296 Chapter 9
Maintaining An Open Mind on Nonnative Species
Ariel E. Lugo, l,Tsiitute of Tt-opícnl Forestrt¡, Puerto Rico
"7iJ ct)Dsidct Lmly tlrc i t adi¡tg species
tltnrcelres in tlcrelop¡ttg ttt nget,]l.ltt pt1)
g1'a s or i¡i rec1¡lutsldiltg regulalt)1'tl
act¡u1s is Latltntln¡fit to ./ü¡itg sytltpto]fit
a¡td ¡'tot tlis¿ds¿."
Eu,el 1986
': Several reasorrs are given for ihe suc-
cess of nonnatñ,e species. The¡, rn¿1, d6
well because they are freed from naturai
enenies, competitors, and parasites.
Corl-cqtrentl);.r \ onrnlon str.lteAt of
control programs is to introduce organ-
isms from the native habitats of target
nonnati\ e piants to harm ihenl; thar is,
ehmrn,lte nonndtiv(,s by inlroJurrng
more nonnatives! Nonnative species
may also find little competition in the
ecosystems they in\.ade; empty niches
may be available to exotics, causing lit-
tle effect on the invaded ecosystem. The
success of exotics has also been attdb-
uted to their being aggressive coloniz
ers, fast growing, and highly fecund
.pecie.. Hor,,ever. Ewel ( lq86) noLeo
that "species invasions often ref'lect the
conditions of the community being
invaded rather than the uniquely
aggressive traits of the invader"
I subscribe to E!ve['s point of view.
When a good match is made between
the genome of .rny sper ie- anel the envi
ronmental conditions that support its
growth, the result can be explosive pop-
ulation increases. This is why organisms
that are rare in theil natural habitats
may suddenly become weedy in a new
situation. The water hyacinth (El.¡io,'-
ni0 crn>,tpt't i. fairl) inconspicrrou' in
its natural Amazonian habitats but
exhibits explosive growth in the slow
moving, highly eutrophic waters of arti-
ficial canals and reservoirs in Florida.
Experiments show that it cannot grow
¿s nell rr hen sr¡bjecteJ lo oligolrophic
waters or fast flow. Close observation of
successful nonnative organisms usually
yields similar results.
Ecological Functions and Services
of Nonnative Species
I have studied monoculture pla¡rtations
of nonnative trees in Puerto Rico and
have compared them ivith native forests
of similar age. My strldies include 73
comparisons of structure, composition,
and fllnction of tllese ecos)/stems an(l I
.,rrnol fin(l ,1 \in8le ( rrmlJr iiorl lh¿l
suggests an ecological anor-nalv in ihe
i,,rosl\.innrinareLl irl nnrt 'alir r, .1.er i,,r
These ecos)¡stems furnction like n¿rti\.e
forcsts, \,\,ith diffcrences lrosti), in the
rnagnitude of rates and state \'¿rriables
(i.e., biorrrass and other'structr¡ral fea-
tures). No t1e¡jative cffects h.rvc bce'n
detected in the water cycle, accuúrüa-
tion of carbon and nutrients, or' i1r any
other site conditiol]. Claims to the con
trar)¡, such as those leveled against
Er.dlypr¡rs trees, cannot be sribstantiated
when evaluated criticaliy.
I have also found that native plant
species gror,r, and develop under '
canopies of nonrrative species estab-
lished in degraded sites, inchrding some
in which succession w¿ls arrested pior
to plantint nonn¿ltives. Native birds are
attracted to and use the native species
understory of these nonnative tree plan-
tations. For these reasons, I (and others)
have suggested that nonnative species
cán be important tools for land ¡ehabili-
taiion and restoiation of biological
diversity in damaged sites where nat¡.r-
ral succession is arrested. For examplc,
nitrogen fixing trees can increase soil
nitrogen for the entire ecosystem, and
promote higher productivity. One
\'\,ould expect that highe¡ productivity
would eventually rcsult in a greater
( Jp.l( il) lo [i\ .,]rbon, Circ l¿te nr¡tri-
ents, a¡d support more species.
W¡ll Nonnative Spec¡es Dominate
the World?
The dominance of nonnative species on
the landscape will be a function of the
clegree of human modification of the
environme rl. ln gerreral, humdn .r. tiv-
ity fragments the landscape, far.ors
establishment of nonnative spccics,
increases envi¡onmental heterogeneity,
may cause species extinctions, and may
augment the total number of specics on
the landscape. The Pacific lslands are
instructive because they rcpresent a
worst-case scenario of the effects of
intensive human activity on small land
areas isolated from sources of biotic
replenishm.,nt. On these islands, partic-
ularly the Hawaii¿n chain, isolation
¿llowed the evolutioñ of a highly
clivcrsr, ancl ende¡nic sr-¡ite of organ
isrrs, oliginally exposecl to a slow rate
of inr,¡sic¡r by nonnative species.
Humans greatly accelerated that rate
.tr-r.l in tht: proccss transformed tl-re
fl,,r.r arrJ ilurra ollL,,- laiifi. l:lands.
T'he plocess has been ongoing fbr some
2000 to 4000 years and the results have
bcen staBflering in telms of species
extinctiolrs and transformation of biotic
composition- The m[nber of species
.icross all iaxonomic groups has
incre¿rsed llom about 9000 to 12,000,
and manv see this trend as an erosion
in g,l, 'b,rl bioJirer:il1 berau.eendemic
forms are lost while pantropical weeds
replace them. There is certainly truth to
this argument, but it is not entirely
accurate because many nonnati\¡e
\fe(re\ ¿re 'lLithHr \aeedy nor pdntrupr-
cal. Some may be rare and endangered
species that find refuge in another,
mo¡e favorable location and then
become weedy. ln Puerto Rico, for
example, Dclonr:r i'egl4 is a common nat-
urJli/ed 5pecic5 in d¿nger of e\tinction
in its native Madagascar habitat.
To better understand why the Pacific
I'l¡rrd-.rppe¡r -o vulnerrble to ¡nvJ.
sJve species, I compared the density of
species in the Caribbean Islands with
those of the Hawaiian Islands. If the
area afld plant species density values of
the H¿w¿iian lsl¿nd5 ¿re used to esti-
mate the numbers of plant species
expected in Cuba and Puerto Rico
(same latitude as Hawaii), the results
underestimate the actual number of
species on the two Caribbean islands.
This means that when area is corrected
fot the density of species in the
Carjbbean is much higher than in the
Hawaiian lriand-. l.rrt of thi. higher
species density in the Caribbean is
caused by the much greater density of
-¡recie. in C uba and Puer(o RicL) than in
tl¡e Hawaiian lslands. This is explained
in part because these islands a¡e close¡
to continental landmasses, but also by
the fact that the Caribbean lslands are
six times older than the Hau,aiian
Islands. Could this mean that the
Hawaiian Islands have a greater capac-
ily lo ab.orb addrtional planl rpccie*
than do Caribbean lslands? The age of
islands as well as their degree of isola
tion influence the density of species

and invasibility of thejr communities.
ln Hawaii, non¡ative species ¡rc the
iLell inraJer. of pl.rnl ,,,mmunilie.
because they are aciively transported
fuom a large rescrvofu of genetic mate-
rial (the wholc world) while the native
species evolve slowly and are con-
straincd by founder effects.
I would expect that in the absence of
significant climate change, environ
rncnts that toda), support high sPecies
richness will do so in the future. but the
ipecies composition [ra1, be different.
And we shc¡uld not lbrget that the
iorces of evolution are not suspendcd
Íor nonn¿rti\.e species. One could argue
that the en chment of islands with
nonn¿rtive tenomes ¡rrovides fuel for
the evolutiona¡v process ¿rnd greatl),
¡¡¡¡e1.e-.r¡1,rplirc p,
^\ihilil:c.. Thi. i-
particular'1y jmportani in light of
hur¡an-induced changes in the atmos-
phere, climerte, geor¡orpholotiy, and
other environmcnial conditions.
Tire change in species composition
takint piace in the world toclay is not a
.hJot'( lro.e.-: iL Is,r p-(\ (\- Ihál i5
'pspundinB ro ft¡ndam. ni¿l (llárl8e5 .n
ihe conditions of the planet. Age olcl
lation-scale impact is relatjvely easy to
,locumenl, ¿rnd m.'r) e\lend to mdjor
chan8es in community structure, meas-
rrred as chdnges in trophic dy'namicr or in-
teraction strengths between species.
INTRODUCED PREDATORS ThC bTOWN
snake, Boigo itcgtLlrrrt.s, is a str-nning exam-
ple of the population- and community-
ler.el impacts of an introduced vertebrate
predator on a naive island population. The
snake's nati\.e range extends from Aus-
tralia through New Guinea to the Solomon
Islands. Shortly after World War II, it ar-
rir.ed c¡¡ the islald of Guam where the only
native snake was a tiny, wormlike insecti-
r.ore. The invader gradually spread across
the island but remained at low densitics for
trvo decades until the early 1960s, when it
began to appear in large numbers. More
and more well-fed snakes were found in
chicken coops; others were carbonized on
power lines, causing elcctrical blackouts
(Figure 9.2). During this time, native birds
bcgan disappearilrg.
Ten species of forest birds followed a
similar pattern of decline: Each disap-
ecological constraints such as ti¡¡c,
energetics, biotic factors, growth condi-
tions, and opportunity are at play, re8u-
laiing which species are successful and
wlich are not in a specific location.
Human activity geñe¡atcs the environ-
mental change that powers the
response of organisms through adapta-
tion, evolution, or formation of new
groupings of species and communities.
Management Strategies for a
Changing World
I have highlighted contradictions in the
r..rr we Jeal \^ ith biñdi\ er.ity i.\ue\ in
Eieneral and nonnative species in partic-
ular. Even in Hawaii, where the¡e is
grcat concern about the degrec of no11
native species invasion and iheir poten-
tiall), negative effects, the government
actively and successfully introduces
hund¡ecls of insect specjes for agricul-
trral pest control. We correctly u,orry
about the netative effects of humalt
activit, but forget th¿i this activity
-L'rted lh¡'U.¿nd. oi ye..rrs d8o.Jt a lime
when people depended directly on the
environment for survival. Even then,
ITCC
3. 200
E
'i rso
'E 100
E.
!co
A,
s- §ñ r9" g$'
Figure 9.2 A¡¡ual electrical outages caused by the ir1vading brown tree
snake t¡n Guam. (Modified from Fritts 2002.)
Species Invasions 297
nonnativei were inlrodured Jnd Per
haps countless numbers of species were
driven to extinction. I point these
things out not to excuse introductions
ol specie. or to . onJonc driving -pecies
to extinction. But, we must maintai¡ an
open mind and analyze the issue of
nonnative specjes jntroductions and
man¡8emenI ds dn inlrin\ic ¿nd cont]n-
uous process in a wo¡ld \^'here our o19n
species is a main driver of change.
It is in our power to iake actions to
mititate the negatives of our activities
and to enh¿n(c lhe positivec. Aciions
that may help are leaming to manage
and control environmental change, rec-
ogrizillg u,hen conditions are obviously
beyond our control, avoiding condemn-
ing species because of successional
stage or ecological function, improving
our capacity to manage biotic resources,
concentrating human activity to allow
more Space for n¿rti\¡e ecosystem§, and
encouraging environmental heterogene-
ity as a mechanism to maximize biodi-
versiry One thing is clear: The world
will continue to change and become less
familiar to those that walked on it o¡
wrote about it centuries ago. .i
g.ob gtb §¡ñ 9''"' 9+ g''b