Multiregional origin of modern humans

The multiregional hypothesis, multiregional evolution

(MRE), or polycentric hypothesis is a scientific model
that provides an alternative explanation to the more
widely accepted "Out of Africa" model of monogenesis
for the pattern of human evolution.

Multiregional evolution holds that the human species first

arose around two million years ago and subsequent
human evolution has been within a single, continuous
human species. This species encompasses all archaic
human forms such as H. erectus and Neanderthals as
well as modern forms, and evolved worldwide to the
diverse populations of anatomically modern humans
(Homo sapiens).

The hypothesis contends that the mechanism of clinal

variation through a model of "Centre and Edge" allowed
for the necessary balance between genetic drift, gene
flow and selection throughout the Pleistocene, as well as
overall evolution as a global species, but while retaining
regional differences in certain morphological features.[3]
Proponents of multiregionalism point to fossil and
genomic data and continuity of archaeological cultures as
support for their hypothesis.
The multiregional hypothesis was first proposed in 1984,
and then revised in 2003. In its revised form, it is similar
to the Assimilation Model, which holds that modern
humans originated in Africa and today share a
predominant recent African origin, but have also
absorbed small, geographically variable, degrees of
admixture from other regional (archaic) hominin
species.[4]
A graph detailing the evolution to modern humans
using the multiregional hypothesis of human
evolution. The horizontal lines represent
'multiregional evolution' gene flow between regional
lineages. In Weidenreich's original graphic,[1] there
were also diagonal lines between the populations,
e.g. between African H. erectus and Archaic
Asians and between Asian H. erectus and Archaic
Africans. This created a "trellis" (as Wolpoff called
it[2]) or a "network" that emphasized gene flow
between geographic regions and within time. It is
important to remember that the populations on the
chart are not discrete – i.e., they do not represent
different species, but are samples within a long
lineage experiencing extensive gene flow.

Contents
History
Overview
"Classic" vs "weak" multiregionalism
Genetic studies
Fossil evidence
Morphological clades
Indonesia, Australia
China
Europe
Genetic evidence
Mitochondrial Eve
Neanderthal mtDNA
Nuclear DNA
Ancient DNA
See also
References
Further reading
External links

History

Hominin timeline
Overview
0— ←Modern humans
The Multiregional hypothesis was proposed in P NeanderthalsHomo sapiens Earliest clothes
– l H. heidelbergensis
1984 by Milford H. Wolpoff, Alan Thorne and e ←Earliest cooking
Xinzhi Wu.[5][6][3] Wolpoff credits Franz -1 — i
s Homo erectus
Weidenreich's "Polycentric" hypothesis of
– t ←Earliest fire use
human origins as a major influence, but cautions o ←Expansion beyond
that this should not be confused with -2 — c Africa
e
polygenism, or Carleton Coon's model that n Homo habilis
–
minimized gene flow.[6][7][8] According to e
Wolpoff, multiregionalism was misinterpreted by -3 —
William W. Howells, who confused ←Stone tools
– Australopithecus
Weidenreich's hypothesis with a polygenic P
"candelabra model" in his publications spanning -4 — l ←Earliest bipedal
five decades: i
Ardipithecus H
–o
c
-5 — e o
How did Multiregional evolution n
–e Hominini m
get stigmatized as polygeny? We
believe it comes from the confusion -6 — Orrorin i ←Chimpanzee split
of Weidenreich's ideas, and
– n
ultimately of our own, with Coon's.
The historic reason for linking Sahelanthropus i
-7 — M
Coon's and Weidenreich's ideas i d
came from the mischaracterizations –o
c
of Weidenreich's Polycentric model Oreopithecus s
-8 — e
as a candelabra (Howells, 1942, n
1944, 1959, 1993), that made his –e
Polycentric model appear much Ouranopithecus
-9 — ←Gorilla split
more similar to Coon's than it
actually was.[9] –
Nakalipithecus
-10 — ←Earlier apes
(million years ago)
Through the influence of Howells, many other
anthropologists and biologists have confused
multiregionalism with polygenism i.e. separate Ice Ages Life timeline
0— Primates ←Earliest apes
 0 Flowers Birds Primates
←Earliest apes
or multiple origins for different populations. Quaternary
P Mammals
Alan Templeton for example notes that this –h Plants Dinosaurs
K aroo a ←Tetrapoda
confusion has led to the error that gene flow A ndean
n
-500 — e Arthropods Molluscs
←Cambrian explosion
between different populations was added to the r ←Ediacaran biota
Multiregional hypothesis as a "special pleading C ryogenian o ←Earliest animals
–z
in response to recent difficulties", despite the o ←Earliest plants
fact: "parallel evolution was never part of the -1000 — i Multicellular
c life
multiregional model, much less its core, whereas ←Sexual reproduction
–
gene flow was not a recent addition, but rather
was present in the model from the very P
-1500 — r ←Earliest fungi
beginning"[10] (emphasis in original). Despite o
this, multiregionalism is still confused with –t Eukaryotes
e
polygenism, or Coon's model of racial origins, r
-2000 — o
from which Wolpoff and his colleagues have z
distanced themselves.[11][12] Wolpoff has also – o
H uronian i ←Oxygen crisis
defended Wiedenreich's Polycentric hypothesis c
-2500 — ←Atmospheric oxygen
from being labeled polyphyletic. Weidenreich
himself in 1949 wrote: "I may run the risk of Photosynthesis
–
being misunderstood, namely that I believe in Pongola
polyphyletic evolution of man".[13] -3000 — A
r
In 1998, Wu founded a China-specific –c
h
Multiregional model called "Continuity with e
-3500 — a ←Earliest oxygen
[Incidental] Hybridization".[14][15] Wu's variant n Single-celled
only applies the Multiregional hypothesis to the – life
East Asian fossil record, and is popular among ←LHB meteorites
Chinese scientists.[16] However, James Leibold, -4000 —
H
←Earliest life
a political historian of modern China, has argued a Water
–d
the support for Wu's model is largely rooted in e
a ←Earliest water
Chinese nationalism.[17] Outside of China, the -4500 — n ←Earliest Earth
Multiregional hypothesis has limited support, (million years ago) (−4540)
held only by a small number of
paleoanthropologists.[18]

"Classic" vs "weak" multiregionalism

Chris Stringer, a leading proponent of the more mainstream recent African origin theory, debated
Multiregionalists such as Wolpoff and Thorne in a series of publications throughout the late 1980s and
1990s.[19][20][21][22] Stringer describes how he considers the original Multiregional hypothesis to have been
modified over time into a weaker variant that now allows a much greater role for Africa in human evolution,
including anatomical modernity (and subsequently less regional continuity than was first proposed).[23]

Stringer distinguishes the original or "classic" Multiregional model as having existed from 1984 (its
formulation) until 2003, to a "weak" post-2003 variant that has "shifted close to that of the Assimilation
Model".[24][25]

Genetic studies
The finding that "Mitochondrial Eve" was relatively recent and African seemed to give the upper hand to the
proponents of the Out of Africa hypothesis. But in 2002, Alan Templeton published a genetic analysis
involving other loci in the genome as well, and this showed that some variants that are present in modern
populations existed already in Asia hundreds of thousands of years ago.[26] This meant that even if our male
line (Y chromosome) and our female line (mitochondrial DNA) came out of Africa in the last 100,000 years or
so, we have inherited other genes from populations that were already outside of Africa. Since this study other
studies have been done using much more data (see Phylogeography).

Fossil evidence

Morphological clades

Proponents of the multiregional hypothesis see regional continuity of

certain morphological traits spanning the Pleistocene in different regions
across the globe as evidence against a single replacement model from
Africa. In general, three major regions are recognized: Europe, China, and
Indonesia (often including Australia).[27][28][29] Wolpoff cautions that the
continuity in certain skeletal features in these regions should not be seen in
a racial context, instead calling them morphological clades; defined as sets
of traits that "uniquely characterise a geographic region".[30] According to Replica of Sangiran 17 Homo
Wolpoff and Thorne (1981): "We do not regard a morphological clade as a erectus skull from Indonesia
unique lineage, nor do we believe it necessary to imply a particular showing obtuse face to vault
angle determined by fitting of
taxonomic status for it".[31] Critics of multiregionalism have pointed out
bones at brow.
that no single human trait is unique to a geographical region (i.e. confined
to one population and not found in any other) but Wolpoff et al. (2000)
note that regional continuity only recognizes combinations of features, not
traits if individually accessed, a point they elsewhere compare to the forensic
identification of a human skeleton:

Regional continuity... is not the claim that such features do not

appear elsewhere; the genetic structure of the human species
makes such a possibility unlikely to the extreme. There may be
uniqueness in combinations of traits, but no single trait is likely
to have been unique in a particular part of the world although it
might appear to be so because of the incomplete sampling
Cast of anatomically modern
provided by the spotty human fossil record.
human Kow Swamp 1 skull
from Australia with a face to
Combinations of features are "unique" in the sense of being found in only vault angle matching that of
one region, or more weakly limited to one region at high frequency (very Sangiran 17 (Wolpoff's
rarely in another). Wolpoff stresses that regional continuity works in reconstruction).
conjunction with genetic exchanges between populations. Long-term
regional continuity in certain morphological traits is explained by Alan
Thorne's "Centre and Edge"[32] population genetics model which resolves Weidenreich's paradox of "how did
populations retain geographical distinctions and yet evolve together?". For example, in 2001 Wolpoff and
colleagues published an analysis of character traits of the skulls of early modern human fossils in Australia and
central Europe. They concluded that the diversity of these recent humans could not "result exclusively from a
single late Pleistocene dispersal", and implied dual ancestry for each region, involving interbreeding with
Africans.[33]
Indonesia, Australia

Thorne held that there was regional continuity in Indonesia and Australia for a morphological clade.[34][35]
This sequence is said to consist of the earliest fossils from Sangiran, Java, that can be traced through
Ngandong and found in prehistoric and recent Aboriginal Australians. In 1991, Andrew Kramer tested 17
proposed morphological clade features. He found that: "a plurality (eight) of the seventeen non-metric features
link Sangiran to modern Australians" and that these "are suggestive of morphological continuity, which
implies the presence of a genetic continuum in Australasia dating back at least one million years"[36] but Colin
Groves has criticized Kramer's methodology, pointing out that the polarity of characters was not tested and that
the study is actually inconclusive.[37] Dr. Phillip Habgood discovered that the characters said to be unique to
the Australasian region by Thorne are plesiomorphic:

...it is evident that all of the characters proposed... to be 'clade features' linking Indonesian Homo
erectus material with Australian Aboriginal crania are retained primitive features present on Homo
erectus and archaic Homo sapiens crania in general. Many are also commonly found on the crania
and mandibles of anatomically-modern Homo sapiens from other geographical locations, being
especially prevalent on the robust Mesolithic skeletal material from North Africa."[38]

Yet, regardless of these criticisms Habgood (2003) allows for limited regional continuity in Indonesia and
Australia, recognizing four plesiomorphic features which do not appear in such a unique combination on
fossils in any other region: a sagittally flat frontal bone, with a posterior position of minimum frontal breadth,
great facial prognathism, and zygomaxillary tuberosities.[39] This combination, Habgood says, has a "certain
Australianness about it".

Wolpoff, initially skeptical of Thorne's claims, became convinced when reconstructing the Sangiran 17 Homo
erectus skull from Indonesia, when he was surprised that the skull's face to vault angle matched that of the
Australian modern human Kow Swamp 1 skull in excessive prognathism. Durband (2007) in contrast states
that "features cited as showing continuity between Sangiran 17 and the Kow Swamp sample disappeared in
the new, more orthognathic reconstruction of that fossil that was recently completed".[40] Baba et al. who
newly restored the face of Sangiran 17 concluded: "regional continuity in Australasia is far less evident than
Thorne and Wolpoff argued".[41]

China

Xinzhi Wu has argued for a morphological clade in China

Replica of Homo erectus ("Peking man")
skull from China.
spanning the Pleistocene, characterized by a combination of 10
features.[42][43] The sequence is said to start with Lantian and
Peking Man, traced to Dali, to Late Pleistocene specimens (e.g.
Liujiang) and recent Chinese. Habgood in 1992 criticized Wu's
list, pointing out that most of the 10 features in combination appear
regularly on fossils outside China.[44] He did though note that
three combined: a non-depressed nasal root, non-projecting
perpendicularly oriented nasal bones and facial flatness are unique
to the Chinese region in the fossil record and may be evidence for
limited regional continuity. However, according to Chris Stringer,
Habgood's study suffered from not including enough fossil
samples from North Africa, many of which exhibit the small
combination he considered to be region-specific to China.[22]
Facial flatness as a morphological clade feature has been rejected by many anthropologists since it is found on
many early African Homo erectus fossils, and is therefore considered plesiomorphic,[45] but Wu has responded
that the form of facial flatness in the Chinese fossil record appears distinct to other (i.e. primitive) forms. Toetik
Koesbardiati in her PhD thesis "On the Relevance of the Regional Continuity Features of the Face in East
Asia" also found that a form of facial flatness is unique to China (i.e. only appears there at high frequency,
very rarely elsewhere) but cautions that this is the only available evidence for regional continuity: "Only two
features appear to show a tendency as suggested by the Multiregional model: flatness at the upper face
expressed by an obtuse nasio-frontal angle and flatness at the middle part of the face expressed by an obtuse
zygomaxillay angle".

Shovel-shaped incisors are commonly cited as evidence for regional continuity in China.[46][47] Stringer
(1992) however found that shovel-shaped incisors are present on >70% of the early Holocene Wadi Halfa
fossil sample from North Africa, and common elsewhere.[48] Frayer et al. (1993) have criticized Stringer's
method of scoring shovel-shaped incisor teeth. They discuss the fact that there are different degrees of
"shovelled" e.g. trace (+), semi (++), and marked (+++), but that Stringer misleadingly lumped all these
together: "...combining shoveling categories in this manner is biologically meaningless and misleading, as the
statistic cannot be validly compared with the very high frequencies for the marked shoveling category reported
for East Asians."[28] Palaeoanthropologist Fred H. Smith (2009) also emphasizes that: "It is the pattern of
shoveling that identities as an East Asian regional feature, not just the occurrence of shoveling of any sort".[4]
Multiregionalists argue that marked (+++) shovel-shaped incisors only appear in China at a high frequency,
and have <10% occurrence elsewhere.

Europe

Since the early 1990s, David W. Frayer

has described what he regards as a
morphological clade in
Europe. [49][50][51] The sequence starts
with the earliest dated Neanderthal
specimens (Krapina and Saccopastore
skulls) traced through the mid-Late
Pleistocene (e.g. La Ferrassie 1) to
Vindija Cave, and late Upper Palaeolithic
Cro-Magnons or recent Europeans.
Although many anthropologists consider
Neanderthals and Cro Magnons Comparison of modern human (left) and Neanderthal (right) skulls.
morphologically distinct, [52][53] Frayer
maintains quite the opposite and points to
their similarities, which he argues is evidence for regional continuity:

"Contrary to Brauer's recent pronouncement that there is a large and generally recognized
morphological gap between the Neanderthals and the early moderns, the actual evidence provided
by the extensive fossil record of late Pleistocene Europe shows considerable continuity between
Neanderthals and subsequent Europeans."[28]

Frayer et al. (1993) consider there to be at least four features in combination that are unique to the European
fossil record: a horizontal-oval shaped mandibular foramen, anterior mastoid tubercle, suprainiac fossa and
narrowing of the nasal breadth associated with tooth-size reduction. Regarding the latter, Frayer observes a
sequence of nasal narrowing in Neanderthals, following through to late Upper Palaeolithic and Holocene
(Mesolithic) crania. His claims are disputed by others,[54] but have received support from Wolpoff, who
regards late Neanderthal specimens to be "transitional" in nasal form between earlier Neanderthals and later
Cro Magnons.[55] Based on other cranial similarities, Wolpoff et al. (2004) argue for a sizable Neanderthal
contribution to modern Europeans.[56]

More recent claims regarding continuity in skeletal morphology in Europe focus on fossils with both
Neanderthal and modern anatomical traits, to provide evidence of interbreeding rather than
replacement.[57][58][59] Examples include the Lapedo child found in Portugal[60] and the Oase 1 mandible
from Peștera cu Oase, Romania,[61] though the Lapedo child is disputed by some.[62]

Genetic evidence

Mitochondrial Eve

A 1987 analysis of mitochondrial DNA from 147 people by Cann

et al. from around the world indicated that their mitochondrial
lineages all coalesced in a common ancestor from Africa between
140,000 and 290,000 years ago.[63] The analysis suggested that
this reflected the worldwide expansion of modern humans as a
new species, replacing, rather than mixing with, local archaic
humans outside of Africa. Such a recent replacement scenario is
not compatible with the Multiregional hypothesis and the mtDNA
results led to increased popularity for the alternative single
replacement theory.[64][65][66] According to Wolpoff and
colleagues:

"When they were first published, the Mitochondrial

Eve results were clearly incongruous with
Multiregional evolution, and we wondered how the
two could be reconciled."[67]

Human mitochondrial DNA tree.

Multiregionalists have responded to what they see as flaws in the "Mitochondrial Eve" is near the top of
Eve theory,[68] and have offered contrary genetic the diagram, next to the jagged arrow
evidences.[69][70][71] Wu and Thorne have questioned the pointing to "Outgroup", and her distance
reliability of the molecular clock used to date Eve.[72][73] from any nonafrican groups indicates
that living human mitochondrial lineages
Multiregionalists point out that Mitochondrial DNA alone can not
coalesce in Africa.
rule out interbreeding between early modern and archaic humans,
since archaic human mitochondrial strains from such interbreeding
could have been lost due to genetic drift or a selective
sweep.[74][75] Wolpoff for example states that Eve is "not the most recent common ancestor of all living
people" since "Mitochondrial history is not population history".[76]

Neanderthal mtDNA

Neanderthal mitochondrial DNA (mtDNA) sequences from Feldhofer and Vindija Cave are substantially
different from modern human mtDNA.[77][78][79] Multiregionalists however have discussed the fact that the
average difference between the Feldhofer sequence and living humans is less than that found between
chimpanzee subspecies,[80][81] and therefore that while Neanderthals were different subspecies, they were still
human and part of the same lineage.
Nuclear DNA

Initial analysis of Y chromosome DNA, which like mitochondrial DNA, is inherited from only one parent,
was consistent with a recent African replacement model. However, the mitochondrial and Y chromosome data
could not be explained by the same modern human expansion out of Africa; the Y chromosome expansion
would have involved genetic mixing that retained regionally local mitochondrial lines. In addition, the Y
chromosome data indicated a later expansion back into Africa from Asia, demonstrating that gene flow
between regions was not unidirectional.[82]

An early analysis of 15 noncoding sites on the X chromosome found additional inconsistencies with the recent
African replacement hypothesis. The analysis found a multimodal distribution of coalescence times to the most
recent common ancestor for those sites, contrary to the predictions for recent African replacement; in particular,
there were more coalescence times near 2 million years ago (mya) than expected, suggesting an ancient
population split around the time humans first emerged from Africa as Homo erectus, rather than more recently
as suggested by the mitochondrial data. While most of these X chromosome sites showed greater diversity in
Africa, consistent with African origins, a few of the sites showed greater diversity in Asia rather than Africa.
For four of the 15 gene sites that did show greater diversity in Africa, the sites' varying diversity by region
could not be explained by simple expansion from Africa, as would be required by the recent African
replacement hypothesis.[83]

Later analyses of X chromosome and autosomal DNA continued to find sites with deep coalescence times
inconsistent with a single origin of modern humans,[84][85][86][87][88] diversity patterns inconsistent with a
recent expansion from Africa,[89] or both.[90][91] For example, analyses of a region of RRM2P4
(ribonucleotide reductase M2 subunit pseudogene 4) showed a coalescence time of about 2 Mya, with a clear
root in Asia,[92][93] while the MAPT locus at 17q21.31 is split into two deep genetic lineages, one of which is
common in and largely confined to the present European population, suggesting inheritance from
Neanderthals.[94][95][96][97] In the case of the Microcephalin D allele, evidence for rapid recent expansion
indicated introgression from an archaic population.[98][99][100][101] However, later analysis, including of the
genomes of Neanderthals, did not find the Microcephalin D allele (in the proposed archaic species), nor
evidence that it had introgressed from an archaic lineage as previously suggested.[102][103][104]

In 2001, a DNA study of more than 12,000 men from 163 East Asian regions showed that all of them carry a
mutation that originated in Africa about 35,000 to 89,000 years ago and these "data do not support even a
minimal in situ hominid contribution in the origin of anatomically modern humans in East Asia".[105]

In a 2005 review and analysis of the genetic lineages of 25 chromosomal regions, Alan Templeton found
evidence of more than 34 occurrences of gene flow between Africa and Eurasia. Of these occurrences, 19
were associated with continuous restricted gene exchange through at least 1.46 million years ago; only 5 were
associated with a recent expansion from Africa to Eurasia. Three were associated with the original expansion
of Homo erectus out of Africa around 2 million years ago, 7 with an intermediate expansion out of Africa at a
date consistent with the expansion of Acheulean tool technology, and a few others with other gene flows such
as an expansion out of Eurasia and back into Africa subsequent to the most recent expansion out of Africa.
Templeton rejected a hypothesis of complete recent African replacement with greater than 99% certainty
(p < 10−17 ).[106]

Ancient DNA

Recent analyses of DNA taken directly from Neanderthal specimens indicates that they or their ancestors
contributed to the genome of all humans outside of Africa, indicating there was some degree of interbreeding
with Neanderthals before their replacement.[107] It has also been shown that Denisova hominins contributed to
the DNA of Melanesians and Australians through interbreeding.[108]
By 2006, extraction of DNA directly from some archaic human samples was becoming possible. The earliest
analyses were of Neanderthal DNA, and indicated that the Neanderthal contribution to modern human genetic
diversity was no more than 20%, with a most likely value of 0%.[109] By 2010, however, detailed DNA
sequencing of the Neanderthal specimens from Europe indicated that the contribution was nonzero, with
Neanderthals sharing 1-4% more genetic variants with living non-Africans than with living humans in sub-
Saharan Africa.[110][111] In late 2010, a recently discovered non-Neanderthal archaic human, the Denisova
hominin from south-western Siberia, was found to share 4–6% more of its genome with living Melanesian
humans than with any other living group, supporting admixture between two regions outside of
Africa.[112][113] In August 2011, human leukocyte antigen (HLA) alleles from the archaic Denisovan and
Neanderthal genomes were found to show patterns in the modern human population demonstrating origins
from these non-African populations; the ancestry from these archaic alleles at the HLA-A site was more than
50% for modern Europeans, 70% for Asians, and 95% for Papua New Guineans.[114] Proponents of the
multiregional hypothesis believe the combination of regional continuity inside and outside of Africa and lateral
gene transfer between various regions around the world supports the multiregional hypothesis. However, "Out
of Africa" Theory proponents also explain this with the fact that genetic changes occur on a regional basis
rather than a continental basis, and populations close to each other are likely to share certain specific regional
SNPs while sharing most other genes in common.[115][116] Migration Matrix theory (A=Mt) indicates that
dependent upon the potential contribution of Neanderthal ancestry, we would be able to calculate the
percentage of Neanderthal mtDNA contribution to the human species. As we do not know the specific
migration matrix, we are unable to input the exact data, which would answer these questions irrefutably.[80]

See also
Archaic human admixture with modern humans
Human evolution
Human origins
Mitochondrial Eve
Phyletic gradualism
Recent African origin of modern humans
Y-chromosomal Adam

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Further reading
Biogeography of Middle Pleistocene hominins in mainland Southeast Asia: A review of current
evidence (https://www.academia.edu/189705/Biogeography_of_Middle_Pleistocene_hominins
_in_mainland_Southeast_Asia_A_review_of_current_evidence), Academia.edu, 23 July 2008.
retrieved on 4, April 2014

External links
From www.conrante.com (https://web.archive.org/web/20051218071948/http://www.corante.co
m/loom/img/Templeton%20tree%20600.jpg): Templeton's lattice diagram showing major gene
flows graphically.
From darwin.eeb.uconn.edu (http://darwin.eeb.uconn.edu/eeb348/lecture-notes/mutation-drift/m
utation-drift.html): notes on drift and migration with equations for calculating the effects on allele
frequencies of different populations.
Human Evolution. (2011). In Encyclopædia Britannica. Retrieved from
http://www.britannica.com/EBchecked/topic/275670/human-evolution
Plural Lineages in the Human mtDNA Genome (http://www.rafonda.com/plural_lineages.html)
Human Timeline (Interactive) (http://humanorigins.si.edu/evidence/human-evolution-timeline-int
eractive) – Smithsonian, National Museum of Natural History (August 2016).

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