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Africa to Aotearoa: the longest migration

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Africa to Aotearoa: the longest

migration
a b
E Matisoo-Smith & C Daugherty
a
Department of Anatomy and Allan Wilson Centre for Molecular
Ecology and Evolution, University of Otago, Dunedin, New Zealand
b
School of Biological Sciences and Allan Wilson Centre for
Molecular Ecology and Evolution, Victoria University of
Wellington, Wellington, New Zealand

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 Journal of the Royal Society of New Zealand
Vol. 42, No. 2, June 2012, 87!92

Africa to Aotearoa: the longest migration

E Matisoo-Smitha* and C Daughertyb
a
Department of Anatomy and Allan Wilson Centre for Molecular Ecology and Evolution, University of Otago,
Dunedin, New Zealand; bSchool of Biological Sciences and Allan Wilson Centre for Molecular Ecology and
Evolution, Victoria University of Wellington, Wellington, New Zealand
(Received 24 September 2011; final version received 2 March 2012)

Modern human origins have been securely located in Africa. Human migrations out of Africa
began approximately 65,000 years ago and ended with the settlement of Aotearoa/New Zealand
Downloaded by [E Matisoo-Smith] at 08:45 28 May 2012

only about 750 years ago. Researchers have used mitochondrial DNA and other molecular
markers to track these migration pathways. This paper describes the archaeological, historic and
mitochondrial DNA evidence tracking the history of the longest human migration, that from
Africa to Aotearoa.
Keywords: molecular anthropology; Polynesia; mitochondrial DNA; origins; Lapita;
New Zealand

Introduction these early recognised clues, many scientists

The primary goal of the first voyage of Captain
James Cook was to observe the transit of Venus
in Tahiti and from there to search for the great
southern continent Terra Australis. Another
major goal for Cook as well as Joseph Banks
and Daniel Solander, the naturalists who ac-
companied him, was to observe and describe
the plants, animals and peoples that they en-
countered in these new lands. Over the course
of his three Pacific voyages, Cook, like many
explorers both before and after, was struck by
the cultural, linguistic and physical similarities
of Polynesian peoples. JR Forster, who tra-
velled with Cook as the naturalist on his second
voyage, correctly recognised the shared char-
acters of the Malayo-Polynesian languages
(now a recognised sub-group of the Austrone-
sian languages*see Gray et al. 2009) and
commented on the likely origins of Polynesians
and on the probable short settlement history of
the islands (Forster et al. 1996 [1778]). Despite
who followed Cook and Forster looked else-
where to identify the origins of the Polynesians,
including suggestions of Aryan and American
origins (Heyerdahl 1952; Tregear 1984 [1885]).
The so-called Polynesian problem was the
focus of much research over the following 200
years (Handy 1920; Groube 1971).
Linguistic and archaeological research con-
firms that Polynesian origins are linked most
closely to the Lapita culture (Kirch & Green
2001). Lapita sites have now been identified
from the Bismarck Archipelago, off the north
coast of New Guinea, as far east as Samoa and
Tonga. The earliest Lapita sites date to ap-
proximately 3350 years before present (yr BP)
(Kirch 2001).
The Lapita people, however, were not the
first to arrive in the Pacific. Archaeological
evidence for the initial human occupation of
New Guinea dates to approximately 50,000 yr
BP (Summerhayes et al. 2010), some of the
earliest archaeological dates associated with

*Corresponding author. Email: matisoo-smith@otago.ac.nz

ISSN 0303-6758 print/ISSN 1175-8899 online
# 2012 The Royal Society of New Zealand
http://dx.doi.org/10.1080/03036758.2012.673495
http://www.tandfonline.com
 88 E Matisoo-Smith and C Daugherty
modern humans outside Africa. This initial
phase of human occupation of the Pacific
included the settlement of New Guinea and
Australia. The earliest sites on islands of the
Bismarck Archipelago date to about 40,000 yr
BP and those of the main Solomon Islands to
29,000 yr BP. This region of earliest occupation
has been called Near Oceania (Pawley & Green
1973). The islands of the rest of the Pacific,
which they defined as Remote Oceania, have a
much shorter settlement history, and much of
the region was first settled by the Lapita people.
Within a few hundred years of the first
appearance of Lapita sites in the Bismarck
Archipelago, the Lapita peoples breached the
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barrier between Near and Remote Oceania,
arriving in Vanuatu, New Caledonia and Fiji
around 3000 yr BP, Tonga by 2900 yr BP and
Samoa by 2700 yr BP. Lapita expansion
stopped there, on the western edge of the Poly-
nesian Triangle, for a period of at least 1000
years. During this period, in Samoa and Tonga,
the Ancestral Polynesian language, culture
and biology were established (Kirch & Green
2001). Further expansion eastward into central
eastern Polynesia began around 1500 yr BP.
Settlement of the Cook, Society and Marquesas
Islands occurred and then, from this central
Polynesian homeland, voyagers discovered and
eventually settled the extremes of the triangle
reaching Hawaii, Rapa Nui/Easter Island and,
finally, Aotearoa/New Zealand between 1000
and 700 yr BP (Kirch & Kahn 2007).
Despite the archaeological and linguistic
evidence for Polynesian origins and settlement,
questions remain regarding the settlement of
East Polynesia and the biological origins of
Polynesian ancestors: what was the genetic
makeup of the Lapita peoples in Samoa and
Tonga, and where were their biological origins?
These are the same questions asked by the
earliest European explorers, yet, over the years,
biologists and anthropologists could still not
answer those using traditional morphological
and blood group analyses.
In the mid 1980s molecular anthropology
was emerging as a discipline. At the centre of
this development were New Zealander Allan
Wilson and his colleagues at the University of
California, Berkeley. Wilson and his students
had recognised the value of a new genetic
marker, mitochondrial DNA (mtDNA), and
were using it for studying population histories.
Unlike nuclear DNA, which is inherited
from both parents and recombines to form
a unique, mixed DNA sequence in the off-
spring, mtDNA is passed down only through
the maternal line. Offspring inherit an exact
copy of their mothers mtDNA. This mtDNA
signature is called a haplotype. Occasionally,
every 100 generations or so, a mutation, or
random change similar to a spelling mistake,
occurs in the mtDNA in a females egg, and this
mistake is then passed on to her descendants.
Each time a mutation occurs and is passed
on, it represents a new branch on the mtDNA
family tree. Major branches on the mtDNA
tree share common ancestral mutations and
are called haplogroups; each major haplogroup
has been assigned a letter in the alphabet.
Haplogroups can be further divided as new mu-
tations occur, and each of the smaller branches
within a haplogroup is given a series of num-
bers and then letters which identify its location
in the mtDNA tree. Thus, for example, in the
human mtDNA phylogeny or evolutionary
tree (Fig. 1) haplogroup B is a lineage that
can be traced back to the R, then the N, and
finally the L3 branch.
Rebecca Cann, Mark Stoneking and Allan
Wilson (1987) published their landmark analy-
sis of mtDNA obtained from 147 women of
European, African, Asian, Australian Aborigi-
nal and Papuan ancestry, providing exciting
and controversial results about human origins
and migrations. They concluded that all hu-
man mtDNA lineages originated from a single
mtDNA haplotype belonging to a woman who
lived approximately 200,000 years ago in Africa.
This maternal ancestor to all living human
populations was quickly dubbed Mitochondrial
Eve. Despite initial academic critiques which
focused primarily on the computer methods
used to reconstruct the phylogenies or family

Figure 1 A simplified human MtDNA phylogeny.
trees (Darlu & Tassy 1987), subsequent analyses
have confirmed that the results of Cann et al.
were remarkably robust. As more populations
have been added to the mtDNA database, the
branches have been pruned, reassigned and fine-
tuned, and the dates for the common ancestor
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and for the migrations out of Africa have been
regularly reassessed, but the general conclusions
stand.
All extant human mtDNA lineages can now
be reliably traced back to Africa. It appears
that at some point around 65,000 yr BP, people
carrying one of the major African lineages,
identified as L3, moved out of Africa. Some-
where in what is now the Middle East, the
L3 lineage split into two branches designated
M and N in the mtDNA tree (see Fig. 2). The
M lineages were carried east, along the south-
Figure 2 Map of human dispersals based on mtDNA with a focus on those lineages that arrived in the
Pacific. Dotted line delineates Near and Remote Oceania. Solid lines indicate the Polynesian Triangle.
Africa to Aotearoa: the longest migration 89
ern coast of Asia (Macaulay et al. 2005).
M lineages and all sub-branches are found
exclusively in Asian and Asian-derived popula-
tions (Friedlaender et al. 2007). Meanwhile,
people carrying the N lineages moved north
and northeast into Europe and Asia (Mellars
2006). Most European populations today have
mtDNA lineages that belong to the larger
N branch of the tree, in particular, those that
derive from the R branch.
M lineages were carried through southern
Asia, into East Asia and what is now Island
Southeast Asia until the people encountered
the water barrier separating the Asian from the
Australian continent. Eventually, these people
managed to cross that barrier*a voyage
requiring open water crossings of at least
70 km. Given the archaeological evidence
 90 E Matisoo-Smith and C Daugherty
from New Guinea, we know that people
reached Sahul (the greater continent created
when lowered sea levels joined New Guinea
and Tasmania to the Australian mainland) by
50,000 yr BP. At some point prior to or around
that time, the people carrying the Asian N
lineages joined the populations carrying the
M lineages and made it to Sahul. Here, new
mtDNA lineages evolved, including hap-
logroup Q, and several new subtypes of hap-
logroup M, all of which are found only in Near
Oceania, sometimes only in specific regions
within Near Oceania. Similarly, new N-derived
lineages emerged, including haplogroup P
(Friedlaender et al. 2007) and S, which is found
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only in Australia (McEvoy et al. 2010).
It is not clear how many ancient migration
events occurred into Sahul and Near Oceania,
but the populations that did arrive appear
to have dispersed quickly across the land-
scape. Archaeological sites dating to 40,000!
45,000 yr BP are found across Australia, and
in New Guinea and the Bismarck Archipelago.
These populations were probably relatively
isolated, and further migrations into the region,
it seems, did not occur until much later. The
mtDNA evidence of unique lineages, not
shared with other populations, is indicative of
a long period with little contact with outside
regions as well as relative isolation of popula-
tions within Near Oceania.
The unsettled climate and resulting envir-
onmental changes associated with the end of
the Pleistocene and beginning of the Holocene
period, around 10,000 yr BP, ceased by about
6000 yr BP. New and rich coastal environments
are likely to have opened up and better, more
stable weather conditions prevailed, providing
ideal conditions for population mobility and
interaction between Near Oceania and Island
Southeast Asia (Terrell 2004). Around this
time, new populations moved into Near Ocea-
nia from East Asia. Lineages belonging to
haplogroup B4 arrived in parts of Near Ocea-
nia during the mid-Holocene period. The
distribution of the haplotype B4a1a and de-
rived lineages (e.g. B4a1a1a) has been widely
associated with the spread of the Lapita
cultural complex through Near Oceania, into
Remote Oceania, ultimately into Polynesia.
The timing and distribution of the mtDNA
B4a1a lineages may be associated with the
spread of Austronesian languages brought
about by the Neolithic population expansion
following the development of rice agriculture
in Asia (Bellwood 2005). Interestingly, there
is currently no evidence of B4 lineages in
Australian Aboriginal populations.
Polynesians, and East Polynesian popula-
tions in particular, have high frequencies of
mtDNA haplotype B4a1a1a, which has been
dubbed the Polynesian motif (Melton et al.
1995). This label, however, is misleading as the
haplotype is found in many non-Polynesian
populations, including some as far away
as Madagascar (Razafindrazaka et al. 2010).
Some communities in Near Oceania have fre-
quencies over 80%, and many in New Ireland
and Bougainville have frequencies over 50%
(Friedlaender et al. 2007). Soares et al. (2011)
suggest that the Polynesian motif actually
originated in the Bismarck Archipelago, and
that it occurred much earlier than the earliest
Lapita sites and prior to the Neolithic expansion
out of Asia (Soares et al. 2011).
The B4a1a1a haplotype is not the only
mtDNA lineage found in Polynesian popula-
tions. Low frequencies of one of the Near
Oceanic lineages, haplotype Q1, have been
identified in Samoa, the Cook Islands, Tahiti,
Mangareva and in New Zealand Maori popu-
lations (Friedlaender et al. 2007; Deguilloux
et al. 2011). The limited amount of mtDNA
variation in Polynesia has been explained as
the result of multiple bottleneck events that
occurred during the long ocean voyages and
relative isolation of Polynesian populations
(Hurles et al. 1998). Analyses of ancient DNA
in East Polynesian skeletal remains, however,
have indicated much higher frequencies of
Q1 in at least some populations in the past
(Deguilloux et al. 2011).
So, while the Polynesian mtDNA haplo-
types belonging to the B4a1a1 lineages can

ultimately be traced back to Southeast Asia,
Polynesian origins lie in both Asia and Near
Oceania. Clearly, admixture was taking place
between more recent Asian-derived populations
and the indigenous peoples of Near Oceania
before Lapita peoples moved out into Remote
Oceania. The mtDNA evidence suggests this
admixture was relatively limited, but analyses
of DNA variation in the male counterpart to
mtDNA, the paternally inherited Y chromo-
some, suggest that a significant percentage of
males carrying indigenous Near Oceanic genet-
ic markers were part of the Lapita expansion.
The pattern of Y chromosome diversity in
Remote Oceania is the opposite of the mtDNA
Downloaded by [E Matisoo-Smith] at 08:45 28 May 2012
data, with a larger percentage of Near Oceanic
as opposed to Asian-derived lineages (Kayser
2010).
Human migration is an on-going process,
with numerous subsequent arrivals adding to
the cultural and biological complexity of a
population after the initial colonising event.
Y-chromosome studies in most Polynesian
populations show that the arrival of the first
European sailors in the Pacific altered the
genetic makeup of populations in the region.
European Y chromosomes occur in high fre-
quency throughout Polynesia, which no doubt
is the legacy of early explorers, whalers and
traders and later colonial settlers of the Pacific
(Hurles et al. 1998). European men were joined
in New Zealand and throughout Polynesia
by Chinese miners, traders and agricultural
workers in the 1860s. In New Zealand this
was followed by the arrival of the Dalmatian
gum diggers in the 1880s. Again, these migrants
tended to be mostly, if not exclusively, men and
they often married local Maori women, con-
tributing significantly to the Y-chromosome
diversity of Aotearoa.
English settlers constituted the bulk of
migrants and settlers from the 1840s through
1914. After the World Wars, refugees and
immigrants from Europe arrived, further en-
hancing the genetic diversity of Aotearoa. This
diversity was supplemented by the Pasifika
migrations of the 1950s and 60s. People still
Africa to Aotearoa: the longest migration 91
arrive in Aotearoa, from countries all over the
world. These new immigrants have taken many
different routes and bring many different
migration stories. Ultimately, however, all
people of Aotearoa share a single shared
maternal ancestor. We have taken many differ-
ent routes, but we, through our ancestors,
all participated in the longest journey*from
Africa to Aotearoa.
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