Chapter 9 Notes

The combination of suitable conditions is a species’ niche, and we might expect to find a species
occupying areas of the landscape in which environmental conditions match the species’ niche
requirements.

9.1 Because all organism has access to limited energy and resources, there are fundamental trade-
offs in how these can be allocated between survival, offspring number, and offspring size.

 Trade-offs in energy allocation influence an organism’s ability to cope with abiotic

stressors, such as low water availability or extreme temperatures. However, the principle
of allocation is just as important in influencing how an organism allocates energy within a
given segment of its energy budges, such as reproduction.

 Size and Number of Offspring

o (Fish Egg Size and Number) More robust patterns of variation can be obtained by
analyzing relationships within closely related species such as within families and
genera.
o Turner and Texler (1998). Were particularly interested in determining the
relationship between egg size and egg number also known as fecundity.
o Fecundity is simply the number of offspring produced by an organism.
o Darters are the most species-rich group of vertebrates in North America. Despite
the fact that darters as whole live in similar habitats and have similar anatomy,
they vary widely in their life histories.
 Of the darters they collected at these locations, they chose 15 species for
detailed study. They found that larger darter species produce larger
numbers of eggs and darters that produce larger eggs produce fewer eggs.
This suggests that there is a trade-off between offspring size and number.
o Turner and Texler also proposed that gene flow would be higher among
populations producing more numerous smaller eggs, that is, among populations
with higher fecundity.
o Strong positive relationship of gene flow with the number of eggs produced by
females.
 Populations of darter species that produce many small eggs showed less
variation in genetic diversity across the study region than did populations
that produce fewer larger eggs.
  Basic of life history trade-offs: offspring size and number has the potential
to influence macro evolutionary processes, such as variation in speciation
rates. (pg 234)
 Seed and Size Number in Plants
o Species producing larger numbers of seeds on average produce fewer seeds.
o They classified seeds with no specialized structures for dispersal as unassisted
dispensers.
 If seeds had hooks, spines, or barbs, they were classified as adhesion-
adapted.
 Meanwhile seeds with wings, hairs, or other structures that provide are
resistance were assigned to a ‘wind dispersed category’
 Elaisome; a structure on the surface of some seeds generally containing
oils attractive to ants, as ant-dispersed.
 Seeds with flesh or with an aril, a fleshy covering of some seeds that
attracts birds and other vertebrates, were classified as vertebrate
dispersed.
 Finally, they classified as scatterhoarded those seeds known to the
gathered by mammals and stored in scattered caches of hoards.
o Plants disperse their seeds in different ways tend to produce seeds of different
sizes.
 Adult Survival and Reproductive Allocation
o Reproductive effort is the allocation of energy, time, and other resources to the
production and care of offspring. Within an individual, reproductive effort
involves trade-offs with other needs of the organism, including allocation to
growth and maintenance. However, delaying reproduction also involves risk. An
individual that delays reproduction runs the risk of dying before it can reproduce.
Consequently, evolutionary ecologists have predicted that variation in mortality
rates among adults will be in association with variation in the age of first
reproduction, or age of reproductive maturity.
 Variation Among Species
o The relationship between mortality, growth and age at first reproduction or
reproductive maturity has been examined in a large number of organisms.
o Many animals continue growing after they reach sexual maturity. In addition,
most vertebrate species begin reproducing before they reach their maximum
body size.
o In addition, most vertebrate species begin reproducing before they reach their
maximum body size. As a result, the energy budgets of these other vertebrate
species, such as fish and reptiles, are different before and after sexual maturity.
 o Before these organisms reach sexual maturity, energy required by an individual is
allocated to one of two competing demands: maintenance and growth.
o After reaching sexual maturity, limited energy supplies are allocated to three
functions: maintenance, growth, reproduction.
o Because of the increase in reproductive rate associated with larger body size,
deferring reproduction would lead to a higher reproductive rate.
o Where mortality rates are high, deferring reproduction increases the probability
that an individual will die before reproducing.
o Gunderson estimated reproductive effort as each population’s gonad somatic
index or GSI. GSI was taken as the ovary weight of each species divided by the
species body weight and adjusted for the number of batches of offspring
produced by each species per year.
 Variation Within Species
o In general, fry that grow rapidly are larger as juveniles and are more likely to
become jacks, while the slower growing individuals tend to become hooknose.
 There are two alternative mating behaviors available to both jacks and
hooknose: “sneaking” and “fighting”. Fighters will typically fight for
proximity to a desired female who is about to spawn, while sneakers will
try to fertilize the female’s eggs while other males are engaged in fighting.
o By observing males of different sizes, Gross (1985) was able to determine that
both very small males and very large males had roughly equal success in getting
close to females, while intermediate sized males were rarely able to get near a
female.
 Large fish were better at fighting
 Small fish were better at sneaking
 Middle sized fish were too large to sneak but too small to fight and thus
they rarely were able to get near females.
o Removing the largest fish from the population should decrease the size at which
fighting is a successful strategy.
o Alternative life histories can coexist within a population, and that behavioral
strategies are intimately tied to the fitness of individuals. If the costs or benefits
of alternative aspects of a life history change this could result in natural selection
toward alternative life-history traits.
 Genetic Control of Life History
o Variation in life-history traits, such as relationship between adult and juvenile
survival, can be due to differences in both the ecological and the environmental
conditions faced by individuals and populations.
 This environmentally induce variation can only influence evolution if there
is also a genetic and heritable component to life-history variation among
individuals within and between populations.
o Pathogenesis – in this form of reproduction, individuals create diploid eggs
through mitosis, rather than producing haploid eggs meiosis, as is the norm for
vertebrates.
 The diploid eggs develop into offspring, resulting in genetic clones of the
mother water flea. As a result, a single female is able to produce
genetically identical males and females! These females will undergo
meiosis, producing haploid diapausing (resting) eggs that require
fertilization by the males to develop. These clones produce diapausing
eggs through mitosis, not meiosis, and thus have no haploid aspect to
their life cycle.

9.2 Life – History Classification

 The great diversity of life histories observed in nature can be classified on the basis of a
few common characteristics.
o These classification schemes should be used to organize the variety that exists in
nature and not to assign a classification to every species.
 R and K Selection
o The terms r and K selection refer to parameters of population growth models. R
represents a measure of population growth rate, with larger values corresponding
to populations that are growing rapidly. K represents the maximum sustainable
size of a population, such that habitats with larger K values could contain larger
populations than areas with smaller K values.
o These suggest that for some species, selection will favor those life-history traits
that cause increases in growth rates, so called “r-selected” species.
o They proposed that in other species, selection will act upon life-history traits
important for when habitats are “full” of a population. These “k-selected species
are likely to have life-history traits that favor efficient utilization of resources such
as food and nutrients, rather than maximizing growth rates.
o K selection would be most prominent in those situations where species
populations are near the “carrying capacity” of the habitat much of the time.
o r selection and K selection are the endpoints on a continuous distribution and
that most organisms are subject to forms of selection somewhere in between
these extremes.
 o r and K selection with attributes of the environment and of populations. He also
listed the population characteristics that each form of selection favors.
o While r selection should be characteristics of variable unpredictable
environments, fairly constant or predictable environments should create
conditions for K selection.
o Development should be rapid under r selection and relatively slow under K
selection.
 Meanwhile early preproduction and smaller body size will be favored by r
selection, while K selection favors later reproduction and larger body size.
o Pianka predicted that reproduction under r selection will tend toward a single
reproductive event in which many small offspring are produced. This type of
reproduction is called semelparity, which occurs in organisms such as annual
weeds and salmon.
o In contrast K selection should favor repeated reproduction or iteroparity, of fewer
larger offspring. Iteroparity which spaces our reproduction over several
reproductive periods during an organism’s lifetime, is the type of reproduction
seen in most perennial plants and most vertebrate animals.
 Plant Life Histories
o Phil Grime proposed that variation in environmental conditions has led to the
development of distinctive strategies or life histories among plants. The two
variables that he selected as most important in exerting selective pressure on
plants were the intensity of disturbance and abiotic stress. Grime also argues that
competition can exert strong pressures on plants, and that competition should be
relatively more important when stress and disturbance are low.
 Grime contrasted four extreme environmental types, which he
characterized by combinations of disturbance intensity and stress
intensity. (Plants only occupy three of these)
 Low disturbance – low stress
 Low disturbance – high stress
 High disturbance – low stress
 High disturbance – high stress
o Grime described plant strategies or life histories, that match the requirements of
the remaining three environments. His strategies were ruderal, stress-tolerant,
and competitive.
 Ruderals are plants that live in highly disturbed habitats and that they may
depend on disturbance to persist in the of potential competition from other
plants.
  Ruderals have a few characteristics which allow them to sustain
themselves under intense disturbance, which he defined as any
mechanisms or processes that limit plants by destroying plant
biomass.
o On the characteristics of ruderals is their capacity to grow
rapidly and produce seeds during relatively short periods
between successive disturbances.
o Ruderals also invest a large proportion of their biomass in
reproduction, producing large numbers of seeds that are
capable dispersing to new habitats made available by
disturbance.
 Stress tolerant – with a definition of stress as “… external constraints which
limit the rate of dry matter production of all or part of the vegetation.” |
Stress is induced by environmental conditions that limit the growth of all
or part of the vegetation. Stress is the result of extreme temperatures,
high or low, extreme hydrologic conditions, too little or too much water,
or too much or too little light or nutrients.
 Some animals are more tolerant to the environmental extremes
that occur than other species. These are species that are stress
tolerant species. Stress tolerant plants are plants that live under
conditions of high stress but low disturbance.
 Generally, stress tolerant plants grow slowly, evergreen, conserve
fixed carbon, nutrients, and water; and are adept at exploiting
temporary favorable conditions. In addition, stress – tolerant
plants are often unpalatable to most herbivores.
 Competitive – Competitive are plants that occupy environments where
disturbance intensity is low, and the intensity of stress is low. Under
conditions of low stress and low disturbance, plants have the potential to
grow well. As they grow they eventually compete with each other for
resources such as light, water, nutrients, and space. Plants living under
such circumstances will be selected for strong competitive abilities.
 Opportunistic, Equilibrium, and Periodic Life Histories
o They called the three endpoints on their surface opportunistic strategy by
combining low juvenile survival, low numbers of off spring, and early reproductive
maturity, maximizes colonizing ability across environments that vary
unpredictably in time or space. Remember that while the absolute reproductive
output of opportunistic species may be low, the percentage of their energy
budget allocated to reproduction is high.
 o The periodic strategy combines low juvenile survival, high numbers of offspring,
and late maturity. Among fish, periodic species tend to be large and produce
numerous small offspring.
 By producing large numbers of offspring over a long-life span, periodic
species can take advantage of infrequent periods when conditions are
favorable for reproduction.
o However opportunistic species differ from the typical r-selected species differ
from the typical r-selected species because they tend to produce small clutches of
offspring. The equilibrium, which combines production of high juvenile survival,
low numbers of offspring, and late reproductive maturity, approaches the
characteristics of typical K-selected species.
 Lifetime Reproductive Effort and Relative Offspring Size: Two Central Variables
o Charnov’s aim for creating a new approach to life history classification is to
develop classification free of the influences of size and time that would facilitate
he exploration of life-history variation within and among groups of closely related
taxa.
o By removing size and time effects, we may be able to more clearly detect life-
history differences among evolutionary lineages.

9.3 Fundamental and Realized Niches

The fundamental niche reflects the environmental requirements of species, while the realized nice
also includes interactions with other species.

 The word “niche” has been used to summarize the factors that influence growth, survival
and reproduction of a species.
o Grinnell’s idea of the niche focused on the influences of the physical
environment, while Elton’s earliest concept included both biological interactions
and abiotic factors.
o Hutchinson defined that niches were n-dimensional hyper volume where n equals
the number of environmental factors important to the survival and reproduction
of a species.
 This hyper volume is the fundamental niche, which specifies the values of
the n environmental factors permitting a species to survive and
reproduced. The fundamental niche defines the physical condition under
which a species may live, in the absence of interactions with other species.
  Interactions among species, such as competition, may restrict the
environments in which a species actually can persist, and he referred to
these more restricted conditions as realized niche.
 The Feeding Niches of Galapagos Finches
o Differences in beak size among small, medium and large ground finches translate
directly into differences in diet.
o The size of seeds that can be eat by Galapagos finches can be estimated by simply
measuring the depths of their beaks. Within its population, individuals with the
smallest beaks fed on the softest seeds and deepest beaks fed on the hardest
seeds.
o The important of beak size to seed use was also demonstrated by the effects of a
1977 drought on the G. fortis population of Daphne Major.
 During a drought mortality did not happen quickly on all segments of
population. As seeds were depleted, the birds ate the smallest and softest
seeds first, leaving the largest and toughest seeds. Following the drought
not only were seeds in short supply, the remaining seeds were also
tougher to crack. Because they could not crack the remaining seeds,
mortality fell most heavily on smaller birds with smaller beaks.
 The Habitat Niche of a Salt Marsh Grass
o Allopolyploidy is a process of speciation initiated by hybridization of two different
species.
o This environmental tolerance is reflected in the distribution of the plant in
northwestern Europe, where it generally inhabits the most seaward zone of any
of the salt marsh plants.
o The local distribution of S. anglica in the British Isles is well predicted by a few
physical variables related to the duration and frequency of inundation by tides
and waves.
o The lower and upper intertidal limits of the grass are mainly determined by the
magnitude of tidal fluctuations during spring tides. Where tidal fluctuations are
greater, both the lower and upper limits are higher up on the shore.