Dispersal: Biogeography
David M Wilkinson, Liverpool John Moores University, Liverpool, UK
Dispersal is one of the fundamental processes in biogeography crucial to understanding
the distribution of organisms and important in understanding the impact of climate
change on distributions.
Introduction
Biogeography is the area of science at the interface between
ecology and geography that attempts to describe and
explain spatial patterns in the distribution of organisms. It
has often proved much easier to describe these patterns
than to explain how they were formed. However, at the
most basic level there are three fundamental processes in
biogeography: evolution, extinction and dispersal (Brown
and Lomolino, 1998). Combinations of these three
processes explain the presence or absence of a species at
any given location (Figure 1). Dispersal is therefore one of
the key processes in biogeography and is best defined as
‘the movement of organisms away from their point of
origin’ (Brown and Lomolino, 1998). The term can be
applied to individuals, species or higher taxa, so that one
can write of the dispersal of an individual seed or the
dispersal (or lack of dispersal) of a whole family of plants.
As well as its central role in biogeography, dispersal is of
importance in other areas of ecology including metapopu-
lation dynamics (Hanski, 1998) and population ecology,
for example, as a possible mechanism for synchronizing
fluctuations in local populations (Sherratt et al., 2000).
Processes of Dispersal, from Passive to
Active
The most basic distinction between types of dispersal is
between organisms that disperse using their own energy
(active dispersal) and those that use energy from the
environment (passive dispersal). Passive dispersal can
make use of energy from both the abiotic environment,
such as wind and water, or energy from the biotic
environment, an example being seed dispersal by birds.
Organism present at site
either
Evolved at the site
or
Dispersed to the site
Figure 1 Role of the three fundamental processes in biogeography.
ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
Secondary article
Article Contents
. Introduction
. Processes of Dispersal, from Passive to Active
. Dispersal, Migration, Vagrancy and Nomadism
. Spatial Patterns of Dispersal; Diffusion and Jump
Dispersal
. The Power of Dispersal, Examining the Tail of the
Distribution
. Centres of Dispersal or Vicariance Biogeography?
. Implications of Dispersal Limitation for Biogeography
Some organisms can use a mix of passive and active
dispersal; for example, a species of mammal could arrive on
an island either by swimming or by floating on a raft of
vegetation. A range of different technical terms is applied
to different mechanisms of dispersal (Table 1), but in most
cases they fail to add any precision to discussions of
dispersal and as such are best avoided, although they have
been widely used in the past.
Passive dispersal
Much of biodiversity is microscopic; for example, approxi-
mately half of all phyla are composed of microscopic
species; organisms this size can easily be dispersed by air or
water currents. This has given rise to the controversial
suggestion that microbial biodiversity is likely to be low
because widespread dispersal leads to limited geographical
isolation and so a low rate of speciation. There is some
limited evidence in favour of this view, mainly from work
on ciliate protists (Finlay and Fenchel, 1999).
Macroscopic species may also have propagules that are
small enough to be widely dispersed without requiring
adaptations other than small size. Examples include many
bryophyte (e.g. moss) spores; this is strikingly illustrated by
some geothermally heated volcanic soils in the Antarctic
that support bryophyte species from warmer areas of
South America (Smith, 1993).
Larger diaspores require specific adaptations to allow
them to float in water (e.g. coconut palm, Cocos mucifera)
Organism absent from site
either
Became extinct at the site
or
Was never at the site (for evolutionary and/or dispersal reasons)
1
 Dispersal: Biogeography
Table 1 Technical terms often applied to the different
mechanisms of dispersal. In most cases they fail to add any
precision to discussions of these topics; for example, ‘wind
dispersal’ is as precise as ‘anemochory’ and is to be preferred
because it is likely to be understood by a wider range of
readers. Definitions follow Van der Pijl (1969)
Endozoochory Dispersal within animals (e.g.
within the gut)
Epizoochory Dispersal on the outside of animals
(e.g. in fur or feathers)
Myrmechory Ant dispersal
Anemochory Wind dispersal
Hydrochory Water dispersal
Autochory Dispersal by the organism itself,
active dispersal
or on air currents (e.g. dandelions, Taraxacum spp.). Even
with such adaptations, many wind-dispersed plants do not
move very far. This is illustrated by a sample (n 5 25) of
wind-dispersed herbs of open habitats, which had a median
maximum dispersal distance of 12 m (Cain et al., 1998).
An alternative adaptation for passive dispersal is to
make use of actively dispersing animals. In a study of
waterfowl in a salt marsh in North America, over 75% of
the birds examined had plant seeds attached to their
feathers or feet (Vivian-Smith and Stiles, 1994). Some
plants have diaspores with hooks specifically adapted for
transport on fur or feathers (e.g. cleavers, Galium aparine);
however, even seeds without such adaptations can be
dispersed by this method (Fischer et al., 1996).
Active dispersal
Many larger animals are capable of dispersal under their
own power. Because of their ability to fly, the greatest
potential dispersals are seen by birds and bats. For
example, wandering albatrosses, Diomedia exulans, can
move over 8000 km from their breeding sites during the
course of a nonbreeding year (Weimerskirch and Wilson,
2000). This gives considerable potential for range expan-
sion; for example, the collared dove, Streptopelia decaocto,
expanded its range into Western Europe from the eastern
Mediterranean around 1950, but the reasons for this
dramatic range expansion are unclear.
Dispersal, Migration, Vagrancy and
Nomadism
In addition to defining types of dispersal according to
mechanisms (active or passive), a range of other terms have
been used to describe different kinds of dispersal. As
described above, dispersal can be defined as the movement
of organisms away from their point of origin and can be
2 ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
applied to both individuals and taxa. The term ‘migration’
is often used as a synonym for dispersal (e.g. Baker, 1982;
Huntley and Webb, 1989), while other workers restrict its
use to the often annual return migrations typical of the
Arctic tern, Sterna paradisaea, or the monarch butterfly,
Danaus plexippus. The least confusing use of these terms is
probably to consider dispersal and migration as synonyms
and use ‘return migration’ for the more restricted definition
of migration in which the individual makes migrations in
both directions during the course of its life. Huntley and
Webb (1989) argued that there are strong analogies
between the annual return migrations of birds and the
movement of plants over a glacial/interglacial cycle, both
being driven by orbital forcing albeit on very different time
scales. They argue that this makes migration an appro-
priate term to use in both cases.
Individual organisms can sometimes appear at a
location well outside their normal range. For example,
birds or butterflies may be blown large distances by storms,
or floating seeds may sometimes cross an entire ocean.
Such individuals are referred to as vagrants (sometimes
called accidentals) and are of biogeographical interest
because of the possibility that they could lead to the
colonization of a new site and so produce an increase in a
species range. Such events, although presumably rare, are
likely to be very important in the colonization of islands.
The concept of vagrant may be restricted to multicellular
organisms. If the idea that most microbes are so small that
they can disperse anywhere is correct, then establishment,
rather than dispersal, will be the key mechanism control-
ling whether a population can develop at a given location.
This idea is encapsulated in an old maxim of microbiology
‘everything is everywhere, the environment selects’.
Even when an organism is capable of active dispersal,
there are large differences in the probability of long-
distance dispersal. Some organisms move very little during
their life, while others cover large distances, sometimes
appearing to wander at random. Such behaviour is called
nomadism and the wandering albatross has been consid-
ered a classic example. It has long been assumed that
during nonbreeding periods these birds wander aimlessly,
circumnavigating the southern ocean. Recent work sug-
gests that this is not the case and individual birds return to
favourite nonbreeding areas (Weimerskirch and Wilson,
2000). This raises the question, are other ‘nomadic’ species
so classified because of lack of knowledge rather than
because they have a random dispersal pattern?
Spatial Patterns of Dispersal; Diffusion
and Jump Dispersal
There are two main patterns of range expansion. Either a
population can slowly expand from the margins of its
geographical range (diffusion, the analogy is with a
 Dispersal: Biogeography

diffusing chemical) or a small number of individuals can The Power of Dispersal, Examining the
disperse to a new location some distance from the current
edge of the species range (jump dispersal); or a combina-
Tail of the Distribution
tion of these two processes can occur (Figure 2). Clearly
Diffusion as an explanation for many range expansions
jump dispersal is by definition important in island
experiences serious problems when compared to data from
biogeography; a classic example is given by the volcanic
Quaternary geology. As pointed out by Clement Reid at
islands of Hawaii, which were never connected to any
the end of the nineteenth century, there has not been
continental land mass but have a diverse flora and fauna
enough time since the last glaciation for plants spreading
(Carlquist, 1981). Jump dispersal is also likely to allow
slowly by diffusion to return to Britain from their refugia in
faster range expansion than would be possible with pure
southern Europe. This result was confirmed by more
diffusion.
formal modelling in a classic paper by Skellam (1951); the
Some organisms are more likely to show jump dispersal
problem is now sometimes called Reid’s Paradox (Clark
than others; for example, birds tend to be more likely
et al., 1998). Most plant seeds disperse only a small distance
candidates than flightless (nonvolant) mammals. Many
from the parent plant (e.g. Cain et al., 1998), so if a statistic
plant species also show evidence of jump dispersal; while
such as mean dispersal distance is used in a model of range
factors such as wind dispersal undoubtedly contribute to
expansion then speeds much slower than those in the
this, several people have stressed the central role of birds
Quaternary fossil record are often obtained. The problem
transporting seeds in plant jump dispersal (Carlquist, 1981;
is that speed of range expansion is largely controlled by a
Wilkinson, 1997). The implication of this is that plant jump
few seeds that move much farther than the mean dispersal
dispersal (and so speed of migration) would have been less
distance, i.e. the extreme right hand of the distribution of
prior to the evolution of birds.
dispersal distances (Figure 3). Because such events are rare,
Many cases of range expansion probably involve both
they are very difficult to quantify. Recent modelling has
diffusion and jump dispersal. A classic example is the
demonstrated the great importance of these rare extreme
spread of the cattle egret, Bubulcus ibis, in the Americas.
dispersal events in determining the rate of migration of a
This bird, a native of Africa, colonized northeastern South
population such as oak, Quercus sp., in Europe at the end
America by jump dispersal towards the end of the
of the last glaciation (Clark et al., 1998). Both Reid and
nineteenth century. It subsequently spread over much of
Skellan implicated birds in these rapid plant migration
South America and also spread north into the southern
events, but for a full understanding more data is required
United States. Maps of this spread (e.g. Brown and
on these rare jump dispersal events.
Lomolino, 1998) suggest a pattern similar to Figure 2a.
Range expansion as a steadily expanding front, albeit with
some additional jump dispersal events, such as the
colonization of many Caribbean islands. In most cases of
range expansion the correct question is not ‘is it diffusion or Centres of Dispersal or Vicariance
jump dispersal?’ but ‘what are the relative contributions of Biogeography?
these two processes?’. The big problem with jump dispersal
is that by its nature it is a rare event and so difficult to study. The reliance of dispersal biogeography on chance events
has been seen as undesirable by some biologists as they
view the resulting explanations as untestable ‘just so

(a) (b) (c)

Figure 2 Types of range expansion. (a) Diffusion. Range expansion as a steady expanding front. (b) Jump dispersal. Founding a new population well
outside the species current range. (c) Combination of diffusion and jump dispersal. Over time the new population founded by a jump dispersal event may
merge with the main population as both expand by diffusion.

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 Dispersal: Biogeography
Frequency
Dispersal distance
Figure 3 Distribution of seed dispersal distances for many organisms has a
very long tail with a few diaspores or individuals dispersing much further
than the average distance. The shape of the curve illustrated is typical for
plant species as different as the herb Asarum canadense and the tree Picea
sp.
stories’. Vicariance biogeography attempted to address
this problem by downplaying the importance of dispersal
and replacing it with the idea that a taxon’s range can be
divided by the formation of physical barriers. For example,
if closely related taxa are found on both sides on an ocean,
dispersal biogeography would point to a dispersal event as
the explanation (as in the cattle egret example), while
vicariance biogeographers would suggest that a formerly
continuous distribution of an ancestral taxon had been
divided by the formation of the ocean. The advantage in
the vicariant position is that the physical division of the
taxon’s range is open to independent geological verifica-
tion. These different traditions can be traced back to the
nineteenth century when Charles Darwin emphasized
dispersal and the importance of ‘occasional means of
distribution’, while his close friend Joseph Dalton Hooker
emphasized the physical dissection of previously contin-
uous distributions. Wilson (1991) tabulated the differences
between dispersal (migration) biogeography and vicar-
iance, along with the closely related ideas of panbiogeo-
graphy, in detail. The problems with the vicariance
approach include the assumption that all speciation is
due to geographical isolation (the alternative, sympatric
speciation, is increasingly the subject of much active
research) and the refusal to consider long-distance
dispersal, despite the many cases described in the literature.
Most modern biogeographers work within the Darwinian
dispersal tradition while allowing that vicariant events (e.g.
due to plate tectonics) can sometimes be important. The
central paradigm of dispersal biogeography is that taxa
usually originate at a single location (a centre of dispersal)
and may then expand their range to produce their present
distribution (Wilson, 1991).
4 ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
Implications of Dispersal Limitation for
Biogeography
It is a common observation that different species are found
in different parts of the world. One explanation for this is
that species are found where the environmental conditions
are suitable for them. This could be true for many free-
living microbes, which are small enough to easily disperse.
However for many larger organisms it is clear that the
distribution of suitable habitats is not the only factor
limiting their distribution. This is demonstrated by the
example of the cattle egret, which after managing to cross
the Atlantic colonized large areas of the Americas. Clearly
suitable habitat was available in America; the limiting
factor was dispersal from Africa. The same point is
underlined by the success of many species that have been
introduced to Britain and survived to produce self-
sustaining populations; these include birds such as the
Canada goose, Branta canadensis, and ruddy duck, Oxyura
jamaicensis, as well as mammals such as the brown rat,
Rattus norvegicus. When humans solved these animals’
dispersal problems they were able to expand their ranges to
include Britain. Humans now play a major role in the
dispersal of many organisms; for example, they have
introduced over 1000 plant species to the British country-
side, of which around 70 have been widely naturalized
(Crawley, 1997).
Past dispersal limitation
The idea of movement out from a centre of dispersal is
nicely illustrated by our own species, Homo sapiens. It
appears to have originated in Africa in excess of 100 000 yr
ago; by approximately 50 000 yr ago, they were in Asia and
even Australia and arrived in the Americas some time later,
although the exact time is still controversial. However,
remote islands such as Hawaii and Easter Island were not
reached until around AD 500 because of dispersal
problems.
In the case of humans on Hawaii, it appears that the
difficulties in dispersal, rather than environmental condi-
tions on the island, were the rate-determine step in the
colonization process. But this is not always the case and
migration rates can be constrained by environmental
factors such as rate of climate change, rather than by the
potential dispersal rate. Consider the migration rate of
European trees after the end of the last glaciation (Table 2):
the speeds reconstructed from analysis of fossil pollen
preserved in peat and lake sediments rule out diffusion
(Reid’s paradox discussed above), and modelling by Clark
(1998) suggests that these migration velocities may not
have reached their maximum, ‘being stalled by rates of
climate change, geography or both’. This has implications
for future migration rates in response to climate change
(discussed below).

Table 2 Maximum rate of migration of some European tree
taxa during the past 10 000 yr
Maximum migration rate
Taxa (m per yr)
Hazel, Corylus 1500
Beech, Fagus 300
Ash, Fraxinus 500
Oak, Quercus 500
Lime, Tilia 500
Elm, Ulmus 1000
On a larger geological time scale, a major constraint on
dispersal has been the changing patterns of the continents
due to plate tectonics. For example, the formation of the
Himalayas as a result of collisions between continental
plates isolated many species on either side of the barrier,
while creating a habitat for others such as the cold-adapted
snow leopard, Panthera uncia.
Future dispersal limitations
There is now great concern over climate change caused by
factors such as the increase in greenhouse gases. One of the
main insights from the study of past dispersal events is that
most species have responded to past climatic changes by
migration rather than by in situ adaptation to the new
conditions (Huntley, 1994). Thus a key question is whether
organisms will be able to disperse at a speed that will allow
them to track the changing climate.
The fact that immobile organisms, such as plants, can
migrate at rates well in excess of those predicted from their
average dispersal distance could hold promise for popula-
tions in the face of future global changes (Clark, 1998).
However, the future changes are unlikely to be identical to
those in the past. For example, it is possible that the rate of
climate change could be greater than any seen in the last 2.4
million yr (Huntley, 1994). In addition, the nature of the
landscape through which organisms disperse has now
changed through agriculture and urban development. It is
clearly vital to preserve the countryside in a state that
allows organisms to migrate through it unimpeded in
response to climate change. This may no longer be the case
in many parts of the world, so that the optimism based on
past migration rates may be misplaced. Modelling studies
by Dyer (1995) have illustrated the way different species
may show variations in rates of response to climate change.
In his model the fastest migration rate observed for wind-
dispersed plants was over 1.5 times slower than that for
bird-dispersed species. However, Wilkinson (1997) has
argued that, at the temporal and spatial scales of interest in
biogeography, such simple classifications of dispersal types
(as listed in Table 1) can break down, so that, for example,
wind-dispersed seeds can be dispersed by birds.
Human influence has had another effect on dispersal,
namely reducing the effects of dispersal barriers by moving
ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
Dispersal: Biogeography
organisms around the world. A historical example of this
effect is the Polynesian colonization of Henderson Island,
in the Pitcairn group of islands in the South Pacific. At least
three species of snails first appear in the Polynesian
occupation horizons. At the same time at least five native
snail species became extinct, probably through habitat
destruction caused by humans, although competition with
the introduced snails cannot be ruled out (Preece, 1998). A
study of the current British flora by Hodkinson and
Thompson (1997) found that many species were now
dispersed by soil carried by motor cars, or as topsoil and by
other human-based dispersal mechanisms. In the marine
environment, transport on the hulls of ships is now a major
dispersal mechanism for some types of bryozoa (Watts
et al., 1998). When a species is dispersed to another part of
the world by human actions, its main parasites and
predators can be left behind; occasionally this can cause
a dramatic increase in its population at the new site and the
introduced organism can become a pest species. A
dramatic illustration of the effects of such a release from
natural enemies is rubber, Hevea brasiliensis, which cannot
be grown as a plantation crop in its native Brazil but is a
major plantation crop in South East Asia where it has been
introduced without many of the species that feed on it in
South America (Crawley, 1997). If an introduced species
becomes a major problem, one possibility is to introduce
some of its natural enemies. This was done to control the
cactus Opuntia ficus-indica, introduced to South Africa
from the Americas. which at one point infested 900 000 ha
of South Africa (Nobel, 1994). Increasingly, human
actions are reducing the effect of barriers to dispersal; this
can confront species of restricted range with new compe-
titors that are often widespread successful species (such as
Opuntia) that can outcompete them, leading to a decline of
biodiversity.
References
Baker RR (1982) Migration, Paths Through Time and Space. London:
Hodder and Stoughton.
Brown JH and Lomolino MV (1998) Biogeography. Sunderland, MA:
Sinauer.
Cain ML, Damman H and Muir A (1998) Seed dispersal and the
Holocene migration of woodland herbs. Ecological Monographs 68:
325–347.
Carlquist S (1981) Chance dispersal. American Scientist 69: 509–516.
Clark JS (1998) Why trees migrate so fast: confronting theory with
dispersal biology and the paleorecord. American Naturalist 152: 204–
224.
Clark JS, Fastie C, Hurtt G C et al. (1998) Reid’s paradox of rapid plant
migration. Bioscience 48: 13–24.
Crawley MJ (1997) Plant Ecology, (chap. 19). Oxford: Blackwell.
Dyer JM (1995) Assessment of climatic warming using a model of forest
species migration. Ecological Modelling 79: 199–219.
Finlay BJ and Fenchel T (1999) Divergent perspectives on protist species
richness. Protist 150: 229–233.
Fischer SF, Poschlod P and Beinlich B (1996) Experimental studies on
the dispersal of plants and animals on sheep in calcareous grassland.
Journal of Applied Ecology 33: 1206–1222.
5
 Dispersal: Biogeography
Hanski I (1998) Metapopulation dynamics. Nature 396: 41–49.
Hodkinson DJ and Thompson K (1997) Plant dispersal: the role of man.
Journal of Applied Ecology 34: 1484–1496.
Huntley B (1994) Plant species response to climate change: implications
for the conservation of European birds. Ibis 137: s127–s138.
Huntley B and Birks HJB (1983) An Atlas of Past and Present Pollen
Maps for Europe 0 – 13000 BP. Cambridge, UK: Cambridge
University Press.
Huntley B and Webb T (1989) Migration: species response to climatic
variations caused by changes in the Earth’s orbit. Journal of
Biogeography 16: 5–19.
Nobel PS (1994) Remarkable Agaves and Cacti. Oxford: Oxford
University Press.
Preece RC (1998) Impact of early Polynesian occupation on the land
snail fauna of Henderson Island, Pitcairn Group (South Pacific).
Philosophical Transactions of the Royal Society of London B 353: 347–
368.
Sherratt TN, Lambin X, Petty SJ et al. (2000) Use of coupled oscillator
models to understand synchrony and travelling waves in populations
of the Field Vole Microtus agrestis in Northern England. Journal of
Applied Ecology 37(supplement 1): 148–158.
Skellam JG (1951) Random dispersal in theoretical populations.
Biometrika 38: 196–218.
Smith RIL (1993) The role of bryophyte propagule banks in primary
succession: case study of an Antarctic fellfeild soil. In: Miles J and
Walton DWH (eds) Primary Succession on Land. Oxford: Blackwell.
Van der Pijl L (1969) Principles of Dispersal in Higher Plants. Berlin:
Springer-Verlag.
Vivian-Smith G and Stiles EW (1994) Dispersal of salt marsh seeds on
the feet and feathers of waterfowl. Wetlands 14: 316–319.
6 ENCYCLOPEDIA OF LIFE SCIENCES / & 2001 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net
Watts PC, Thorpe JP and Taylor PD (1998) Natural and anthropogenic
dispersal mechanisms in the marine environment a study using
cheilostome bryozoa. Philosophical Transactions of the Royal Society
of London B 353: 453–464.
Weimerskirch H and Wilson RP (2000) Oceanic respite for wandering
albatrosses. Nature 406: 955–956.
Wilkinson DM (1997) Plant colonization: are wind dispersed seeds really
dispersed by birds at larger spatial and temporal scales? Journal of
Biogeography 24: 61–65.
Wilson JB (1991) A comparison of biogeographic models: migration,
vicariance and panbiogeography. Global Ecology and Biogeography
Letters 1: 84–87.
Further Reading
Baker RR (1978) The Evolutionary Ecology of Animal Migration.
London: Hodder and Stoughton.
Cox CB and Moore PD (2000) Biogeography, 6th edn. Oxford:
Blackwell.
Pitelka LF, Garden RH, Ash J et al. (1997) Plant migration and climate
change. American Scientist 85: 464–473.
Thornton I (1996) Krakatau. Cambridge, MA: Harvard University
Press.
Whittaker RJ (1998) Island Biogeography. Oxford: Oxford University
Press.
Wilkinson DM (1999) Plants on the move. New Scientist 2178(suppl.):
S1–S4.