Population

- group of interacting (or potentially interacting) individuals of the same species

living in the same geographic area (or space) at the same time
Distribution
- the pattern of where organisms live
- the way in which individuals of a population are physically dispersed over an area
- includes the size, shape, and location of the area that a population occupies
Abundance
- population size; related to density
- may be differentiated as follows:
1. Population size is the actual number of individuals in a population.
2. Population density is a measurement of population size per unit area (i.e.,
population size divided by total land area).
3. Abundance/relative abundance refers to the relative representation of a
species in a particular ecosystem.

KEY CONCEPTS IN DISTRIBUTION AND ABUNDANCE

1. The environment limits the geographic distribution of species.
→ Law of Tolerance explains the distribution limits of populations
→ While there are few environments on Earth without life, no single species can
tolerate the full range of Earth’s environments.
→ Because of energy constraints (remember the concept of trade-offs; losses and
gains need to be balanced), the physical environment places limits on the
distributions of populations.
→ Environmental factors known to limit species distributions are as follows:
✓ Climatic factors
o temperature, precipitation, humidity/moisture, light intensity
o the effect of climate is sometimes indirect, affecting other factors such
as food availability, water supply, habitat, incidence of other biotic factors
✓ Biotic factors
o parasites, pathogens, competitors, etc.
✓ Abiotic factors
o salinity, pH, intertidal exposure, nutrients, soil/substrate conditions, etc.
2. On small scales, individuals within populations are distributed in patterns
that may be random, regular, or clumped.
- Distribution patterns may also change as individuals of the population grow/mature,
or depending on intensity of interactions (e.g. level of aggression)
 3. On large scales, individuals within a population are clumped.
- On a large scale, there is significant environmental variation.
4. Population density declines with increasing organism size.
Density
o the size of a population in relation to a definite unit of space
a. Crude density
✓ measure of the number of individuals per unit area but populations don’t
occupy all the space within a unit
✓ more general area
b. Ecological density
✓ measured in terms of the amount of area available as living
space/habitat space (e.g., number of birds per km of hedgerow rather
than per hectare)
✓ specific area
NOTE:
➢ It is generally & usually very difficult, if not outright impossible, to know the actual
population size, especially of small, minute &/or mobile organisms at any given time.
So, estimate from a sample (e.g., Capture-Mark-Recapture)
➢ Inverse relationship between density and size
o size-density relationship changes with maturity (e.g. self-thinning in plants)

5. Rare species are often vulnerable to extinction, while abundant species are
seldom so.
- D. Rabinowitz (1981) devised a classification of commonness and rarity based on
combinations of 3 factors:
1. the geographic range of species (extensive vs. restricted)
2. habitat tolerance / range of conditions tolerated (broad vs. narrow)
3. local population size (large vs. small)
→ There are 8 possible combinations of these factors (see figure in next page), 7 of
which include at least 1 attribute of rarity.
→ Most species are uncommon; 7 combinations of range, tolerance, and population size
each create a kind of rarity.
→ In nearly all cases, the key to a species’ survival is increased distribution and
abundance. This is often the key goal of programs aimed at preserving endangered
species.
Population dynamics
- expansion, decline, or maintenance of populations
- one of the most important areas of ecology as it holds the key to understanding
and, hopefully, preventing the decline and extinction of endangered species,
the control of pests, including parasites and pathogens, and the maintenance of
economically and culturally important populations.

DEMOGRAPHIC FACTORS affecting changes in population size through time

1. Natality or Births (B)
o add new individuals to the populations and therefore increase population size 2.
2. Mortality or Deaths (D)
o subtract individuals from the population leading to a decrease in population size
3. Immigration (I)
o individuals coming into and adding to the population size
4. Emigration (E)
o individuals going out and subtracting from the population size

Change in population size (N)

ΔN = (B-D) + (I-E)

KEY CONCEPTS IN POPULATION DYNAMICS

1. Dispersal can increase or decrease local population densities.
→ Immigration (dispersal into the local population) & Emigration (dispersal out of the
local population)
• Passive (esp. plants) – aided by a dispersal agent; distance traveled depends
on the quality of the dispersal agent
• Active (mobile animals) – young vs adults vs sub-adults; male vs female
✓ Natal dispersal – juvenile moves away from the place it was born
✓ Breeding dispersal – adult moves away from one location to breed
elsewhere
• Dispersal decreases during periods of population decline
• Dispersers seek vacant habitat – distance depends on surrounding environment
• Go on exploratory forays before finally leaving the natal site
• Rule of dispersal: move to the 1st uncontested site and no further
• Organisms have dispersing and sedentary stages
→ Migration
- dispersal with a return to the place of origin in response to an evolutionary or
environmental adaptation or pressure
• Repeated - Daily, seasonal (short- or long-range)
• Only 1 return trip (e.g., pacific salmon)
• Only offsprings return (e.g., monarch butterfly)
→ Expanding populations are in the process of increasing their geographic range
→ Range changes in response to climate change, to changing food supply
2. Ongoing dispersal can join numerous subpopulations to form a
metapopulation.

Metapopulation
• A group of distinct, partially isolated subpopulations in widely separated
discrete patches of breeding habitat within a matrix of unsuitable habitats that are
linked only by dispersal / migration
• each subpopulation has its own dynamics (independent); local extinctions; re-
colonizations; rescue effect; highly applicable to problems of habitat fragmentation
and conservation of species
3. A survivorship curve summarizes the pattern of survival in a population.
• Patterns of survival vary from one species to another; dictated by environment
Type I (Convex)
✓ Humans and most primates
✓ organisms tend not to die when they are
young or middle-aged but, instead, die
when they become elderly
✓ Species with Type I curves usually have
small numbers of offspring and
provide lots of parental care to make
sure those offspring survive.
Type II (Straight)
✓ Many bird species
✓ organisms die more or less equally at
each age interval
✓ Organisms with this type of survivorship curve may also have relatively few
offspring and provide significant parental care.
Type III (Concave)
✓ Trees, marine invertebrates, and most fish
✓ very few organisms survive their younger years. However, the lucky ones that
make it through youth are likely to have pretty long lives after that.
✓ Species with this type of curve usually have lots of offspring at once—such as a
tree releasing thousands of seeds—but don't provide much care for the
offspring.
 • Recorded as part of life tables; can be estimated by:
a) Age distribution/population profiles
b) Cohort life tables
c) Static life tables
4. The age distribution of a population reflects its history of survival,
reproduction, and potential for future growth.

Age distribution
- proportion of individuals belonging to each age group
- Gives an echo of past events influencing the population
- May predict future outcomes
- 3 types: (1) Expanding, (2) Stable, (3) Declining

- An age-sex pyramid is a "snapshot" of a population in time showing how its

members are distributed among age and sex categories.
Sex ratio
- proportion of males to females
5. A life table combined with a fecundity schedule can be used to estimate net
reproductive rate (R0), geometric rate of increase (λ), generation time (T), and
per capita rate of increase (r).

Life Tables
- summarize birth and death rates for organisms at different stages of their lives.
Fecundity
- an organism’s potential for reproduction
- measured by the number of gametes, seed set, propagules
o Fecundity schedule
✓ the tabulation of birth rates for females of different ages in a population
TYPES OF LIFE TABLES
1. Dynamic (Age-specific)
- Cohort life table
- follows the fate of each individual in a cohort from birth to death
Cohort
✓ a group of individuals born at the same time or over a short period
✓ Used to determine age- or stage-specific fecundity and mortality rates,
survivorship, and basic reproductive rates, enables an analysis of their annual
variation.
2. Static (Time-specific)
- contains the age groups in a population at one particular period of time (a
snapshot).
✓ Cohorts are not followed in time → reconstructed using one-time observations.
 ✓ The assumption is that the mortality experienced by the cohort at any age
stays constant in time.
EXAMPLE

→ R0 is the average number of offspring each individual leaf.

o A value greater than 1 indicates a growing population.
→ T is the average time from one generation to the next (e.g., from egg to egg or seed
to seed of the next generation).
→ A negative r value would indicate a declining population
→ A positive r indicates a growing population
→ r=0 indicates a stable population
→ r can be used as a sensitive indicator of the ecological effects of environmental
change.
Population Growth
- increase in the number of individuals in a population

KEY CONCEPTS OF POPULATION GROWTH

1. In the presence of abundant resources, populations can grow at geometric or
exponential rates.

Geometric growth
- successive generations differ in size by a constant ratio; populations with pulsed
reproduction
Exponential growth
- continuous growth in an unlimited environment; natural populations may grow at
exponential rates for short periods of time in the presence of abundant resources
- important during establishment in new environments or during the process of recovery
- cannot continue indefinitely, must eventually slow down and population size level
off

Both are J-shaped growth curves

2. As resources are depleted, population growth rate slows and eventually stops;
this is known as logistic population growth.

Logistic Growth
- as population size increases, growth rate eventually slows and then ceases as
population size levels off; sigmoidal or S-shaped growth curve
Carrying Capacity (K)
- the number of individuals of a particular population that the environment can
support
- the population size at which growth stops
- b=d; r=0
o If N<K, r is positive and the population grows
o if N = K, r=0 and population growth stops
o if N>K, r is negative and the population declines
Growth Curves (see figure):
A. Exponential
B. Logistic
C. Exponential growth with discrete generations
D. Logistic growth with medium time lag
(dampened oscillations)
E. Logistic growth with large time lag

3. The environment limits population

growth by changing birth and death
rates.

Density-dependent factors
- environmental factors that are often influenced by population density (e.g., biotic
factors such as disease, predation, competition, etc.)
Density-independent factors
- environmental factors that exert their influence on the population independent of its
density (e.g., abiotic factors such as flood, extreme temperature, etc.)
Life History
- the series of changes undergone by an organism during its lifetime
KEY CONCEPTS OF LIFE HISTORIES

1. Because all organisms have access to limited energy and other resources, there
is a trade-off between the number and size of offspring.
- those that produce larger offspring are constrained to produce fewer, while those
that produce smaller offspring may produce larger numbers. (~large but few vs.
small but many)
o Animals that produce larger eggs and plants that produce larger seeds
lay fewer eggs and produce fewer seeds, respectively.
2. Where adult survival is lower, organisms begin reproducing at an earlier age
and invest a greater proportion of their energy budget into reproduction; where
adult survival is higher, organisms defer reproduction to a later age and
allocate a smaller proportion of their resources to reproduction. (~high adult
survival leads to delayed reproductive maturity)
o Reproductive effort – the allocation of energy, time, and other
resources to the production and care of offspring
o In organisms, energy supplies are allocated to three functions:
maintenance, growth, and reproduction.
o The exact amounts allocated for each depends on the organism and life
stage.
3. The great diversity of life histories may be classified on the basis of a
few population characteristics.
- Examples include fecundity or number of offspring (mx), survival (lx), relative
offspring size, and age at reproductive maturity (α).
o There are different ways of classifying life history patterns but only one
will be discussed, which is the r and K selection of MacArthur and
Wilson, 1967 and Pianka, 1970.
r selection – selection favouring a higher population growth rate
K selection – favours more efficient utilization of resources
o Classification into r and K selection never captures the full diversity of
nature, but makes understanding it easier
o Most species fall somewhere in between the extreme types.
Life history information plays a key role in conservation and restoration of ecosystems and
endangered species.
 Characteristics favoured by r versus K selection

Population attribute r selection K selection

Intrinsic rate of increase, r max High Low

Competitive ability Not strongly favoured Highly favoured

Development Rapid Slow

Reproductive maturity Early Late

Body size Small Large

Reproductive events Single, semelparity Repeated, iteroparity

Offspring Many, small Few, large

Survivorship curve Type III Type I or II

Description of population r strategist; K strategist;

ruderals (in plants) Stress-tolerant (in plants)
 POPULATION ECOLOGY: Population Responses to
Climate
- Major determinant of where organisms can live
- The niche space in which species can survive, grow, and reproduce is determined in
large part by suitable climatic conditions.
- When climates change, population attributes, dynamics, and interactions change as
well.

A. Species Range Shifts

Species’ range
- area in which it is found, including both its extent (extent of occurrence) and the
locations within that extent that are actually occupied by the species (area of
occupancy)
- determined by the spatial distribution of individual populations.
- Abundance, which is often governed to some degree by climate, determines
whether populations endure.
• When populations are lost on the range periphery, or when new populations
appear that expand the range, a range shift occurs.
Species’ ranges shift over time to track suitable climate.
- When climate changes in a location, some species may find themselves in suddenly
hostile conditions (recall Shelford’s Law of Tolerance). Others will find that
previously unsuitable climates have changed in their favor.
o Individuals in unsuitable conditions will die or fail to reproduce, gradually
disappearing from the location, whereas individuals near newly suitable
habitat will gradually occupy areas in which they have not occurred
previously.
o In other words, species are able to track suitable climatic conditions, occupying
new areas and leaving unsuitable locations as climate changes, a
process sometimes termed “niche tracking”.
o The usual trend is a poleward and upslope shift in a wide variety of settings
and regions.
Drives of Range Shifts
1. By long-term changes in mean climate state
- e.g., coral bleaching
2. By short-term climatic extremes such as freezing
3. By interactions with other species being driven by climate change
- e.g., arctic fox and red fox.
Managing these movements is one of the great challenges for conservation in
this century.
Tropics - are sensitive to climate change.
- Tropical organisms generally have narrower fitness tolerances to temperature
than temperate ones.
o This is because tropical species are evolved to thrive in a relatively narrow
temperature range. Thus, although the magnitude of climate change may be
 highest at high latitudes and near the poles, sensitivity may be greatest in the
tropics.
The impact of climate change is the product of the magnitude of change and
sensitivity.
- Expressed from the standpoint of the species, exposure to change multiplied by
sensitivity equals vulnerability.
- Vulnerability can be reduced by conservation actions. However, for the most part,
impact will scale with sensitivity and exposure.
- Physical change may occur first in high latitudes where warming is
greatest, but the greatest biological damage may occur in the tropics, where
sensitivity is greatest.

B. Extinction
- When climate changes become too severe for species to keep up through
range shifts, extinctions can occur.
- One example is the extinction of the golden toad (Bufo periglenes) caused by
warming and drying of the cloud forest.
- Another extinction event is that of the harlequin frog (Atelopus sp.). Warming
enhanced the growth of a pathogen that caused the death of the frogs
resulting to a rapid loss of the species in its limited range.
- As climate changed, these 2 amphibian populations had nowhere else to go
because they are both mountaintop species.
C. Changes in Phenology
Changes in timing of biological events due to climate change (some examples):
1. Earlier spring timing of events
a. Earlier budburst and flowering
b. Earlier caterpillar development BUT young hatchlings have not shifted sufficiently
to keep pace (lead to decline in bird abundance)
c. Earlier spring migrants (for birds and butterflies) and first flights of those in
diapause
d. Earlier spring vocalization in frogs
e. Earlier nesting and laying of bird eggs
2. Earlier ice breakup in temperate lakes and rivers leading to…
a. Algal blooms
b. Changes in stratification of lakes
c. Changes in trophic structure
3. Spring ahead + fall behind = longer growing season
4. Changes in tropical forest phenology – changes the whole trophic
structure
5. Changes in marine seasonal variations – changes the processes occurring
at the base of the marine food web (phytoplankton) leading to changes in the
whole marine food web
6. Accelerated life cycles
→ leading to trophic mismatches, changing species interactions in the process
Thermal-cued responses change more rapidly in response to climate change but
photoperiod responses are evolutionary so they don’t respond to warming. This leads to
trophic mismatches and timing mismatches between interacting species.