HURLBERT THE EXPERIMENTAL APPROACH
ABSTRACT 5 COMPONENTS TO AN EXPERIMENT
Pseudoreplication
 (In manipulative experiments) Defined as
the use of inferential statistics to test for
treatment effects with data from
experiments where:
o either treatments are not
replicated (though samples may be)
or
o replicates are not statistically
independent
 In ANOVA terminology:
o The testing for treatment effects with
error term inappropriate to the
hypothesis being considered
 Refers to not a problem with experimental
design, but to a particular combination of
experimental design (or sampling) and
statistical analysis which is inappropriate
for testing the hypothesis of interest
Nondemonic intrusion
 The impingement of chance events on an
experiment in progress
 As safeguard against this and other
preexisting gradient, INTERSPERSION is
considered an obligatory feature of a good
design.
 Randomization and interspersion can
sometimes be in conflict with one another,
especially in small experiments.
Comprehension of this conflict can be aided
by:
o Distinguishing pre-layout
(conventional) and layout-specific
alpha (the probability of a type I
error)
 Type I error = when the null
hypothesis is rejected, when it
is in fact true
 When it was
concluded that there
is a significant
difference, when there
really is none
 Hypothesis
o Of primary importance
 If it is not good, the
experiment will be of little
value
 Experimental design
o The logical structure of the
experiment
o Based on the objectives, and the
objectives should describe or specify
the:
 Nature of experimental units
to be employed (ex. the type
of glassware)
 The number and kinds of
treatments
 Properties or responses of
the experimental units that
will be measured (ex.
accuracy and precision)
o It should specify:
 How treatments are assigned
to the available experimental
units
 The number of replicates per
treatment
 Physical arrangement of
experimental units
 Temporal sequence in which
treatments are applied
 Measurements made on
different experimental units
 Experimental execution
o Success requires avoiding systematic
error and minimizing random error
 Systematic error renders an
experiment invalid or
inconclusive
o Errors in experimental execution are
often subtler and more likely to
happen than design errors + they
are more difficult to detect
 Statistical analysis
o To increase the clarity, conciseness,
and objectivity with which results are
presented and interpreted
  Interpretation
 Interpretation and statistical analysis are the
least critical aspects of experimentation, as
the data can be reanalyzed.
 The only complete remedy for design or
execution errors is repetition of the
experiment.
MENSURATIVE EXPERIMENTS
 Involve only the making of measurements at
one or more points in space or time
 Space or time is the only experimental
variable or treatment.
 Usually do not involve the imposition of an
external factor, thus, every experimental
units are treated identically
QUESTION:
EXAMPLE:
 Objective: to determine the decomposition
rate of maple leaves in a 1-m isobath
COMPARATIVE MENSURATIVE EXPERIMENTS
 Measures a property of the system at two
points within it and asking whether there is
a real difference
EXAMPLE:
 Objective: to determine the decomposition
rate of maple leaves in a 1 m isobath vs. 10
m isobath
 Hypothesis: the decomposition rate will be
different
IMPORTANT:
 The dispersion of the replicate samples in a
manner appropriate to the hypothesis being
tested is the most critical aspect of a
mensurative experiment.
PSEUDOREPLICATION IN MENSURATIVE
EXPERIMENTS
 Often a consequence of the actual physical
space over which samples are taken, or
 Measurements being made smaller than
the inference space implicit in the
hypothesis being tested
o Inference space is the space in which
the samples occupy. It should be
congruent with the sampling domain,
which is based on the objectives of
the experiment.
MANIPULATIVE EXPERIMENT
 Involves two or more treatments
 The different experimental units receive
different treatments and the assignment of
these treatments can be randomized
Difference with Mensurative Experiments:
 Mensurative: comparing photosynthetic
rates of naturally established oak and maple
trees
o Treatment variable: species
o Randomized assignment of
treatments to locations WOULD
NOT BE POSSIBLE
 Manipulative: comparing decomposition
rate of maple and oak leaves
o Treatment variable: species
o Randomized assignment of
treatments to locations IS POSSIBLE
CRITICAL FEATURES OF A CONTROLLED
EXPERIMENT
Critical Features of Experimental Design:
1. Controls
o “Any treatment against which one or
more other treatments is to be
compared”
o May be an:
 Untreated treatment
 No imposition of an
experimental variable
 Procedural treatment
 Simply a different treatment
o In experimentation with biological
systems, controls are required
because these systems exhibit
temporal change.
o Controls for temporal change and
procedure effect
 o Regulation of the conditions under
which the experiment is conducted
 The homogeneity of
experimental units
 Precision of treatment
procedures
 Regulation of the physical
environment in which the
experiment is conducted
o ^ However, the above definition is
unfortunate usage, as the adequacy
of true controls is independent of
this regulation of physical
conditions, thus the validity of the
experiment is unaffected by this
regulation.
__
 Replication and randomization both
improve estimation and permit testing.
2. Replication
o Controls for the stochastic factor,
which is the variability among
replicates arising from:
 The material itself
 The experimenter
 Nondemonic intrusion
o Reduces the effects of random error
and increases precision
o With respect to testing, its main
purpose is:
 supply an estimate of error
by which the significance of
these comparisons is to be
judged
3. Randomization
o Controls for potential experimenter
bias in the assignment of
experimental units to treatments and
the execution of the experimental
procedure
o Increases accuracy of estimates
o With respect to testing, its main
purpose is:
 To guarantee the validity of
the significance test based on
the error estimate from
replication
Why is this important?
 Randomization confers validity because it
guarantees that, on the average, errors are
independently distributed.
o The lack of independence of errors
prohibits us from knowing alpha, the
probability of a type I error.
4. Interspersion
o Controls for regular spatial variation
in properties of the experimental
unit
 The physical layout of the
experiment and how
experimental units should be
distributed in space
o Most of the time, when you have
randomization, you also have
interspersion, but that is NOT always
the case.
 Interspersion is the most
critical concept, and
randomization is simply a
way of achieving
interspersion  eliminate
bias  allow accurate
specification of the alpha
o For preliminary assessment of the
adequacy of experimental designs,
interspersion is more practical to
consider than randomization.
Sources of Confusion in Manipulative Experiments:
 MODES OF SPATIAL INTERSPERSION AND
SEGREGATION
A-1 COMPLETELY RANDOMIZED DESIGN
 Most basic and straightforward way of
assigning treatments to experimental units
 Not frequently employed in ecological field
experiments (when experimental units are
large)
 Has a good chance of producing treatments
that are segregated rather than spatially
interspersed
A-2 RANDOMIZED BLOCK DESIGN
 Commonly used design in ecological field
experiments
 Reduces the probability of chance
segregation of treatments
 Helps prevent pre-existing gradients and
nondemonic intrusion
A-3 SYSTEMATIC DESIGN
 Achieves a very regular interspersion of
treatments but runs the risk that the spacing
interval coincides with the period of some
periodically varying property of the
experimental area
 In both systematic and randomized block
designs, we can base the assignment
process NOT ON THE LOCATIONS but on
the internal properties of the experimental
units prior to imposition of treatments.
o Risk: spatially segregated treatments
 The magnitude of this risk decreases with
increasing number of replicates.
B-1 AND B-2 SIMPLE AND CLUMPED
SEGREGATION
 Simple segregation
o Only one layer of treatments, but the
placement of a treatment group is
adjacent to the same group
 Rarely employed in ecological field
experiments
 More commonly found in laboratory
experiments
 Dangers lead to spurious treatment effects.
Causes:
o Pre-existing difference in the
“locations” of two treatments
o Non-demonic intrusion
 Difference between locations
can become greater during
the experiment
independently of any true
treatment effect
B-3 ISOLATIVE SEGREGATION
 Poses all dangers of simple segregation but
in a more extreme form
 Further increase the likelihood of a spurious
treatment effect, as within-treatment
variances are less likely to be increased
B-4 PHYSICALLY INTERDEPENDENT REPLICATES
 Example:
o 4 aquaria in each set sharing
common heating, aeration, filtration,
etc.
 The replicates are dependent on each
other.
RANDOMIZATION VS. INTERSPERSION
 POSSIBLE SOLUTION (Cox)
1. Reject highly segregated layouts and
rerandomize
o Risk: precludes the probability of
knowing the exact value of alpha
Pre-layout and Layout-specific Alpha
 αPL
o Conventional alpha
o The probability, averaged over all
possible layouts of a given
experiment, of making a type I error
  αLS
o the probability of making a type I
error if the layout was used
o will usually be less than or greater
than the pre-layout alpha
 When not adhering to strict randomization
procedures, the αPL will have marginal error.
However, it does not mean that it cannot be
done. It is up to experimenter to determine
whether knowing the aPL exactly is better
than having an idea of a possible upper
bound to aLS.
Biased estimation of treatment effects
 The greatest bias in estimating a treatment
effect will result from some particular
nonsystematic design and not from a
systematic one.
PSEUDOREPLICATION IN MANIPULATIVE
EXPERIMENTS
 If treatments are spatially or temporally
segregated (B1-3), if all replicates of a
treatment are interconnected (B-4), or if
“replicates” are only samples from a single
experimental unit (B-5), then it is
pseudoreplication.
o At best, they can only demonstrate a
difference between locations
SIMPLE PSEUDOREPLICATION
 Involves only a single replicate per
treatment
 The validity of using unreplicated treatments
depends on the experimental units being
identical at the time of manipulation and
after manipulation, except as there is a
treatment effect.
o The lack of significant difference
prior to manipulation cannot be
interpreted as evidence of
identicalness, as it can only be a
consequence of a small number of
samples taken from each unit.
 Two experimental units are different in
every measurable property
o Thus, if we increase the number of
samples taken from each unit, our
chances of finding a significant
premanipulation difference will
increase (alpha will approach 1.00)
 THIS IS NOT WHAT WE NEED.
 Replicability is a false issue. It pertains to
the similarity that can be obtained once the
experimental design is repeated.
o The question to be asked is NOT:
 Are experimental units
sufficiently similar for one to
be used per treatment?
o Rather, it is:
 Given the observed or
expected variability among
experimental units, how
many should be assigned to
each treatment?
TEMPORAL PSEUDOREPLICATION
 Multiple samples from each experimental
unit are taken sequentially over several
dates, which are then taken to represent
replicated treatments
SACRIFICIAL PSEUDOREPLICATION
 Results when an experimental design:
o Involves true replication of
treatments but data are pooled prior
to statistical analysis, or
o Where two or more samples taken
from each experimental unit are
treated as independent replicates
(the concept of subsamples)
 Example:
o In any field situation, two replicate
plots or ponds in the same
treatment are NOT identical, so a
significance test of the difference is
not relevant.
CHI-SQUARE AND PSEUDOREPLICATION
Employing a Chi-square analysis pn subsamples
rather than replicates.
IMPLICIT PSEUDOREPLICATION
 When authors seem to regard paired and
non-overlapping 95% confidence intervals
as equivalent to significance tests, and if
 they offer no specific disclaimer that their
data is inadequate in assessing treatment
effects

QUESTIONS
 Can mensurative experiments have control
treatments?
o Mostly, no.
 Which is better, to randomize the glassware
or to randomize the people who measure?
o
 Where will we need subsamples? And where
would subsamples be a waste of time?
o Sdsd
 EVOLUTIONARY ECOLOGY
 Seeks to understand evolutionary
connection between traits of populations
and species and their biotic and abiotic
environment.
 An essential part of introductory ecology
because the proximate explanations for
ecological phenomena are environmental,
but the ultimate explanations are typically
rooted in evolutionary theory.
EXPLANATIONS FOR ECOLOGICAL PHENOMENA
 Proximate explanations
o Immediate cause of a phenomenon
o Ex.
 Birds lay fewer eggs, have
smaller clutches during times
of food scarcity
 Proximate explanation
for fewer eggs is food
scarcity
 Temperature, day length,
moonlight intensity, time
after sunset
 Ultimate explanations
o More general reason; rooted in
evolutionary theory
o Ex.
 As the bird in the example
evolved, those birds that lay
fewer eggs in times of food
scarcity get to pass on their
genes in future generations.
Overtime, this trait became a
common feature of the
species.
 Seasonal temperature cycles
that may optimize
physiological performance
for sexual reproduction in
corals, wind speeds and tidal
phases that may maximize
fertilization success and
dispersal, and diurnal cycles
that may allow for predator
avoidance.
 Why we have to study evolution in an
ecology class
EVOLUTION AND ADAPTATION
 Evolution
o A genetic change in a population of
organisms over time
o Organism’s ecological situation
directs its evolution
o The organism’s response to its
ecological situation may be
evolutionary
o Population
 The basic evolutionary unit
 A group of conspecifics
(members of the same
species) inhabiting a
specified geographic area.
o Occurs over several hundreds,
thousands, etc. of lifetime
 Adaptation
o A genetically determined
characteristic (whether behavioral,
morphological, or physiological) that
improves an organism’s ability to
survive and reproduce in a
particular environment
 This characteristic must be
expressed as a phenotype.
o Verb: Evolutionary process whereby
organisms become better suited to
their environments
o It is driven by natural selection
 Acclimatization
o Phenotypic differences based on
how genes are expressed as a
function of the environment
EVOLUTION IN POPULATIONS
HARDY-WEINBERG THEOREM
 Mathematical algorithm that detects and
measures genetic change, that might be the
result of evolution
Assume a diploid organism polymorphic for A (A, a)
with following genotype frequency:
 AA: 30 individuals (30/100 = 0.3)
 Aa: 20 individuals (20/100 = 0.2)
 Aa: 50 individuals (50/100 = 0.5)
Genotype frequency – the proportion of individuals  It is the variety of alleles that creates
in the population bearing a specific genotype. variation in a population.

0.3 AA + 0.2 Aa + 0.5 aa = 1

For haploid:
 Allele frequency for A:
o p = (30 +30 + 20)/200 = 0.4
 Allele frequency for a:
o q = (50 + 50 + 20)/200 = 0.6

What does it mean?

 40% of their gametes will carry A allele.
MECHANISMS OF EVOLUTION
 60% of gametes will carry the a allele.
Genetic drift
 Random shifts in allele frequencies
 Occurs in small and large populations but
the effect is most apparent in the former
 Determined by effective population size
(Ne)
 Two mechanisms:
o Founder effects
 When a subset of the original
population becomes a new
population
o Population bottlenecks
 Can occur when large
portions of a population are
killed by a chance event
unrelated to genotype

Gene flow
 Loss or gain of alleles resulting in change in
allele frequencies
 Occurs via emigration and immigration

Natural selection

NATURAL SELECTION
 the differential representation of genotypes
in future generations, resulting from
 The equilibrium genotype frequencies is in heritable differences in survival and
the first row. “Hardy-Weinberg reproduction among these genotypes
equilibrium”
 Elements:
 Equilibrium o Inherited variation exists among
o Means that the population is not individuals in a population
evolving or the genetic makeup is  Traits
not changing
 o Not all individuals have the same
phenotype
 Phenotype – what the
environment sees; expressed
as a character
o Resources are limiting, hence there is
competition among organisms for
scarce resources
o Phenotypes (and corresponding
genotype) which can garner more of
the scarce resources leave more
offspring than others.
 X-axis – histogram bins (i.e. beak depth)
 Y-axis – frequency
Natural selection results from differences in fitness,
which is based on adaptation.
 Stabilizing selection
o Selection against the extreme and
for the average
o The average beak depth did not
change
 Directional selection
o Shift in the average condition
o Will lead to either one of the
extreme (but not both)
 Disruptive selection
o Selection against the average and for
the extremes
o Will lead to the prominence of 2
extremes
FITNESS
 The contribution by an individual’s
descendants to future generations
o It doesn’t matter if the individual has
a lot of muscles or controls a lot of
resources; what matters is the ability
to reproduce.
 Absolute fitness
o The expected number of offspring
produced by a particular genotype in
a population
 Relative fitness
o The relative ability of a genotype to
obtain representation in the next
generation.
o Expressed as percentage
 o Small beak depth = small food
resource

 Imagine: The ancestors of birds first

colonized Charles, Chatham. From here, they
colonized the area of Daphne.
o Directional selection, along with
stabilizing selection (because one
extreme are no longer represented
in the population in Daphne).
 Imagine: From Charles, Chatham to
Crossman
o Disruptive selection

TOLERANCE AND ECOLOGICAL NICHE

 Coevolution leads to divergence of
phenotypes in two species.
o Energy used supposedly for
reproduction will be used for
competition, thus, it will reduce
fitness.

Abiotic factors and Limits

 Physical resources
o Abiotic factors that an organism
must assimilate if it is to live and
prosper
 Physical factor denotes other kinds of
abiotic parameters whose physical or
In the exams: chemical effects may delimit a zone in which
life is possible. When these determine the
presence or absence of a species, they are
referred to as limiting factors.

Shelford’s Law of Tolerance

 There are upper and lower bounds to the
physical factors that an organism can
tolerate (which can still change as a result of
evolution)
o This tolerance is the result of
adaptation by natural selection or
acclimatization.

Ecological Niche
 The ecological role of a species
 A hypervolume in multidimensional space
 The differences in beak depth has to do with
defined by the intersection of resource
food sources.
utilization and tolerance curves
 o Defined by niche optima, niche
breadth (how wide the niche is), and
niche separation
 Higher niche overlap indicates higher
competition
EVOLUTIONARY ECOLOGY IN TEXT BOOK
 Phenotypic variation among individuals in a
population results from the combined
effects of genes and environment
o Ex,
 Differences among
populations in the three
common garden sites
indicate genetic differences.
 Differences within
populations in the three
common garden sites
indicate environmental
differences among the
common garden sites, NOT
genetic differences.
 Phenotypic plasticity
o When organisms change the
expression of traits in response to
the environment
o Phenotypic variability that might not
actually be caused by genotypic
variability
o If one has phenotypic plasticity, the
environment leading to that change
in phenotype is NOT affecting the
genetic makeup of the population,
which insulates the genome from
changing, and hence, will inhibit
selection
o Related to “acclimatization”, because
it happens in ONE life span
 With reduced gene flow, populations could
diverge genetically. Such genetic divergence
would increase the genetic variation among
populations.
 Populations with the highest levels of
inbreeding or with the lowest
heterozygosity had the highest probabilities
of extinction.
SPECIES INTERACTIONS AND COMPETITION IN
TEXT BOOK
 Symbiosis
o Interaction in which most or all of
the life cycle of one organism occurs
inside or on another
 Under commensalism is:
o Inquilinism – when one organism
uses another for residence
 If the impact is negative, it means there is
energy expended or injury incurred.
 In general, competition will be negative for
both parties because even the winner will
have expended more energy to gain that
resource than it would have if there was no
competition.
COMPETITIVE INTERACTIONS
 Interference competition
o Involves direct interactions between
individuals
 E.g. excessive defense of
territories, cannibalism (which
plays into how interference
may okay a role in
intraspecific competition,
even in the absence of
obvious resource limitation.)
 Intraspecific competition
o Competition with members of their
own species
o Self-thinning rule
 Plants will decrease in
population density as the
total biomass of the
population increases
 Consequence of intraspecific
competition for limited
resources
 o May also happen in the absence of
obvious resource limitation
 Interspecific competition
o Competition between individuals of
2 species that reduces the fitness of
both
 This reduction in fitness may
not be equal
 Resource or exploitative competition
o Competition involving the use of
limited resources
Other ideas:
 Resource limitation
o When there is a limited supply of
resources that causes competition
COMPETITION AND NICHES
 Competitive Exclusion Principle
o Proposes that 2 species with
identical niches cannot coexist
indefinitely, which leads to the
prediction that coexisting species
will have different niches
o Realized within the context of
interspecific competition
o Competitive exclusion can result
from competition for food
 Niche partitioning
o Done by organisms of different
species with similar niches to reduce
competition
o An example of strong selective force
competition
 Priority effect
o Under intermediate conditions, the
species establishing itself in greater
numbers first generally wins out in
competition
 Competition can have short-term ecological
effects on the niches of species by
restricting them to realized niches, yet
these species may retain their capacity to
inhabit the fuller range of environments we
call the fundamental niche. However, if
competition persists, they might have an
evolutionary response that would alter their
fundamental niches.
 Long-term competition between two
species can lead to evolutionary divergence
in morphology.
 Because interspecific competition reduces
the fitness of competing individuals,
individuals that compete less should have
higher fitness than individuals that compete
more.
Interspecific competition  character displacement
 Character displacement
o Circumstance in which 2 species
differ more from each other in
geographic areas where they occur
together than where their
distributions do not overlap
Six criteria to build a definitive case for
character displacement:
1. Morphological differences between a pair
of sympatric species are statistically greater
than the differences between allopatric
populations of thee same species
2. The observed difference have a genetic
bases
3. Differences between populations must have
evolved in place (they evolved from the
same island)
4. Variation in character must have a known
effect in use of resources
5. There must be demonstrated competition
for resource and competition must be
directly correlated with similarity in the
character
6. Differences in the character cannot be
explained by differences in the availability of
resources to sympatric and allopatric
populations
 POPULATION ECOLOGY
INTRODUCTION
Populations
 Group of conspecifics inhabiting a specific
place in a specific time
o Conspecific – individuals belonging to
the same species
 A human construct, i.e. they are not
normally discernible in nature
Parameters that Describe Populations
 Population density
o number of individuals per unit area
or volume, or as percentage cover
o can be expressed as crude density or
ecological density

Boundaries of populations are defined by many

means, mostly arbitrary:
 Relative to species ranges
o Relative to where it is found
 Relative to life-span of individuals
 To maximize important processes (e.g.
reproduction, dispersal) within boundaries
compared tot hose between the study area
and other regions
 Genetically; e.g., demes – locally defined
population characterized by random mating
within the group  Dispersion
o Spatial distribution of organisms

 Age structure
o The distribution of individuals in the
various age classes

 Sex ratio
o The proportion or percentage of the
population that is either male or
female
 POPULATION DENSITY
Crude density
 Number of individuals per unit area or
volume
Ecological density
 Number of individuals per habitable unit
area or volume
 May not be appropriate for clonal
organisms, or those whose individuals or
modules are difficult to distinguish
o Ex. corals
 Organisms may be unitary or modular
o Genet – A genetic individual arising
from a zygote
o Modules produced asexually by the
genet are ramets
 They may be physically linked
to the parent or separate
 They are clones or exact
copies of the parent genet
 Population density declines with increasing
organism size.
o Self-thinning
 Species living at low population densities
generally have small, restricted distributions.
 3 factors important for the Classification of
Commonness and Rarity by Rabinowitz
o Geographic range (extensive vs.
restrictive)
o Habitat tolerance (broad vs. narrow)
o Local population size (large vs. small)
DISPERSION
 In dispersion, the patterns are not easily
discernible. There should be quantitative
ways to distinguish one from the other.
Patterns of Dispersion
These patterns are produced by (1) social interactions
within populations, (2) structure of the physical
environment, (3) a combination of both.
a. Random patterns (Random dispersion)
o An individual has an equal
probability of occurring anywhere in
an area
o Processes:
 Neutral interactions between
individuals and between
individuals and local
environment
o Individuals are randomly spaced
within the community
 Not that much common in
nature
 This occurs as there are no
strong interactions between
the individuals
 Show neutral interactions
b. Regular patterns (Uniform dispersion)
o Processes:
 Antagonistic interactions
between individuals or local
depletion of resources
o Individuals are uniformly spaced
within the population
 Mainly results from the
competition for resources
 Therefore there are
aggressive interactions
between the neighbors
c. Clumped patterns
o Attraction between individuals or
attraction of individuals to a
common resource
o “contagious dispersion”
o Most common pattern that can be
observed in populations
o Individuals are bunched into groups
within the population
  Mainly results from the
response to the unevenly
distributed resources in
their environment
 also happens if there are
positive interactions
(mutual attraction) between
individuals
o Advantages:
 Provides protection from
predators
 Facilitates finding mates and
reproduction.l
 Decrease the energetic cost
of food search
 Ecological concepts are very scale specific. A
pattern in 1x1 meter can be very different
from a patter in 20x20 meter
 On small scales, individuals within
populations are distributed in patterns that
may be random, regular, or clumped.
 On large scales, individuals within a
population are clumped.
o Why?
 Because the environment
varies and individuals
aggregate in areas where the
environment in favorable.
Dispersal can increase or decrease population
densities.
 Density of predator populations can
increase with increased prey density.
 Ongoing dispersal can join numerous
subpopulations to form a metapopulation.
o Subpopulation = a part of a larger
population that sustains a limited
exchange of individuals through
immigration and emigration.
o Metapopulation = group of
subpopulations living on habitat
patches connected by exchange of
individuals among patches
AGE STRUCTURE AND SEX RATIO
 Observations:
o Kenyan population is fairly young
and is growing
The effect of sex ratio
NE = (4NmNf)/(Nm + Nf)
LIFE TABLES
 Used to calculate and summarize important
parameters relating to the growth of
populations, such as survivorship, mortality,
and life expectancy
 Proportion surviving = survivorship
 Age-specific/Cohort Life Tables
o Cohort is followed over time
o Requires frequent censuses
 Time-specific Life Tables (Static Life
Tables)
o Useful for long-lived organisms
o Assumes stationarity and number
born each year is the same
Birth rate
 Number of young born per female in a
period of time
Fecundity schedule
 Tabulation of birth rates for females of
different ages in a population
 mx = the number of offspring of a given age
class
 lxmx = allows for a number of applications
for management of populations
o ex.
 Age class with the most
contribution to reproduction
is 6-7, therefore, to make
sure the population does not
die out, we have to take care
of this population.
 If this life table is a life table
for fishes, we can infer that
those with lxmx that, when
subtracted to R0, still
generates a difference
greater than 1, then catching
fishes of these age classes
should not affect the growth
of the fish population. In
contrast, if we catch fishes of
age class 6-7, then that
would cause the population
to shrink because it
contributes the most to the
population’s reproduction.
 R0 = over the average life span of a typical
organism, it will produce this number of
offspring on average
o The population of lions in this life
table increases its population by 33%
for every generation (roughly every 3
years).
o If R0 is 1, then the population is not
growing. If <1, the population is
shrinking.
 Life tables are for females, mostly.
 POPULATION GROWTH
 The life table will ultimately allow us to
compute net reproductive rate, which can
be converted to r, the intrinsic growth
rate, which can then be used in modeling
the pattern of population growth.
 Population growth accelerates.
Two patterns of growth:
 exponential growth (J-shaped curve)
o growth that does not occur in
discrete generations, but is instead
continuous
o rate of population growth increases
over time
 logistic growth (S-shaped curve).
o Growth pattern that happens when
resources are depleted and
population growth rate declines then
eventually stops
 Carrying capacity
 The population size at
which growth stops
 Represents the
number of individuals
that an environment
can support
o Population growth is highest when
N = K/2 and ceases when N = K. N is
population size.
o As N gets closer to K, environmental
factors increasingly impede further
population growth.
 Despite the name, life history strategies
followed by species or populations are
outcomes of evolution, not conscious
choice. The classical scheme assigns
organisms to either r-selected or K-
selected strategies. Newer schemes divide
organisms to ruderals, stress-tolerators, and
competitive strategies. Few species can be
unambiguously assigned to a strategy in
both schemes. Most species exhibit
combinations of traits of alternative
schemes. Why do you think that is so?
SURVIVORSHIP CURVES
 Describe the intrinsic properties of a species
with regards to life history strategy
Types:
 Type I
o Most individuals lost when older
(Higher mortality in elderly)
o Found in vertebrates especially those
with parental care
 Type II
o Constant mortality rate across age
groups
o In Hydra and many birds
 Type III
o Most individuals lost when young
(Higher mortality in young
individuals)
o Found in many invertebrates and
plants
 Age (x-axis)
o Maximum longevity
 Logarithmic scale of nx (y-axis)

Two Types:
LIFE HISTORY STRATEGIES
 Life history
o Any aspect of the developmental
pattern and mode of reproduction of
an organism that results from
evolution (and not “conscious
decision”)
o Adaptations of an organism that
influence aspects of its biology, such
as reproduction, survival, size, and
age at reproductive maturity.
o Aspects include:
 Body size
 Developmental stages
 Instars
 Metamorphosis
 Senescence – the process of
timing, aging, degeneration,
and death
 Reproductive patterns
 Ability to disperse
 Organisms that produce larger offsprings
are constrained to produce fewer.
o Why? Organisms take in energy at a
limited rate due to external
environment conditions and internal
constraints. These constraints are
fundamental to the principle of
allocation, which states that if an
organism uses energy for one
function such as growth, it reduces
the amount of energy available for
other functions such as
reproduction.
 When adult survival is lower, organisms
begin reproducing at an earlier age and
invests a greater proportion of their
energy budget to reproduction.
 When adult survival is higher, organisms
defer reproduction to a later age and
allocate a smaller proportion of their
resources to reproduction.
o High adult survival = delayed
maturity
 r-selected
o observed in opportunistic small
organisms with good dispersal
capabilities
 strongest in species that
often colonizes new or
disturbed habitats
o Characteristic of unpredictable or
variable environments
o have short life spans, rapid
development
o early reproductive age
o Semelparous
 Reproduce only once and
typically die after that
o fast population growth; variable
population size below K
 K is carrying capacity
o Type III survivorship, no parental
care
 K-selected
o Large organisms with poor
dispersal capability but good
competitive ability
 Favors more efficient
utilization of resources such
as food nutrients
 Most prominent in
populations near carrying
capacity most of the time
o Characteristic of predictable
environments
o Long-lived, slow development
o Late reproductive age
o Iteroparous
 Can reproduce many times
o Slow population growth; population
size around K
  Likely because of late rate of growth becomes slower until
reproductive age it stabilizes at the carrying capacity.
o Type I or II survivorship
TRADE-OFFS IN REPRODUCTIVE STRATEGIES
 Number of offspring per reproductive event:
o Large litters or clutches do not
increase fitness since these are offset
by higher mortality from predators,
lower body weight of offspring,
higher feeding costs, etc.
 Present vs. Future Reproduction
o Selection favors age of reproduction
that occurs before survivorship
declines markedly
o Semelparity vs. Iteroparity
 Age at Sexual Maturity

 Most species are a mix of these two

strategies.

Growth patterns are related to life history

strategies.

 Describes 3 types of life history strategies

 r-selected  J-shaped
o Exceeding the carrying capacity K
means the number of individuals in
the population often exceed the
ability of the environment to support
these numbers  the population will
increase, then crash, then increase
again, then crash.
 Evident in pests
 K-selected  S-shaped
o Population grow fastest at about
halfway the carrying capacity
o As the slope population size
approach the carrying capacity, the
o Ruderal – small, shortlived; allocate
more energy to reproduction,
dispersal
 Persist in locations
experiencing high
disturbance, meaning that
organisms with this life
strategy have several
characteristics that endure
any mechanism that limits
them by destroying their
biomass (predation ???)
 High disturbance, low stress
life history strategy
o Stress-tolerant = allocate more
energy to maintenance
 Large body sizes, longer lives
 Rarely reproduces
 Low disturbance, high stress
 Stress – induced by
environmental
 conditions that limit
an organism’s growth
 Grow slowly
o Competitors = allocate more energy
to growth, resource acquisition, and
competition
 Faster growth but not as fast
as the ruderal
 Low disturbance, low stress
 Organisms have the
capacity to grow well,
but this results to
competition among
them.
 Irruptions
o When populations that typically have
low densities and stable numbers
suddenly explode in numbers
o Example:
 Outbreaks of coral-eating
starfish
DENSITY DEPENDENT VERSUS DENSITY
INDEPENDENT FACTORS
 Density dependent factors

POPULATION REGULATION o In populations that are regulated

 Stems from observations that show that more when their densities are higher
different species have different patterns with  Density increases = birth rate
regards to the sizes of their populations decreases = death rate
increases
Patterns of population fluctuation o Found in K-selected species
 Small-magnitude irregular fluctuations o Usually lead to increased mortality
o Year to year random changes of one percentage as population density
order of magnitude (base 10) or less increases
 Large-scale irregular fluctuations
o Peaks may be several orders of  Density-independent factors
magnitude higher than the lows
o Typical of r-selected species like
most insect pests
 Cycles
o Regular changes with fixed periods,
magnitude can be large or small
o Best known examples:
 4 year cycles of microtine
rodents (lemmings and voles)
 10 year cycles of hare,
grouse, and lynx
o Typically k-selected species
 o Density’s impact on birth and death  And this depends on the
rates are somewhat fixed, fairly level of productivity, that is,
stable how much food is available,
how much photosynthesis is
occurring in the ecosystem
that they are exploiting

QUESTIONS
 Density-dependent or density-
independent?
o Density of humans and their
mortality due to typhoons?
 Density-independent
o Density of ant colonies and their
mortality due to (paghuhukay XD)
 Density-independent
Barnacles o Density of invertebrates and their
 Logs are density independent mortality due to predation
o Why?  Density-independent
o Extinction
 Abiotic factors are density-independent??  ???
 Biotic factors are density-dependent?? o Mortality of plants due to herbivores
o ^ Not a good generalization.  Density-independent
o Mortality ng ipis dahil sa baygon
spray
 Inverse density dependent?
 If we assume na the
strength of baygon
spray is at maximum
strength if mas
mataas yung
concentration sa
isang ipis
 Density dependent
 If we assume na one
 Ecosystem Exploitation Hypothesis baygon spray can kill
o “The factor that regulates the one ipis and 4 ipis at
population of one species is usually equal efficiency
either its own kind or a predator or
herbivore of that species.”
o Mortality of humans due to COVID-
19
 Density-dependent
o Mortality of bad bacteria sa body
 Inverse density dependent
 SYSTEMS ECOLOGY  Steady state of ecosystems
 System o When total input = total output
o Any part of the universe that can be o Dynamic equilibrium
isolated for the purposes of o Homeostasis = when environment is
observation and study kept constant

Ecosystem vvv closed system

 any unit that includes all organisms that
function together (the biotic community) in
a given area interacting with the physical
environment so that the flow of energy
leads to clearly defined biotic structures and
cycling of materials between living and non-
living parts (Odum 1983).
 Basic functional unit of ecology
 Components and processes of an ecosystem:
o Biotic community
 Group of populations that
interact with one another
o Flow of energy
o Cycling of materials
o Feedback control loops

 Autotrophs – produce their own food

 Heterotrophs – eat autotrophs for energy
needs

 Ecosystems are living systems difficult to

isolate without affecting its properties
o BOUNDARIES are best defined in
terms of flows
 There should be less of these
crossing boundaries than
staying with them.
o Open vs. Closed system
 IMBALANCES IN NATURAL CYCLES Important Notes to Supplement the Slides.
SUMMARY OF LESSONS:

Regarding flow of energy:

 Flow of energy tends to be one-way and is
eventually lost (most times as heat). We see
this in the trophic level concept.
 As energy moves from one trophic level to
the next, most of the useful energy is given
off as heat. Only 10% is retained in biomass.
 In land, there are 3-5 trophic levels.
 In seas, there are 6-9 trophic levels.

Regarding open and closed systems:

 Open system
o Energy flows outside of the defined
boundaries.
o Because energy cross boundaries,
some energy is lost from the system.
 Closed system
o No material/energy crossing the
boundary of the system.
o The system is in a steady state.

Regarding Nutrient Cycling (Yung pail-water-

The associated urbanization is cutting (?) and
pump illustration):
releasing heat, manifested as convection current,
 Changes associated with bifurcation will
which is the turbulence you feel when a plane
lead to changes that will affect other
crosses over a city.
compartments.
o Whether the input is decreased or
-------
increased, there will be a cascade of
changes in all compartments.
 If no bifurcation:
o If flow rate in is increased
 Water level rises in one
compartment, decreases in
the other, but there are no
visible changes on the water
level of other compartments.
o If flow rate in is decreased
 Water level drops in one
compartment, which will
lower its flow rate to the next
compartment, and so on,
until one compartment has
most of the water and all
other compartments nearly
empty.
  However, the pail will never
run dry, as this is a closed
system.
 Higher content = Export will remain the
same
o No change is observed in other
compartments.
 Lower content = Export will decrease after
the content nearly empties
o Other compartments will change
 Is the water stagnant?
o NO. The water level is stable, but
they don’t have permanent
residence in one compartment, as
they are still flowing from one
compartment to another.
Regarding Biogeochemical cycles:
 Turnover rate
o # of times the contents of a
compartment is replaced
 Residence time
o The amount of time before the
contents are completely replaced
 Carbon cycle
o Amount of carbon in Ocean > C in
soil > C in atmosphere
o Has the fastest turnover rate and
shortest residence time
o The pipes and pumps assumption
does NOT apply here because the
amount of C in the atmosphere
affects how much C flows to the
ocean (a simple dissolution
process), and such, the rate of
import to one container affects the
rate of its export, which should not
be the case in the illustrated
example of a system using the pails
and water and pumps.
 Phosphorus cycle
o Sedimentary cycle (so predominantly
stored in sediments under the
ocean) and tied to the rock cycle,
hence, it has the longest residence
time of a thousand years and the
slowest turnover rate
o Phosphorus is usually the limiting
nutrient in life processes (remember
the Redfield ratio of 106 C: 16 N: 1
P), so the increase in P distribution
usually leads to cultural
eutrophication
 Nitrogen cycle
o Predominantly a gaseous cycle, so it
is predominantly stored in the
atmosphere
o The pipes and pumps assumption
applies in this cycle, as the amount
of N in the atmosphere does not
affect the rate of nitrification done
by bacteria and lightning.
Regarding biodiversity ecosystem function:
 Interaction strengths vary in every
ecosystem, and the bigger the system is, the
less important the role every little
interaction plays.
o More species with similar
abundances  more stable network
of interactions.
 Ex. a predator will not go
extinct if a prey of theirs go
extinct, as it can prey on
other organisms. Compare it
to when a predator only has
one species as a prey, then its
survival is solely dependent
on the existence of that prey.
o A species that has a big role in the
ecosystem (e.g. whales in aquatic
ecosystems) can cause a collapse if
its role is disturbed.
 SAMPLE QUESTIONS FROM QUIZZES, REVIEW
SESSIONS BY SIR AL, MODIFICATIONS + POINTS
FOR DISCUSSION
LE1
1. Which of the following types of
experimental designs has a regular
interspersion of treatments but is prone to
the risk of periodically varying properties of
the experimental area within the space
intervals?
o Systematic
2. Lydia works in a flower shop after school.
She notices that the hydrangeas in
Container 1 have pink blossoms, while the
hydrangeas in Container 2 have blue
blossoms. She makes sure they all receive
the same amount of light and water. Her
boss tells her that the flowers were all
grown from the same seed variety. Lydia
discovers, after testing the soil, that
although the same type of soil was used, the
soil in Container 1 has a pH of 6.0, and the
soil in Container 2 has a pH of 5.0. Lydia
conducted her investigation to answer
which question about hydrangeas?
o How does the chemistry of soil
affect the color of hydrangea
blossoms?
3. Professor Z sets up a lab for her biology
students using a culture of the small
crustacean Daphne, obtained from a pond
that was 20 degrees C. The students are to
investigate the effect of temperature on
Daphnia. The students will observe the
crustacean’s heartbeat under the
microscope, at the ff. temps: 0, 10, 20, 30,
and 40C. They will count the number of
heartbeats per sec. To avoid
pseudoreplication, which would be the best
experimental design?
o Placing 5 of each identical cultures
of Daphnia at different
temperature
o Placing 5 cultures, each with a
different type of crustacean, at a
different temp
4. T or F: All experimental designs should have
a control group
o False
5. Identify the no. of true replicates and no. of
subsamples in the mensurative study aimed
at comparing the rate of leaf decomposition
at high (the “treatment” group) and low (the
“control” group) altitude.
Design: 10 randomly selected leaf samples
at the peak of the hill, another 10 atleast
100 m below the peak of the hill
o 10 replicates, no subsamples per
treatment
6. During randomization for sampling or
experimental lay-outs, ensuring
interspersion means:
o All of the choices given are
appropriate
o Resorting to haphazard
arrangements instead of
randomization
o Accepting any randomization
outcome
o Repeating the randomization to
get a satisfactory interspersion
o Making your own lay-out if the
randomization results are not
satisfactory
7. The following was done in an experiment to
determine if noise (from window-type air
conditioners) in classrooms affects the
performance of students in standardized
intelligence tests: 120 students were
randomly divided into eight groups of 15
students each, randomly assigned a
classroom per group and given 90 minutes
to complete the same test. Four of the
rooms had noisy AC units and four didn't.
The rooms are randomly interspersed. How
many independent samples per treatment
did this experiment have?
o 4
8. Where is RCB more advantageous?
 o When the environment can
introduce bias.
There were five separate experiments conducted to
determine if noise (from window-type air
conditioners) in classrooms affects the performance
of students in standardized intelligence tests taken
in those classrooms:
Experiment A: 120 students were randomly divided
into four groups of 30 students each and randomly
assigned to a classroom per group and given 90
minutes to complete the same test. Two of the
rooms had noisy AC units and two didn’t. The
rooms are randomly interspersed.
Experiment B: 120 students were randomly divided
into 8 groups of 15 students each and randomly
assigned to a classroom per group and given 90
minutes to complete the same test. Four of the
rooms had noisy AC units and four didn’t. The
rooms are randomly interspersed.
Experiment C: 120 students were randomly divided
into four groups of 30 students each and randomly
assigned to a classroom per group and given 90
minutes to complete the same test. All four rooms
had noisy AC units. The following day, the
procedure was repeated except all the four rooms
used had quiet AC units, and the exam used was
different (from that of the first day), but is otherwise
equivalent.
Experiment D: 120 students were randomly divided
into four groups of 30 students each and randomly
assigned to a classroom per group and given 90
minutes to complete the same test. Two of the
rooms had noisy AC units and two didn’t. The
rooms are randomly interspersed. The following
day, the procedure was repeated except those who
were in quiet rooms in the previous run were
assigned to noisy rooms and vice versa, and the
exam used was different (from that of the first day),
but is otherwise equivalent.
Experiment E: 120 students were randomly divided
into 8 groups of 15 students each and randomly
assigned to a classroom per group and given 90
minutes to complete the same test. All 8 rooms had
noisy AC units. The following day, the procedure
was repeated except all the 8 rooms used had quiet
AC units, and the exam used was different (from
that of the first day), but is otherwise equivalent.
9. Which experiment will be most affected by
genetic differences among students?
o Experiments A and B, because the
variability will not cancel out.
However, if a choice between the
two is needed, the more appropriate
would be Experiment B because it
had less number of subsamples.
10. Least affected by genetic differences?
o Experiment D, RCB design.
11. Which experiment got completed earlier?
o Experiments A and B
12. Most biased by repetitive testing?
o Experiments C and E, they took the
test twice + segregated design
13. Most affected by potential long term effects
of air conditioning
o Experiments C and E
IMPORTANT NOTES:
 Something is manipulative when you’re
making decisions based on levels.
o Mensurative = when you’re
determining levels
 The “randomization” in RCB should be
within the blocks.
 Standard error as a basis for determining
significant differences in graph (error bars)
would have narrower differences compared
to when standard deviation is used.
Therefore, if the error bars are close to each
other and SD was used, it’s likely not
significantly different with each other.
LE2
BIOLOGICAL EVOLUTION of the NATURAL
SELECTION:
A. Populations grow faster than their resources
can support. Random mutation creates
variation in genotype (and consequent
 phenotype). These variations are passed on
to succeeding generations.
B. Individuals in the population compete
depending on crowding, resource supply.
Both lead to different biological conditions
over time and space. Physical and chemical
conditions in the environment differ over
time and space.
C. Some phenotypes (and corresponding
genotypes) confer an advantage during
competition; others confer a disadvantage
to those competing individuals that have
this trait.
D. Those genotypes that confer an advantage
get passed on to more individuals in the
succeeding generations as their bearers are
more successful in having more offspring.
E. The genetic composition and consequent What type of selection occurred:
general characters of the populations
change over time. 4. Jervis  Daphne
o Directional
1. A hypothetical population was found to
have a genotype frequency of AA=25%, 5. Chatham  Jervis (G. fulliginosa)
Aa=50%, and aa=25%. What is the o Directional
frequency of the A allele in proportions?
o 0.5 6. G. magnirostris in Jervis
o Directional
2. A hypothetical population was found to
have a genotype frequency of AA=20%, 7. Which of the following statements is wrong?
Aa=40%, and aa=40%. Which observed o The basic unit of selection is the
genotype is/are the farthest from population
equilibrium? o Adaptation is the result of genetic
o Aa change
o Biological factors can be selective
3. A hypothetical population was found to agents
have a genotype frequency of AA=10%, o Niches are exclusively defined by
Aa=80%, and aa=10%. What type of physical and chemical factors
selection is this trait likely experiencing? o Migration can occur without gene
o Stabilizing selection flow

LE3
1. T or F: Competition between different
species can lead to the expansion of the
distribution limits of one or both species.
o True

2. Why are individuals in some population

more clumped in dispersion than others?
 o Due to availability of limited
resources
3. Which of the following factors does not
likely influence the dispersion of plants in a
population?
o Genetic diversity
Density-dependent or density-independent?
4. Mortality of plants due to limited nutrients
in the soil?
o Density-independent
5. Mortality of farmers due to drought?
o Density-independent
6. Bamboo and sunlight
o Density-independent (?)
7. Complete the life table for the following: dx
of 500, 375, 109, 15, 1; a 5 year life span,
fecundity schedule (mx) of 0.00, 0.9, 1.9, 4.0,
0.0 respectively for each of the five years.
What is the next reproductive rate (R0) of
this population?
o 0.75
8. Generation time?
o Dfsdf
13. What is the effective population size if 40%
of the panmictic population of 100 humans
are polygamous males and the rest are
polygamous females?
o 96
14. What is the effective population size if 40%
of the panmictic population of 100 humans
are monogamous males and the rest are
monogamous females?
o 80
15. What is the effective population size if 40%
of the panmictic population of 100 humans
are polygamous males and the rest are
monogamous females?
o 96
16. What is the effective population size if 40%
of the panmictic population of 100 humans
are monogamous males and the rest are
polygamous females?
o 80
17. Cohort life tables require that the
stationarity assumption be made:
o False

9. Is the population from above growing?

o No

10. What type of survivorship curve does this

hypothetical organism have?
o Type I

11. Assuming this species is a commercially

important finfish from the sea, which is the
youngest age class that could be harvested
without endangering the survival of this
population?
o This population cannot support
harvesting.

12. If this is a pest, which age class will you

target to impact its growth rate?
o 1-2 year olds