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What is ecology?
Study of the environment; scientifically, the study of interactions among organisms in an environment; interdisciplinary (biology, genetics, evolution, phys. sciences, earth sciences, social sciences)
Not environmentalism → not a moral stance
Scales of ecology
Individual ←→ pop. ←→ community ←→ ecosystem ←→ biosphere; all impact each other; hierarchical
Spatial scale → globe to bacteria (m) (the amount of space in which something occurs)
Temporal scale → glacial retreats to bat echolocation (days) (the amount of time in which something occurs)
Climate vs weather
Weather → day-to-day variation (in temp., rainfall, humidity, etc.)
Climate → longterm average description of weather
Why do we care about climate?
Impacts species distribution and abundance → differs according to species (Rufous crowned vs House sparrows); species dist. are related to constraints imposed by the env.
Climate - what do we mean?
Averages, extremes (heat and coral reefs, drought and borer beetles), variability, timing
All impact species distribution and abundance; a matter of which of these factors are doing what
Why does climate vary across the earth?
Bands of wet and dry (Hadley Cells), uneven solar input, tilt and orbit of the earth, winds and Earth’s rotation (Coriolis Effect), topography, ocean currents
Hadley Cells
Large-scale atmospheric convection cells in which air rises at the equator and sinks at medium latitudes
How?
Intense solar heating at the equator → air expands and rises as it warms → rising cools it down, forming clouds → cold air holds less moisture → moisture falls as rain → more air rises → dry air is pushed towards the poles → gets denser; sinks → dry bands created → warm air picks up moisture and rises → completes the cycle
Solar radiation
Ultimate source of energy that drives global climate
Uneven input from the sun → at the equator, solar radiation is concentrated over a small area (warmer); in the northern hemisphere, solar radiation is concentrated over a larger area (cooler)
Tilt and orbit of the earth
Creates the seasons → different in northern vs. southern hemis.; no season at the equator (wet and dry seasons, but no actual change in day/night length)
Coriolis Effect
Differences in velocity mean that as air or water moves over Earth’s surface, their trajectories appear to be deflected (moving faster at the equator; think of throwing a ball from Costa Rica to Anchorage → it’ll end up in front of them)
Topography
Going up a mountain = colder temperature and more rain
Rainshadow → the dry area on the leeward side of a mountain (opposite of wind’s direction); air warms and travels up the mountain, cools to form clouds, drops moisture on wayward side, dry by the time it reaches the leeward side → creates a wetter climate on one side, and a dryer climate on the other
Ocean Currents
Move the cold and warm water around
Biomes
Based on the growth form of the dominant vegetation; a combination of precipitation and temperature; line up with the bands of wet and dry
Examples:
Tropical rainforest → diverse; temp. doesn’t really change; precip. can change (but there’s lots of it)
Tropical dry/seasonal forest → temp. consistent; more severe dry seasons (trees will drop leaves)
Temperate grassland → big swings in temp. and precip.
Temperate seasonal/deciduous forest → big swings in both (but lots of rain)
Temperate evergreen → huge amounts of rain; little bit of seasonal temp.
Deserts (hot or cold) → very little rainfall; temp. fluctuations
Boreal forest → huge swings in temp.; decent amt. rainfall
Tundra → basically freezing year round
Physiological tolerance
Limits on environmental conditions that an organism can tolerate (Rufous crowned vs. House sparrow)
Tolerance and species ranges
Tolerance defines species ranges
There’s an optimum (of many factors and conditions) that a species can do its best in → differs between species
Lizards → temp. and sprint speeds (Goldilocks zone)
Dr. Janzen → tropical species have a limited range of temperatures they can handle; nontropical species can typically tolerate a wider range
Environmental stress strategies
Avoidance - migratory birds; ephemeral annuals; hibernators
Tolerate - polar fishes; succulents
Shifts in species ranges
Some species track changes in temp. and precip. → Grinnel Resurvey Project (84% of birds change their ranges across 100 years)
Evolution
Before genetics: a change in distribution of a pop. from one generation to the next; pop.-level shift in trait dist.
Now: change in allele frequencies
Can happen rapidly (native anole lizards and arboreal traits)
Mechanisms of evolution
Natural Selection
Gene flow
Genetic drift
Mutation
Natural Selection
The process by which individuals with certain heritable characteristics survive and reproduce better than individuals with other heritable characteristics
Non-random
Tenets of evolution by natural selection
Individuals vary in their traits
Traits are inherited from their parents
Individuals with certain traits are better at surviving and reproducing than individuals with other traits
Types of natural selection
Directional selection
Stabilizing selection
Disruptive selection
Directional selection
Changes the average value of a trait (a small or large trait is favored) → e.g., Darwin’s finches
Effect on heterozygosity depends

Stabilizing selection
Central trait value is favored; no change in the average trait value over time (selection against the extremes) → e.g., human birth rate (lower weight = hard to maintain body heat; higher weight = birth complications)
Decreases heterozygosity

Disruptive selection
Extremes are favored; no change in the average trait value over time (selection against the middle); can lead to reproductive isolation and thus speciation → e.g., seedcracker finches (finches with larger mandibles could handle hard seeds; finches with smaller mandibles could handle soft seeds; finches in the middle were the masters of none)
Increases heterozygosity

Genetic Drift
Due to chance; random with respect to fitness (survival and reproduction); greater in small populations
In small populations, the loss of alleles and fixation of alleles can lead to an increased frequency of harmful alleles → shrimp-spined wolves; prairie chickens
E.g., a landslide randomly wipes out the majority of a population → bottleneck; drastically reduces the size of a population
Decreases heterozygosity
Gene flow
The movement of alleles between populations; can introduce new alleles to a population; can make different populations with gene flow resemble each other
E.g., hatchery fish (wild x captive caught spices up diversity [new population resembles both parent pops.])
Increases heterozygosity
Mutation
A change in the DNA of a gene; ultimate source of genetic variation; leads to the formation of new alleles (beneficial [or otherwise])
Increases heterozygosity
Genetic diversity
The total number of genetic characteristics within a pop.; influences the ability of a population or species to respond to changes in the environment (higher g.d. = greater ability to respond)
Founder event
When a small number of individuals set out from an ancestral population to establish a new, isolated colony (new colony only has a fraction of the g.d. of old colony)
Foraging
How organisms select, search for, and consume food within their natural environment
Foraging - costs and benefits
Costs - decrease fitness → often energetic or predation risk
Benefits - increase fitness for survival and repro. (finding the energy required to grow or be reproductively active)
Organisms must find the balance
Optimal foraging theory
Evolution has favored behaviors that maximize an individual’s rate of food harvest and/or while minimizing costs
Prediction: If organisms are foraging optimally, they’re going to maximize the net benefit while minimizing costs
Predators change the costs (not benefits); changes in food resources change the benefits (not costs)
Max benefit: closest, highest peak
Costs: increase linearly
B=C → max distance an organism should forage away from the den before costs outweigh benefits
OFT - diet choices
Energy content = E (1 or 2)
Handling time - time it takes to process one prey item (h)
Profitability - energy gained per unit of time (E/h)
Critical search time
((E1 • h2)/E2) - h1
Compare to the search time of the preferred prey item
If > CST, organism should invest in a mixed diet
If < CST, organism should only eat preferred prey
Factors such as time of day, temperature, season, etc. can impact all of the above
Social behavior
Any direct interaction among a group of individuals of the same species that is organized in a cooperative manner—extending beyond sexual and parental care
Costs and benefits of group living
As group size increases, more competition within group; as group size decreases, more competition between groups (and a higher risk of predation)
Must find optimal group size
Evolution of cooperation
Mutual benefit
Reciprocity
Kin selection
Delayed direct fitness benefits
Cooperative breeding
Mutual benefit
Defense against predators; protection; acquisition of resources (food, shelter, etc.)
Schools of fish, nesting colonies of seabirds (predation risk lowered)
African wild dogs
Birds (larger the group, the fewer head jerks [more time to eat]; as flock gets bigger, handling time decreases)
BUT, the bigger the group, the quicker resources are depleted and the more often they have to move to look for food
Costs → dilution of resources, predator attraction, increase in parasites/pathogens, cheating (on effort)
Reciprocity
If you scratch my back, I’ll scratch yours
Allogrooming
Vampire bats and sharing food (more likely to share with those who shared before)
They must be able to recognize each other; must be able to remember who helped them before
Kin selection
Between related individuals; individuals will gain direct and indirect fitness benefits by helping
Direct fitness benefit → fitness through one’s own reproduction
Indirect fitness benefit → fitness through helping kin
Inclusive fitness → sum of direct and indirect fitness
Coefficient of relatedness (r ) → proportion of an individual’s total genotype that is identical to another due to shared ancestry
50% for mother, father, sibs → r = 0.5
25% for grandparents, nieces → r = 0.25
12.5% for cousins → r = 0.125
Unrelated individuals → r = 0
Identical twin → r = 1.0
Hamilton’s Rule
Equation to decide whether cooperation occurs due to kin selection
r = B - C > 0
r = coefficient of relatedness
B = benefit to the recipient (usually # offspring/dominant male - # offspring/solo male)
C = cost to the actor (usually # offspring/solo male - # offspring/subordinate male)
If r > 0, net positive; kin selection occurring
If r < 0, net cost; kin selection not occurring
Delayed direct fitness benefits
Gain direct fitness benefits later on → common in lekking groups (learning to attract mates); manakins (Michael Jackson birds)
Cooperative breeding
Helping raise others’ young → extreme helping (helping 💯)
Sexual selection
Differential reproductive success due to variation among individuals and success at getting mates → might result in survival deficit (but, hey, at least they had sex)
Male and female differences → females get to be choosier (an egg is a lot more energetically expensive than sperm)
Skewed mating success → a few males might get most of the matings
Lekking species → 80% of the matings go to a few males
Two forms: intrasexual and intersexual
Intrasexual selection
Male - male competition; fighting within the same sex; leads to the development of exaggerated weapons (antlers, horns, tusks, claws, songs)
Intersexual selection
One sex (usually female) chooses a mate from among members of the opposite; leads to the development of exaggerated ornaments or dances
Direct benefits → parental care, resources, protection
Indirect benefits → good genes, sexy sons (sons who are more likely to find mates and thus spread genes)
Peacocks → males with more eyespots have more matings and a better immune system; bigger eyespots = higher probability that the offspring will survive
Alternative reproduction strategies
Sneaker males → rush in and release sperm on females’ eggs
Satellite males → help males
Males that look like females
When sexual and natural selection are at odds with each other
What’s good for the ladies might not be what’s best for, you know, living
Ravens
Populations have increased by 300%; ravens are generalists → can take advantage of human infrastructure; can regularly impact species of concern (can push small, isolated groups to extinction; anthropogenic features mean there’s no limitation of prey abundance)
Managed by
Lethal removal, egg removal, nest removal, resource management
BUT native species; Migratory Bird Act
Habitat selection
The process by which an organism chooses specific environmental patches/living areas based on factors that influence its survival and repro. rates
Space use - how organisms distribute themselves, move, and utilize the physical areas within their environments
Life histories
How and why organisms survive and reproduce in different ways → strategy of the major events in an organism’s life
Examples
Size and # of offspring
Body size
Specialization (vs. being a generalist)
Age of maturity
Size at maturity
Lifespan
Life history schedules
Different for each species
Black rhino (K-selected): “If I can survive until next year, I can reproduce again.”
Slow pace of life → steady, predictable environments
Long lifespan
Reproduce at long intervals; long gestation
1 offspring/pregnancy
Investing in growth for future fecundity
House mouse (r-selected): “I probably won’t live until next year, anyway. Live hard, die hard.”
Fast pace of life → variable, unpredictable environments
Short life spans
Short reproduction interval; short gestation
Lots of offspring/pregnancy
Investing in immediate reproduction
How many young should an organism have?
Lack’s hypothesis: Clutch size is determined by the max # of offspring an organism can feed
As clutch size increases, the probability of each offspring surviving decreases (becomes a curve)
Parental cost → bigger broods decreased survival and reproduction of the parents (it takes a lot of energy)
Those with smaller broods probably don’t have a lot of energy to go around
Reproductive decisions
Timing of breeding
# offspring/attempt
# young/year
Amt. of parental care → early growth period = critical for later survival
Everything’s a tradeoff
Allocating limited energy to growth and reproduction in a way that’s efficient within evolutionary and environmental constraints
Classifying life histories
r-selected species: high population growth rate, not very strongly competitive, rapid development, small, fast maturation, semelparitous (single) reproduction, lots of (small) young
K-selected species: low population growth rate, high competitive ability, slow development, large, slow maturation, iteroparous (many) reproductions, few (large) young
Life history traits
Fast-slow gradient → spectrum of ways organisms live their lives
Slow-fast life history continuum → resource availability, unpredictability of an environment (variability)
Fast life histories in the temperate zone
Competitive - Stress - Ruderal (Grime’s) Triangle

Trade-off examples
Migration → better environment, but lots of energy
Being flashy → better chances at repro., but also of predation
Offspring # vs. size
Current reproduction vs. future reproduction/survival
Higher reproductive effort is correlated with higher mortality
Greater costs for “poorer quality” individuals: how good is the organism at surviving and reproducing (more likely to be affected by other trade-offs)?
Greater costs following harsher winters → resources are not always limiting for all individuals or in all conditions (rich = cheating at life)
Conservation implications
Reproduction will vary more than survival for slow life histories (often trade off repro. outlet for survival)
Ducks → when habitat was better, females had higher natural mortality (more energy towards repro.; more likely to die) → reproduction has become more costly for adult females
Is managing for repro. still the most effective management strategy—even if ducks have a slow life history?
Population ecology
What influences where species are found?
Density
Abundance
Population
A group of interacting individuals of one species living in a specific area
Interacting = reproducing sexually → group of interbreeding individuals; reproducing asexually; other kinds of interactions (competition, etc.)
Characteristics of populations
Size, density, age structure, sex ratio, genetic structure, dispersion
Size → # individuals in an area
Density → # of individuals per unit area (pop. size/habitat size)
How to determine size and density?
Count them all (geographically restricted populations [e.g., island])
Sample populations
Area-based count
Distance sampling
Mark-recapture
Population abundance
Relative representation of a species in a particular area; measured via population sampling methods
Area-based count
Quadrats → counting # individuals in a given area or volume
For organisms that don’t move much and can be counted accurately
Assumption: area is representative of the entire population you’re sampling
Distance sampling
Measurements of distance from a point/straight line
For organisms that move a lot and can be counted accurately → birds, etc.
Mark-recapture methods
Capture, mark with individual bands, release, recapture (at a different point in time)
For organisms that are mobile that you can capture and recapture → organisms that are hard to detect (nocturnal, etc.)
Simple population growth models
r = b • d
r = estimated annual pop. growth rate
b = per capita birth rate
d = per capita death rate
b > d → pop. increasing
b < d → pop. decreasing
b = d → pop. stable
Simple population growth models cont.
r = b • d / t • Navg.
t = time
Navg. = avg. pop. size over period of time (average of two [or more] pop. sizes provided)
Simple population growth models assume that b and d are consistent over time (closed pop. → no migration, etc.)
Immigration → movement into pop. → increase abundance
Emigration → movement out of pop. → decrease abundance
Age structure
Proportion of pop. within each age class; whether pops. are increasing or decreasing → not every age class dies or reproduces at the same rate
Abundance is good, but not always sufficient → loggerhead sea turtle example; turtles don’t start reproducing until adulthood (takes a long time); large juveniles were the most important stage (not eggs)

Life tables
Display how survival and reproduction vary with age, size, or the life cycle stage; 2 types:
Static
Cohort
Static life tables
Calculate age-specific survival and fecundity at one period in time
Cohort life tables
Follow individuals from birth to death (dynamic); work great for short-lived species
Cohort - group of individuals born at the same time
Life table variables
Nx → age group
lx → survivorship (proportion of individuals that survive from birth - given time [x]) → Nx/N0 (starting pop.)
qx → mortality rate (how many are dying between age classes)
1 - (Nx+1/Nx) → Nx+1 = next age class down from age of interest (Nx) → nothing fancy; don’t panic
Fx → fecundity (total fecundity of entire pop. at age x)
Mx → per capita fecundity → per-female fecundity for age class x → Fx/Nx
lx • mx → survivorship • fecundity → R0 → net reproductive rate per generation
How many offspring each female has
R0 > 1 → pop. increasing
R0 < 1 → pop. decreasing
R0 = 1 → pop. stable
T → generation time → (sum of x • lx • mx column [age • lxmx])/R0
r vs. R0
r is relative to 0; R0 is relative to 1
r = ln (R0)/T
Population growth models
Geometric
Exponential
Logistic
Geometric growth
Populations are increasing; unlimited resources; discrete time intervals
Pops. reproduce in synchrony (at discrete times)
Pops. grow by constant proportions; not limited in growth → algae, invasive species, disturbed environments
Lambda → annual rate of change → N(t+1)/N(t)
> 1 → pop. increase
< 1 → pop. decrease
= 1 → pop. stable
Can predict pop. size in future:
Nt = Lambda^t • N0
Exponential growth
For species that reproduce at all different times (continuous reproduction), but pops. still change by constant proportions and are unlimited in their growth
dN/dt (slope) → change in pop. over time
dN/dt = rN (intrinsic growth rate • pop.)
Predict future pop. size
Nt = N0e^rt
N0 = starting pop size
r = intrinsic growth rate
t = time
Logistic growth
Populations cannot continue unlimited growth forever → pops. grow until K (carrying capacity) is reached
K = max # of individuals that an environment will sustainably support
dN/dt = rN(1 - N/K)
What drives logistic growth and influences pop. size?
Density dependence
Density independence
Density dependence
Initial rates change as a function of the density of a pop.
Disease, food, light, space
Negative density dependence → logistic growth patterns (pop. growth decreases as density increases)
Positive density dependence → Allee effects (pop growth decreases and density decreases [pack hunters, etc.])
More regular fluctuations around K
Density independence
Vital rates are not a function of pop. density
Fires, pollution, floods, severe weather (not caused by the # of individuals)
Larger, more severe and sudden fluctuations around K
Type I survivors
Low mortality at young age; higher mortality at the end of life (old age); K - selected species → humans, slow species, most larger mammals
Type II survivors
Mortality rate constant throughout life (linear) → birds, lizards, small mammals
Type III survivors
Higher mortality at younger ages (very low chances of survival); mortality rate decreases as age goes on → faster species; r-selected species; invertebrates
Survivorship curves and extinct species
Aged (at time of death) fossils and survivorship curves can help us understand the lifestyle of extinct species
Population dynamics
The ways in which populations change in abundance over time
Annual K is rarely constant → changes based on resources
Fluctuations are the rule
Variation in the density of a population; can be influenced by how much the environment fluctuates (tropics vs. temperate) and the inherent stability of a population (slower species vs. faster species)
Irregular fluctuations
Stochasticity → random events
Periodic fluctuations
Delayed density dependence → population performance reflects density in previous years → species responding to what happened in the past
Time lags → delays in response of birth and death rates to changes in the environment
Delayed density dependence and logistic growth
Time lags and DDD → dN/dt = rN (1-N(t-†)/K)
† = time lag
Overshooting K and then plummeting below K (degraded K) is common
Extinction risk and variable growth
Greater variability → more likely to go extinct (> variability in lambda > extinction)
Ecological Traps
When an individual mistakenly thinks that a bad quality habitat is good quality habitat → environments are falsely attractive → mayflies
Can be used as a management strategy (goats in the Galapagos)
Mismatch in evolutionary cues
Undervalued resources
When an individual mistakenly thinks that good quality habitat is bad quality habitat → environments are falsely unattractive
Mismatch in evolutionary cues