Language Phylogenies Reveal Expansion Pulses and Pauses in Pacific Settlement

Author(s): R. D. Gray, A. J. Drummond and S. J. Greenhill

Source: Science , Jan. 23, 2009, New Series, Vol. 323, No. 5913 (Jan. 23, 2009), pp. 479-
483
Published by: American Association for the Advancement of Science

Stable URL: http://www.jstor.com/stable/20402911

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 * A A
IL uk EmIL

evolves at a rapid rate (

(25) previous parsimony
sian lexical data found su

Language Phylogenies sequence predicted by t

but limitations of the d
R
Expansion Pulses and meant that the predictio
Pacific settlement could P
n
Lexical data. The Aus
Pacific Settlement around 1200 languages s
family is the one of the la

R. D. Gray,1 A. 1. Drummond,2 S. J. New Greenhill

Zealand and Madag
We have constructed a la
Debates about human prehistory often centerbasic
nesian on the role that
vocabulary
shaping biological and cultural diversity.
210Hypotheses on the
items of basic orig
vocab
the Pacific are divided between a recent "pulse-pause"
guage, expansio
including words for
"slow-boat" diffusion from Wallacea. We used
simple lexical
verbs, data and
colors, and
construct a phylogeny of 400 languages. In is
lary agreement with the
both relatively sta
trees place the Austronesian origin in ally
Taiwan approximately
less likely 5
to be borro
of settlement pauses and expansion pulses
(27). linked
From thisto technologi
database, a
results are robust to assumptions about the rooting
tified the setsand calibra
of homolo
the combined power of linguistic scholarship,
followingdatabase technol
the linguistic co
phylogenetic methods for resolving questions
We extractedabout human pre
the cogn
attested languages for an
fundamental Polynesia
goal of the humancoinciding with the
sciences
comprise developmen
a third of the en
A is to understand the major factors
pre-Polynesian society that
representative
(6, 10), beforesample
a secondo
pansion phase
have shaped the diversity of our into tronesian subgroup.
Eastern Polynesia
species. W
betw
1200 and 1800
At one extreme, innovationist modelsyears B.P.,
Austronesian
argue settling
that Tahiti,
languages
Cook and
advances in technology Islands,
socialTuamotu,
organization
the trees: Marquesas,
an archaicHawvari
Rapanui,
have driven population and New
expansions and Zealand.
shaped
language Chinese that wa
In contrast,
the patterns of cultural and biological and
proponents 2900
diversity years
of the B.P. and
slow-boat sc
(1, 2). At the othernario
extreme, that theBuyang
argue diffusionist/wave (28). These
Austronesians langu
emerged f
models (3) argue thatan innovations and population
extensive sociocultural
ally part network of marit
of the Austrones
exchange
expansions are not critically in Wallacea
linked, andofnewcognates
(in the region
tech have been
of mo id
day Sulawesi
nologies difiuse between and settlement
societies. The the
nate Moluccas) around
sets for all 210 me 13
of the Pacific ocean
to by years B.P. languages
Austronesian
17,000 speakers
based were
on the encoded
dating of m
(hereafter we will chondrial
use the termlineages (11, 12). to
Identified
"Austronesian" This Wallacean sl
"borrowings"
refer to these people)
boat
is one
scenario
of the removed
differs
most remark
from anfrom frither
alternate ana
slow-b
able prehistoricmodel
human that, in agreement
expansions. Thehave
innova with
shown the the
that pulse-p
amo
tionist"pulse-pause" scenario
scenario, posits rowing
postulates that the
an East needs to be very
Asian/Taiwanese
Austronesians originated in 14).
gin (13, Taiwan around
According 5500
to the bias
substantially Wallacean s
either t
years ago and boat
spread scenario,
through the the spread
Pacific in
date aof
se the Austronesia
estimates (30). The res
quence of
expansion pulses by
was driven and settlement
the a total of 34,440
submerging of the charact
Sunda s
at the end
pauses (2, 4-6). According of the
to this whole
last mitochondrial
ice the
scenario, age g
(15). These flo
first pause occurredtriggered
after population
the settlement of expansions
these characters
Taiwan fromwere the
par A
and was followed
by a rapid homeland
tronesian expansionin Language
pulse as
Wallacea tree topology
in a two-pron
expansion.
the Austronesians spread over 7000Onekm of from
thesethe
tions prongs
about the moved
origin, no
s
Philippines to Polynesiathroughin lessthe Philippines
than 1200
the years.and into Taiwan.
Austronesian expansio
As the Austronesians second
spreadexpansion
through using
prong Bayesian
thesespread
re eastphyloge
along
gions, they integrated New Guinea
with coast
existing and into
populations
number ofOceania
models of andcogP
and innovated newnesia technologies,
(following including
same routethe
the best-performing described
model for
ha
Lapita cultural complex (5). The
pulse-pause archaeological
scenario). The pulse-pause
cognate and sl
gains and losses
evidence suggests the boat Austronesians
scenarios differ reached
specific therate variation
substantially in whereus th
previously uninhabited locate the of
islands where
Austronesian
the Reefs/Santa characters
homeland, incould
the exs
Cruz around 3000 to sion3200 sequence
years beforethey postulate,
slow
the rates at
present and in the age
different br
(B.P.) (7), New Caledonia,
timing and of thisVanuatu around
Early
expansion. attempts to estim
Genetic studies o
3000 years B.P., and cificTonga, Samoa(13,
settlement and
guage Fijirelationships
16-18) inhave been hampe using
Western Polynesia by in problems
the period in between
ods (32)
separating 2900produced
ancient fromtrees
rec
to 3200 years B.P. (8, 9). This (19)
admixture initial
andrapid pulse from
different
difficulties those obt
in precisely da
was followed by athe second pause in the
mitochondrial Western
and Y chromosome
comparative methodha
groups found in the Bayesian
Pacific (20,phylogenetic
21). t
We used phylogenetic analyses
we obtained of languag
from our bas
'Department of Psychology, University of Auckland, Private
to trace the history congruent with the trad
of human populations bec
Bag 92019, Auckland 1142, New Zealand. 2Department of
language
Computer Science, University of Auckland, fied
is linkedPrivate
to by
other
Bag phonological
cultural traits and (m
92019, Auckland 1142,contains
New Zealand.large amounts such ofas information
the loss of the Pr
(23),

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I RESEARCH ARTICLE
in the Central Pacific subgroup (34) or the low change (East Formosan). The remaining five lowing these are the languages of the Philippines,
ering of high vowels in morphemes identifying (Westem Oceanic, Malayo-Chamic, Greater Cen Bomeo/Sulawesi, Central Malayo-Polynesia, South
Central-Eastem Malayo-Polynesian (35). The trees tral Philippines, Greater Barito, and Barrier Halmahera/West New Guinea, and the Oce
support 26 of the 34 putative Austronesian lan Islands/North Sumatra) may be obscured in our anic languages. This chained topology is pre
guage subgroups and linkages discussed in (28). analyses because of conflicting signals caused cisely the structure predicted by the pulse-pause
Of the remaining seven unsupported groups, two by undetected borrowing between neighbor scenario.
are linkages that lack exclusively shared innova ing languages. Our results place the Formosan One potential problem with this analysis is
tions (Central and Western Malayo-Polynesian), languages of Taiwan at the base of the trees that Old Chinese may be too distantly related to
and one is only supported by a single sound immediately after the outgroups (Fig. 1). Fol Austronesian to reliably root the tree, whereas

~Japan < North America

South-East Asia

* P Sa~w au I Hawaii

Micronesia

New Guinea

Polynesia
-o..Madagascar V~u !
New nia
Australia Rapanui

New Zealand

7000

6000

l '00 ause 1
- IPulse 1
a 4000

&_ 300
0 ~~~~~~Pulse 3
co

>_ ~~~~~~~~~~~~~~~~~~~~~~~~Pa
2000
4Pulse 2

0
a cd ef a hil ait4 A B C
Fig. 1. Map and maximum cLade j,
Sangiric; credibility tree
Celebic; k, of
Bima
languages. The Central
tree shows Maluku;
four major n, Timor;
expansion o
pulses
Schouten
Pacific settlement. Branches colored(North
red areNew
thoseGuin
ide
significant increases in Melanesian);
language diversification
s, North New Gurat
subgroups are color-codedv, and
Meso-MeLanesian; w, Temo
labeled as follows: a, O
Old Chinese); b, Formosan; c, Sama-Bajaw;
Loyalties/New d, Goront
Caledonia; A,
e, Philippine; f, Barito; g, Polynesian.
Malayo-Sumbawan; h, Greater

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 RESEAR
Buyang and the other Tai-Kadai languages may stochastic-Dollo model
the base of of co
the t
be a sister group to Malayo-Polynesian (29). To languages
(28), which can estimatewith 10
wher
investigate the reliability of the Taiwanese root Age of Austrone
be without specifying outgrou
ing, we conducted a separate analysis using a ence
tional analysis between
placed the
Old Chine
of the Austrones
pause scenario pre
between 5000 to
1400 -
Wallacean slow b
age of between 1
test between thes
1200 - age of Proto-Aus
likelihood rate-sm
than assuming co
(n 1000 - ment, this metho
the observed rates
trees. We calibrat
archaeological da
800 - ment times (28).
E Old Javanese wer
z they were spoken
& 600 calibrated their to
languages were tr
The divergence t
S 400 - the Austronesian
pulse-pause scena
age of Austrones
trees has a mean
200 - posterior density
years B.P.). The d
robust across a r
0 ent models (28). In
age of Austrones
0 2 4 6 8 10 12 14 16 18 20 Polynesian constr
estimated age (m
Age (Thousands of years Before Present)
4730 to 5790 yea
the impact
Fig. 2. Histogram of the estimated age of the Austronesian of diff
language family. The l
the (5000
shows the age range predicted by the pulse-pause scenario agetoof 6000 Proto-
years B.P.),
bar shows that predicted by the slow-boat scenarioClade
(13,000 toCredibility
17,000 years B.P.).
combinations. Th
age of Proto-Aus
years (28).
A 20 Pause 1: Proto-Malayo-Polynesian Our estimates for the age of the Austronesian
80 expansion are considerably younger than the
40 deep age estimates of the slow-boat scenario
(11, 12, 15). One possibility is that these deep
estimates are artifacts due to problems with ac
curately dating genetic change. There is increas
B ing evidence that rates of genetic change estimated
1 120 Pause 2; pre-Polynesian
over thousands of years are substantially higher
0 80 than the long-term substitution rate (21). This

40 violation of the molecular clock leads to the sys

tematic overestimation of recent divergence times.
The difficulties of obtaining accurate molecular
01 Overall dates are probably compounded by the use of the
error-prone rho dating method (20), especially
15000 j when it is applied to sequences of high-rate hetero
geneity such as hypervariable region 1 (HVR-l).
10000
Another possibility is that the genes and lan
5000 guages have quite different histories. However,
0j both Austronesian expansion scenarios envisage
a ooupling between genetic and linguistic histo
0 0.5 1.o 1.5 2.0 ries. In the pulse-pause scenario, the considerable
Branch length (Thousands of
diversity of Fonnosan languages reflects the Tai yea

Fig. 3. Histograms of the wanese origin of Austronesian.

branch length In contrast, distribu
the
pause, (B) the distribution Wallacean
of the slow-boat pre-PoLynesian
scenario argues that Taiwan

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I RESEARCH ARTICLE
is an "Austronesian backwater" (12) and that the branch in the tree, we used an indicator variable to pause in Western Polynesia is that the final pulse
initial diversity of Malayo-Polynesian languages model whether the rate of language diversifica into the far-flung islands of Eastern Polynesia
has been obscured by language-leveling as a tion changed below the branch and a relative rate required further technological advances. These
result of extensive socioeconomic networks (11). variable to specify the new rate of diversification might have included the ability to estimate lat
Recent genetic studies of complete mitochondrial relative to the rate on the parent branch. If no itude from the stars, the ability to sail across the
sequences (18) and genome-wide autosomal change is indicated on a given branch, the diversi prevailing easterly tradewinds, and the use of
markers (14, 16) also show that, despite consid fication rate of the parent language is inherited. double-hulled canoes with greater stability and
erable admixture in Near Oceania (38), there is a We employed a full language-diversification carrying capacity (4, 42). Alternatively, the vast
clear signature linking Austronesian speakers from model in which the number, phylogenetic loca distances between these islands might have re
Taiwan to Polynesia. Even at a fine geographic tion, and magnitude of the changes in diversifi quired the development of new social strategies
scale on the east Indonesian island of Sumba, cation rate were all estimated directly from the data for dealing with the greater isolation found in
there are strong correlations between languages, by using Bayesian stochastic variable selection Eastern Polynesia (42). These technological and
genes, and geography (39). The Austronesian within a Markov chain Monte Carlo method (28). social advances in Eastern Polynesia may also
expansion has therefore produced a close initial The posterior estimate of the number of underlie the fourth pulse into Micronesia.
coupling between genes and languages that has changes in diversification was 4.3 (95% credible The language phylogenies reveal the rapidity
subsequently broken down in some regions such set: 1 to 7) with a total of 10 branches showing of major cultural development in the Pacific. As
as Near Oceania (38). strong evidence of changes [Bayes Factor (BF) > the Austronesians spread along New Guinea and
Pulses and pauses. If language diversification 20] in diversification rates (Fig. 2 and fig. S4). into the Solomons, they developed the Lapita
(cladogenesis) is linked to population expan The pulse with the highest posterior probability cultural complex through interaction with the
sions, then expansion pulses should leave a series occurred in three of the branches leading to existing populations in Near Oceania (5, 10).
of short branches in the phylogenies because Proto-Malayo-Polynesian (BF = 397, 79, and 33, This complex includes distinctive and often elab
there will be little time for linguistic changes to respectively). The second identified pulse oc orately decorated pottery, adzes/axes, shell or
accumulate before speech communities fragment. curred in two of the branches after the pre naments, tattooing, and bark-cloth (10). The
In contrast, when the geographic spread of cul Polynesian stage (BF = 29 and 36). The location phylogenies show that there was only a very
tures is constrained by physical or social bound of these two pulses is in agreement with those small time-window for this complex to develop.
aries, the rate of linguistic diversification should predicted by the pulse-pause scenario. Changes Based on the age of the Eastern Malayo
decrease leading to longer branches (anagenesis). in diversification rate were also evident in three Polynesian clade the Austronesians entered the
The pulse-pause scenario predicts the existence of other locations. The third pulse was found in the South Halmahera/West New Guinea region at
two settlement pauses: the first occurring before branch leading to the Philippines languages around 3680 years B.P. (95% HPD, 3640 to 3,710
the settlement of the Philippines and correspond (BF = 38) after another lengthy pause. Our results years B.P.), and had reached Remote Oceania by
ing with the development of the Proto-Malayo place the age of this pulse around 2500 years B.P. 3575 years B.P. (95% HPD, 3560 to 3590 years
Polynesian language around 3800 to 4500 years This is consistent with arguments that the Greater B.P.). The high levels of male-biased admixture
B.P. (4, 6), and the second occurring after the Central Philippines subgroup expanded at the detected in Polynesian genetic studies (13, 14)
settlement of Western Polynesia by 2800 years expense of other lineages between 2000 and must either have occurred over this very short
B.P., before the expansion into Central and Mar 2500 years B.P., reducing linguistic diversity in time span (approximately four generations), with
ginal Eastern Polynesia (4, 6, 10). This Western the region (40). A fourth pulse was evident in Melanesian males actively incorporated into the
Polynesian settlement coincides with the devel three of the branches leading to the Micronesian Austronesian expansion, or there was extended
opment of the temporally brief Proto-Central languages (BF = 66, 29, and 23). Within this post-settlement contact between Near Oceania
Pacific dialect network in Fiji, Tonga, and Samoa Micronesian group the Trakic languages contain and Polynesia. The results presented here show
(10), with the Polynesian languages emerging the fastest (single-population) rates in the entire the combined power of Bayesian phylogenetic
from the eastern part of this dialect network some family. The final branch to show a significant methods and large lexical databases to resolve
time later. This second pause is therefore harder increase in diversification rates is that leading to questions about human prehistory. Just as molec
to place cleanly on a tree, but should correlate both the Micronesian and Polynesian subgroups ular phylogenies provide the fundamental frame
with the development of a pre-Polynesian stage. and suggests that there might be a common under work for studies of biological evolution, language
To test for the predicted signature of settle lying factor between the subsequent pulses into phylogenies open up the exciting possibility of a
ment pauses, we extracted all the internal branch Polynesia and Micronesia (Fig. 1 and fig. S4). Darwinian approach to cultural evolution based
lengths from the posterior distribution of the Discussion. Our results show that the diversi on rigorous phylogenetic methods (43).
dated trees. We compared the branches corre fication of Austronesian languages was closely
sponding to the Proto-Malayo-Polynesian and coupled with geographic expansions. The avail References and Notes
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An Entanglement Filter
Ryo Okamoto,l 2* Jeremy L. O'Brien,3* Holger F. Hofmann,4 Tomohisa Nagata,12
Keiji Sasaki,1 Shigeki Takeuchi1'2t
The ability to filter quantum states is a key capability in quantum information science and technology,
in which one-qubit filters, or polarizers, have found wide application. Filtering on the basis of
entanglement requires extension to multi-qubit filters with qubit-qubit interactions. We demonstrated
an optical entanglement filter that passes a pair of photons if they have the desired correlations of their
polarization. Such devices have many important applications to quantum technologies.
Filters, which pass the desired and reject the
unwanted (material, signal, frequency, or
polarization, for example), are one of the
most important scientific and technological tools
available to us. Quantum information science and
technology is concerned with harnessing quan
tum mechanical effects to gain exponential im
provement of and new functionality for particular
tasks in communication (1), computation (2), mea
surement (3), and lithography (4). Perhaps the
most distinctive of these quantum mechanical
features is entanglement. Filters that act on the
'Research Institute for Electronic Science, Hokkaido University,
Sapporo 060-0812, Japan. 2The Institute of Scientific and
Industrial Research, Osaka University, Mihogaoka 8-1, Ibaraki,
Osaka 567-0047, Japan. 'Centre for Quantum Photonics, H. H.
Wills Physics Laboratory and Department of Electrical and
Electronic Engineering, University of Bristol, Merchant Ven
turers Building, Woodland Road, Bristol BS8 1UB, UK.
4Graduate School of Advanced Sciences of Matter, Hiroshima
University, Hiroshima 739-8530, Japan.
*These authors contributed equally to this work.
tTo whom correspondence should be addressed. E-mail:
takeuchi~es.hokudai.ac.jp
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quantum correlations associated with entangle
ment must operate nonlocally on multiple quan
tum systems, typically two-level qubits. Such a
device has been proposed for photonic qubits (5);
however, the technical requirements to build such
a device, an optical circuit with two ancillary
Fig. 1. The function of a
polarization filter and an
AW o o W
B~~~~~ _, I_
entanglement filter. (A)
Polarization filters pass
only the certain polariza
tion component of single
photons. (B) Entangle
ment filters pass a pair
of photons only if they
share the same horizontal
or vertical polarization.
(C) Because the quantum
coherence between these
two possibilities is pre
served during the pro
cess, the output state is
entangled when two diagonally polarized photons are input.
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41. A. Pawley, M. Pawley, in Austronesian Terminologies:
Continuity and Change, A. Pawley, M. Ross, Eds.
(Australian National University, Canberra, 1994),
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42. G. Irwin, ;. Polyn. Soc. 107, 111 (1998).
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(2007).
44. We thank R. Blust, ]. Lynch, ]. Marck, C. Nickelsen,
M. Ross, L. Sagart, and all contributors to the Austronesian
Basic Vocabulary Database for their assistance with lexical
data and cognate judgments. We gratefully acknowledge
BeSTGRID (www.bestgrid.org) for Web hosting and
A. Meade, M. Pagel, and the Centre for Advanced
Computing and Emerging Technologies at the University of
Reading for computational support. R. Blust, M. Corballis,
R. Green, F. Jordan, M. Pagel, R. Ross, and M. Suchard
commented on the manuscript. Funding was provided by
the Royal Society of New Zealand Marsden Fund and a
Bright Futures doctoral scholarship to S.G.
Supporting Online Material
www.sciencemag.org/cgi/content/futll323/5913/479/DC1
Materials and Methods
SOM Text
Figs. S1 to S5
Tables S1 to S3
References
6 October 2008; accepted 10 December 2008
10.1126/science.1166858
photons and multiple quantum gates, requiring
both quantum interference and classical interfer
ence in several nested interferomters, have been
lacking.
We demonstrate an entanglement filter made
by combining two key recent technological ap
proaches: a displaced-Sagnac architecture (6) and
partially polarizing beam splitters (PPBSs) (7-9).
The entangling capability of the filter was veri
fied, distinguishing it from classical ones. Be
cause our entanglement filter acts on photonic
qubits, it is promising for quantum technologies;
photons are the logical choice for communication
(1, 10), metrology (6, 11), and lithography (12)
and are a leading approach to information pro
cessing (13). The filter can be used for the cre
ation as well as the purification of entanglement
(14, 15), which will be important in realizing quan