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Ecology ofInvasive Species
Ecology 2013
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But we can fight back!!!!
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Zebra mussels
Clog water intakes forindustry, drinking water ($2billion/yr)
Alter food webs by filter-feeding and
cleaning
water
Monopolize substrate andeven encrust livingorganisms
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Garlic mustard( Alliaria petiolata )
Invades moist easterndeciduous forests
Very common in WesternPennsylvania May threaten many
understory herbs
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West Nile Virus
First US occurrence in Fall 1999in NYC
Has been detected in humans,horses and birds
Mosquito-borne Green areas = wild bird cases in
2001
Some speculation this was a typeof terrorist attack
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Asian Tiger Mosquito( Aedes albopictus )
Imported toSoutheastern US in ashipment of usedtires
Vector for denguefever and yellowfever outside of US
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Purple Loosestrife(Lythrum salicaria )
Invades fertile marshesthroughout the US
Threatens many plant and
animal species Forms dense, nearlymonospecific stands
Vectors: Ship ballast,medicinal plant trade, beekeepers
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-Paul Ehrlich, 1989
Ecologists cannot accurately predictthe results of a single invasion or
introduction event
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Outline
Magnitude of theinvasive speciesproblem
Costs of invasivespecies
Anatomy of aninvasion
Introduction Establishment Growth and Spread
Invasion Theory Escape from natural
enemies
Evolution of increasedcompetitive ability Hybridization Novel weapons Traits
Resource competitionand invasion by PurpleLoosestrife
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Anatomy of an Invasion
1. Introduction to a new environment2. Establishment in the new environment3. Population growth and spread into undisturbed habitats
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Stage 1. Introductions
Natural pathways Air and water
currents. Wildlife
movements.
Human pathways Intentional
Domesticated
animals Horticultural escape
Unintentional Ballast water: common vector
for aquatic organisms Shipping and commerce
Fruit and vegetable imports Emerald Ash Borer
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Control during the introduction stage
Intentional Introductions Many introduced species are beneficial (crops,
ornamentals, biocontrol species) Only 0.1% of introduced species will become invasive. Risk assessment must be used to determine which
species pose little dangerUnintentional introductions
Never desirable Careful inspection of imported fruits and vegetables Ballast water: force cargo ships to change ballast in
open ocean
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Anatomy of an Invasion
1. Dispersal to new environment2. Establishment in new environment3. Population growth and spread into undisturbed habitats
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Stage 2: Establishment
Control Options Control is far easier when populations are small Yet there are still too many species to control all of
them. Identification of potential invaders is critical Mechanical control
Mowing, hand removal Chemical control
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Anatomy of an Invasion
1. Dispersal to new environment2. Establishment in new environment3. Population growth and spread into undisturbed
habitats
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Stage 3. Growth and Spread
The cat is now out of the bag Identification of invaders is easy! Once a species is widespread, control is very
difficult Control at local sites is expensive Re-invasion from surrounding area is likely
Effective control requires broad-based, coordinatedeffort
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Stage 3. Growth and Spread
Control options Mechanical and chemical control are poor long-
term solutions Expensive,
global
control is required Biological control is the only cost-effective long-
term option
Many problems associated with biological control
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Biological Control
Assumes that invaders are successful because theyleave their natural enemies behind
Involves seeking and testing many possible agents
Agents chosen are usually highly species specificand heavily damage the host
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Pros and cons of biocontrol
Pro: Long term solution Inexpensive after
agents are established Highly targeted and
species specific
Con: Most efforts fail despite
successful introductions:
insects running amok. Identification of agents is
expensive Agents MAY NOT be species
specific or may evolve andattack species we like
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Why do some species become
invasive?(or, how can we predict which non-
natives pose the greatest threat andthereby target them for control?)
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Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits
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Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits
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Escape from natural enemies
Hypothesis: Escape from natural enemies increases theresource competitive ability of invaders
Rationale: Herbivores, predators and pathogens limit the competitive
ability of otherwise dominant species Natural enemies are not introduced with native plants Invaders then can out-compete natives which still have a
herbivore loadEvidence: Biological control Soil-borne pathogens
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St. Johns Wort and Chrysolinabeetles
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Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits
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Evolution of increasedcompetitive ability
Hypothesis: Invaders succeed because they evolve lowerallocation to herbivore defense when released fromherbivory
Rationale: Darwin
s theory of evolution by natural selectionpredicts that: if there is variability in allocation to defense,and if that allocation is costly, then invaders which haveescaped their natural enemies will evolve lower allocation todefense and thereby become better competitors
Evidence: Little. Only a few tests, all mostly negative.Example: None. However, this remains an active area of
research
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Hybridization
Hypothesis: Invaders succeed because hybridize andthereby become better competitors
Rationale: The invasion process brings together
related species which are not naturally co-occurringin their native habitats; hybrids often show
hybridvigor
; thus these new hybrids become super-competitors
Evidence: Found in several invaders, including hybridcattail and salt-cedar. Probably not common.
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Example: Salt cedar
Salt cedar: invades riparian
areas throughoutthe arid west
displaces adiversestreamside shrubcommunity
threatens severalspecies includingthe endangeredWillow Flycatcher
At least eight salt cedar species havebeen introduced, including Tamarixramosissima and T. chinensis
The Nature Conservancy
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Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits
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Example: Spotted knapweed
Spottedknapweed(Centaureamaculosa)invades shrublandsthroughout theNorthern
Rockies Converts
diversegrasslands tomonocultures
The Nature Conservancy
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Example: Spotted knapweed
Knapweed producescatechin, which is toxic toplant roots
Catechin: is common in soil ofEuropean but not North
American ecosystems causes widespread death
of plant root cells reduces germination and
growth of North Americangrassland species
(Bais etal. 2003)
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Example: Spotted knapweed
Knapweed and catechin Poorly tested in the field Is unlikely a common mechanism of invasion
because invaders should be susceptible to the
novel weapons
produced by natives The one major field test that found a negative
impact of catechin could not be duplicated in follow-up experiments.
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Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits
f l d f l
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Traits of plant invaders facilitatetheir dominance
Hypothesis: Invaders possess traits which make them goodcompetitors in specific environments
Rationalization: Invaders succeed because they are bettercompetitors for limiting resources than the native species intheir new environment.
Evidence: UnknownExample: Resource competition theory
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Resource competition theory(Tilman
s R*) provides clear, testable
predictions for both invaders andbiocontrol:
1. A successful invader must deplete limitingresources (light or nitrogen) to lower levels thannative species
2. A successful biocontrol agent must limit theinvader
s ability to deplete limiting resources
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Resource Competition models
0
1
Invader Native
R e s o u r c e a v a
i l a
b i l i t
y
0
1
0 25 50 75 100Insect density
r e s o u r c e
a v a
i l a
b i l i t
y
InvaderNative
Invasion predictedEffective biocontrol at
50 insects/m 2
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Resource competition theory
Can predict which species are likely to become invasive(stages 1 and 2)
Can predict whether biocontrol will succeed BEFORE spending millions to introduce another non-nativespecies, hoping it will control the invader (stage 3)
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Model System Purple loosestrife (Lythrum
salicaria ), an herbaceousperennial from Europe, invadesfertile marshes and wetlandsthroughout the US
Broad-leaved cattail ( Typhalatifolia ) is commonly out-competed and displaced byloosestrife
A leaf-feeding beetle(Galerucella calmariensis ), iscurrently being introduced in aneffort to control loosestrife
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Methods II:
Grow invasive andnative in competition
Winners and losersshould be predictedby resourcecompetitive abilitiesdetermined above
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Preliminary results
0%
10%
20%
30%
40%
50%
60%
70%
80%
90%
0 20 40 60 80 100Percent loosestrife leaf area damaged
P e r c e n t
l i g
h t a v a i
l a b i l i t
y
Lythrum Low FertilityTypha Low Fertility
Our data show that 17% leafarea damage is required toraise loosestrife's R* abovethat of cattail (arrow). Thissuggests that if we do not seethese levels of damage in thefield, then biocontrol will not besuccessful.
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Potential applied benefits
If successful, this methodology will allow ecologists to: Predict where potential invaders might successfully invade Predict the efficacy of biological control efforts
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