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Wildlife Research, 2007, 35, 180–184 www.publish.csiro.au/journals/wr
The invasion ecology of mammals: a global perspective
Mick N. CloutA,B and James C. RussellA
ACentre for Biodiversity and Biosecurity, School of Biological Sciences, University of Auckland,
Private Bag 92019, Auckland 1142, New Zealand.
BCorresponding author. Email: m.clout@auckland.ac.nz
Abstract. Of the extant species of land mammals, 124 (2.6%) can be classed as ‘successful invaders’, whereas 1038
(21.6%) are classed as threatened, according to the 2006 IUCN Red List. Relatively high proportions of successful
invaders are found among Artiodactyla, Carnivora, Lagomorpha and Perissodactyla. Compared with other organisms,
mammals seem relatively likely to become established when introduced outside their natural range. Studies of determi-
nants of invasion success indicate that the number of individuals released, the size of the natural range of the introduced
species, and the temperateness of climate in the new range can all increase the probability of establishment of introduced
mammals. Negative impacts of invasive mammals on native biodiversity include direct effects such as predation, browsing
and competition, but can extend to disruption of patterns of nutrient flow, and trophic cascades. Eradication of several
species of invasive mammals from increasingly large areas is now possible. In this context, it is important to better under-
stand ecological interactions between such mammals (and between them and other species) to avoid unwanted
consequences such as mesopredator or competitor release, after the removal of particular species. Finally, it is increasingly
apparent that research is needed on the behaviour of dispersing and invading individuals, to improve the early detection of
new mammal invasions or reinvasions.
Introduction
The deliberate or accidental introduction of species beyond their Methods
native range has been a historical consequence of human migra- We searched the comprehensive book by Long (2003) for
tion and trade, and has accelerated greatly in recent times. mammal species that have been recorded as being introduced
Several introduced species (including most crops and domestic and establishing a self-sustaining wild population in at least one
livestock) are largely beneficial, but many others have become location outside their natural range. Such species were classed
weeds or pests. Some have proved to be highly invasive, modi- as ‘successful invaders’, and included probable ancient intro-
fying natural ecosystems and threatening native species, ductions and introductions for conservation purposes outside
The process of biological invasion is now widely accepted to the natural range. We excluded unconfirmed prehistoric intro-
consist of a series of stages: introduction (or transport), estab- ductions, reintroductions for conservation purposes within the
lishment, and spread (Mack et al. 2000; Kolar and Lodge 2001). natural range, scientific trials, and all instances where there is
Terminology surrounding the phenomenon of biological inva- doubt (according to Long 2003) that a self-sustaining wild
sions has, however, been a subject of recent debate (Colautti and population was established. We included cases where a self-
MacIsaac 2004). sustaining wild population was subsequently eradicated by
Mammals were among the first organisms to be introduced human actions, but excluded those where populations died out
by humans: as livestock (sheep, goats, cattle), as companions naturally or where Long (2003) expresses any doubt about their
(dogs) or as commensals (rodents). More recently, many other fate. For analysis of the proportions of the various mammal
species of mammals have been deliberately introduced as sport- groups that are successful invaders, we used the global list of
ing animals, for novelty reasons, and (ironically) to control pre- wild mammals compiled by Ceballos et al. (2005), which incor-
viously introduced species that had proved to be pests. Overall, porates data from the IUCN Red List. For completeness, we
mammals are perhaps the best documented of all introduced added to this list eight domesticated mammal species – goat
organisms and have been the subject of two global compilations (Capra hircus), sheep (Ovis aries), horse (Equus caballus),
(Lever 1985; Long 2003) and some excellent regional records of
attempted and successful introductions (Thomson 1922; Myers donkey (Equus asinus), cattle (Bos taurus), water buffalo
1986; Jeschke and Strayer 2005). (Bubalus bubalis), dog (Canis familiaris), and cat (Felis catus)
The purpose of our review is first to examine differences – that are recorded by Long (2003) as having established truly
between mammal groups in their propensity to contain suc- unmanaged wild (feral) populations. For comparison with the
cessful invaders or threatened species. We then review the lit- proportions of the various mammal groups that are classed as
erature on determinants of invasion success by mammals and threatened (i.e. vulnerable, endangered or critical, according to
consider overall implications for the management of invasive 2006 IUCN Red List criteria), we also followed the global
mammals. mammal list compiled by Ceballos et al. (2005). For both
© CSIRO 2008 10.1071/WR07091 1035-3712/08/030180
The invasion ecology of mammals: a global perspective
analyses we excluded marine mammals (Cetacea, Pinnipedia,
Sirenia) and all extinct species.
The taxonomic distribution of mammalian invaders
Our dataset contains 4816 species of extant land mammals,
including domesticated species. Of these extant land mammals,
124 (2.6%) can be classed as ‘successful invaders’, according to
our criteria and the data compiled by Long (2003), whereas
1038 (21.6%) are classed as threatened, according to the 2006
IUCN Red List. Overall proportions of successful invaders
range from 0% (e.g. Chiroptera) to 14.7% (Artiodactyla),
whereas proportions of threatened species range from 0% (e.g.
Edentata) to 100% (Proboscidea) (Fig. 1). The orders
Artiodactyla (pigs, camels, deer, cattle, sheep, goats and ante-
lope), Carnivora (canids, bears, mustelids and cats),
Lagomorpha (rabbits and hares), and Perissodactyla (equines)
all contain high proportions of successful invaders. The
mammal family with the highest proportion of successful
invaders is Cervidae (29.2%). Only 1.8% of the order Rodentia
can be classed as successful invaders, but 13% of the genus
Rattus have this dubious distinction.
Wildlife Research 181
The absence of any successful invaders among over 1000
species of bats, compared with (for example) 32 successful
invaders among 217 species of Artiodactyla suggests that there
are differences in the propensity of different groups to be trans-
ported and establish outside their natural range. Contrasts in the
proportion of successful invaders between major taxonomic
groups seem to be largely due to differences in propensity for
deliberate introduction of species valued for hunting, as fur-
bearers, as domestic animals, as biological control agents, or as
easily transported novelties. For example, bats and insectivores
are less favoured than deer and rabbits. Beyond this initial
human ‘filter’, other biological determinants of success
become important.
A relatively small number of mammal species have success-
fully established at more than 30 locations around the world
(Long 2003). They include feral domestic animals (horses,
sheep, goats, cattle, pigs, donkeys, cats, dogs), European rabbits
(Oryctolagus cuniculus), red deer (Cervus elaphus), American
mink (Mustela vison), Indian mongoose (Herpestes javanicus)
and six species of rodents. The rodents include two deliberately
introduced species – coypu (Mycocastor coypus) and muskrat
Fig. 1. Percentage of successful invaders, threatened species and those in neither category, for each mammalian order. The number
of species in each order is shown above each bar. Nomenclature follows Long (2003) for orders Edentata, Insectivora and Marsupialia.
182 Wildlife Research M. N. Clout and J. C. Russell
(Ondatra zibethicus) – and four very widespread commensal
species that have been accidentally introduced to many locations
(Rattus rattus, R. norvegicus, R. exulans and Mus musculus).
Determinants of invasion success
Although less than 3% of the world’s mammals can be classed
as ‘successful invaders’, the data compiled by Long (2003)
reveal that many of these species have also been the subject of
unsuccessful introductions. Some other mammal species
(excluding those reintroduced for conservation purposes) have
never been successfully introduced, despite efforts to do so.
These observations raise the question of what determines inva-
sion success, a topic that has been the subject of several recent
studies.
From an analysis of the invasion success of mammals, fish
and birds in Europe and North America, Jeschke and Strayer
(2005) demonstrate that introduced vertebrates have a relatively
high probability (~50%) of establishing and spreading. They
conclude that the ‘tens rule’ of Williamson (1996) (that ~10% of
species will make each of these transitions) does not apply to
vertebrates. They also show that there is no clear difference in
invasion success of vertebrates introduced from Europe to
North America or vice versa.
The excellent records kept by Acclimatisation Societies in
New Zealand (Thomson 1922) have provided a fertile source of
data for analyses of factors affecting introduction success. In a
study of the outcomes of introductions of 14 ungulate species,
Forsyth and Duncan (2001) showed that the 11 successful
species had shorter maximum life spans and were introduced in
greater numbers than the unsuccessful ones. For all independent
introductions there was a highly significant relationship
between the number of individuals introduced and introduction
success, with an apparent threshold introduction size of about
six individuals, above which success was likely. A more recent
analysis by the same authors (Duncan and Forsyth 2006) exam-
ined population persistence of 164 introductions of six mammal
species (rabbit, goat, sheep, cat, pig and brushtail possum) to
85 islands in New Zealand. They show that small populations
were initially at greatest risk of extinction; those that survived
for 25 years were likely to persist for much longer; and the
probability of mammal populations persisting on islands
declined markedly with increasing latitude. A study of the
species richness of 17 large and small introduced mammals on
297 islands in New Zealand (Russell et al. 2004) revealed that
human transportation, facilitated by wharves and permanent
inhabitants, increased the likelihood of introduced mammals
being present, although small mammals were also dispersing by
swimming to many islands.
Forsyth et al. (2004) analysed data on 40 mammal species
introduced to mainland Australia, 23 of which successfully
established. They found that successful species had a greater
area of climatically suitable habitat available to them, had larger
overseas ranges, and were introduced more times. Overall
results were similar to those obtained for introduced birds in the
same region (Duncan et al. 2001). In both groups, established
species that had spread to occupy a large range in Australia also
tended to be species of small body masses with more offspring
per year. Among mammals, these more widespread species
tended to be carnivores or omnivores, rather than herbivores
(Forsyth et al. 2004). Burbidge and Manly (2002) demonstrated
a relationship between native mammal extinctions on Australian
Islands and the presence of introduced European foxes (Vulpes
vulpes) and feral cats.
Implications for the management of invasive mammals
Some introduced mammals have not only successfully estab-
lished self-sustaining populations, but have also spread and have
caused significant impacts in the ecosystems that they have
invaded. Such species can be classed as invasive (‘widespread
and locally dominant’), according to the definition of Colautti
and MacIsaac (2004). Examples of indisputably invasive
mammals are the three widespread Rattus species (Amori and
Clout 2003), European rabbits, feral cats, several species of
deer, several mustelids and viverrids, and some other top preda-
tors such as the European fox and Arctic fox (Alopex lagopus).
Further introductions (or range extensions) of such invasive
species should be prevented wherever possible, and their eradi-
cation undertaken where feasible. The eradication of introduced
mammals has progressed substantially in recent decades (Veitch
and Clout 2002), especially from islands in countries such as
New Zealand (Clout and Russell 2006). Removal of a range of
mammals from increasingly large areas is now possible using a
range of techniques. Successful eradications have now included
the removal of rats from islands as large as 11 300 ha (Towns and
Broome 2003), and the eradication of coypu from a large area of
south-eastern England (Gosling and Baker 1989).
Changes to ecosystem processes
Losses of native biodiversity (including the extinction of
endemic species) as a consequence of invasion by new mammal
species have now been widely documented, especially in iso-
lated ecosystems such as New Zealand (Atkinson 2006) and
other oceanic archipelagos. It is clear that invasive mammals are
capable of causing radical change to ecosystems that they
invade, not only by extinguishing native prey species and alter-
ing plant communities, but also by disrupting patterns of nutri-
ent flow. An example of such effects is the trophic cascade
generated by the introduction of Arctic foxes to the Aleutian
archipelago (Croll et al. 2005). By preying on seabirds and
greatly reducing their abundance, the foxes also reduced the
input of marine nutrients (imported by seabirds to their breed-
ing colonies) to invaded islands. This subsequently changed the
soil nutrient regime and caused vegetation to change from grass-
land to tundra. Similar indirect effects have been obtained in
studies of the impacts of Rattus species on offshore islands of
New Zealand (Fukami et al. 2006). Comparison of rat-free and
rat-invaded islands revealed that predation of seabirds by rats
reduced forest soil fertility by disrupting nutrient transport by
the seabirds. This fertility reduction led, in turn, to effects on
below-ground organisms and the ecosystem processes driven by
them. Carbon sequestration in live plants was also indirectly
enhanced on the rat-invaded islands (Wardle et al. 2007). In a
similar vein, Wardle (2006) has suggested that introduced
browsing mammals not only directly affect vegetation but also
indirectly alter the decomposer subsystem.
The ultimate ecological impacts of the many introductions of
species that have occurred worldwide in recent centuries and
decades have yet to be fully felt or understood (Strayer et al.
The invasion ecology of mammals: a global perspective
Fig. 2. Eradications and reinvasions of rats on New Zealand islands,
1960–2005. Data extracted from Clout and Russell (2006).
2006). Many introduced mammals are yet to attain the full
extent of their distributions, and global climate change may
influence the potential range that species could occupy. Local
genetic adaptation and even divergent evolution are also possi-
ble over long periods (Gleeson et al. 2006). It is nevertheless
clear that the risk of new invasions should be minimised and the
removal of existing introduced mammals should be seriously
considered where possible.
Interactions between invaders
A topic that merits greater attention when planning eradications
is ecosystem responses to species removals. Ecosystems that
contain multiple invaders, have lost native species along with
their functional roles, or have suffered long-term change to
soils, can respond to eradications in unexpected ways (Zavaleta
2002). Interactions between introduced mammals and other
invasive species may generate positive feedbacks whereby inva-
sion by different species is enhanced (e.g. Pitman et al. 2005), a
process known as ‘invasional meltdown’ (Simberloff 2006).
Interactions between introduced mammals need to be properly
understood before eradication or control of one or all of the
introduced mammal species at a site is undertaken (Courchamp
et al. 2003). For example, ‘mesopredator release’ (Courchamp
et al. 1999) may become evident if a top predator is removed
whilst leaving other predators over which this predator may have
exerted some previous control. ‘Competition release’ may simi-
larly occur when an introduced competitor is removed from a
system (Caut et al. 2007). More research is needed to under-
stand such potential effects.
Risks of reinvasion
A final subject that merits more attention in relation to planning
the eradication of introduced mammals is the risk of reinvasion.
For example, along with recent successes in eradicating rodents
from islands there have been several reinvasions. This is illus-
Wildlife Research 183
trated by data extracted from Clout and Russell (2006), plotting
the number of rat eradications from islands in New Zealand
alongside the number of rat reinvasions (Fig. 2). Eradications
peaked in the early 1990s, but reinvasions have risen since then,
with an apparent lag of ~10 years. Given the current substantial
investment in eradications of introduced mammals from islands
around the world, it is vital that we achieve better understanding
of how mammals (especially highly dispersive species such as
rodents) may be transported or disperse back to sites from which
they have been cleared. It is particularly crucial that we research
the behaviour and detectability of individual dispersers that are
likely to be the founders of new populations. Some experimental
work has been undertaken on rats in this context (Russell et al.
2005, 2008) but more is needed.
Conclusion
Although only a small proportion of the world’s land mammals
can be classed as ‘successful invaders’, their ecological impacts
are sometimes immense. Many introduced species are yet to
attain the full extent of their global and regional distributions.
Understanding what determines invasion success, and how
introduced mammals interact with each other and with other
components of invaded ecosystems, is of fundamental scientific
interest (Sax et al. 2005; Cadotte et al. 2006). Such research is
also vital in underpinning continued attempts to reverse some of
the negative effects of introduced mammals on the world’s
native biodiversity.
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