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G U E S TE D I T O R I A L

Dispersal is fundamental to

biogeography and the evolution of

biodiversity on oceanic islands

Robert H. Cowie* and Brenden S. Holland

INT RO DUCT IO N

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 tooffer 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.

A consequence of this thinking seems to have been a focus

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 henceuninteresting.

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

Center for Conservation Research and

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: [email protected]

ABS T RACT

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 beingrecognized, 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.

KeywordsBiodiversity, dispersal, endemism, historical biogeography, hot spot islands,

land snails, oceanic islands, Pacific Ocean.

Journal of Biogeography ( J. Biogeogr .) (2006) 33, 193–198

ª

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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;

Sanmartın & 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.

V ICARIANCE AND DIS P E RS AL IN O CE ANIC

IS LAND S Y S T E M S

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

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.

GE NE RAL P AT T E RNS O F DIS P E RS AL

Strict vicariance biogeography has essentially considereddispersal 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

R. H. Cowie and B. S. Holland

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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.

FUT URE DIRE CT IO NS IN O CE ANIC IS LAND

BIO GE O GRAP HY

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 mostmarine 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

subgenus of Pacific blackflies (Simuliidae) (e.g. Craig et al.,

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, thecombined land area ofall oceanicislands 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 theHawaiian 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

thesecomprehensive phylogeniesis a wide-open opportunity for

research that will lead to major advancesin ourunderstanding 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 pathwaysvia 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

Kauai (5.1)auai 5.1)

Oahu (3.7)ahu 3.7)

Molokai (1.9)olokai 1.9)

Maui (1.3)aui 1.3)

Lanai (1.3)anai 1.3

Hawaii (0.4)awaii 0.4)

100 km00km

N

i u n i u a M i

i

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.

Guest Editorial

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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).

CO NCLUS IO N

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.

ACK NO W LE DGE M E NT S

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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Guest Editorial

Journal of Biogeography 33, 193–198 197ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd

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B I O S K E T C H E S

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, focusingprimarily on molluscs, both marine and non-marine.

Editors: David Richardson and Robert Whittaker



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