(>740 species). Angophora and Corymbia are often treated
as subgenera of Eucalyptus, sensu lato. The genus Eucalyp-
tus (sensu stricto) is currently divided into ten subgenera,
six of which are monotypic (having only one species).
Naturalized eucalypts belong almost exclusively to the
two largest subgenera, Eucalyptus (Monocalyptus, >140
species) and Symphyomyrtus (>360 species) (Table 1).
Most eucalypts are trees (10 to >50 m in height), some
are “mallees” (multistemmed from ground level, usually
<10 m in height), and a few are shrubs. Eucalypts are the
tallest nonconiferous trees in the world. Several species
(E. regnans, E. grandis, E. deglupta) reach more than 70 m
in height. The tallest known specimen of E. regnans was
110 m. The tallest known living specimen of this taxon in
Tasmania is 99.6 m tall. The maximum age of eucalypt
species, estimated from dendrochronological (tree-ring)
and radiocarbon measurements, is between 400 and 600
years. Interestingly, in spite of the otherwise enormous
range of adaptations among eucalypts, shade-tolerant
subcanopy species are not known.
The eucalypt breeding system is one of mixed mating
with preferential outcrossing. A reduction in fruit (cap-
sule) production, seed set, and seedling vigor has been
demonstrated after self-pollination. However, eucalypts
generally do not need any special pollinators. They are
pollinated by many species of bees and wasps and, to a
lesser extent, by birds, mammals, and wind.
Most eucalypts are well adapted to frequent ﬁ res. The
most common adaptations are lignotubers and epicormic
buds. The lignotuber is a woody swelling at the base of
the stem; most eucalypts sprout from lignotubers. The
epicormic buds (buds present in the outer bark) allow the
sprouting of new branches from stems (after a ﬁ re or after
a severe winter). Some species are able to sprout from both
lignotubers and stems (combination sprouters). Another
adaptation to ﬁ reprone environments is serotiny (most
seeds are kept in fruits and released only after ﬁ re). Some
successful invaders among eucalypts are obligate seeders,
depending completely on seed production (E. conferru-
minata, E. grandis). Seeds can be shed in large numbers
(up to 4,000 seeds per m2). However, eucalypts produce
very small seeds, usually 1–3 mm long and less than 2 mg.
Some species have seeds even lighter than 0.5 mg (e.g.,
E. grandis and deglupta). Eucalypts produce seeds with no
obvious endosperm (tissue that surrounds and nourishes
the embryo). Therefore, the newly emerged seedlings are
sustained by cotyledon photosynthesis, and their roots
have to reach suitable substrate very soon.
Eucalypts are the dominant species of the wet coastal
and near coastal parts of Australia. However, some species
SEE ALSO THE FOLLOWING ARTICLES
Early Detection and Rapid Response / Endangered and Threatened
Species / Herbicides / Invasion Economics / Islands / Lag Times /
Risk Assessment and Prioritization
Clout, M. N., and P. A. Williams, eds. 2009. Invasive Species Management:
A Handbook of Techniques. Oxford: Oxford University Press.
Genovesi, P. 2007. Limits and potentialities of eradication as a tool for
addressing biological invasions (385–400). In W. Nentwig, ed. Biological
Invasions. Ecological Studies, Vol. 193. Heidelberg: Springer-Verlag.
McNeely, J. A. 2001. The Great Reshufﬂ ing: Human Dimensions of Invasive
Alien Species. Gland, Switzerland: IUCN.
Myers, J. H., D. Simberloff, A. M. Kuris, and J. R. Carey. 2000. Eradi-
cation revisited: Dealing with exotic species. Trends in Ecology and
Evolution 15: 316–320.
Veitch, C. R., and M. N. Clout, eds. 2002. Turning the tide: The eradication
of invasive species. Occasional Paper 26. Gland, Switzerland: IUCN.
University of California, Davis
DAVID M. RICHARDSON
Stellenbosch University, Matieland, South Africa
Over 800 species of eucalypts (Angophora, Corymbia, and
Eucalyptus) are native to Australia and a few Paciﬁ c islands.
These genera include some of the most important solid
timber and paper pulp forestry trees in the world. Besides
pines, eucalypts are the most commonly and widely cul-
tivated exotic trees. Almost 20 million ha (200,000 km2)
of eucalyptus plantations exist in tropical, subtropical, and
temperate countries. In many countries, eucalypts are the
most common and conspicuous nonnative trees. Over 70
species are naturalized (reproduce and maintain their pop-
ulations) outside their native ranges. However, given the
extent of cultivation, eucalypts are markedly less invasive
than many other widely cultivated trees and shrubs. Rea-
sons for this relatively low invasiveness are still not com-
pletely understood. Conclusions about positive or negative
environmental and economic impacts of eucalypts are often
anecdotal, highly controversial, and context-dependent.
TAXONOMY, GROWTH FORMS, REGENERATIVE
STRATEGIES, AND HABITATS
Eucalypts (family Myrtaceae, subfamily Leptosper-
moideae) are currently classiﬁ ed into three genera: Ango-
phora (14 species), Corymbia (113 species), and Eucalyptus
05_Simberloff10_E_p169-222.indd 20305_Simberloff10_E_p169-222.indd 203 9/13/10 10:02:33 AM9/13/10 10:02:33 AM
From Daniel Simberloff and Marcel Rejmánek, editors, Encyclopedia of Biological Invasions,
Berkeley and Los Angeles: University of California Press, 2011.
Naturalized Species of Corymbia and Eucalyptus
calophylla C, LS Australia (Western Australia), Hawaii?, New Zealand?
citriodora B, LS Australia (Victoria, Western Australia), California, Hawaii?, India, South Africa, Zimbabwe
ﬁ cifolia C, LS Hawaii, New Zealand, South Africa
maculata B, LS Australia (Victoria, Western Australia), South Africa?
torelliana B, LS Australia (Queensland), China, Florida
botryoides S, CS Australia (Norfolk Island, South Australia, Victoria, Western Australia), Hawaii, New Zealand
bridgesiana S, LS Hawaii, South Africa?
camaldulensis S, CS/SS Argentina?, Australia (Western Australia) Bangladesh, California,a,b Cyprus, France, Greece,
Hawaii,a India, Israel, Italy, Pakistan, Portugal, Spain,a,b South Africa,a,b Zimbabwe
cinerea S, LS Hawaii, New Zealand, South Africa
cladocalyx S, SS Australia (Victoria, South Australia, Western Australiab), California, Hawaii?, South Africaa
conferruminata S, OS, often confused Australia (Victoria, Western Australia), California, South Africaa,b
with E. lehmanii
crebra S, LS Hawaii, South Africa?
deanei S, LS Hawaii, South Africa?
deglupta S, OS Hawaii, Malaysia?
delegatensis E, OS New Zealand, South Africa?
elata E, LS New Zealand, South Africa?
fastigata E, OS California, New Zealand, South Africa
globulus S, CS, incl. E. maidenii Australia (Western Australia), Azores, California,a,b Canary Islands, Chile,a China,a France,
Hawaii,b India, Italy?, Jamaica, New Zealand,a,b Pe ru,b Portugal,a Spain,a South Africa,a
gomphocephala S, SS Australia (Victoria), Cyprus, Hawaii, South Africa, Spain
grandis S, OS Argentina?, California, Florida, New Zealand, Nigeria?, South Africaa,b
gunnii S, LS New Zealand, Portugal, Spain
leucoxylon S, LS Australia (Victoria), New Zealand?, South Africa
longifolia S, LS Australia (New South Wales), South Africa?
macarthurii S, LS California, New Zealand, South Africa?
marginata E, LS Hawaii, South Africa?
microcorys S, LS Australia (Western australia), Hawaii,a Sri Lanka, South Africa, Zimbabwe
muelleriana E, LS Australia (Western Australia), New Zealand?, South Africa?
nitens S, SS New Zealand, South Africa?a
obliqua E, LS New Zealand, South Africa
occidentalis S, LS Australia (Victoria), South Africa?
ovata S, LS California, New Zealand
paniculata S, LS Hawaii,a South Africa,a Zimbabwe
pilularis E, OS Hawaii,a New Zealand, South Africa
polyanthemos S, LS Australia (Western Australia), California, South Africa?
pulchella E, LS California, New Zealand
regnans E, OS New Zealand, South Africa?
resinifera S, LS Hawaii, Mexico?, New Zealand
robusta S, OS Brazil?, California, Florida, France, Hawaii,a La Réunion Island, Madagascar,a Malaysia?,
New Zealand, Portugal, South Africa, Spain, Sri Lanka, Zimbabwea,b
rudis S, LS California?, Hawaii
saligna S, LS Australia (Western Australia), Florida, Hawaii,a New Zealand, South Africa?, Sri Lanka,?,
salubris S, OS Australia (Queensland), South Africa
sideroxylon S, CS Botswana?, California, Hawaii, New Zealand, Portugal, South Africa,a Spain
sieberi E, SS New Zealand, South Africa
tereticornis S, LS California, Cyprus, Hawaii, India,a Mexico (reported as E. resinifera), New Zealand?, South
viminalis S, LS California, Hawaii, New Zealand, South Africa?
note: Naturalized (reproducing and maintaining populations without human help) species of Corymbia and Eucalyptus. Based on published records, labels on herbarium
specimens, information from experienced botanists and foresters, and personal observations. Question marks indicate lack of certainty as to whether the species is already
naturalized, or is just a casual resident. Only species reported as naturalized from at least two countries are included. Subgenera: B – Blakella, C – Corymbia, E – Eucalyptus
(Monocalyptus), S – Symphyomyrtus. Regenerative strategies: LS – lignotuber sprouter, SS – stem sprouter, CS – combination sprouter, OS – obligate seeder.
a Extensive planting.
b Species is classiﬁ ed as invasive in a particular state (spreading spontaneously away from points of introduction).
05_Simberloff10_E_p169-222.indd 20405_Simberloff10_E_p169-222.indd 204 9/13/10 10:02:33 AM9/13/10 10:02:33 AM
area of more than 52 million ha. The most commonly cul-
tivated eucalypt is probably E. globulus (Tasmanian blue
gum, with 2.3 million ha worldwide in 2008). Eucalyptus
globulus is the primary source of global eucalyptus oil pro-
duction, with China being the largest commercial producer.
The other three most commonly cultivated eucalypts are
E. camaldulensis (river red gum), E. grandis (ﬂ ooded gum),
and E. tereticornis (forest red gum). Current estimates of
areas of eucalyptus plantations on individual continents
and in Oceania are summarized in Table 3. The American
statistic is dominated by Brazil, with over 3.5 million ha of
plantations, while in Asia the leader is India, with over 5
million ha, including small cultivations on farms.
Eucalypt planting has accelerated recently in many trop-
ical countries. However, extensive planting in the humid
tropics has been inhibited by the incidence of pathogens
and insect pests. Only a few species, such as E. deglupta, E.
pellita, and E. urophylla, appear to be adapted to hot, humid
environments. Eucalypts, because of their rapid growth and
capacity for producing biomass, have been recently widely
mentioned as feedstock for biofuels (e.g., E. globulus, E.
grandis, E. robusta, E. saligna, E. urophylla). Widespread use
for this purpose could substantially increase propagule pres-
sure and increase the probability of local invasions.
ARE EUCALYPTS INVASIVE?
Eucalypts have enjoyed a history of widespread planting
similar to that of pines. Given this, and the many spe-
cies involved (with representatives of most of the major
taxonomic and functional groups in the genus), we would
expect to ﬁ nd the full range of outcomes: everything from
taxa that are highly invasive and cover large areas as invad-
ers to species that fail to recruit any offspring. Indeed, euca-
lypts, like pines, are prominent species on many national
or regional weed lists in many parts of the world. How-
ever, they have been orders of magnitude less successful as
(E. rameliana, E. pachyphylla) extend to arid regions with
annual precipitation below 350 mm. Within their native
range, most eucalypt species grow in tall forests, wood-
lands, and savannas. Seasonal ﬂ ooding is essential for suc-
cessful regeneration of some species (E. camaldulensis).
Over 20 species of eucalypts can grow on saline soils (e.g.,
E. robusta in swampy estuarine habitats or E. camaldulensis
in valleys of old river systems). Some small trees or shrubs
(e.g., E. coccifera or E. pauciﬂ ora subsp. niphophila) are
adapted to subalpine areas (900–1,400 m) in Australia.
When Australia was settled in the eighteenth century,
eucalypts were used for farm buildings, fencing, and ﬁ re-
wood. The ﬁ rst eucalypt species to be cultivated from seeds
outside its native range was E. obliqua in Royal Botanic
Gardens at Kew, United Kingdom, in 1774. Other species
were soon cultivated in botanical gardens and arboreta
Areas of Eucalyptus Plantations beyond Their Native Range in 2008
Asia 8.3 (India: >5.0; China: >2.0; Vietnam:
Americas 6.4 (Brazil: >3.6; Chile: >0.3; Argentina:
>0.2; Peru: >0.2)
Africa 2.2 (South Africa: >0.4; Angola: >0.2;
Europe 1.3 (Portugal: >0.6; Spain: >0.5;
cultivated.html and other sources.
Naturalized Corymbia and Eucalyptus by Regenerative Strategies
Number Regenerative Number
Subgenus of Species Strategy of Species
Blakella 3 Lignotuber 28
Corymbia 2 Stem sprouters 4 (1 invasive)
Eucalyptus 10 Combination 4 (2 invasive)
Symphyomyrtus 30 (6 invasive) Obligate seeders 9 (3 invasive)
note: Numbers of naturalized Corymbia and Eucalyptus species by subgenera
and by regenerative strategy. Only species reported as naturalized in at least two
countries are included. Derived from Table 1.
in Europe as botanical curiosities and ornamentals. Once
in cultivation, eucalypts appealed to foresters because
of their fast growth (even on nutrient-poor soils) and
because they yielded a variety of timber and nontimber
products. Productivity of properly designed plantations is
20–70 m3 ha–1 year–1. Today, eucalypts provide saw tim-
ber, plywood, ﬁ berboard, pulp for paper, poles, ﬁ rewood,
charcoal, essential oils, honey, and shelter. They are also
considered to be suitable trees for biofuel production.
To date, the total area of eucalyptus plantations has
been increasing exponentially. The global extent of euca-
lypt planting outside Australia reached 15.6 million ha in
the 1990s—more than four times the global total in the
1970s. In 2008, the total area of eucalyptus plantations was
estimated at 19.6 million ha. This is an area larger than the
state of Washington, or the nations of Austria and Hungary
together. Only one genus of trees is more extensively culti-
vated than eucalypts: Pinus (pines), with a global plantation
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2.0 and 5.5 m s–1. Lower terminal velocity values mean that
seeds can be carried by winds for longer distances.
HIGH MORTALITY OF SEEDLINGS. As noted above, euca-
lypts produce very small seeds (usually <2 mg) with no
obvious endosperm. Therefore, the newly emerged seed-
lings are sustained by cotyledon photosynthesis, and their
roots have to penetrate into suitable wet substrate very
quickly. As a result, eucalypts can successfully regenerate
from seeds only on wet, bare soil free of litter. However,
seedlings in wet environments frequently die because of
damping off caused mainly by parasitic fungi (Botrytis,
Colleotrichum, Cylindrocladium, Fusarium) and water
molds—oomycetes (Phytophthora, Pythium). Moreover,
if there is any dense vegetation around, tiny eucalyptus
seedlings are necessarily losers. Therefore, the window of
opportunity for eucalyptus seedlings is rather narrow.
LACK OF COMPATIBLE ECTOMYCORRHIZAL FUNGI. It
seems that the majority, if not all, of ectomycorrhizal
fungi (EM) associated with eucalypts outside Australia are
Australian species that have spread with their hosts. How-
ever, the importance of EM for establishment, growth,
and spread of introduced eucalypts is not clear. There has
been no example reported from exotic eucalypts that can
compare with the dramatic response to mycorrhizal inoc-
ulation reported from exotic pine seedlings in Australia
or South Africa in the early years of their introduction.
With their more ﬁ nely branched root systems and evo-
lutionary adaptation to low phosphorus soils, eucalypts
seem to be less dependent on EM than pines. The lack
of EM may be not important for seedlings transplanted
from nurseries, but, as some ecologists have suggested, it
may be crucial for spontaneous establishment of seedlings
away from plantations. However, colonization of eucalypt
roots usually does not start with EM, but with ubiquitous
nonspeciﬁ c vesicular-arbuscular endomycorrhizae (AM)
(Gigaspora spp., Glomus spp.). It is possible that AM play
a more important role in initial eucalypt seedling estab-
lishment than EM.
Finally, we may ask whether some eucalypt species are
inherently more invasive than others. Or is it only propagule
pressure (the extent of planting) that makes some species
more often naturalized, and some of them even somewhat
invasive (spreading spontaneously over 100 m from points
of introduction)? Tables 1 and 2 show that the majority of
species naturalized in at least two countries (30) belong to
the subgenus Symphyomyrtus. This seems to be just propor-
tional to the size of this subgenus (>360 species). Still, all
six species that can be classiﬁ ed as invasive in at least one
invaders than pines and several other widely planted trees,
including many ﬂ eshy-fruiting trees (e.g., Elaeagnus angus-
tifolia, Ligustrum spp., Psidium spp., Morella (Myrica) faya)
and legumes (e.g., Acacia spp., Prosopis spp., Leucaena leu-
cocephala). Where eucalypts have invaded, they have very
seldom spread considerable distances from planting sites,
and their regeneration is frequently sporadic (Fig. 1). Given
that many eucalypts produce very large quantities of seeds,
and in light of their diverse adaptations for dealing with
disturbance (notably ﬁ re), their poor (or at best mediocre)
performance as invaders is enigmatic. Many other Austra-
lian trees, including taxa that evolved under the same con-
ditions as Eucalyptus, are much more invasive in other parts
of the world (e.g., Melaleuca quinquenervia, Hakea spp.,
and many species of Acacia). What makes this difference?
Are eucalypts inherently less invasive, or are they just a tick-
ing time bomb? There seem to be three major reasons for
the limited invasiveness of eucalypts: (1) relatively limited
seed dispersal, (2) high mortality of seedlings, and (3) lack
of compatible ectomycorrhizal fungi.
RELATIVELY LIMITED SEED DISPERSAL. Seeds of planted
eucalypts are very small, but they have no adaptations for
dispersal (wings or ﬂ eshy tissues) that would help them to
proceed from local establishment (naturalization) to inva-
sion. The passive release of seeds is undoubtedly aided by
wind. However, all rigorous studies of eucalypt seed disper-
sal and seedling spatial distribution show that in general,
seeds are dispersed over quite short distances. This is in
agreement with measurements of terminal descent veloci-
ties of their seeds. While terminal velocities of seeds of
invasive pine species are between 0.7 and 1.5 m s–1, and for
invasive maples (Acer) between 0.9 and 1.2 m s–1, terminal
velocities of seeds of all tested eucalypt species are between
FIGURE 1 A stand of Eucalyptus globulus on Santa Cruz Island,
California. Saplings of eucalypts are usually found only close to planta-
tions. (Photograph courtesy of M. Rejmánek.)
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Of all widely used plantation species, eucalypts have
attracted by far the most criticism. There are four main
concerns: (1) excessive water use and suppression of food
crops growing nearby, (2) suppression of ground vegetation
(possible allelopathic effects) and resulting soil erosion,
(3) increased ﬁ re hazard, and (4) generally poor wildlife
value. There is some substance to each of these concerns.
country belong exclusively to the subgenus Symphyomyr-
tus. There seems also to be a somewhat larger proportion
of subgenus Eucalyptus species among commonly planted
eucalypts with no evidence of naturalization (Table 4). On
average, seeds of species in the subgenus Symphyomyrtus
are smaller (a notable exception, however, is E. globulus)
than seeds of species in the second largest subgenus—
Eucalyptus (>140 species). Therefore, terminal velocities
of Symphyomyrtus seeds should be less, and they could be
dispersed somewhat longer distances by wind. Also, none
of the species classiﬁ ed as invasive in at least one country
is a straight lignotuber sprouter (LS). This could also point
to the primary importance of seed dispersal in invasive-
ness. Nevertheless, the more straightforward explanation is
that species classiﬁ ed as invasive have been planted more
extensively (see notes in Table 1). Then, however, reasons
for their extensive planting may be correlated with their
growth and timber characteristics that are associated with
particular regenerative strategies.
ENVIRONMENTAL CONTROVERSIES AND
Considering the amount of planting, eucalypts are rela-
tively noninvasive species. If their potential spread is
the only concern, then eucalypts should not be planted
near rivers and streams. Temporarily ﬂ ooded or eroded
banks are suitable habitats for spontaneous establishment
of their seedlings (Fig. 2). Moreover, their seeds can be
dispersed for long distances by running water. However,
there are other concerns.
FIGURE 2 Eucalyptus camaldulensis
is invasive along hundreds of kilome-
ters of rivers in South Africa’s West-
ern Cape province. The images show
(A) a general view of E. camdulensis
established in the Berg River, (B) the
extreme persistence of adult plants
due to their ability to sprout from the
roots, and (C) microsites for seedling
establishment. (Photographs cour-
tesy of D. M. Richardson.)
Eucalyptus Species with No Conclusive Evidence of Naturalization
Species Subgenus Regenerative Strategy
acmenoides Eucalyptus (Monocalyptus) Lignotuber sprouter
bosistoana Symphyomyrtus Lignotuber sprouter
erythrocorys Eudesmia Lignotuber sprouter
fraxinoides Eucalyptus (Monocalyptus) Obligate seeder
jacksonii Eucalyptus (Monocalyptus) Lignotuber sprouter
macranda Symphyomyrtus Lignotuber sprouter
pauciﬂ ora Eucalyptus (Monocalyptus) Lignotuber sprouter
preissiana Eucalyptus (Monocalyptus) Lignotuber sprouter
pulverulenta Symphyomyrtus Lignotuber sprouter
punctata Symphyomyrtus Lignotuber sprouter
radiata Eucalyptus (Monocalyptus) Lignotuber sprouter
smithii Symphyomyrtus Lignotuber sprouter
spathulata Symphyomyrtus Obligate seeder
torquata Symphyomyrtus Lignotuber sprouter?
note: Commonly cultivated Eucalyptus species with no conclusive evidence of
naturalization. Species listed here are relatively commonly cultivated or tested,
but the extent of their cultivation is still very limited when compared with many
major plantation species listed in the Table 1. Therefore, “no conclusive evidence of
naturalization” does not necessarily mean conclusive noninvasiveness.
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Because of accumulated litter, dense eucalypt stands are
extremely ﬂ ammable. The situation is exacerbated after
winter freezes, when trees drop dead branches and foliage.
During the last two weeks of 1990, a mass of frigid arctic air
moved into California, and temperatures plunged to record
lows along the Paciﬁ c Coast. It is very likely that fuel accu-
mulation in unmanaged eucalyptus stands contributed to
the intensity of the tragic ﬁ re in the Berkeley–Oakland Hills
area in October 1991 (Fig. 3).
In arid and semiarid countries, where shortage of water
is a big concern, beneﬁ ts of eucalypt plantations may be
outweighed by their negative environmental impacts:
namely, their high water consumption. In South Africa,
invasive eucalypts have been cleared over large areas as
part of a national restoration program called Working for
Water. In most areas, standing trees are felled and, where
it is practically possible, their timber is harvested. Where
recovery of the timber is impractical, felled plants are often
stacked and burned. The most challenging management
operations involve clearing river banks in large parts of
the Western Cape province of dense stands of invasive E.
camaldulensis. Clearing alone often causes destabilization
of the river banks, and research is under way to determine
the most effective ways of thinning the invasive stands
gradually while simultaneously reintroducing key native
plants to stabilize the sites and launch succession toward a
sustainable community dominated by native plants.
The fact that eucalypt seeds do not have dormancy, with
seed storage in the soil lasting less than a year, makes local
Nevertheless, it is important to realize that in many tropi-
cal countries, where eucalypts are grown on degraded soils
unsuitable for continuing to support native trees (usually
abandoned agricultural land), fuel and other products of
resprouting eucalypts can greatly reduce the increasing
human pressure on the remnants of natural forests. Even
here, however, deleterious human practices associated with
consecutive cutting cycles may eventually lead to yield
decline and forest site degradation on a long-term basis.
For long-term site quality and sustainability of biomass
production, prolonging the cutting cycles and prohibiting
or controlling litter raking appears to be imperative.
Eucalypts may be a major source of both nectar and
pollen for honeybees. Because ﬂ owering of many euca-
lypts is abundant and lasts for long periods, some spe-
cies (particularly E. camaldulensis and E. cladocalyx) are
very valuable for the honey industry. When compared
with native vegetation, usually signiﬁ cantly lower spe-
cies diversity of arthropods, small mammals, and birds is
reported from eucalyptus stands. For example, in Angel
Island State Park, California, 41 species of birds were
observed in native vegetation, but only 30 species in the
eucalyptus forest. However, there may be also some other
trends: approximately three times more California slender
salamanders (Batrachoseps attenuatus) were found in euca-
lyptus vegetation than in native. Even more importantly,
in California, eucalypts are major providers of shelter and
nectar to the migrating monarch butterﬂ y (Danaus plex-
ippus) during winter months.
Allelopathic effects of eucalypts on native species are
widely reported. Such reports are mostly based on labo-
ratory bioassays. However, some ﬁ eld trials also point to
decline of seed germination and increase of seedling mor-
tality of some native species. If not chemical inhibition,
then at least accumulation of dead material on the ﬂ oor of
eucalypt plantations retards regeneration of native species.
Mixed-species plantations of eucalypts with native (mainly
nitrogen-ﬁ xing) species have the potential to increase pro-
ductivity while maintaining soil fertility and biodiversity.
Tasmanian blue gums (E. globulus) were planted in the
San Francisco area of California as early as the second half
of the nineteenth century. Having been in this landscape
for such a long time, many old eucalypts are now treated
as trees with “historical value” or as “heritage trees.” Many
people feel that eucalypts give California a “unique exotic
ﬂ avor” lacking in other parts of the United States. This is
the reason why removal of eucalypts on Angel Island in the
San Francisco Bay (1990–1996) sparked a raging contro-
versy. In a very balanced way, the history of this episode was
described by Peter Coates.
FIGURE 3 It is undeniable that unmanaged stands of some eucalypt
species can accumulate highly ﬂ ammable dead material. To what
extent Tasmanian blue gum (Eucalyptus globulus) groves contributed
to the intensity of the tragic Berkeley–Oakland Hills ﬁ re in 1991 remains
a subject of bitter discussions. This hazy but dramatic photograph of
the 1991 ﬁ re was shot by former ﬁ re captain Wayne Drager through a
plexiglass window, using a disposable camera on a bumpy helicopter
ﬂ ight. (Photograph courtesy of Wayne Drager.)
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EUTROPHICATION, AQUATIC 209
contain less than 5–10 µg L-1 phosphorus and less than 250–
600 µg L-1 nitrogen. These nutrient concentrations are at
least 2 to 10 times as high in eutrophic waters and can have
major effects on biotic communities, including the loss of
biodiversity and the invasion of nonnative species (Fig. 1).
NATURAL AND ANTHROPOGENIC
Eutrophication is a slow natural process. The slow accu-
mulation of nutrients is especially prevalent in deposi-
tional environments, such as lakes and wetlands, where
nutrients and sediments derived from a watershed are
collected in a basin and permanently or temporarily
immobilized and stored. For example, estuaries are natu-
rally eutrophic, and hence very productive, because they
receive nutrients derived from watersheds and tidal ﬂ ows.
Lakes accumulate sediments and organic matter and over
time convert into productive and nutrient-rich shallow
lakes and emergent marshes. Other naturally eutrophic
systems are areas along the coast where upwelling con-
veys nutrient-rich water to the surface. As an ecosystem’s
nutrient status changes over millennial time scales, so
does its community structure, with local extinction and
colonization of new species working in concert to pro-
duce species-rich and productive ecosystems.
However, the process of eutrophication can be greatly
accelerated by human activities, such as runoff of excess fer-
tilizer, sewage efﬂ uent, and stormwater runoff. In Australia,
for example, sites affected by human activity have mean lev-
els of 780 µg L-1 N and 95 µg L-1 P compared to 300 µg L-1
N and 21 µg L-1 P at less impacted sites. Because estuaries
eradication an achievable goal. However, resprouting of cut
trees from stumps or lignotubers, which is advantageous in
some situations, makes control of eucalypts difﬁ cult. Con-
tinuously cutting back the regrowth can eventually kill
the tree, but this is a labor-intensive and expensive control
method. Herbicide applications (triclopyr or glyphosate) to
freshly cut stumps can greatly reduce resprouting. Because
eucalypts are valued as timber and ornamental trees in many
settings, biological control is very unlikely as an option.
SEE ALSO THE FOLLOWING ARTICLES
Allelopathy / Fire Regimes / Forestry and Agroforestry /
Invasiveness / Mycorrhizae / Propagule Pressure / Trees and Shrubs
Coates, P. 2006. American Perceptions of Immigrant and Invasive Species.
Berkeley: University of California Press.
Díez, J. 2005. Invasion biology of Australian ectomycorrhizal fungi intro-
duced with eucalypt plantations into the Iberian Peninsula. Biological
Invasions 7: 3–15.
Doughty, R. W. 2000. The Eucalyptus: A Natural and Commercial History
of the GumTree. Baltimore: The Johns Hopkins University Press.
Keane, P. J., G. A. Kile, F. D. Podger, and B. N. Brown, eds. 2000. Dis-
eases and Pathogens of Eucalypts. Collingwood, Australia: CSIRO
Nicolle, D. 2006. A classiﬁ cation and census of regenerative strategies in
the eucalypts (Angophora, Corymbia and Eucalyptus—Myrtaceae), with
special reference to the obligate seeders. Australian Journal of Botany
Poore, M. E. D., and C. Fries. 1985. The Ecological Effects of Eucalyptus.
FAO Forestry Paper 59: 1–87. Rome: FAO.
Rejmánek, M., D. M. Richardson, S. I. Higgins, M. J. Pitcairn, and E.
Grotkopp. 2005. Ecology of invasive plants: State of the art (104–161).
In H. A. Mooney, R. N. Mack, J. A. McNeely, L. E. Neville, P. J. Schei,
and J. K. Waage, eds. Invasive Alien Species: A New Synthesis. Washington,
DC: Island Press.
Ritter, M., and J. Yost. 2009. Diversity, reproduction, and potential for
invasiveness of Eucalyptus in California. Madroño (in press).
Slee, A. V., M. I. H. Brooker, S. M. Duffy, and J. G. West. 2006. Euclid:
Eucalypts of Australia, 3rd ed. Collingwood, Australia: CSIRO.
Williams, J., and J. Woinarski, eds. 1997. Eucalypt Ecology: Individuals to
Ecosystems. Cambridge: Cambridge University Press.
KATHARINA A. M. ENGELHARDT
University of Maryland Center for Environmental Science,
Eutrophication is the natural or anthropogenic accumula-
tion of nutrients in soil or water (from Greek eu = “well” and
trophe = “nourished”). Oligotrophic (low-nutrient) waters
FIGURE 1 The impacts of eutrophication on aquatic ecosystems.
(Figure developed by the Integration and Application Network of the
University of Maryland Center for Environmental Science.)
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