BREVIA

Two external fossil calibrations were used

EVOLUTION in separate ML molecular clock calculations

Flight of the Dodo

after tests found no violation of clocklike
behavior (3). The analyses indicate that the
dodo/solitaire and Caloenas diverged in the
Beth Shapiro,1,2 Dean Sibthorpe,1 Andrew Rambaut,1 mid/late Eocene, around 42.6 million years
Jeremy Austin,3 Graham M. Wragg,1 Olaf R. P. Bininda-Emonds,1 ago (Ma) (95% confidence interval ⫽ 31.9 to
Patricia L. M. Lee,1 Alan Cooper1,2* 56.1 Ma), whereas the dodo and the solitaire
separated in the late Oligocene, about 25.6
The evolutionary history of the dodo is very tris (originally named for its dodolike beak) is Ma (17.6 to 35.9 Ma). The latter date is
poorly understood. Like many avian island en- the basal member of this strongly supported biogeographically interesting as it is consid-
demics, a high degree of morphological change clade of large, generally ground-dwelling, is- erably older than the islands of Mauritius and
associated with flightlessness and gigantism has land endemics. Furthermore, the phylogeo- Rodrigues. Geological evidence suggests that
obscured phylogenetic relationships, and histor- graphic distribution of this morphologically di- Mauritius emerged in a series of volcanic
ically the dodo has been linked with avian verse clade suggests that the dodo and the sol- events, the earliest of which occurred around
groups ranging from the ratites to 6.8 to 7.8 Ma, whereas Rodrigues
the raptors (1). Since the mid- did not emerge until 1.5 Ma (5).
1800s, morphological studies have Therefore, it seems highly unlike-
linked the dodo (initially Didus in- ly that the large genetic distance
eptus, now Raphus cucullatus) and between the dodo and the solitaire
its presumed close relative the sol- resulted from isolation on the two
itaire (Pezophaps solitaria) with islands. Drilling projects have es-
the Columbiformes ( pigeons and tablished that ridges surrounding
doves), but their exact position is the Mascarene Plateau were
unresolved and they have been above sea level in the late Oligo-
placed in many positions within the cene and have subsided slowly
cosmopolitan Columbidae or in thereafter (5). The similarity be-
their own family, Raphidae, out- tween the timing of the dodo/sol-
side the Columbidae but within itaire divergence and the first
Columbiformes (1, 2). To answer geological evidence of land in the
these questions, we extracted DNA Mascarene island chain is striking
and amplified 1.4 kb of mitochon- and suggests that island stepping-
drial sequence [360 – base pair (bp) stones may have been used before
12S rRNA and 1050-bp cyto- the two species eventually found
chrome b] from 37 species of pi- their way to Mauritius and Ro-
geons and doves, including the drigues. Whether such dispersal
dodo, the solitaire, and multiple was flighted or not cannot cur-
members of each of the five extant rently be determined, although
subfamilies (3). Short overlapping the relatively isolated position
fragments were amplified by poly- and young age of Rodrigues
merase chain reaction (PCR), with would suggest that the solitaire, at
museum specimens limited to 120 least, may have retained a capac-
to 180 bp. Strict ancient DNA pro- ity for flight up until at least 1.5
cedures were used to minimize Ma.
contamination and detect nuclear
mitochondrial copies (4), PCR References and Notes
1. A. Janoo, Bull. Mus. Natl. Hist. Nat. 18,
products were cloned, and the sol- 57 (1996).
itaire sequences were independent- 2. D. Gibbs, E. Barnes, J. Cox, Pigeons and
ly replicated at the Natural History Doves: A Guide to the Pigeons and
Fig. 1. ML tree depicting the phylogenetic relationships of 37 species of Doves of the World (Pica Press, Sussex,
Museum, London. UK, 2001).
The maximum likelihood (ML) pigeons and doves. Geographic distributions of the pigeons are as follows: 3. Detailed information is available on
Australian (red), African/Madagascan (green), Asian/Southeast Asian ( pur-
tree for the combined data set (Fig. ple), Mascarene Island (blue), and New World (yellow) taxa. The concentra- Science Online at www.sciencemag.
1) reveals that the dodo and the tion of New World taxa at the base of the tree suggests that the initial org/ cgi/ content/ full/ 295/ 5560/
1683/DC1.
solitaire are sister taxa. Despite the radiation occurred in the New World. ML bootstrap consensus values for 4. A. Cooper, H. Poinar, Science 289,
substantial morphological differ- relationships with ⱖ70% support (italicized) and Bayesian posterior support 1139 (2000).
ences between the dodo, the soli- values (bold) are given. Detailed phylogenetic analyses of potential out- 5. I. McDougall, F. H. Chamalaun, Bull.
Geol. Soc. Am. 80, 1419 (1969).
taire, and other pigeons, the ML groups identified Laridae, and consequently two species (Great Skua, Ca-
tharacta skua, and Parasitic Jaeger, Stercorarius parasiticus) were used (3).
analysis shows them to be nested
within the family Columbidae. The
closest living relative is the monotypic Nicobar itaire dispersed from Southeast Asia to the Mas- 1
Department of Zoology, University of Oxford, Ox-
pigeon (Caloenas nicobarica) from the Nicobar carenes at some point in the past. Figure 1 ford, OX1 3PS, UK. 2Henry Wellcome Ancient Biomol-
Islands and nearby Southeast Asia, and the indicates that a similar route was used by the ecules Centre, University of Oxford, Oxford, OX1 3PS,
UK. 3Department of Zoology, The Natural History
sister taxa to this clade are the crowned pigeons Madagascan endemic Alectroenas madagas- Museum, London SW7 5BD, UK.
(Goura), from New Guinea. The unusual Sa- cariensis, which is closely related to another *To whom correspondence should be addressed. E-mail:
moan tooth-billed pigeon Didunculus strigiros- Indo-Australian radiation. alan.cooper@zoology.oxford.ac.uk

www.sciencemag.org SCIENCE VOL 295 1 MARCH 2002 1683