MANAGEMENT OF INVASIVE PLANTS IN THE WESTERN USA
  • Defining the Problem
    • What is a Weed? >
      • Federal Definitions of Noxious Weeds
    • Costs of invasive plants
    • Human Factor
    • Challenges of Invasive Plants
    • Wildfires in the Western USA >
      • Forest Fires: Structure
      • Bark Beetles & Forest Ecosystems
      • Rangeland Fires
    • Climate Change Impacts on Plants >
      • Climate Change: CO2, NO, UV, Ozone Impacts on Plants
      • Climate Change Impacts on Crops
      • Climate Change Impacts on C4 Plants
      • Climate Change Impacts on Rangeland
    • What are we doing?
  • Focus of this Project
    • Why Western States? >
      • Audience for these reports
    • History: Are we doomed to repeat it? >
      • Dust Bowl Re-visited >
        • China: Past & Present
        • UN Biodiversity Report
    • Policy vs. Practice
    • Ecosystems & Economics >
      • Reductionist Approach to science
      • Ecology & Feminism
      • Systems View of Life
      • Ecosystems Health
      • Economic Growth
      • Impact of the Petrochemical Industry
      • Interrelation of Economics & Ecology
    • Federal Agencies >
      • Federal Agencies and Invasive Species
      • History of Coordination with States
      • Challenges of Coordination between Federal Agencies
      • Collaboration or Confusion
    • Organizations to assist landowners
    • Federal Legislation on Invasive Species >
      • 1930s Federal Laws on Invasive Species
      • Federal Seed Act 1939
      • 1940s-1960s Federal Laws on Invasive Species
      • 1970s Federal Laws on Invasive Species
      • 1980s Federal Laws on Invasive Species
      • 1990s Federal Laws on Invasive Species
      • 2000-2010 Federal Laws on Invasive Species
      • 2011-2022 Federal Laws on Invasive Species
      • Federal Bills on Invasive Species not passed
      • Executive Orders on Invasive Species
      • Federal Excise Taxes
    • State Laws and Lists of Noxious Weeds
    • My Inspirations
  • Why we need plants
    • Native Plants
    • Plant Resources
  • Invasive Success Hypotheses
    • Unified Framework
    • Role of Diversity >
      • How Ecosystems Maintain Diversity
      • Fluctuation Dependent Mechanisms
      • Competition-based coexistence mechanisms
      • Niche Differences
      • Species Richness
    • Enemy Release Hypothesis
    • Constitutive Defense Mechanisms
    • Evolution of Increased Competitive Ability
    • Role of Microbes
    • Indirect Defense Mechanisms
    • Novel weapons hypothesis
    • Evolutionary Shifts
    • Resource Allocation
    • Evolutionary Dynamics >
      • Pre-introduction evolutionary history
      • Sampling Effect
      • Founder Effect
      • Admixture, hybridization and polyploidization
      • Rapid Evolution
      • Epigenetics
      • Second Genomes
    • Role of Hybridization
    • Role of Native Plant Neighbors
    • Species Performance
    • Role of Herbivory
    • Evolutionary Reduced Competitive Ability
    • Summary Thoughts on Research
  • Historical Record
    • Regional Conferences
    • Timeline
  • Innovative Solutions
    • Agricultural Best Practices >
      • Ecologically based Successional Management
      • Perennial Crops, Intercropping, beneficial insects
      • Soil Solarization
      • Natural Farming
      • Permaculture
      • Organic Farming
      • Embedding Natural Habitats
      • Conservation Tillage
      • Crop Rotation
      • Water Use Practices
      • Tree Planting: Pros & Cons
    • Grazing Solutions >
      • Sheep and Goat Grazing
      • Cattle & Sheep Grazing
      • Cattle and Bison Grazing
      • Grazing and Revegetation
    • Rangeland Restoration >
      • Federal Goals for Rangelands
      • Novel Ecosystems
      • Prairie Restoration >
        • Prairie Restoration Workshop
        • Weed Prevention Areas
        • California grassland restoration
        • Selah: Bamberger Ranch Preserve
      • Sagebrush Steppe Restoration >
        • Low Nitrogen in Sagebrush Steppe
      • Revegetation with Native Plants
      • Dogs as detectors of noxious weeds
    • Nudges
  • Biological Control
    • Insects as Biocontrol >
      • Impacts of Biocontrol Agents on Non-Target Species
      • Indirect Impact of Biocontrol on Native Species
    • Challenges of Using Biocontrols >
      • DNA studies on Biocontrol Insects
      • Biocontrol takes time
    • Prioritization process for Biocontrol Programs
    • Evolutionary changes impact Biocontrol
    • Vertebrates as Biocontrol Agents
  • Herbicides: History and Impacts
    • Effectiveness of Herbicides in Agricultural Lands
    • Effectiveness of Herbicides in Rangelands
    • History of Use of Herbicides and Pesticides Prior to and During WWII
    • Herbicide use during and post-World War II >
      • 2,4-D Herbicide Use
      • 2,4-D and 2,4,5-T, post-World War II
    • Modern use of Herbicides >
      • Atrazine Herbicide
      • Dicamba Herbicide
      • Glyphosate Herbicide
      • Paraquat Dichloride
      • Picolinic acid family of herbicides >
        • Picloram (Tordon 22K) Herbicide
        • Triclopyr Herbicide
    • Herbicide Resistance in Invasive Plants >
      • Herbicide Resistant Crops
      • Controlling herbicide-resistant weeds in herbicide-resistant crops
      • Best Management Practices
    • Myth of the Silver Bullet
    • Myth of Eradication
    • Merging of Agrochemical Companies
    • Impacts of Pesticides on Environment and Human Health >
      • Pesticide Drift
      • Impacts of Pesticides on Biological Diversity
      • Impacts of Herbicides on Native Plants
      • Pesticide Impacts on Insects >
        • Butterflies: The Impacts of Herbicides
        • Monarch Butterflies: Impacts of Herbicides
      • Impacts of Pesticides on Wildlife >
        • Reptiles & Amphibians: Pesticide Impacts
      • Pesticide Residue in Foods
    • Funding for Research on Pesticides
    • Commentary on Herbicide Use
  • Interviews
    • Interviews Biocontrol >
      • Biocontrol Wyoming
      • Montana Biocontrol Interview Maggio
      • Montana Biocontrol Interview Breitenfeldt
    • California Interviews >
      • Robert Price
      • Doug Johnson
    • Colorado Interviews >
      • George Beck Interview
      • Scott Nissen Interview
    • Idaho Interviews >
      • Purple Sage Organic Farms in Idaho
    • Montana Interviews >
      • Jasmine Reimer Interview Montana
      • Organic Farms Montana Interviews
    • Texas Interviews
    • Washington Interviews >
      • Ray Willard
    • Wyoming Interviews >
      • Slade Franklin Interview
      • John Samson Interview
    • Wyoming Weed and Pest Districts >
      • Josh Shorb Interview
      • Slade Franklin Interview 2
      • Lars Baker Interview
      • Steve Brill Interview
      • George Hittle Interview
      • Peter Illoway Interview
      • Robert Jenn Interview
      • Sharon Johnson Interview
      • Larry Justesen Interview
      • Gale Lamb Interview
      • Stephen McNamee Interview
      • Allen Mooney Interview
      • Rob Orchard Interview
      • Robert Parsons Interview
      • Dick Sackett Interview
      • Comments by Delena
    • NRCS Interviews: Wyoming
  • Western Weed Control Conference 1940s Minutes
    • 1942 Conference
    • 1945 Conference
    • 1946 Conference
  • Who am I?
    • My Work
    • My Adventures
    • Contact Page
  • Road Logs
    • Colorado Road Logs
    • Idaho Road Logs
    • Montana Road Logs
    • New Mexico Road Logs
    • Texas Road Logs
    • Wyoming Road Logs
  • Bibliography

How Ecosystems Maintain Diversity

Photo: Sagebrush habitat, Yellowstone NP. © 2016 Delena Norris-Tull

How ecosystems maintain diversity

Summary of the research and commentary by Dr. Delena Norris-Tull, Professor Emerita of Science Education, University of Montana Western, July 2020.
 
In a review of research on how ecosystems maintain species diversity, Chesson (2000) stated that, “The most common meaning of diversity maintenance is coexistence in the same spatial region of species having similar ecology… Another meaning of diversity maintenance refers not to coexistence of fixed sets of species but to the maintenance of species richness and evenness over long timescales, necessitating consideration of speciation and extinction rates, and infrequent colonizations.”
 
Chesson, 2000, points out that “species coexistence… may be stable or unstable. Stable coexistence can be quantified by the long-term rates at which community members recover from low density… Coexistence mechanisms function in two major ways: They may be (a) equalizing because they tend to minimize average fitness differences between species, or (b) stabilizing because they tend to increase negative intraspecific interactions relative to negative interspecific interactions. Stabilizing mechanisms are essential for species coexistence.”
 
Chesson, 2000, found that, “Models of unstable coexistence, in which species diversity slowly decays over time, have focused almost exclusively on equalizing mechanisms. These models would be more robust if they also included stabilizing mechanisms… Models of unstable coexistence invite a broader view of diversity maintenance incorporating species turnover.” Chesson describes the Lotka-Volterra competition model, the gold standard for mathematical calculations of species coexistence. “The defining feature of Lotka-Volterra competition is that per capita growth rates are linear decreasing functions of the densities of each of the species… ‘Species j cannot competitively exclude species i if the effect that species j has on itself is more than the effect that species j has on species i….’ [In other words], The criteria for species coexistence in a two-species system… can be read very simply as ‘intraspecific competition must be greater than interspecific competition.’”
 
Chesson, 2000, points out that, “models with nonlinear per capita growth rates can be written [using the Lotka-Volterra equations]… by making the competition coefficients functions of density…, and the results…remain true, provided the competition coefficients are evaluated for the resident [species] at equilibrium and the invader at zero. Lotka-Volterra models and their nonlinear extensions may be thought of as models of direct competition: Individual organisms have immediate direct negative effects on other individuals.”
 
Chesson, 2000, then explains models with explicit resource dynamics, which “are not defined by a mechanism of competition. A mechanistic understanding of competition usefully begins with Tilmans’ resource competition theory in which species jointly limited by a single resource are expected to obey the R* rule. For any given species, R* is the resource level at which the species is just able to persist. The winner in competition is the species with the lowest R* value. A species’ R* value, however, reflects not the ability of members of the species to extract resources when they are in low concentration, but their ability to grow and reproduce rapidly enough, at low resource levels, to compensate for tissue death and mortality, which are affected by such factors as grazing and predation… It is the overall fitness of a species that leads to it's R* value.”
 
Chesson, 2000, points out that, “Tradeoffs play a major role in species coexistence: Advantages that one species may have over others are offset by compensating disadvantages. However, examined in relation to the R* rule there is clearly more to stable coexistence. With just a single resource as a limiting factor, tradeoffs may make the R* values of different species more nearly equal, but that would not lead to stable coexistence... Similarity in average fitness does not lead to stable coexistence,… [as] only one [species] can increase from low density in the presence of the other.”
 
“Coexistence with resource partitioning contrasts with this. Using MacArthur’s mechanistic derivation of Lotka-Volterra competition… The two species coexist if the stabilizing term is greater in magnitude than the fitness difference term because then both [species] have positive growth as invaders… The assumptions of the model entail that doing well on some resources means doing less well on others. Each species has density-dependent feedback loops with its resources that limit it intraspecifically and limit other species interspecifically. However, limited resource overlap and tradeoffs in resource benefits means that intraspecific limitation is enhanced relative to interspecific limitation. This concentration of intraspecific effects relative to interspecific effects is the essence of stabilization.”
 
Chesson, 2000, points out that MacArthur’s equation “generalizes to multispecies communities involved in diffuse competition, that is, where competition between species involves comparable interaction strengths for all pairs of species… [In the model’s equation] the first term is an average fitness comparison and the second term is a stabilizing term. Without this term, the first term of necessity leads to loss of all species… However, if the stabilizing term is larger in magnitude than the relative average fitness term for the worst species, then all species coexist. These two general terms may involve different mechanisms. Those [terms] reducing the magnitude of the fitness difference term will be referred to as equalizing mechanisms, while those increasing the magnitude of the stabilizing term will be referred to as stabilizing mechanisms. In the absence of the stabilizing term, equalizing mechanisms can, at best, slow competitive exclusion; but in the presence of stabilizing mechanisms, equalizing mechanisms may enable coexistence.”
 
Chesson, 2000, reviewed research on niche space, “conceived as having four axes: resources, predators (and other natural enemies), time, and space.” For example, “A species consumes resources, and therefore has an effect on resource density. Individuals of a species may also reproduce, grow, or survive in response to resources. The essential way in which stabilization occurs is most clearly seen with resource competition. If a species depends most on a particular resource (strong response), and also reduces that resource (strong effect), then it has a density-dependent feedback loop with the resource and is limited by it. If a second species has a similar relationship with a different resource, then even though the species each consume some of the resource on which the other depends most strongly (limited resource overlap) each species depresses its own growth more than it depresses the growth of other species. The result is stable coexistence.”
 
Chesson, 2000, points out that “stable coexistence mechanisms may be fluctuation dependent or fluctuation independent… Fluctuation-independent mechanisms are resource partitioning and frequency-dependent predation.” Two examples of resource partitioning are spatial partitioning of soil moisture and grazing. “These mechanisms can function in the presence of environmental fluctuations… We can think of the operation of the mechanism as independent of the [environmental] fluctuations in the system.”

Reference:
  • ​Chesson, P. (2000). Mechanisms of maintenance of species diversity. Annual Review of Ecological Systems, 31, 343-366.

Next Sections on the role of diversity:
  • Fluctuation Dependent Mechanisms​
  • Competition-based coexistence mechanisms
  • Niche Differences
  • Species Richness

Next Sections on the research on the success of invasive species:
Enemy release hypothesis
Constitutive defense mechanisms
Copyright: Dr. Delena Norris-Tull, July 2020. Management of Invasive Plants in the Western USA.

These webpages are always under construction. I welcome corrections and additions to any page.
​Send me an email, and I can send you the original Word format version of any page you wish to correct.
contact Dr. Norris-Tull
Bibliography
who am i?
My work
my inspirations
my adventures
  • Defining the Problem
    • What is a Weed? >
      • Federal Definitions of Noxious Weeds
    • Costs of invasive plants
    • Human Factor
    • Challenges of Invasive Plants
    • Wildfires in the Western USA >
      • Forest Fires: Structure
      • Bark Beetles & Forest Ecosystems
      • Rangeland Fires
    • Climate Change Impacts on Plants >
      • Climate Change: CO2, NO, UV, Ozone Impacts on Plants
      • Climate Change Impacts on Crops
      • Climate Change Impacts on C4 Plants
      • Climate Change Impacts on Rangeland
    • What are we doing?
  • Focus of this Project
    • Why Western States? >
      • Audience for these reports
    • History: Are we doomed to repeat it? >
      • Dust Bowl Re-visited >
        • China: Past & Present
        • UN Biodiversity Report
    • Policy vs. Practice
    • Ecosystems & Economics >
      • Reductionist Approach to science
      • Ecology & Feminism
      • Systems View of Life
      • Ecosystems Health
      • Economic Growth
      • Impact of the Petrochemical Industry
      • Interrelation of Economics & Ecology
    • Federal Agencies >
      • Federal Agencies and Invasive Species
      • History of Coordination with States
      • Challenges of Coordination between Federal Agencies
      • Collaboration or Confusion
    • Organizations to assist landowners
    • Federal Legislation on Invasive Species >
      • 1930s Federal Laws on Invasive Species
      • Federal Seed Act 1939
      • 1940s-1960s Federal Laws on Invasive Species
      • 1970s Federal Laws on Invasive Species
      • 1980s Federal Laws on Invasive Species
      • 1990s Federal Laws on Invasive Species
      • 2000-2010 Federal Laws on Invasive Species
      • 2011-2022 Federal Laws on Invasive Species
      • Federal Bills on Invasive Species not passed
      • Executive Orders on Invasive Species
      • Federal Excise Taxes
    • State Laws and Lists of Noxious Weeds
    • My Inspirations
  • Why we need plants
    • Native Plants
    • Plant Resources
  • Invasive Success Hypotheses
    • Unified Framework
    • Role of Diversity >
      • How Ecosystems Maintain Diversity
      • Fluctuation Dependent Mechanisms
      • Competition-based coexistence mechanisms
      • Niche Differences
      • Species Richness
    • Enemy Release Hypothesis
    • Constitutive Defense Mechanisms
    • Evolution of Increased Competitive Ability
    • Role of Microbes
    • Indirect Defense Mechanisms
    • Novel weapons hypothesis
    • Evolutionary Shifts
    • Resource Allocation
    • Evolutionary Dynamics >
      • Pre-introduction evolutionary history
      • Sampling Effect
      • Founder Effect
      • Admixture, hybridization and polyploidization
      • Rapid Evolution
      • Epigenetics
      • Second Genomes
    • Role of Hybridization
    • Role of Native Plant Neighbors
    • Species Performance
    • Role of Herbivory
    • Evolutionary Reduced Competitive Ability
    • Summary Thoughts on Research
  • Historical Record
    • Regional Conferences
    • Timeline
  • Innovative Solutions
    • Agricultural Best Practices >
      • Ecologically based Successional Management
      • Perennial Crops, Intercropping, beneficial insects
      • Soil Solarization
      • Natural Farming
      • Permaculture
      • Organic Farming
      • Embedding Natural Habitats
      • Conservation Tillage
      • Crop Rotation
      • Water Use Practices
      • Tree Planting: Pros & Cons
    • Grazing Solutions >
      • Sheep and Goat Grazing
      • Cattle & Sheep Grazing
      • Cattle and Bison Grazing
      • Grazing and Revegetation
    • Rangeland Restoration >
      • Federal Goals for Rangelands
      • Novel Ecosystems
      • Prairie Restoration >
        • Prairie Restoration Workshop
        • Weed Prevention Areas
        • California grassland restoration
        • Selah: Bamberger Ranch Preserve
      • Sagebrush Steppe Restoration >
        • Low Nitrogen in Sagebrush Steppe
      • Revegetation with Native Plants
      • Dogs as detectors of noxious weeds
    • Nudges
  • Biological Control
    • Insects as Biocontrol >
      • Impacts of Biocontrol Agents on Non-Target Species
      • Indirect Impact of Biocontrol on Native Species
    • Challenges of Using Biocontrols >
      • DNA studies on Biocontrol Insects
      • Biocontrol takes time
    • Prioritization process for Biocontrol Programs
    • Evolutionary changes impact Biocontrol
    • Vertebrates as Biocontrol Agents
  • Herbicides: History and Impacts
    • Effectiveness of Herbicides in Agricultural Lands
    • Effectiveness of Herbicides in Rangelands
    • History of Use of Herbicides and Pesticides Prior to and During WWII
    • Herbicide use during and post-World War II >
      • 2,4-D Herbicide Use
      • 2,4-D and 2,4,5-T, post-World War II
    • Modern use of Herbicides >
      • Atrazine Herbicide
      • Dicamba Herbicide
      • Glyphosate Herbicide
      • Paraquat Dichloride
      • Picolinic acid family of herbicides >
        • Picloram (Tordon 22K) Herbicide
        • Triclopyr Herbicide
    • Herbicide Resistance in Invasive Plants >
      • Herbicide Resistant Crops
      • Controlling herbicide-resistant weeds in herbicide-resistant crops
      • Best Management Practices
    • Myth of the Silver Bullet
    • Myth of Eradication
    • Merging of Agrochemical Companies
    • Impacts of Pesticides on Environment and Human Health >
      • Pesticide Drift
      • Impacts of Pesticides on Biological Diversity
      • Impacts of Herbicides on Native Plants
      • Pesticide Impacts on Insects >
        • Butterflies: The Impacts of Herbicides
        • Monarch Butterflies: Impacts of Herbicides
      • Impacts of Pesticides on Wildlife >
        • Reptiles & Amphibians: Pesticide Impacts
      • Pesticide Residue in Foods
    • Funding for Research on Pesticides
    • Commentary on Herbicide Use
  • Interviews
    • Interviews Biocontrol >
      • Biocontrol Wyoming
      • Montana Biocontrol Interview Maggio
      • Montana Biocontrol Interview Breitenfeldt
    • California Interviews >
      • Robert Price
      • Doug Johnson
    • Colorado Interviews >
      • George Beck Interview
      • Scott Nissen Interview
    • Idaho Interviews >
      • Purple Sage Organic Farms in Idaho
    • Montana Interviews >
      • Jasmine Reimer Interview Montana
      • Organic Farms Montana Interviews
    • Texas Interviews
    • Washington Interviews >
      • Ray Willard
    • Wyoming Interviews >
      • Slade Franklin Interview
      • John Samson Interview
    • Wyoming Weed and Pest Districts >
      • Josh Shorb Interview
      • Slade Franklin Interview 2
      • Lars Baker Interview
      • Steve Brill Interview
      • George Hittle Interview
      • Peter Illoway Interview
      • Robert Jenn Interview
      • Sharon Johnson Interview
      • Larry Justesen Interview
      • Gale Lamb Interview
      • Stephen McNamee Interview
      • Allen Mooney Interview
      • Rob Orchard Interview
      • Robert Parsons Interview
      • Dick Sackett Interview
      • Comments by Delena
    • NRCS Interviews: Wyoming
  • Western Weed Control Conference 1940s Minutes
    • 1942 Conference
    • 1945 Conference
    • 1946 Conference
  • Who am I?
    • My Work
    • My Adventures
    • Contact Page
  • Road Logs
    • Colorado Road Logs
    • Idaho Road Logs
    • Montana Road Logs
    • New Mexico Road Logs
    • Texas Road Logs
    • Wyoming Road Logs
  • Bibliography