. The figure is a simplification of the tree presented

http://dx.doi.org/10.1016/j.gene.2016.03.018
0378-1119/© 2016 Elsevier B.V. All rights reserved.
10 Letter to the Editor
Fig. 1. Phylogenetic relationship among Denisovans, Neanderthals (Hsn, Homo sapiens
neanderthalensis) and recent humans (Hss, Homo sapiens sapiens) as recovered in studies
of mitochondrial DNA (mtDNA). Denisova is sister to a branch (Hs, Homo sapiens) that
splits into Neanderthals and recent humans. Krause et al. (2010) estimated the
divergence time between Denisova and Hs at ≈1 MYBP (million years before present)
and that between Neanderthals and recent humans at 0.5–0.3 MYBP. The split between
San and Han represents the deepest divergence within Hss. Molecular datings based on
mtDNA place Denisova within the temporal realm of Eurasian Homo erectus and related
taxa (e.g. Ferring et al., 2011; Lordkipanidze et al., 2013), but its mtDNA identity as to a
defined taxon remains unknown. H? indicates an unspecified Homo outgroup.
by Krause et al. (2010) and those of Meyer et al. (2014) and
Sawyer et al. (2015). The earliest bifurcation in the tree is between a
branch that includes Denisovans and a branch that splits into Homo sapi-
ens sapiens (Hss) and Homo sapiens neanderthalensis (Hsn). As a conse-
quence the branch leading to Hss/Hsn is designated Hs for H. sapiens. In
comparison, studies of nuclear DNA, Fig. 2, place Denisovans as the sister
taxon of Neanderthals (Reich et al., 2010; Meyer et al., 2012; Sawyer
et al., 2015). The effect of this discrepancy vis-à-vis the mtDNA tree is ap-
parent in that Hsnn and Hsnd (d for Denisova) join on a common branch,
Hsn, as the sister group of Hss. It should be noted that the length of the Hs
branch leading to the split between Hss and Hsn remains unknown in the
absence of a molecularly definable sister group of that branch.
Regarding the phylogenetic incongruity that may occur in analysis of
nuclear and mitochondrial DNA it is noteworthy that as long as 25 years
ago Pickford (1991) drew attention to this circumstance, with particular
reference to OOAH.
Fig. 2. Phylogenetic studies of nuclear DNA place Denisovans and Neanderthals as sister
groups on a branch, Hsn, that is sister to recent humans, Hss. The relationship among
recent humans is in accord with Reich et al. (2010). The star introduced in the tree
indicates the deepest potential position for genomic transfers between Neanderthals/
Denisovans (Hsnn/Hsnd) and the ancestors of recent humans (Hss). The position does
not require, however, the presence of Neanderthal and/or Denisovan DNA in the
genomes of all recent Eurasians. The location of the star implies, in conjunction with
limited occurrence of Neanderthal/Denisovan DNA in African genomes, that the ancestors
of San and Yoruba (both African) split from the Eurasian ancestors of recent humans prior
to the genomic exchanges between the Eurasians and Neanderthals/Denisovans. With ref-
erence to Nordic mythology (Sturluson, ≈ 1220) the names Askur and Embla denote the
biparental nature of the nuclear DNA of the last common ancestors of Hsn and Hss.
2. Discussion
The availability of sequences of ancient DNA, both mitochondrial
and nuclear, from Neanderthals and Denisovans and progress in Homo
palaeontology allow examination of OOAH in a manner that could not
have been envisaged in the 1980's. Thus, when OOAH became accepted,
the Homo/Pan calibration point was commonly placed at 4.5–5 MYBP.
This age became significantly revised, however, with the description
of Orrorin tugenensis (Senut et al., 2001) and Sahelanthropus tchadensis
(Brunet et al., 2002), two fossils that in an instant required moving
the Pan/Homo split to N7 MYBP.
The time of divergence between Artiodactyla (as represented by ru-
minants) and Cetacea (whales) set at 60 MYBP (Arnason and Gullberg,
1996; Ursing and Arnason, 1998) has become the primary molecular
standard (A/C-60) for calculating evolutionary divergences among
mammals in cases where the fossil record is inconclusive. A/C-60 was ex-
amined critically by van Tuinen and Hadly (2004) and shown to outper-
form other mammalian standards of this kind. Application of A/C-60 to
the sequences of complete mtDNA molecules places the Pan/Homo diver-
gence at ≥7.5 MYBP and the deepest divergences among recent humans
at ≈1/30 of this time, i.e. at ≥250,000 YBP (Arnason et al., 2008). This
dating is somewhat earlier than most other estimates of this divergence,
a circumstance that may be related to differences in the applied ap-
proaches, not least the dating allocated to the divergence between Pan
and Homo, which is still placed at ≈6.5 MYBP by some authors despite
its conflict with established palaeontology. The 250,000 YBP estimate is
consistent with the marked progressive Palaeolithic exploitation of
small animals in the Levant ≥200,000 YBP (Stiner et al., 1999) as a conse-
quence of enduring human presence and faunal exploitation outside
Africa. A much earlier and enduring presence of advanced Homo in the
Levant (e. g. Acheulian Technocomplex) has also been documented ar-
chaeologically (Goren-Inbar et al., 2000; Goren-Inbar, 2011).
Based on analysis of the first complete Denisovan mtDNA, Krause
et al. (2010) concluded that Denisova derived from a migration out of
Africa ≈ 1 MYBP, followed by an exodus of early Neanderthals (also
out of Africa) between 500,000 and 300,000 YBP and the ancestors of
non-African modern humans ≈ 50,000 YBP. The study did not clarify
how finds in the Altai Mountains (Denisova Cave) could indicate migra-
tion out of Africa 1 MY earlier. Similarly, the exclusive location of Nean-
derthal fossils in Eurasia is problematic for the traditional OOAH. Hublin
(2009) referred to that crucial Neanderthal issue in the following
manner: “… none have been documented in North Africa”.
It should be borne in mind that Krause et al. (2010), at the time of
the publication of their study, were incognizant of the phylogenetic
discrepancy between the nuclear and mitochondrial DNA of Denisova
mentioned above. They could therefore not examine the Neanderthal/
Denisova relationship in a perspective that otherwise had made it difficult
to reconcile OOAH with the absence of Neanderthals in Africa. Martinón-
Torres et al. (2011) examined the study of Krause et al. (2010) from a
palaeontological/archaeological point of view and expressed reservations
regarding the latter's conclusions that Denisovans and Neanderthals de-
rived from migrations out of Africa.
Groucutt et al. (2015) addressed OOAH in a recent study by simulat-
ing different OOAH scenarios. The authors maintained that current ge-
netic, palaeontological and archaeological data indicated that Hss had
originated in Africa and dispersed from that continent. The authors
expressed, however, a particular qualification regarding this view, viz.:
“Future fossil discoveries in Southern Asia have the potential to radically
transform our understanding of that dispersal” (i.e. OOAH). This reser-
vation became actualized most recently at the palaeontological descrip-
tion of a collection of human teeth, with an age of 80,000–120,000 years,
in Hunan Province, southern China (Liu et al., 2015) and soon thereafter
by the demonstration of hominin presence in Sulawesi from
N200,000 YBP until ≈100,000 YBP (van den Bergh et al., 2016).
The basal mtDNA relationship of recent humans that Cann et al.
(1987) obtained by midpoint-rooting has been recovered in virtually
 Letter to the Editor
all later studies that have applied rooting with an outgroup. Hence the
concern with the results of Cann et al. (1987) is not related to rooting pro-
cedures as such but rather to the superimposed placement of the root of
the tree in Africa. Although that geographical placement of the Hss tree
appeared reasonable at the time of its presentation, recent scientific ad-
vances in palaeontology and genomics, notably palaeogenomics, and ap-
plication of the principle of last common ancestor (LCA) show limited
compatibility with that proposal. Instead, recognition of these advances
suggests that early Hss evolution took place in a largely Asian biogeo-
graphic area that included NE Africa, the Arabian Peninsula, the Levant,
Asia Minor, parts of SE Europa and areas extending far eastward
into Asia. It appears that the scientific progress mentioned above
harmonizes far better with such an interpretation than with the African
location upon which OOAH was based some 30 years ago. It is therefore
more logical to suggest that the basal African group(s) identified by
Cann et al. (1987) split off in the western part of this largely Asian biogeo-
graphic region while the remaining human ancestors evolved within Eur-
asia (primarily Asia) and spread gradually from there into other parts of
the world. A scenario of this kind is in accord with the occurrence of
both Neanderthal (Green et al., 2010; Wall et al., 2013; Sankararaman
et al., 2014; Vernot and Akey, 2014) and Denisovan (Reich et al., 2011;
Skoglund and Jakobsson, 2011; Meyer et al., 2012) nuclear DNA in the ge-
nomes of recent Eurasians and the limited degree of these genetic com-
ponents in African populations (Wall et al., 2013; Sankararaman et al.,
2014). Fig. 2 depicts the underlying genetic transfers in a phylogenetic
context, showing that they took place after the separation of the African
and Eurasian human populations. With this molecular information in
mind, together with the absence of Neanderthal fossils in Africa it be-
comes increasingly problematic to place the initial separation among
the ancestors of recent humans (Hss) and those of Neanderthals (Hsn)
outside Eurasia.
Prüfer et al. (2014) also addressed the relationship between and
within Hsn and Hss in the context of gene flow events among Neander-
thals, Denisovans and early modern humans. The authors maintained,
consistent with traditional OOAH, that the ancestors of both Neander-
thals and Denisovans had left Africa long before the emergence of mod-
ern humans. Letting alone the circumstance that Neanderthal ancestry
is not traceable to Africa the authors might have considered that Hsn
and Hss constitute sister groups that share common ancestry (Fig. 2).
Therefore, and in compliance with the principle of LCA, the authors
should have recognized that the ancestors of Neanderthals (Hsn) and
these of recent humans (Hss) could arise neither at different times nor
in different parts of the world.
Kuhlwilm et al. (2016) described the occurrence of gene flow from
early Hss into Neanderthals of the Altai Mountains ≥ 100,000 YBP, i.e.
much earlier than the Hsn/Hss genetic admixture noted by Wall et al.
(2013), Sankararaman et al. (2014) and Vernot and Akey (2014).
Kuhlwilm et al. (2016), reported furthermore that the genomic contri-
bution of Hss to the Altai Neanderthals shared characteristics with the
genomes of Africans such as these represented by the San branch in
Fig. 2, thereby indicating an early Hss dispersal out of Africa. In order
to provide additional support to this conclusion the authors stated at
the end of their paper: “The recent demonstration that modern humans
may have been in China as early as 120,000 years ago (Liu et al., 2015)
also suggests that modern humans migrated early out of Africa.” This in-
terpretation of the Daoxian finds is diametrically opposed to what the
authors (Liu et al., 2015) maintained themselves, namely that the
study showed “that fully modern [human] morphologies were present
in southern China 30,000–70,000 years earlier than in the Levant and
Europe”.
When considered together the studies of Liu et al. (2015) and
Kuhlwilm et al. (2016) may allow a coherent conclusion, namely that
a northern fraction of E Asian Hss became incorporated in the Altai Ne-
anderthals while the main portion of west migrating Hss diverged in
Asia and spread from there into Africa and Europa, with the lineage rep-
resented by San in Fig. 2 becoming the earliest offshoot into Africa along
11
a route which at that time did not cross Neanderthal territories. The
probable route into Africa, a northern one or one going across the Red
Sea to E Africa, is irrelevant for this proposal. The occurrence of limited
amounts of Neanderthal DNA in some recent Kenyans (Wall et al., 2013;
Sankararaman et al., 2014) might suggest a Red Sea route for that ad-
mixture, however.
3. Conclusions
In the light of a series of genomic and palaeogenomic molecular re-
sults the model presented here suggests an Asian split between the an-
cestors of recent humans (Hss) and Neanderthals (Hsn). Similarly, in a
palaeontological/archaeological sense a scenario of this kind is consistent
with the recently described dental finds in Hunan Province, China (Liu
et al., 2015) and the presence of archaic hominins in Sulawesi from
N200,000 YBP until ≈100,000 YBP (van den Bergh et al., 2016). Taken to-
gether these recent studies render the assumptions behind OOAH highly
problematic, since persisting with that hypothesis would require the pre-
sentation of African evidence that is of a comparable level with these re-
cent Eurasian molecular and palaeontological results. Until such African
evidence is presented and critically compared with this Eurasian evi-
dence it appears appropriate to attach the names Askur and Embla
(Sturluson, ≈ 1220) – with their Eurasian connotations – to the
hypothetic last common Eurasian ancestors of Hsn (Neanderthals/
Denisovans) and Hss (recent humans). The inclusion of both a male
(Askur) and a female (Embla) relates to and underlines the biparental na-
ture of the DNA that allows this suggestion.
Conflict of interests
The author declares no conflict of interest.
Acknowledgments
The author thanks Prof. Eric H. Harley, Prof. Kent Larsson, Dr. Maria
Nilsson-Janke, Prof. Axel Janke and Prof. Martin Pickford for their valu-
able comments at different stages during the writing of the paper. The
positive input of the reviewers is also acknowledged.
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