Biological Journal of the Linnean Society, 2003, 80, 699–715. With 6 figures
© 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 699–715 699
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Original Article
DIVERSIFICATION OF PHILIPPINE RODENTS
S. J. STEPPAN
ET AL
.
*Corresponding author. E-mail: steppan@bio.fsu.edu
Molecular phylogeny of the endemic Philippine rodent
Apomys (Muridae) and the dynamics of diversification in
an oceanic archipelago
SCOTT J. STEPPAN
1
*, CHRISTOPHER ZAWADZKI
1
and LAWRENCE R. HEANEY
2
1
Department of Biological Science, Florida State University, Tallahassee, FL 32306–1100, USA
2
Field Museum of Natural History, 1400 S Lake Shore Drive, Chicago, IL 60605, USA
Received 21 January 2003; accepted for publication 22 May 2003
We analysed the phylogenetic relationships of ten of the 13 known species of the genus Apomys using DNA sequences
from cytochrome b. Apomys, endemic to oceanic portions of the Philippine archipelago, diversified during the Pliocene
as these oceanic islands arose de novo. Several of the speciation events probably took place on Luzon or Mindanao,
the two largest, oldest, and most topographically complex islands. Only one speciation event is associated with vicar-
iance due to Pleistocene sea-level fluctuation, and a Pleistocene diversification model in which isolation is driven by
sea-level changes is inconsistent with the data. Tectonic vicariance is nearly absent from the Philippines, in which
tectonic coalescence plays a significant role. Most speciation events (about two-thirds) are associated with dispersal
to newly developed oceanic islands. The data imply that the species have persisted for long periods, measured in mil-
lions of years after their origins; further implications therefore are that faunal turnover is very slow, and persistence
over geological time spans is more prominent than repeated colonization and extinction. Neither the equilibrium nor
the vicariance model of biogeography adequately encompasses these results; a model incorporating colonization,
extinction, and speciation is necessary and must incorporate long-term persistence to accommodate our
observations. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 699–715.
ADDITIONAL KEYWORDS: biogeographical models – cytochrome b – dispersal – island biogeography –
Pleistocene – speciation – vicariance.
INTRODUCTION
The process of diversification - the increase in number
of species within a given lineage over time - lies at the
root of biology, and has a long history of investigation
(Heaney & Vermeij, in press). Nevertheless, the num-
ber of case studies in which the history and processes
of diversification is well documented is small, even
among vertebrates (e.g. Otte & Endler, 1989; Ricklefs
& Schluter, 1993; Grant, 1998; Magurran & May,
1999). Further, such studies have not usually figured
in analysis of the dynamics of species-richness pat-
terns under either of the current dominant models,
the equilibrium model of island biogeography and
vicariance biogeography (Whittaker, 1998; Heaney,
2000; Lomolino, 2000). Much recent progress on some
issues involving diversification has been made under
the rubric of phylogeography (e.g. Riddle, 1995, 1996;
Avise, 2000; Arbogast & Kenagy, 2001), but most such
studies deal with single species, not extensive radia-
tions, and we know of only two that have been placed
in the context of long-term dynamics of patterns of
insular species richness (Moritz et al., 2000; Ricklefs
& Bermingham, 2001). Here we test hypotheses of
speciation and explore patterns of diversification in
the Philippine islands using a molecular phylogeny of
an endemic rodent clade.
The Philippine Islands, comprising c. 7000 islands
ranging up to 100 000 km
2
, represent a natural the-
atre in which to investigate such issues. With the
exception of one small set (the Palawan group), all of
the islands are oceanic in origin, having emerged
because of a complex but now well-documented set of
tectonic events (Hall, 1996, 1998, 2002). Periods of
lowered sea level during the many Pleistocene ‘ice
ages’ caused some sets of current islands to coalesce
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