Chapter Thirteen:

● Types of species interactions

○ Competition
■ Negative and Negative interactions between two individuals (-, -)
○ Parasitism/Predation/Herbivory
■ Negative and Positive interactions between two individuals (+, -)
○ Mutualism
■ Positive and Positive interactions between two individuals (+, +)
○ + or - is influence of populations on each other in terms of influencing vital rates
○ Vital rates include growth, survival, reproduction
■ Competition, mutualism, herbivory, etc. can all occur at simultaneously and
interact irl
● Ecological Niche and Species Interactions
○ Hutchinson niche
■ Full complement of environmental conditions and resources that define
requirements of a species to practice its way of life and to persist
■ Described the niche as "n-dimensional hypervolume"
○ Fundamental and Realized niche (popularized by Hutchinson)
■ Fundamental niche: defined by abiotic factors
■ Realized niche: fundamental niche modified by biotic/species interactions
(competitors, predators, or pathogens)
● Mutualisms can expand realized niche
○ Resource Utilization Curve
■ Expressions of the resource niche
■ Strong competitor is not modified when overlap, weak competitor is pushed out
● Competition Defined and Limiting Resources
○ Competition is mutually adverse effects of organisms on one another within a local
community (Barbour et al.)
■ Brought about by shared requirement for limited resources
● Food
● Nutrient
● Light
● Space
● Nesting sites
● Mates
● Etc.
■ Leads to reduction in vital rates
● Survivorship
● Growth
● Reproduction
● Biomass production
○ Limiting Resource
 ■ Essential resource that constrains vital rates of an organism or population due to
the scarcity of the resource in the environment relative to its need by the
organism
■ A resource is limiting if a change in the availability of a resource produces a
change in vital rates
● Slope is not 0 (Functional response of a predator to abundance of its
prey)
● Corroborated by paramecia lab experiments
○ Lebig’s Law of the Minimum
■ Crop yield is not limited by total resources available, but by the scarcest resource
● “Lowest stave”
● Stave height indicates supply relative to need, not absolute abundance of
the resource
● Specificity, Forms, Scales of Competition
○ Specificity
■ Intraspecific competition is among individuals of the same species
■ Interspecific competition is between individuals of two or more species
○ Forms
■ Interference competition involves direct interactions. Occurs by obstructing
access to a resource (e.g. territoriality in animals).
■ Exploitative (resource) competition involves indirect negative interactions. Occur
via joint consumption of a shared resource in short supply.
○ Scale
■ Among Individuals, can measure:
● Growth, Biomass, Reproduction of individuals
■ Among Populations, can measure:
● Population Growth; Births and Deaths; dN/dt etc.
● Intraspecific Competition and Self-Thinning
○ Monocultures (growing same species in plot of land)
■ Logistic population growth
● Population size, K is density dependent
■ Resource level curve
● R* is Equilibrium resource level: minimum level of resource required to
sustain the population.
○ The level to which a population will draw down a limiting
resource when the population reaches K
○ Optimal Planting Density
■ Over time:
● t0 is initiation of planting
● t1 is increase in average individual plant weight
● t2 is when mortality occurs, thinning occurs
○ Occurs consistently until reaching biomass carrying capacity
(constant yield)
○ Slope of -3/2
 ■ -3/2 Self-Thinning Rule
● At lower planting densities, thinning commences later in time and at a
greater average plant size
● Applies when plotting AVERAGE DRY MASS PER INDIVIDUAL IN
THE MONOCULTURE
○ Law of Constant Yield
■ Plants in monoculture will grow to a constant yield regardless of initial density.
● Introduction to interspecific competition, the competitive exclusion, and coexistence
○ Importance of Interspecific Competition
■ Can influence the realized niche of a species
■ Can affect the distribution and abundance of a species (where a species is present
or absent; how abundant it is where it present)
■ Can influence species coexistence and diversity of communities
○ Outcomes
■ Species 1 outcompetes and excludes Species 2 from the community
■ Species 2 outcompetes and excludes Species 1 from the community
■ Despite competing, species and 1 and 2 coexist in the community indefinitely in
some sort of stable way
○ Coexistence
■ Each species is limited only by itself (by intra-specific competition)
■ Both species increase up to their carrying capacities expected in monoculture
○ Exclusion
■ Large overlap in resource requirement
● Strong Interspecific competition
■ Species 2 excludes species 1 and increases up to its carrying capacity expected in
monoculture
○ Two co-occurring species that share some limiting resources
■ Coexistence: Small overlap in resource requirement
■ Intra >>> Inter
● Each species limits itself much more than it is limited by the other
species
■ Stable coexistence: Both species persist indefinitely and equilibrate below their
monoculture carrying capacities
○ Limiting similarity
■ Concept in theoretical ecology that proposes the existence of a maximum level of
niche overlap between two given species that will still allow continued
coexistence
● Lotka-Volterra competition equations
○ Simultaneously model population growth of two competing species as affected by both
intra- and interspecific competition.
■ Having 2 N terms per equation accounts for Intra and interspecific competition
■ Competition Coefficient (alpha) – Expresses the per capita effect of one species
on another (a simple conversion factor).
 ● alpha12: effect of one individual of species 2 on one individual of
species 1. (converts species 2 into units of species 1)
● alpha21: effect of one individual of species 1 on one individual of
species 2. (converts species 1 into units of species 2)
■ alpha12= 0: Species 1 experiences no competition from species
■ alpha12= 1: Individuals of species 1 see individuals of species 2 as equivalent
competitors.
● Interspecific competition = Intraspecific competition
■ 1 > alpha12 > 0: Individuals of species 1 see individuals of species 2 as weaker
competitors than they do other individuals of species 1.
● Interspecific competition < (weaker than) Intraspecific competition
■ alpha12>1: Individuals of species 1 see individuals of species 2 as stronger
competitors than they do other individuals of species 1.
● Interspecific competition > (stronger than) Intraspecific competition
○ This model assumes strength of interspecific competition (per capita) is equivalent to
intraspecific competition (Individuals of each species see individuals of the other species
as equivalent competitors)
Chapter Fourteen: (LOOK AT SLIDES 30, 31
● Exploitative interactions encompass a huge diversity of interactions nature
○ Exploitation (interaction) enhances fitness of the “exploiter” (predator, parasite,
parasitoid, pathogen, herbivore) while reducing fitness of the “exploited” (prey, host,
plant).
■ Examples of Exploiters:
● Classic predators: kill and consume other organisms
● Parasitoids: Insect whose larva consume its host, eventually killing it
● Parasites: live on the living tissues of their hosts (usually do not kill the
host immediately).
● Pathogens induce disease in their hosts
● Herbivores: consume living plant material
● Exploiter species can influence the fitness, population dynamics, distribution and abundance of
the species they exploit
○ Exploiters impact realized niche (reduce fundamental niche)
■ Overgrazing can result in impeding of plant regrowth
■ Following mange outbreak that killed red foxes, hare population quickly
increased
● Rats that predate on bird eggs and young reduce population of bird
● Reducing predator density increases prey density
● Population dynamics of the exploiter and the exploited are often cyclical and can be modeled
using simple systems of differential equations
○ Prey increase in lack of predators → Predators increase when prey increases → Prey
declines due to excess predators → Predators decline due to lack of prey
■ Both favored to increase when rare, prevents extinction
○ Lotka-Volterra Predator-Prey/Host model:
■ Population Growth dNh /dt =
 ■ Predator Population Growth dNp /dt =
○ Assumptions of the Lotka-Volterra Predation Model
■ Growth of prey limited only by predation
■ The predator is a specialist on one prey species
■ Predator and prey encounter each other randomly in an homogeneous
environment (No spatial structure, no refuge for prey, no opportunity for
immigration).
■ Unchanging environmental conditions
■ c, p and d don’t change
○ Gause’s predator/prey experiment
■ He need both a prey “refuge” and predator immigration to occur for cycling
● Refugia, Avoidance, and Predator-Prey Coexistence
○ Spatial Refuges
■ Environmental complexity and oscillations in lab populations of an
herbivorous mite and a predatory mite *REVIEW*
■ Coral reefs provide structural Refugia for prey species
■ European perch uses spatio-temporal refugia to avoid predators
○ Avoidance mechanisms
■ Mobility
■ Defenses
● Structural
● Chemical
■ Crypsis (camoflauge)
■ Mimicry
○ Natural selection on Prey should favor a reduction in predation rate (p)
■ Favors quickness and elusiveness
○ Natural Selection on Predators should favor an increase in predation rate (p).
■ Favors quickness and agility
○ Red Queen Hypothesis
■ Changes in environments to stay in the same place relative to each other
■ Example is Toxic Sierra Newt and Sierra Garter Snake
● Newt becomes more toxic, snake gains more resistance to toxins
Chapter Fifteen:
● What is Mutualism? How can we distinguish mutualism from other types of interspecific
interactions?
○ Mutualism (+/+) Interactions between individuals of different species that provide Net
benefits to both partners (fitness: growth, survivorship, reproduction).
■ Not to be confused with symbiosis, that is any sort of close, long-term
relationship between two organisms (also parasitic and commensalistic (0/+))
■ Facilitation is positive interactions in general (0/+) and (+/+)
○ Classifying Mutualisms
■ Symbiotic versus non-symbiotic
● Facultative mutualism – gains benefit but does not depend entirely on
partnership
 ● Obligate mutualism – must form partnership to live
● Generalists – can engage in mutualism with many partner species
● Specialists – can engage in mutualism with a select few or only single
partner species
○ Classifying interactions
■ Weakly negative, non-symbiotic, exploitative, low duration and specificity:
herbivory, mosquito bites (generally)
■ Very negative, low duration and specificity, exploitative: true predation
■ Highly negative, high duration and specificity, exploitative, symbiotic: Parasites
● Parasitoids are similar, but VERY HIGHLY negative and specific
■ Nonsymbiotic, weakly positive, low duration and specificity: pollinator
■ True mutualisms are symbiotic, very positive and specific/durtaion
○ Classifying more mutualisms:
■ Obligate Specialists
● Both partners cannot survive without one another and do not have many
different partners.
○ Most well studied, but likely rare in nature!
○ Examples:
■ Leaf cutter ants and fungal gardens
■ Hawk moths and long corolla flowers
■ Ant acacia mutualisms
■ Corals and zooxanthellae
■ Facultative Generalist
● Neither species depends on mutualism, and can get benefits from many
different partners
○ Many cleaner mutualisms are considered facultative and
generalist (i.e. Aspen sunflower-ant mutualism)
■ Facultative Specialist
● Partners do not need one another for survival, but are highly restricted in
who they do associate with.
○ Clownfish and anemone
■ Obligate Generalist
● Both partners need mutualism generally but can associate with many
different partner species to obtain benefits.
○ May be most common type (i.e. Bee and many aster species –
Honey bees have some of the broadest number of partners)
○ Many (but not all ) mycorrhizal relationships (Differences in
specificity between plant hosts and types of mycorrhizal fungi)
● Mycorrhizae and Nitrogen-fixation symbioses
○ Widespread symbiosis between fungi and plant roots
■ Mycorrhizae are widespread (>90% of vascular plants)
■ Ancient relationship
■ Usually Mutualism, but range over broad spectrum (+/-, - /+, +/+)
○ Relationship itself
 ■ Fungus receives carbs and access to organic carbon
■ Plant receives nutrients: enhanced absorption of nutrients (particularly
Phosphorus) and water; protection from soil pathogens
○ Fungal hyphae
■ Long, branching filamentous structures that function as the main mode of
vegetative growth
■ Collectively called a mycelium
■ Mycorrhizal hyphae extend the root systems of plants
■ Increased surface area for absorption
■ Penetrate root cell, intimately allow nutrient exchange and acquisition
● Nitrogen Fixation Symbioses
○ Conversion of atmospheric Nitrogen (N2 ) to NH3 that occurs via root-microbe
symbioses
■ Microbes receive C and energy and plants obtain N in a useable form
○ Major groups
■ Rhizobia (4 genera): Forms nodules on the roots of legumes (fabaceae)
● Benefit to Plant is enhanced access to inorganic nitrogen
● Benefit to Bacteria is access to organic carbon (photosynthate)
● Nitrogen Fixation → Bacteria convert atmospheric forms of Nitrogen
to a form that plants can use for growth.
■ Frankia (1 genera): Forms nodules on the roots of a diverse array of plant in 17
different families.
Chapter Sixteen: (Slide 59)
● Conceptions of the ecological community
○ Community is an assemblage of potentially interacting species that occur together within
in the same defined area
○ The boundaries of some communities are clearly defined
○ The boundaries of other communities are less certain
○ The boundaries of some communities are defined by the investigator for practical
purposes
○ Because of their complexity, community ecologists have to restrict their studies to certain
groups
● Components of community structure (emergent properties)
○ Species Composition
■ Species composition is the collective list of species present in the community or
sample
○ Species Abundance and Abundance Distributions
■ Abundance:
● Absolute abundance is Raw abundance (numbers, biomass, other).
● Relative abundance is Proportional abundance. Useful for comparing
communities or samples that differ in the overall abundance of
organisms.
■ Measuring Species Abundance Distributions
 ● Frequency Distribution (helpful with many rare species, few highly
abundant)
○ Reverse J
○ Our impressions of species abundance distributions can be
strongly colored by sampling effort
■ Logistic species accumulation curve
○ Species distributions generally conform to a log-normal
frequency distribution (bell shaped curve), that can only be fully
revealed by extensive sampling
■ Preston’s veil is the point at which rare species cease to
be sampled
○ Species may be partitioned this way because of:
■ Niche/resource partitioning
■ Random processes (product of many random variables
acting at once)
● Rank Abundance Curve
○ Pi of log scale vs rank
○ Biological Diversity
■ BIODIVERSITY: Variety of life in all its forms, levels and combinations.
Includes habitat diversity, species diversity, functional trait diversity and genetic
diversity
■ SPECIES DIVERSITY: Diversity of taxonomic species in the community, has
two components:
● Species richness (S) is the number of species present in the community.
● Species evenness (E) measures equitability in the relative abundances of
species in the community
● Shannon-Diversity (H’) combines species richness and evenness into a
single measure of species diversity
● Patterns of species diversity
○ Biological diversity varies greatly from place to place; along major environmental
gradients; and in response to environmental change
● Environmental Complexity and Species Diversity
○ Principle of Limiting Similarity
○ Niche Separation/Differentiation
○ Environmental Heterogeneity
● Disturbance and Diversity
○ Disturbance: A perturbation of an ecological community that harms and/or kills
individuals and disrupts population and ecosystem processes
■ Diversity often peaks at intermediate disturbance intensity and/or frequency
● Few species are highly disturbance tolerant, high mortality
● Low diversity because of competitive exclusion, community dominated
by few competitive species
● Moderate disturbance reduces competitive exclusion, allows persistence
of less disturbance-tolerant species
 ○ Community composed of mixture of species – varied
competitive abilities, varied life history strategies
○ Types of Disturbance
■ Abiotic: fire, hurricanes, tornadoes, ice storms, floods, landslides etc.
■ Biotic: disease outbreaks, severe overgrazing, heavy predation human impacts
(deforestation, pollution, agriculture etc.) etc.
○ Disturbance Severity
■ Frequency: how often disturbance occurs
■ Intensity: how severe a single disturbance event is
■ Extent: how much of the habitat is disturbed
Chapter Seventeen:
● Ecological Network
○ An ecological network is a representation of the biotic interactions in a community, in
which species (nodes) are connected by pairwise interactions (links).
■ Interaction links may be trophic, exploitative, competitive, mutualistic or
commensal.
○ Can be viewed from three different perspectives
■ Feeding relationships: Who is eating who, Trophic structure.
■ Material and Energy Flows: How do interactions among species regulate the
movement of matter and energy throughout ecosystems? Trophic dynamics
● Carbon, Nitrogen, Phosphorus. Energy
■ Population and Community regulation: How do the interactions among species
regulate population dynamics and stability, species relative abundances, and
diversity?
● dN/dt, Species Diversity, Community stability
○ Representations
■ Food webs (trophic webs) describe feeding or energetic relationships among
trophic levels (who is eating who)
● Vertical energy arrows
■ Interactions webs describe which species are interacting (who is affecting who)
● Vertical arrows are trophic interactions, horizontal arrows are non-
trophic interactions
■ Bipartite networks are a particular class of interaction networks, whose nodes are
divided into two sets X and Y
● Strong interactors in interaction networks
○ Interaction strength: magnitude of the direct effect of one species on another (often
represented by the width of the arrows)
■ Strong Interactors: The subset of species in the network whose interactions have
the greatest impact on community structure (Paine 1980).
■ A focus on the strong interactors simplifies the analysis of complex
food/interaction webs
● Indirect Interactions
○ Indirect interactions: effect of one species on another via one or more intermediaries
 ■ Trophic cascade: a chain of indirect effects that propagate across multiple trophic
levels
● Keystone Species
○ Keystone Species is a species with a strong influence on the composition, structure, and
diversity of a community that is greater than expected based on its abundance. Types
Include:
■ Predators (wolves in Yellowstone)
■ Modifiers (Ecosystem engineers, such as beavers)
● Ecosystem Engineers: An ecosystem engineer is a species that creates,
modifies, and maintains habitat used for itself and by many other species.
■ Mutualists (acorn banksia and honeyeater)
○ Foundation species (Dominant species) is a species whose influence on the composition,
structure, and diversity of a community is proportional to its abundance
● Keystone predators and species diversity
○