Journal of Biogeography (J. Biogeogr.

) (2006) 33, 193–198

GUEST Dispersal is fundamental to

EDITORIAL
biogeography and the evolution of
biodiversity on oceanic islands
Robert H. Cowie* and Brenden S. Holland

Center for Conservation Research and ABSTRACT

Training, University of Hawaii, Honolulu,
HI, USA
*Correspondence: Robert H. Cowie, Center for
Conservation Research and Training, University
of Hawaii, 3050 Maile Way, Gilmore 408,
Honolulu, Hawaii 96822, USA.
E-mail: cowie@hawaii.edu
Vicariance biogeography emerged several decades ago from the fusion of cladistics
and plate tectonics, and quickly came to dominate historical biogeography. The
field has since been largely constrained by the notion that only processes of vica-
riance and not dispersal offer testable patterns and refutable hypotheses, dispersal
being a random process essentially adding only noise to a vicariant system.
A consequence of this thinking seems to have been a focus on the biogeography of
continents and continental islands, considering the biogeography of oceanic islands
less worthy of scientific attention because, being dependent on stochastic dispersal,
it was uninteresting. However, the importance of dispersal is increasingly being
recognized, and here we stress its fundamental role in the generation of biodiversity
on oceanic islands that have been created in situ, never connected to larger land
masses. Historical dispersal patterns resulting in modern distributions, once con-
sidered unknowable, are now being revealed in many plant and animal taxa, in large
part through the analysis of polymorphic molecular markers. We emphasize the
profound evolutionary insights that oceanic island biodiversity has provided, and
the fact that, although small in area, oceanic islands harbour disproportionately
high biodiversity and numbers of endemic taxa. We further stress the importance of
continuing research on mechanisms generating oceanic island biodiversity, espe-
cially detection of general, non-random patterns of dispersal, and hence the need to
acknowledge oceanic dispersal as significant and worthy of research.
Keywords
Biodiversity, dispersal, endemism, historical biogeography, hot spot islands,
land snails, oceanic islands, Pacific Ocean.

A consequence of this thinking seems to have been a focus

INTRODUCTION
Since the dispersal versus vicariance debate of the 1970s and
early 1980s and the broad acceptance and melding of the
theories of cladistics and plate tectonics, vicariance approaches
to historical biogeography have dominated the last two or
three decades. Only vicariant mechanisms were considered to
offer testable patterns and refutable hypotheses, dispersal being
a random process essentially adding only noise to a vicariant
system (Morrone & Crisci, 1995; Humphries & Parenti, 1999).
Dispersalist explanations were treated as simply narrative,
appealing to individual explanations and hence ungeneraliz-
able; and a priori assumption of dispersal was considered
unnecessary (Rosen, 1976; Peake, 1981; Lynch, 1989; Morrone,
2005). Dispersal was therefore of little interest and hence given
but ‘footnote acknowledgment’ (Lynch, 1989) by many
vicariance biogeographers.
on the biogeography of continents and continental islands.
Acknowledgement that the distribution of the plants and
animals of oceanic islands is strongly moulded by dispersal,
while at the same time considering that dispersal cannot be
rigorously modelled or tested, meant that the biogeography of
oceanic islands became almost by definition random and hence
uninteresting.
Long-distance dispersal, however defined, certainly occurs
frequently and is often directional (Nathan, 2005; and other
papers in Diversity and Distributions volume 11, number 2).
Although some historical biogeographers have explicitly
acknowledged the importance of dispersal and incorporated
it into their analytical approaches (e.g. Brooks & McLennan,
1991, 2001; McLennan & Brooks, 2002; Halas et al., 2005), for
the most part they have not been searching for general non-
random patterns of dispersal that can explain biogeographical

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R. H. Cowie and B. S. Holland
diversity. They have rather simply acknowledged that dispersal
happens in particular scenarios and should be part of a
combined vicariant-dispersal explanation of the distributions
of the taxa in question.
Recently, a number of articles, both in this journal and
others (Givnish & Renner, 2004; Renner, 2004; Cook & Crisp,
2005; Halas et al., 2005; McGlone, 2005; de Queiroz, 2005),
have argued that dispersal must be seriously considered as an
important process in evolution and speciation, and that its role
has been underestimated in historical biogeography. We agree
wholeheartedly. However, much of this commentary has been
focused on dispersal in a continental or continental island
context, and while trans-oceanic dispersal has been considered,
this has been primarily in the context of explanations for the
occurrence of related taxa on widely separated continents,
notably in the Southern Hemisphere (Givnish & Renner, 2004;
Sanmartı́n & Ronquist, 2004; Cook & Crisp, 2005). In this
editorial we want to take these arguments further and to
emphasize strongly that in oceanic island situations in
particular not only is dispersal important but it is the critical
initiating step in the generation of endemic biodiversity,
without which vicariant evolution within these islands could
not happen, and furthermore that general patterns of direc-
tional dispersal can be detected. The emphasis over the last few
decades on almost exclusively vicariant scenarios as providing
the only rigorously testable hypotheses of historical biogeog-
raphy, with dispersal considered as just random noise, has
stifled important research on the role of dispersal in oceanic
biogeography.
VICARIANCE AND DISPERSAL IN OCEANIC
ISLAND SYSTEMS
The majority of the myriad islands of the Pacific have been
formed in situ and not by the fragmentation of large land
masses. As the Pacific tectonic plate moves steadily north-
westwards, island arcs, such as the Marianas and some of the
Fijian islands, are created at its western edge as it meets the
tectonic plates to the west (Polhemus, 1996). However, most of
the islands of the central Pacific have been formed as the plate
moves over stationary hot spots in the underlying mantle,
which from time to time send magma up through the plate,
forming long chains of volcanoes, each volcano sequentially
younger than the one that preceded it, and which will have
moved north-westwards away from the hot spot (Price &
Clague, 2002). This is how the Hawaiian Islands, the islands of
French Polynesia (Tahiti, etc.), the Samoan Islands, and many
others have been formed. Only few Pacific islands are of
continental origin, for example New Caledonia and New
Zealand. In the Atlantic, similar geological forces have created
volcanic island chains such as the Canary Islands and the
Madeiran archipelago.
Thus, the biogeography and endemic biodiversity of these
oceanic islands, both arc and hot-spot islands, which have
never had a connection to a continental land mass, are
fundamentally a product of oceanic dispersal. That vicariance
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ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
has played a role in the radiation of lineages is not to be
denied, as for instance among the Hawaiian islands
of Molokai, Maui, Lanai and Kahoolawe, the so-called
‘Maui-nui’ island group. Because of a combination of fluctua-
tions in sea level and the dynamic processes of building,
subsidence and erosion of these sequentially produced islands,
the islands of Maui-nui have at various times been emergent
volcanic peaks separated by sea-water channels, then combined
into larger land masses, and finally broken again into a number
of smaller islands (Price & Elliott-Fisk, 2004). Within certain
lineages, intra-island speciation may also have been driven by
various selective forces, such as sexual selection (Kaneshiro &
Boake, 1987) and reproductive isolation brought about by
fragmentation as deep valleys formed through erosion of
previously continuous habitat (Holland & Hadfield, 2002).
However, in the most fundamental sense, oceanic dispersal is
the key to evolutionary radiation on these islands.
GENERAL PATTERNS OF DISPERSAL
Strict vicariance biogeography has essentially considered
dispersal as noise in the system, stochastic in nature, and
therefore exhibiting no general patterns and permitting no
refutable hypotheses, hence being ultimately uninteresting
(Humphries & Parenti, 1999). But there are indeed general
dispersal patterns in oceanic systems, related to ocean currents,
predominant wind patterns (trade winds, hurricane tracks),
the geographical arrangement of islands (facilitating use as
stepping stones) and bird migration routes (Peake, 1981;
Ballard & Sytsma, 2000; Hoskin, 2000; Givnish & Renner,
2004; Renner, 2004). Asymmetry of dispersal – the predom-
inance of dispersal in one direction rather than another – has
recently received particular attention as a potentially con-
founding factor for biogeographical reconstructions that
ignore it (Cook & Crisp, 2005; McGlone, 2005). In addition,
there are well-documented patterns in which taxa with
inherently worse dispersal abilities are unable to colonize
more geographically isolated islands (Peake, 1981; Whittaker,
1998), and disharmonic biotic distributions arise as a result.
Furthermore, to reinforce the notion that dispersal within
hot-spot archipelagos is far from a stochastic process, we can
point to many examples of studies in numerous plant and
animal groups that have detected common patterns of
dispersal that generally conform to the so-called ‘progression
rule’ (Fig. 1) that dispersal and colonization, frequently
accompanied by lineage bifurcation, proceeds from older to
younger islands (Wagner & Funk, 1995; Roderick & Gillespie,
1998; Juan et al., 2000; Hormiga et al., 2003; Nepokroeff et al.,
2003; Holland & Hadfield, 2004). As an island first appears
above the surface of the ocean it is available for colonization,
and its most likely colonizers come from the nearest land mass,
which is the next youngest island in the chain. As the plate
continues to move, new islands form and the process is
repeated. Sometimes colonizing species pass over one or more
islands in colonizing the newest island, sometimes there are
back-colonizations from younger to older islands; although
Journal of Biogeography 33, 193–198

N subgenus of Pacific blackflies (Simuliidae) (e.g. Craig et al.,
Kauai (5.1)
Oahu (3.7) Ma
u 100 km
in
ui
Molokai (1.9)
Lanai (1.3)
Maui (1.3)
Hawaii (0.4)
Figure 1 An example of a non-stochastic dispersal pattern
observed for many plant and animal lineages, the progression rule
pattern of island colonization, here depicted for a hypothetical
Hawaiian island lineage. The volcanoes of the six main Hawaiian
islands arose over a hot spot in the south-east. As the Pacific plate
moves northwestwards it carries with it each sequentially formed
island (island ages shown in parentheses, in millions of years).
According to the progression rule, the initial colonization event
occurs on the oldest island, Kauai, accompanied by subsequent
lineage splitting as individuals disperse down the island chain from
volcano to volcano (black circles). More complex patterns,
involving radiations within islands, back-colonizations and dis-
persal that passes over an intermediate island, are often super-
imposed on the basic progression rule pattern.
stochastic dispersal patterns have been observed (Wagner &
Funk, 1995), particularly for species with a high dispersal
ability such as flying insects, birds and plants with wind-
dispersed seeds (Holland & Hadfield, 2004), the progression
rule pattern holds broadly for many taxa.
FUTURE DIRECTIONS IN OCEANIC ISLAND
BIOGEOGRAPHY
Nonetheless, we still know rather little about the dispersal
processes and mechanisms that bring about the initial
colonizations of these isolated archipelagos. We know, for
instance, that there have been multiple colonizations of the
Hawaiian Islands from more than one region of the Pacific rim
(Gillespie et al., 1994; Rundell et al., 2004). In the marine
realm, coastal organisms, especially those lacking a dispersal
phase, behave rather like terrestrial organisms; but most
marine dispersal in the Pacific has been outward from the
centres of diversity in the East Indies (Briggs, 1999). While
there have been numerous studies of within-archipelago
phylogenetic diversification and biogeography, especially in
Hawaii and the Canary Islands (see references above), there
remain few comprehensive ocean-wide reconstructions of the
phylogeny of any major widespread group of plants or animals,
and, at least in the Pacific, little research addressing both
geographical origins and routes of colonization ocean-wide.
Some of the few examples include morphological research on a
Journal of Biogeography 33, 193–198
ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd
Guest Editorial
2001; Spironello & Brooks, 2003) and molecular work on the
Pacific tree subgenus Metrosideros (Wright et al., 2000).
The downplaying of the role of dispersal may have arisen in
the past in part because of difficulty in resolving the underlying
evolutionary histories within and among island species. Recent
studies that have revealed non-stochastic dispersal and colon-
ization pathways have done so largely as a result of the
development of new laboratory techniques and multivariate
statistical methods, and in particular the availability of
polymorphic molecular markers. DNA-based approaches are
revolutionizing our ability to tease apart and reconstruct
otherwise cryptic biogeographical pathways.
Although the oceans cover nearly three-quarters of the earth’s
surface, the combined land area of all oceanic islands represents a
miniscule fraction of the earth’s total land area. For instance, the
islands of the Pacific (excluding continental New Guinea and
New Zealand) have a total area of about 106,000 km2, approxi-
mately the area of Guatemala and less than 0.1% of the earth’s
total land area. Yet the terrestrial biotas of these islands are
immensely diverse: for example the diversity of land snails in the
Hawaiian Islands (Cowie, 1996a) is comparable to that of the
whole of North America north of Mexico (Pilsbry, 1938–1948)
and exhibits unrivalled endemism, exceeding 95%. This oceanic
island biodiversity has provided biologists since Darwin with
fundamental insight into the processes and patterns of evolution
(Emerson, 2002). Evolution of the biotas of oceanic islands is
therefore of great interest and significance, and it is important
that the major force that has shaped these biotas, namely
dispersal, should not be dismissed but acknowledged and
researched all the more intensely. Phylogenies are the prere-
quisite for ascertaining the detailed pattern of dispersal and
colonization that has resulted in the huge diversity among many
widespread groups of organisms on oceanic islands. Developing
these comprehensive phylogenies is a wide-open opportunity for
research that will lead to major advances in our understanding of
the biogeography of an important component of the earth’s
biodiversity. Also, by re-emphasizing the role of dispersal in
historical biogeography, we see an opportunity, although not an
easy one (McGlone, 2005), to test hypotheses of its non-
randomness, thereby refining our biogeographical interpreta-
tions with a more holistic approach that perhaps will better
reflect biological reality.
Our current research on succineid land snails aims to do
this, by developing and testing phylogenetic hypotheses for the
entire family in the Pacific, permitting us to infer the pathways
via which these snails have colonized the islands from
continental regions, by large trans-oceanic jumps or in a
stepping-stone manner from island group to island group,
gradually making their way across the ocean. The patterns
probably do not follow oceanic currents, as most land snails
would be unlikely to survive such journeys faced with long-
term exposure to salt water. But they may follow bird
migration routes – succineids have been recorded attached to
birds travelling long distances (Anonymous, 1936; Rees, 1965;
Boag, 1986). And they may follow generalized hurricane
195
R. H. Cowie and B. S. Holland
pathways – snails can fly (Kirchner et al., 1997), perhaps
especially if attached to a leaf. Such dispersal mechanisms have
often been suggested for snails (e.g. Vagvolgyi, 1975; Peake,
1981; Hausdorf, 2000).
An estimated 29 successful colonizations are necessary to
explain the diversity of Hawaiian land snails (Ziegler, 2002),
which translates into roughly one every million years, if the
oldest colonization was to the island of Kure, or one every
175,000 years if the oldest colonization was to Kauai (Price &
Clague, 2002). On average, hurricanes with maximum wind
speeds of 64 knots (c. 119 km h)1) (minimum hurricane
intensity) and maximum wind speeds of 125 knots
(c. 232 km h)1) come within 250 nautical miles (463 km) of
the Hawaiian Islands every 7 and 137 years, respectively, with
hurricanes of intermediate wind speed coming at intermediate
frequencies (Chu & Wang, 1998). That snails, with some
exceptions that have evolved in situ (Cowie, 1996a), tend to be
smaller the more isolated the island offers support to the idea
that birds and the wind are the more important mechanisms
(Peake, 1981; Cowie, 1996b).
CONCLUSION
Thus, the immense diversity of land snails and other organisms
on oceanic islands (e.g. Ziegler, 2002) is fundamentally
dependent on dispersal, and that dispersal probably exhibits
patterns both on a small intra-archipelago scale and on a larger
ocean-wide scale. These patterns remain poorly understood
but deserve considerable research attention.
We hope, therefore, that the trend identified by de Queiroz
(2005) – the resurrection of oceanic dispersal as important in
historical biogeography – is real and that the straightjacket of
strict vicariance biogeography is being loosened to include
once again the plurality of mechanisms and processes that
make evolutionary biology the exasperating but ever fascin-
ating discipline that it is. At least in the Pacific, there are
excellent opportunities with numerous plant and animal
groups not only to address the origins and diversification of
this important component of the earth’s biodiversity but also
to contribute profoundly to the advancement of the discipline
of biogeography.
ACKNOWLEDGEMENTS
Our research on Pacific island biogeography is currently
supported by the US National Science Foundation, grant DEB-
0316308. We thank Richard Field for helpful comments on a
draft of this article.
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R. H. Cowie and B. S. Holland

BIOSKETCHES

Robert H. Cowie is an Associate Researcher in the Center for Conservation Research and Training at the University of Hawaii. His
research addresses issues related to the origins and determinants of biodiversity and its conservation, as well as the origins and
impacts of introduced species. His primary focus is on non-marine snails in the islands of the Pacific and freshwater snails in South
and Central America.
Brenden S. Holland is a Junior Researcher in the Center for Conservation Research and Training at the University of Hawaii. His
research uses molecular techniques to address phylogenetic and phylogeographical questions in Pacific island biodiversity, focusing
primarily on molluscs, both marine and non-marine.

Editors: David Richardson and Robert Whittaker

198 Journal of Biogeography 33, 193–198

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