Ecology of Invasive Species Ecology 2013

But we can fight back!!!!

Zebra mussels
Clog water intakes for industry, drinking water ($2 billion/yr) Alter food webs by filterfeeding and cleaning water Monopolize substrate and even encrust living organisms

Garlic mustard (Alliaria petiolata)

Invades moist eastern deciduous forests Very common in Western Pennsylvania May threaten many understory herbs

West Nile Virus

First US occurrence in Fall 1999 in NYC Has been detected in humans, horses and birds Mosquito-borne Green areas = wild bird cases in 2001 Some speculation this was a type of terrorist attack

Asian Tiger Mosquito (Aedes albopictus)

Imported to Southeastern US in a shipment of used tires Vector for dengue fever and yellow fever outside of US

Purple Loosestrife (Lythrum salicaria)

Invades fertile marshes throughout the US Threatens many plant and animal species Forms dense, nearly monospecific stands Vectors: Ship ballast, medicinal plant trade, bee keepers

The Scope of the Problem

10% of wild plant species in the US are exotics Invasive species are a primary or contributing threat to nearly half of all endangered and threatened species Invasive species are estimated to cost the US economy $137,000,000,000 per year
Direct losses Control costs

Ecologists cannot accurately predict the results of a single invasion or introduction event
-Paul Ehrlich, 1989

Outline
Magnitude of the invasive species problem Costs of invasive species Anatomy of an invasion
Introduction Establishment Growth and Spread

Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits

Resource competition and invasion by Purple Loosestrife

Anatomy of an Invasion
1. 2. 3. Introduction to a new environment Establishment in the new environment Population growth and spread into undisturbed habitats

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

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 danger Unintentional introductions Never desirable Careful inspection of imported fruits and vegetables Ballast water: force cargo ships to change ballast in open ocean

Anatomy of an Invasion
1. 2. 3. Dispersal to new environment Establishment in new environment Population growth and spread into undisturbed habitats

Stage 2: Establishment
Successful invasions require establishment in a new environment These environments are often highly disturbed urban or agricultural areas Many species remain in the establishment stage for long periods before spreading and we really have little understanding as to why this is true.

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

Anatomy of an Invasion
1. 2. 3. Dispersal to new environment Establishment in new environment Population growth and spread into undisturbed habitats

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, coordinated effort

Stage 3. Growth and Spread

Control options Mechanical and chemical control are poor longterm solutions
Expensive, global control is required

Biological control is the only cost-effective longterm option Many problems associated with biological control

Biological Control
Assumes that invaders are successful because they leave their natural enemies behind Involves seeking and testing many possible agents Agents chosen are usually highly species specific and heavily damage the host

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 and attack species we like

Why do some species become invasive?

(or, how can we predict which nonnatives pose the greatest threat and thereby target them for control?)

Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits

Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits

Escape from natural enemies

Hypothesis: Escape from natural enemies increases the resource 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 load Evidence: Biological control Soil-borne pathogens

St. Johns Wort and Chrysolina beetles

Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits

Evolution of increased competitive ability

Hypothesis: Invaders succeed because they evolve lower allocation to herbivore defense when released from herbivory Rationale: Darwins theory of evolution by natural selection predicts that: if there is variability in allocation to defense, and if that allocation is costly, then invaders which have escaped their natural enemies will evolve lower allocation to defense and thereby become better competitors Evidence: Little. Only a few tests, all mostly negative. Example: None. However, this remains an active area of research

Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits

Hybridization
Hypothesis: Invaders succeed because hybridize and thereby become better competitors Rationale: The invasion process brings together related species which are not naturally co-occurring in their native habitats; hybrids often show hybrid vigor; thus these new hybrids become supercompetitors Evidence: Found in several invaders, including hybrid cattail and salt-cedar. Probably not common.

Example: Salt cedar

Salt cedar: invades riparian areas throughout the arid west displaces a diverse streamside shrub community threatens several species including the endangered Willow Flycatcher At least eight salt cedar species have been introduced, including Tamarix ramosissima and T. chinensis

The Nature Conservancy

Example: Salt cedar: Tamarix spp

Morphological identification of invaders is difficult, and hybridization has been suspected Gaskin and Schaal (2002) demonstrate that the most common genotype among invasive salt cedar is in fact a hybrid between T. ramosissima and T. chinensis
Unresolved: the degree of competitive advantage of the hybrid in either greenhouse or the field

(Gaskin and Shaal 2003)

Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits

Novel weapons hypothesis

Hypothesis: Invaders produce allelopathic chemicals which are novel to the invaded ecosystem, giving the invader a competitive advantage Rationale: Plant species coevolve to tolerate allelochemicals produced by co-occurring species. When a species invades a new range, the established species are nave, and thus susceptible to the invaders allelochemicals Evidence: Demonstrated in the greenhouse for one invasive species

Example: Spotted knapweed

Spotted knapweed (Centaurea maculosa) invades shrub lands throughout the Northern Rockies Converts diverse grasslands to monocultures

The Nature Conservancy

Example: Spotted knapweed

Knapweed produces catechin, which is toxic to plant roots Catechin: is common in soil of European but not North American ecosystems causes widespread death of plant root cells reduces germination and growth of North American grassland species
(Bais etal. 2003)

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 followup experiments.

Invasion Theory
Escape from natural enemies Evolution of increased competitive ability Hybridization Novel weapons Traits

Traits of plant invaders facilitate their dominance

Hypothesis: Invaders possess traits which make them good competitors in specific environments Rationalization: Invaders succeed because they are better competitors for limiting resources than the native species in their new environment. Evidence: Unknown Example: Resource competition theory

Resource competition theory (Tilmans R*) provides clear, testable predictions for both invaders and biocontrol:
1. A successful invader must deplete limiting resources (light or nitrogen) to lower levels than native species A successful biocontrol agent must limit the invaders ability to deplete limiting resources

2.

Resource Competition models

Invasion predicted
1

Effective biocontrol at 50 insects/m2

1 Invader Native

Resource availability

resource availability
0 Invader Native

0 0 25 50 75 Insect density 100

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-native species, hoping it will control the invader (stage 3)

Model System
Purple loosestrife (Lythrum salicaria), an herbaceous perennial from Europe, invades fertile marshes and wetlands throughout the US Broad-leaved cattail (Typha latifolia) is commonly outcompeted and displaced by loosestrife A leaf-feeding beetle (Galerucella calmariensis), is currently being introduced in an effort to control loosestrife

Methods I: Measuring resource competitive ability

Grow invasive and native in monoculture with and without biocontrol agents Measure resource availability (light and nitrogen)

Methods II:
Grow invasive and native in competition Winners and losers should be predicted by resource competitive abilities determined above

Preliminary results
Our data show that 17% leaf area damage is required to raise loosestrife's R* above that of cattail (arrow). This suggests that if we do not see these levels of damage in the field, then biocontrol will not be successful.
90% 80%
Lythrum Low Fertility Typha Low Fertility

Percent light availability

70% 60% 50% 40% 30% 20% 10% 0% 0 20 40 60 80 100 Percent loosestrife leaf area damaged

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

Summary
What are the likely mechanisms controlling invasions? Traits which make species good competitors Escape from natural enemies (which enhances these traits) What can we do? Apply theory Vigilance: monitor establishing species Screen established but not yet invasive species for threat Use biocontrol only when it can work