ENVS 230

Lecture 9
October 20, 2015

• Summarize intraspecific competition

– Review for Midterm
• Life History Patterns
• Life History Theories
• Go over worksheet 1
• Go over quiz 1
 Beyond Scope of
Course Population Growth Later in Course

One level One Level

Density-Independent; Density Independent;
No age structure With Age structure

Discrete Continuous Discrete Continuous

Stable Age Transient cycles due to

Exponential
Distribution variation in age class
Growth
distribution
One Level One Level
Other Density Dependent;
Density-Dependent;
Mechanisms With Age structure
No age structure
Contest
Continuous Evolution of
Discrete Competition
Life History

Scramble Logistic Model Stable equilibrium

Competition Herbivore- Overshoots/cycles
Resource Model Equilibrium
Other models Two Level Models
No Age Structure Predator-Prey, Interspecific Competition
 One level
Density-Independent;
No age structure

• dN/dt =N*r = N*(per capita growth rate)

– Differential equation; continuous
– Identical to compound interest
• Nt+1 = Nt +dN
• Nt+1 = Nt*(1+ per capita growth rate)
– Difference equation; discrete
– Identical to periodic (e.g., annual) interest
• Both result in exponential growth
 Per Capita Growth Rate
• G = r Density-independent
• G = f(N) Density-dependent
• Density-Independent
– dN/dt = r*N Nt = N0ert
– Nt+1 = Nt(1+s) Nt+1 =N0*(1+s)t
 One Level
Density Independent;
With Age structure

• As population increases,
– b (birth rate) and/or i (immigration) decreases; and/or
– d (death rate) and/or e (emigration) increases;
– These are density-dependent factors;
• Instraspecific competition for resources major
mechanism
• Mechanisms not independent
– E.g., Competition increases exposure to predators
– Or Avoiding predators increases competition
 Density-Dependent Per capita Growth Rate
• dN/dt =N* f(N) = N * r * g(N)
– g(N) is density-dependent part of equation
– If g(N) = 0, get exponential growth
• If g(N) decreases as N increases, then population tends
toward equilibrium
– Caused by resource competition, etc.
• g(N) increases as N increases: Allee effect, depensation
– Unlikely for all N
– Various mechanisms cause g(N) to increase with N up to point,
after which it decreases
 Intraspecific Competition
• Scramble Competition
– All individuals have equal access (symmetric)
– Current growth rate based on current resource
levels
– Decrease in resource levels occurs later; this
regulates population
 Contest Competition
• Includes, but not limited to “contests”
• Asymmetric, where some individuals limit
access of others to resources
• Regulation occurs at current population
level, which determines resource restriction
 Modify Growth Equation
• Can change growth equation to include density-
dependence
– Pearl-Verhulst (logistic) is a simple form
– rN (population growth rate) multiplied by term
which decreases with density
 g(N) decreases as N increases

• Simple form is line g(N) = 1 – aN

– Identical to g(N) = 1-N/K
• Then dN/dt = r*N*(1-N/K)
 Logistic Population
Growth rate
• dN/dt = rN(1-N/K)
• Per capita growth rate+
– (dN/dt)/N = r(1-N/K)
• dN/dt = f(N) = parabola
• Maximum at K/2
• At low N, per capita growth
rate is very high, but N small
• At high N, per capita growth
rate is very small, but N small
• Maximum at intermediate N
and per capita growth rate
N (population size)
Integrate to
Logistic Curve
get N(t)

Perturbation:
Population
decreases to K
 Phase Diagrams
• Will come into play more when we discuss
predation and competition
• Will discuss then
– Translation: won’t be on midterm
 Worksheet 1: All Results
r
0.02 0.04
stoch 2 10 20 2 10 20
n0 46 2 0 46 0 0
n1 1 0 0 2 0 0
n2 0 0 0 0 0 0
n3 0 0 0 0 0 0
n4 0 1 0 0 0 0
n5 0 0 0 0 0 0
n6 0 11 0 0 2 0
n7 0 6 1 0 1 0
n8 0 15 1 0 1 2
n9 0 5 2 0 2 3
n10 0 8 42 0 40 42

ave/10 0.02 7.44 9.85 0.04 9.67 9.85

 Life History
• With age-structured populations, we
projected population size based on age-
specific birth and death rates
• Life History Theory is about control of
those age-specific birth and death rates
– Proximate: genetically fixed; plastic; variability
within and among species;
– Ultimate: evolutionary causes
 Life History Traits: How Do These Evolve?

• Semelparity (breed once) vs iteroparity

• Univoltine (one generation per year) vs. multivoltine
• Age at maturation
• Age-specific reproductive investment of adults (how
much at each age); senescence
• Offspring size (a few large, many small)
• Level of parental care
• Spawning period within season (short or protracted)
 Four Groups of Theories of
Life History Evolution
1. Demographic: builds on life table
– Different reproductive values of different life stages lead to
different allocations of fecundity-risk across life stages
2. Density-dependence
– Different life histories favored in situations where population is
often growing rapidly (unpredictable environment) versus
situations where population is near carrying capacity
(predictable environment)
3. Environmental variability
– Relative variability of adult or juvenile mortality or fecundity
controls selection for early or late reproduction, etc.
4. Case-by-case analysis of response to specific environments
Environmental Determinants of LH
• In single environment, typically
see a range of life histories
• Thus, need to explain variety of
life histories
• Fine-scale environmental
variation
• Same environments are different
to different organisms (scale,
etc.)
• There are multiple successful
strategies
 Trade-offs
• All explanations depend on trade-offs
among life history traits
• Evident in comparisons across species
• Evolution of LH traits depends on similar
trade-offs within species
• Use of interspecific variation to explain
evolution, which is intraspecific, is common
in evolutionary ecology
 Cost of Reproduction:
energy allocation
• Acquired energy split among
– Maintenance
– Activity
– Reproduction
– Growth
– Reserves (e.g., fat in animals, root/tuber starches in
plants; since reserves can be used to supply yolk or
fruit, some reserves may be treated as reproductive)
• Increasing allocation to reproduction means
decreasing one or all of the others, or increasing
energy acquisition (see below)
 Trade-offs
Activity
Loss
Growth

• Among reproduction,
Reproduction growth, reserves,
Maintenance
maintenance, activity
Activity Loss
• Decreases lead to
decreased survival,
Growth future reproduction
Maintenance
Reserves

Reproduction
 What is Parental Investment?
• Should include all costs (e.g., energy, nutrient)
associated with reproduction
• One problem with testing ideas has been difficulty
of accounting for and measuring all components
• Development of eggs, sperm
– Most easily measured
– Proportion of total egg weights to body weight crude
measure
• Spawning movements and associated costs
• Display and spawning territory costs
• Parental care
 Trade-offs among the
number and size of offspring
• Typically many small offspring with little or no parental
care vs. few larger offspring (sometimes with
considerable parental care)
 Plasticity in Amount of Parental Care
Some birds use asynchronous hatching and siblicide to regulate the
number of young.

In times of food shortage, the younger (later-hatched) are either

ignored by the parents or killed by their older siblings.
 Timing of Reproduction (among-year)
• Semelparity (semelparous reproduction);
Single reproductive event
– Most insects
– Many other invertebrates
– A few vertebrates (Pacific salmon,
American eel, lampreys)
– Many plants, including annuals, biennials,
bamboos
Long
• Iteroparity (Iteroparous reproduction); prereproductive
period
Multiple reproductive events with fewer
offspring
Annuals Biennial
– Most vertebrates
– Perennial plants

Latin itero; to repeat; as in

iterate, iteration
Iteroparity Under Partial Environmental Control
• American shad
– Iteroparous in north (lower metabolism in cool water)
– Mixed in mid-Atlantic
– Semelparous in south (higher metabolism in warm water)
– In Delaware River, affected by longer migration route
because of downstream dams, pollution
 Timing of Reproduction (within year)

• Single bout of reproduction

– Short
– Long
• Batch
– Increases production for given size
– Uses repeating events (e.g., phase of moon;
rains)
 Variability among/within years:
Predator Swamping
• Young stages vulnerable to predators
• Large number of young at one time may be
more than predators can consume
• Masting: nut crops differ among years; may
be synchronization among species
• Fish larvae
Henry Ford and the Model T
Henry Ford supposedly checked junkyards to see which parts had failed
or not.
Why?

To see which parts needed improvement in order to last

longer.
To see which parts he had built too well so that he could
make them shoddier and save money.

Easton, PA 1935
 Demographic Explanation: Balance of
Survivorship and Future Reproduction

• High Adult Mortality selects for

– Early maturation
– High early fecundity
• High Juvenile Mortality selects for
– Later age at maturity
– Lower reproductive investment per year
– Longer reproductive life
• Correlational and Experimental Approach
 Harvesting often Increases Adult Mortality
• Harvesting larger fish may
select for
– slower growth (Atlantic
silversides)
– Earlier age of maturity
(Atlantic Cod, hypothesized
for Brook Trout)
 Note Potential Positive Feedback

• Higher adult mortality

– Selects for early reproduction, which increases
later mortality
• Higher juvenile mortality
– Selects for lower reproductive investment per
year, which decreases adult mortality
 Example: Guppies
(Reznick, Endler, etc.)

• In Trinidad, guppies occur in high-predation (pike-

cichlid) and low-predation (killifish or none)
environments
 Example: Guppies (Reznick, Endler, etc.)

• Transplant experiments from

high-predation sites to sites
without guppies or major
predators showed selection for
later maturation time
• Response time differed among
populations and between males
and females
--------- control group (high adult mortality)
_______ experimental group (juvenile mortality)

After introduction into a stream with

fish that preyed on juvenile guppies,
across 11 years:
• Increased age at maturity
• Larger size at maturity for females
• Decreased allotment to reproduction
• Fewer offspring produced
• Larger size of offspring

Why is this everyone’s favorite example?:

• Mix of field and lab studies
• Careful measurement of genetic contribution
• Long period of study
• Lots of nuances, complications
 Density-Dependent
Explanation
(r- and K- strategists)
• Different life histories may be better at low-population,
high-growth phases (called r-selection) than at high-
density, near-equilibrium phases (K-selection)
• Refers to logistic equation, but justification really depends
on interspecific competition models (more later)
• For now, think of this as intuitive idea

Partly grew out of

dispute between
bird and bug
people
 Density-Dependent Explanation
(r- and K- strategists)
• Low-density environments: r-selected
– E.g., Short generation time, early maturation, high
reproductive investment, small offspring
– Also, dispersal (growth phase may be transient)
– Low investment in “expensive” traits for obtaining food
(e.g., structural, detoxification), since other food
available
• Species frequently near equilibrium: K-selected
– Late maturation, high parental care, large offspring
– Also, characteristics to out-shade, hold territories, etc.
 Pros and Cons of r & K gradient
• Conceptual (con)
– Life history traits associated with r, K selection those of age-
structured population, i.e., outside scope of model which produced
them
– r and K themselves are not individual traits open to selection
– Misleading: selection is not on “r” and “K”, but other LH parameters
• Explanatory power (pro and con)
– Doesn’t explain full variation in life histories
• E.g., Cod (large adults, many small larvae)
– Mixed strategy (?) of many organisms with high dispersal of early
stages
– Rarely is direct study made of link between traits and typical
population conditions
• Concept explains some variation in life history traits, so even if
flawed and incomplete, is still useful (and very often used) (pro)
 Environmental Variability
• Bet-hedging theories
• In variable environments, traits balance risks
– E.g., with highly variable among-year variation in survival of newborns,
optimal to spread reproduction over a number of years, i.e., lower investment
in each year, leading to higher adult survival, etc.
– With variable survival within season, protracted spawning better than pulse
spawning
– May be multiple optima: e.g., conservative (“pessimizing”) strategies which
do ok in most conditions but can’t take advantage of extremes vs flexible
strategies which can respond to good times (and bad)
• Note links to economics: Bet-hedging; risk-taking, risk aversion; similarity is not
a coincidence
 Life History as Adaptation to Specific
Environment

• Most salamanders
– Aquatic larval stage

• Redback salamander
– Direct development egg to
terrestrial adult, so species
not tied to aquatic
reproduction
– May necessitate larger egg
and low clutch size
 C. ladon

Insect
Voltism
• Univoltine: one brood per year, with C. idella

diapause of various stages rest of year

• Multivoltine: multiple broods per year

Celastrina neglecta
 Univoltism vs Multivoltism
• Related to length of season: many species
multibrooded in south; univoltine in north
• Related to phenology (timing of various events) of
host plant and tissue
Variety of
host
Celastrina plants
neglecta
Multivoltine
summer

C. idella Eats
Univoltine Holly flowers
Early spring
 Correlational Analysis of Fish Life Histories
(Winemuller and Rose)

• Compiled information on life history traits

• Calculated correlations among traits
• Found correlations (“strategies”)
• Statistically analyzed gradients in groups of life history traits
• Two major gradients in life histories
– Adult size, fecundity, number of spawning bouts
– Investment per offspring
• These gradients show many of the trade-offs previously discussed,
but the correlation among traits somewhat different
 R% + C% + S% = 100%

100 0

100
0

100 Stress 0

Grime’s model of life history variation in plants based on three strategies:

• R = ruderals (disturbed places; annual herbs): Reproduction

• C = competitive (predictable environments): Resource acquisition
• S = stress-tolerant (limited resources; lichens): Maintenance

Note rough correspondence of Ruderals and r-selected;

Competitive with K selected
 Back to Predicting “Best” Strategy
• Earliest approaches estimated optimal strategies, and
assumed populations would reach these
– General selection research program of period, growing from
other (engineering, economic) optimization approaches
• Later approaches more dynamic
– Evolutionarily-stable strategies (ESS); traits which can
invade a population with different traits; game theory
approach
– Direction of selection (dynamic approach)
– Elaboration of physiological or phylogenetic constraints
– Can lead to frequency-dependent changes, with co-existence
of different strategies or oscillations
 Other Life History Traits Outside
Standard LHE models
• Resting stages
• Parthenogenesis, sexual or mixed
reproduction
• Sex change
• Complex life histories
– Multiple stages
• Paedomorphosis
Questions?
 Review
• Unpopular subjects back by popular
demand
 Water as Solvent
• Because of slight polarity, many compounds are highly
soluble (dissolve) in water
– E.g., Ionic compounds (NaCl), alcohol, sugar
• Low solubility
– Nonpolar covalent bonds (e.g., oxygen)
– Many organics
• Solubility function of temperature, pressure, pH
• Dissolution of less soluble substances can be important
– E.g., Calcium carbonate (limestone) slowly dissolved
 Why We Care

• Aquatic systems
– Determines availability and transport of
nutrients, oxygen
– Affects pH
– Determines osmotic pressure of water requiring
energy to keep solutes in tissues (in fresh
water) or out (in salt water)
 Why We Care: Terrestrial Systems

• Water moves through terrestrial systems

• Picks up (leaches) or deposits minerals in soil
– Depends on nature of soil (soil exchange capacity) and
bedrock composition and rate of weathering
– Exchanges minerals with soil
• E.g., Al replacing Ca (increasing soil water toxicity)
• Ca replacing Na in highly saline soils (remediation technique)
– Precipitates from saturation (e.g., stalactites)
– Left following evaporation (many saline, alkali soils)
• Big effects on soil toxicity, fertility, structure
• Many organisms eat clay Soil
• E.g., macaws and other parrots
– Thought that clay binds to and neutralizes
toxins in nuts and fruits
 Low (thin)
Viscosity
• Fluid’s internal resistance to flow
– Varies with temperature
– Different concept than density
– Among common liquids
Water is low viscosity
Why we care:
• Think about
Swimming through High
Honey (thick)
• Effect of viscosity
Depends on size, shape of
organism; for small organism,
water is viscous
• Affects swimming, sinking, feeding
 Possible Explanation For
• Small organisms (larvae,
Microcrustacea, etc.)
• Superbly not streamlined
– Doesn’t matter
– Predators do
• Appendages have “fingers”,
not “paddles”
 If decrease
• Maintenance (often treated as constant in
energetic models, but treat as variable here),
– increasing chance of disease, injury, etc.)
– e.g., grooming, resting metabolism, detox
 If decrease
• Activity; Decreased activity could mean
– Increased risk (less vigilance, preparation for or
movement away from risks)
– Lower future growth or reproduction (less
foraging, or movement to favorable feeding,
resting, or breeding areas);
 If Decrease Growth
• Often means
– Lower fecundity in future years
– Higher predation risk (outgrow
predators)
– Lower ability to win contests
(food, territories, mates);
– (Decreasing growth may also
decrease maintenance costs,
may also decrease future risk of
starvation, etc.)
Growth-Reproduction
Tradeoff

Douglas fir

Round sardinella
Tradeoff between present and future
Reproduction
 If Decrease Reserves
• Often means
– Higher risk of starvation
– More need to forage in high risk conditions
– For warm-blooded animals, decreases insulation
and cold tolerance
 If increase energy acquisition
• Increased risk
• Time allocation issues and decreased time
for other activities
 Life History

• Interaction of environment and genetics

• E.g., Age of Maturation controlled by
amount of food individual gets, as well as
genetics
 Cost of Reproduction:
energy allocation
• Acquired energy split among
– Maintenance
– Activity
– Reproduction
– Growth
– Reserves (e.g., fat in animals, root/tuber starches in
plants; since reserves can be used to supply yolk or
fruit, some reserves may be treated as reproductive)
• Increasing allocation to reproduction means
decreasing one or all of the others, or increasing
energy acquisition (see below)
 Types of Parental Care
Mouth brooding
by cardinalfish
• Gestation
• Nest Building
• Feeding, grooming, cleaning, teaching
• Fanning
• Guarding (attack, distraction, mouth
brooding)
• Opportunity costs (feeding)
• Dead salmon fertilize nutrient-poor
streams

Marty Snyderman

Distraction
displays (feigned
broken wings) by
plovers
Reproductive Risks

• Risk of injury and disease

– Direct injury in combat with resultant disease risk
– Aggregation can increase disease or parasite
transmission
• Vulnerability to predation
– During territorial advertisement and defense, mate
seeking, parental care
– Spawning migrations
• Decreased feeding time and opportunity
 Reproductive Risks
• Decreased feeding time
• High energetic costs for migrating in warm
water
Review of Age Structured
Population Growth
 Age-Structured: Life Table

• Like discrete (difference equation), except

fecundity and survival differ among ages
• Life table shows these schedules
• Need to be careful whether table shows age
specific survival or mortality, or cumulative
mortality up to age

Age (Years) Age Specific Fecundity age-spec mortality age-specific survival

0 0 0.75 0.25

1 2 0.2 0.80

2 3 0.2 0.80

3 3 0.2 0.80

4 0 1 0.00
 Projection
• Like non-age structured, add net births and deaths
over time period
• Except need to do for each age class and add up at
end
Life Table Number at age
Age Specific age-specific
Age (Years) age-spec mortality 1 2
Fecundity survival
0 0 0.75 0.25 600
1 2 0.2 0.80 300
2 3 0.2 0.80 200
3 3 0.2 0.80 100
4 0 1 0.00 0

Example from worksheet (gray squirrel)

• Relatively short life
• Relative even fecundity
• Start near stable age distribution
• Looks like exponential
 Previous
example
Different Example

Average Age Specific age-spec age-specific

AGE (Years) Number
length (cm) Fecundity mortality survival
0 3 1.89 0 0.9993 0.0007 7217.9
1 9 51.03 0 0.7 0.30 5.9
2 16 286.72 0 0.5 0.50 2.1
3 22 745.36 0 0.3 0.70 1.2
4 30 1890 0 0.1 0.90 1.0
5 34 2751.28 0 0.1 0.90 1.1
6 37 3545.71 2000 0.1 0.90 1.1
7 39 4152.33 3000 0.08 0.92 1.2
8 40 4480 4000 0.08 0.92 1.3
9 41 4824.47 5500 0.08 0.92 1.4
10 42 5186.16 9000 1 0.00 1.5

Similar to second example on worksheet

• Long-lived, long age until maturity
• High variability in fecundity
• Start far from Stable Age Distribution
• Large, damped oscillations
• Trending, but long time until exponential
 Consequence

• Chance event (stochasticity) of high/low mortality

or fecundity pulls population away from SAD
• Long time to return
• Large Individual age class important
 Chesapeake
Bay Striped
Bass

Young-of-Year Index

From Secor, 2000, ICES

More Recent Data
MD DNR