Mechanisms of Evolution

Mechanisms of Evolution

Christopher J. Paradise, PhD

A. Malcolm Campbell, PhD
Mechanisms of Evolution
Copyright Christopher J. Paradise and A. Malcolm Campbell. 2016.

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 Abstract
Three of the four major mechanisms of evolution, natural selection, ge-
netic drift, and gene flow are examined. There are 5 tenets of natural selec-
tion that influence individual organisms: Individuals within populations
are variable, that variation is heritable, organisms differ in their ability
to survive and reproduce, more individuals are produced in a generation
than can survive, and survival & reproduction of those variable individ-
uals are non-random. Organisms respond evolutionarily to changes in
their environment and other selection pressures, including global climate
change. The importance of spatial structure of a population in relation to
how it affects the strength of gene flow and/or genetic drift, as well as the
genetic variation and evolution of populations, is shown. Gene flow tends
to reduce variation between populations and increase it within popula-
tions, whereas genetic drift tends to reduce genetic variation, especially
in small, isolated populations. The mechanisms of evolution can lead to
speciation, which requires both time and genetic isolation of populations,
in addition to natural selection or genetic drift.

Keywords
evolution, population, gene flow, genetic drift, adaptation, natural
selection, phenotypes, behavior, predation, selective agent, heterozygous,
homozygous, heritability, dispersal, genotype, gene flow, genetic isola-
tion, genetic distance, non-adaptive evolution, population bottleneck,
inbreeding
 Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 Selection Acts on Individuals with Variable
Characteristics....................................................................1
Selection can Act on Behaviors...........................................1
Natural Selection on a Discrete Trait..................................6
Chapter 2 Species May Evolve in Response to Climate Change........11
Range Expansion.............................................................12
Evolutionary Response to Changing Rainfall...................14
Ethical, Legal, Social Implications: Data are
Needed to Formulate Policy, but Science is
Often Misused in the Process........................................17
Chapter 3 Two Seemingly Isolated Populations may not
Actually be Isolated..........................................................23
Some Populations Have Limited Dispersal.......................23
Dispersal Links Geographic and Genetic Distance...........27
Chapter 4 Populations can Evolve in the Absence of
Natural Selection..............................................................33
Population Isolation Affects Genetic Diversity.................33
A Population Bottleneck Reduces Genetic Diversity.........37
Conclusion............................................................................................41
Glossary................................................................................................43
Index....................................................................................................45
 Preface
This book about mechanisms of evolution is part of a thirty book se-
ries that collectively surveys all of the major themes in biology. Rather
than just present information as a collection of facts, the reader is treated
more like a scientist, which means the data behind the major themes are
presented. Reading any of the thirty books by Paradise and Campbell
provides readers with biological context and comprehensive perspective
so that readers can learn important information from a single book with
the potential to see how the major themes span all size scales: molecular,
cellular, organismal, population and ecologic systems. The major themes
of biology encapsulate the entire discipline: information, evolution, cells,
homeostasis and emergent properties.
In the twentieth century, biology was taught with a heavy emphasis
on long lists of terms and many specific details. All of these details were
presented in a way that obscured a more comprehensive understanding.
In this book, readers will learn about mechanisms of evolution and some
of the supporting evidence behind our understanding. The historic and
more recent experiments and data will be explored. Instead of believing
or simply accepting information, readers of this book will learn about the
science behind the mechanisms of evolution the way professional scien-
tists dowith experimentation and data analysis. In short, data are put
back into the teaching of biological sciences.
Readers of this book who wish to see the textbook version of this
content can go to www.bio.davidson.edu/icb where they will find
pedagogically-designed and interactive Integrating Concepts in Biology
for introductory biology college courses or a high school AP Biology
course.
 Acknowledgments
Publishing this book would not have been possible without the generous
gift of Dr. David Botstein who shared some of his Breakthrough Prize
with co-author AMC. Davids gift allowed us to hire talented artists (Tom
Webster and his staff at Lineworks, Inc.) and copyeditor Laura Loveall.
Thanks go to Kristen Mandava of Mandava Editorial Services for project
management and guidance. In particular, we are indebted to Katie Noble
and Melissa Hayban for their many hours and attention to detail.
Kristen Eshleman, Paul Brantley, Bill Hatfield and Olivia Booker
helped us with technology at Davidson College. We are grateful to ad-
ministrators Tom Ross, Clark Ross, Carol Quillen, Wendy Raymond,
Verna Case, and Barbara Lom who had confidence in us and encouraged
us to persist despite setbacks along the way.
Thanks to my wife Amy Brooks for her constant support during the
development of this textbook, and my daughter Evelyn for her endless
energy. Thanks to Malcolm Campbell for his steadfast resolve and opti-
mism. Without him, this book would not exist. Thanks to collaborator
Laurie Heyer for taking my sometimes half-baked math ideas and turn-
ing them into powerful and elegant Bio-Math Explorations. I learned a
lot from both of them. While the math is largely absent from this
book, our collaboration with her made this a better book. Nancy Stamp
at Binghamton University, and Bill Dunson and Richard Cyr at The
Pennsylvania State University influenced me greatly in how I think as
a scientist and approach my teaching. Finally, I thank my students in
Integrated Concepts in Biology II, who enthusiastically participated in
our experiment to redesign introductory biology, starting with the text
and ending with a new approach to teaching biology.
 Introduction
Although the unit of study in evolution is most often the population,
evolution can be and is studied at the levels of the molecule, cell, and
organism. In this book, evolution will be studied at the level of the in-
dividual organism and the population, that is all the individuals of the
same species living in the same place at the same time, by examining how
natural selection acts on individuals that possess certain traits, the evolu-
tionary consequences of rapid environmental changes, and the evolution-
ary consequences of gene flow and genetic drift, which are all manifested
at the population level. The main themes of the evolution will be evident
throughout the book. For instance, natural selection is the mechanism
that accounts for adaptation to the selective pressures of interacting spe-
cies. Life continues to evolve as the environment changes, and humans
are a major contributor to changes in our 21st century environment.
 CHAPTER 1

Selection Acts on
Individuals with Variable
Characteristics

Variation of phenotypes among individuals in a population was an im-

portant concept to Darwins description of natural selection. The pres-
ervation of favourable individual differences and variations, and the
destruction of those which are injurious means that within the range
of variation of phenotypes, some will be maintained and some will be
eliminated. Individuals vary in their expression of phenotypes in a popu-
lation, and both genes and the environment influence the variation of
those phenotypes. Variation in alleles leads to variation in proteins, which
can lead to variation in phenotypes like blood pressure, but also in ana-
tomical and behavioral characteristics.

Selection can Act on Behaviors

In a study on the evolution of behavior, Lee Dugatkin studied variation
in the behavior of guppies (Poecilia reticulata) when exposed to a preda-
tor. Some guppies, but not all, perform an inspection behavior, where the
individual leaves its school and swims slowly toward a potential predator.
Dugatkin wanted to determine the costs and benefits of such behavior.
Dugatkin collected 60 male guppies from a river in Trinidad, West Indies.
The area of the river where guppies were collected was known to harbor
several different species of fish that preyed on guppies.
Dugatkin first assessed each of the 60 guppies for their tendency to ap-
proach a predator fish. The guppies could see the predator in an adjacent
aquarium, but the predator could not eat the guppy. Dugatkin measured
2 MECHANISMS OF EVOLUTION

how close the guppy came to the predator-containing aquarium. Based

on how close each guppy was to the predator aquarium, Dugatkin classi-
fied it as being bold, ordinary, or timid. The bold guppies had high ten-
dency to inspect and move close to the predator, the timid guppies had a
low tendency to inspect and typically hid behind a plastic plant, and the
ordinary fish exhibited an intermediate level of boldness.
Dugatkin then ran trials where groups of guppies were placed in one
of ten aquaria with a predator (Figure 1). The groups were made of six
fish: two bold, two ordinary, and two timid guppies. Guppies have indi-
vidual variation in their color patterns, which Dugatkin used to tell indi-
viduals apart. After 36 and 60 hours, he collected all the fish and noted
how many and which ones had survived to that point.
Bold individuals do not survive very long in the presence of a preda-
tor. This is an example of how natural selection works: In the context
of the experimental conditions, the timid behavior is favored, whereas
the bold behavior is not. Some fish exhibiting each type of behavior
were consumed by the predator. The bold fish moved closer than timid
fish, leading to their faster demise. Over time, predation acts as a selec-
tive agent, selecting against bold individuals. Selective agents are factors

80

= 36 hour survival
60 = 60 hour survival
percent survival

40

20

0
timid ordinary bold
type of guppy

Figure 1 Total survival (pooled for all ten trials) of guppies with
different behavioral tendencies to inspect potential predators. Survival
was measured at two time points during the experiment.
Source: Data from Dugatkin 1992 in text.
 Selection Acts on Individuals with Variable Characteristics 3

that lead to differences in survival or reproduction and cause natural

selection.
Natural selection, through the agent of predation, should eliminate the
bold behavior from the population. But the boldness behavior is still found
in the population, because many of the guppies that Dugatkin collected
from the wild were bold. The phenotype has not been eliminated from natu-
ral populations of guppies. To further explore the consequences of behavioral
variability, Dugatkin, along with Jean-Guy Godin ran another series of ex-
periments in which guppies were examined for their variability in behavior
and another characteristic, brightness of color (Godin and Dugatkin 1996).
Their first experiment tested the hypothesis that brightly colored
males were bolder than drab males. They used an aquarium with two
compartments separated by a clear partition. In one compartment they
placed a predator fish or nothing, and in the other they placed two male
and two female guppies. One male was brightly colored, and the other
was drab. After a 2 hour acclimation period, the scientists removed an
opaque barrier between the compartments so that the guppies and the
predator could see into the other compartment, but the predator was still
prevented from preying on guppies because of a second clear partition.
The scientists recorded the number of times in 30 minutes that each male
guppy approached the compartment for many pairs of males.
Bright males made an average of 8 (2.1) inspections in 30 minutes
when a predator was present on the other side of the clear partition and
only a little over 2 (1.8) inspections in 30 minutes when no predator
was present. Drab males inspected only about 2 times in 30 minutes,
regardless of whether a predator was present or not (2.2 0.8 with preda-
tor present and 1.7 0.5 with predator absent).
In their next experiment, the scientists used a similar setup, but they
varied the presence of females and used a model of a predator instead of
a live predator. They suspended the model along a movable track so that
they could remotely move the predator toward the guppies. The scientists
created a color index related to the average size and brightness of color
patches on a male. In the first set of trials, paired bright and drab males
were placed in one compartment with or without a female.
The scientists measured the number of approaches each male made
toward the model predator and found that bright males made on average
4 MECHANISMS OF EVOLUTION

about 6 inspections every 15 minutes whether females were present or

absent. Drab males also inspected at about that rate when females were
absent but when females were present drab males inspected only about
half as much as when they were absent.
The scientists determined the relationship between the number of in-
spections that an individual made and the quantitative color index score
of that male and found that there was a statistically significant positive
correlation between the two variables using a Spearman rank correlation
test (rS = 0.43; P = 0.005).
The next trial included a lone male in one compartment and the
model predator in the other compartment. Dugatkin and Godin remotely
moved the model from the far end of its compartment toward the lone
guppy and measured how close the model came before the guppy swam
away. They then determined the relationship between that distance and
the color index score. Again, they found a statistically significant positive
correlation between the two variables using a Spearman rank correlation
test (rS = 0.35; P = 0.02).
In one final experiment, Dugatkin and Godin let females observe pairs
of males, one bright and one drab, in the presence or absence of a preda-
tor. They simulated boldness and timidity by placing males in clear plastic
tubes and holding them still, or moving them closer to the predator. In
half of the trials, boldness was simulated in the bright male, and timidity
was simulated in the drab male. In the other half of the trials, boldness
was simulated in the drab male, and timidity was simulated in the bright
male. Males were then placed with the female, and they observed court-
ship behavior. When males were simulated to be bold by moving them
closer to a predator, more bold males were chosen than timid males, re-
gardless of color (simulated bold-bright: 14/20 (p = 0.058); simulated
bold-drab: 16/20 (p < 0.01)). However, when males were simulated to
be bold by moving them, but no predator was present, bright males were
always preferred to drab males by females (simulated bold-bright: 15/20
(p < 0.05); simulated timid-bright: 16/20 (p < 0.01)).
All of these results can be integrated to draw conclusions regarding
the evolution of morphology and behavior. In the presence of female gup-
pies, bright guppies are more likely than drab guppies to swim toward
and inspect potential predators. The bright guppies are also more likely to
 Selection Acts on Individuals with Variable Characteristics 5

flee from predators before the predator gets too close. This may enhance
their survival, and may be something they were unable to do in the first
experiment, where survival of bold guppies was zero. The variation in be-
havioral tendencies is associated with variation in appearance, as colorful
males were generally bolder, especially in the presence of females. Drab
males spent less time inspecting predators, because they spent more time
near females when females were present. However, in choice tests, fe-
males preferred to mate with bold males when a predator was nearby and
bright males when predators were absent. Inspection behavior was not an
issue when there was no predator nearby, but because of the correlation
between brightness and boldness, females may be able to assess male bold-
ness by examination of their color alone.
In Dugatkins first experiment, bold males were selected againstthat
is, their survival was very low because of their tendency to inspect preda-
tors. A scientist may wonder then how individuals with that behavior
remained in the population, or why the population did not consist of all
timid males. Because boldness is preferred by females that counters preda-
tion selection against the boldness.
In addition, variation in color pattern, which is associated with be-
havior, is used by females as a signal for potential boldness when they
cannot actually observe how a male would behave if there were a predator
present. Bold males inspect more often, which may signal to the predator
that it has been spotted, and this behavior allows bold individuals to flee
an approaching predator earlier than a timid individual. Females may
prefer these types of males because bright color indicates their boldness,
and boldness is related to their ability to detect predators. Bold behavior
may be associated with other aspects of success, such as higher rates of
feeding. The value of this to the female is that bold, healthy males may
contribute more advantageous genes to the females offspring. Phenotypes
that remain in a population are maintained by providing an advantage to
the possessor, whereas phenotypes selected against reduce the ability to
survive or reproduce.
If boldness and brightness are then favored in the environment, why
are there any timid, drab males left in the population? Even though fe-
males prefer bold, bright males, timid males are still able to mate, which
helps explain why this characteristic remains. Dugatkin and Godin found
6 MECHANISMS OF EVOLUTION

that the correlation between brightness and boldness was not perfect; not
all bold males were brightly colored. Multiple genes may code for proteins
involved in those characteristics, and the environment in which a guppy
develops may also contribute to that variation. Natural selection is not
perfect, and there are many factors that affect the success or failure of any
one individual. Timid males may have higher survival than bold males
under some conditions. If all timid, drab males were eliminated from the
guppy population, the remaining population of bright, bold males will be
less variable, and this may have negative consequences in an environment
with a high abundance of predators. Such a population of bold and bright
guppies could be selected against and face extinction.

Natural Selection on a Discrete Trait

Most sexually reproducing populations contain individuals that display
variable behaviors, structures, molecules, and other phenotypes, like the
guppies just studied. Doug Schemske and Paulette Bierzychudek used
another variable population and investigated whether that variability was
maintained by natural selection. Schemske and Bierzychudek worked
with desert snow (Linanthus parryae), a small annual plant that has a
flower color dimorphism; some plants have blue flowers and others
have white flowers. A dimorphism refers to a population that contains
individuals that are of one of two phenotypes. This plant lives in the
deserts of southwest North America. Flower color is determined by a
single gene, with blue dominant to white. Populations of this plant can
contain all blue flowered individuals, all white flowered individuals, or a
mix of the two.
Schemske and Bierzychudek asked how selection acted on individu-
als that had different flower color. Individual populations of desert snow
tend to be stable in flower color; that is, the frequencies of phenotypes
found in one area remains constant over time. They found a shallow ra-
vine where the plants on one side were predominantly blue flowered and
the plants on the other side were predominantly white flowered. Over a
period of 7 years, they sampled plots along two lines that crossed from
one side of the ravine to the other, one at the northern end of the ravine
and one at the southern end. The flower color data were averaged over
 Selection Acts on Individuals with Variable Characteristics 7

many annual censuses, and what they found was that the proportion of
white flowers was consistently at or close to 0 on the west side of the
ravine and consistently at or close to 1.0 on the east side of the ravine.
The researchers hypothesized that there was intense local selection for
flower color, such that blue flowers were favored on the west side of the
ravine and white on the east. To support this hypothesis, the researchers
tested four other genes, predicting that other genetic loci would show
no pattern across the ravine if selection were only on flower color. To
determine the frequencies of alleles of the four genetic loci, the scientists
collected individual desert snow plants across the ravine. They extracted
enzymes and separated allozymes, variants of an enzyme, for the four
genes using electrophoresis. Electrophoresis is the process in which large
molecules can be separated according to size and electrical charge by ap-
plying an electric current to them in a gel. The scientists found exactly
what they predicted; the other genes that do not influence flower color
were not selected for or against.
The scientists also planted both white and blue flowered plants in
plots on both sides of the ravine to determine their seed production suc-
cess in the two habitats. Note that 1995 was wetter than average, and
1996 was drier than average. In 1995, there was no difference in seed
production between blue and white plants on the west side but on the
east side, where white flowers historically dominated, seed production in
white flowers was significantly greater, by about 30%, than seed produc-
tion of blue plants. The trend was reversed in 1996, the drier than aver-
age year, where blue plants produced significantly more seeds than white
plants on the west side, but both white and blue plants produced equal
numbers of seeds, on average, on the east side.
Finally, Schemske and Bierzychudek collected data on environmental
factors on the two sides of the ravine to determine what selective factors
there might be in the two habitats. They looked at the other plants in the
community, which are potential competitors, as well as soil properties.
Nine out of ten plant species had differences in area covered on the west
side versus the east side, with six covering more area on the west side than
the east, and three covering more area on the east side than the west. The
researchers also found significant differences in soil chemistry, with five
out of ten variables being different on one side than the other.
8 MECHANISMS OF EVOLUTION

The scope of Schemske and Bierzychudeks study, in terms of the area

studied and the number of desert snow populations studied, was small.
Within the two small populations studied, however, the researchers es-
tablished that there was a clear difference in flower color on each side of
the ravine and that the observed differences persisted over time. That is
suggestive of two populations adapted to two specific habitats, although
limited dispersal could also explain the distribution of flower types.
Because flower color is determined by one gene, examination of flower
color can lead to estimates of allele frequencies at that genetic locus. On
the east side of the ravine, where white flowered plants predominate, al-
most 100% of the alleles in the population were for white flower color.
This can be concluded easily because that characteristic is recessive. On
the west side, if all the blue-flowered individuals were heterozygous, the
frequency of the white flowered allele would be 50%, and if all blue flow-
ered individuals were homozygous dominant, that frequency would be
0%. The exact percentage in the blue flowered populations is unknown,
but because the flower color is relatively stable and almost always blue, it
can be speculated that most of those blue-flowered individuals are homo-
zygous. Importantly, it is clear that the frequency of one allele changed
across the ravine, something that alleles of the other genes tested do not
do. That suggests no selection for or against those enzymes and that only
flower color is selected for.
The differences in seed production, although not consistent from year
to year, offer a clue to the maintenance of flower color on either side
of the ravine. In 1995, a year with greater precipitation, white flowered
plants on the white flowered side of the ravine produced more seeds per
plant than blue flowered plants. On the blue flowered plant side, blue
and white flowered plants produced equal numbers of seeds per plant.
In 1996, a drier than average year, blue flowered plants produced more
seeds per plant than white flowered plants on the blue flowered side of the
ravine. White flowered plants are typically more successful in wet years
and the blue in dry years. Plant success is also tied to other environmental
conditions, evidenced by the different species of plants present and the
different soil conditions that plants experience on either side of the ravine.
How the soil differences came to be so great over such a small spa-
tial scale is unknown. The soil environment or some other unknown,
 Selection Acts on Individuals with Variable Characteristics 9

unmeasured factor could have then given rise to variation in plant com-
munity composition. Either of these factors, soil or the other species of
plants present on either side of the ravine, could be the source of selection
for flower color on the two sides of the ravine, although Schemske and
Bierzychudek did not test individual factors in the soil or in the plant
community. However, Schemske and Bierzychudek showed that ecologi-
cal factors can and do vary, and this variation leads to natural selection
on a local scale over short periods of time. Natural selection can eliminate
certain characteristics from a population, thereby reducing variation. But
natural selection can also maintain variable characteristics by favoring
certain types in different local habitats. Variation among individuals in a
population can also lead to descent with modification over much longer
periods of time.

Bibliography
Dugatkin LA: Tendency to inspect predators predicts mortality risk in the
guppy (Poecilia reticulata), Behav Ecol 3:124127, 1992.
Godin J-GJ, Davis SA: Who dares, benefits: predator approach behaviour
in the guppy (Poecilia reticulata) deters predator pursuit, P Roy Soc
Lond B Bio 259(1355): 193200 1995.
Godin J-GJ, Dugatkin LA: Female mating preference for bold males
in the guppy, Poecilia reticulata. Proc Natl Acad Sci USA 93(19):
1026210267, 1996. Available online: http://www.jstor.org/stable/
40373.
Schemske DW, Bierzychudek P: Spatial differentiation for flower color
in the desert annual Linanthus parryae: was Wright right? Evolution
61(11):25282543, 2007.
 Index
Allozymes, 7 DNA, 28
Alpine willowherb, 3437 Dominant allele, 6
Altered land use, 11 Dugatkin, Lee, 15

Bees, 25
median flight distance, 2627
Behavior, selection acting on, 16
Bierzychudek, Paulette, 69
Black grouse, 3740
Bladder campion
allele combinations of, 28
genetic distance and geographic
distance, 2830
widespread dispersal of, 30
Brassica rapa. See Wild mustard
Bush cricket, 1213
Business-as-usual policy, 19
Butterflies, 25
median flight distance, 2627
Electrophoresis, 7
Environmental Protection
Agency, 18
Epilobium fleischeri. See Alpine
willowherb
European starlings, 3031
Evolution
non-adaptive, 33
of population, 3340
range expansion, 1213
response to rainfall, 1417
of species. See Species evolution,
in response to climate
change
Executive Orders, 18

Cabe, P. R., 3031 Flower color determination, 68

Campanula thyrsoides. See Yellow
bellflower
Climate change
human-induced, 1718
policy-makers of, 17
species evolving in response to
rainfall, 1417
range expansion, 1213
Color index, 3, 4
Confidence interval, 39
Deforestation, 11, 17
Desert snow, 69
Dimorphism, 6
Discrete trait, natural selection on,
69
Dispersal, 24
geographic and genetic distance,
2731
populations and, 2327
Fossil fuels, burning of, 11, 17
Fungus, 24
Gene adaptation, 31
Gene flow
lack of, 25
between populations, 31
Gene mutation, 31
Genetic distance
of black grouse, 38
of bladder campion, 2830
of European starlings, 3031
Genetic diversity, 31
population bottleneck reducing,
3740
population isolation affecting,
3337
Genetic drift, 36, 40
Genetic isolation, 25
Genotype, 25
46 INDEX

Geographic distance Phenotypes, variation of, 1

of bladder campion, 2830 Poecilia reticulate. See Guppies
of European starlings, 3031 Policy makers
Geum reptans. See Rose, species of for climate change, 1718
Global climate change. See Climate denying global climate
change change, 19
Godin, Jean-Guy, 35 responded to IPCC, 18
Greenhouse gases, 11, 17 using scientific methods, 19
Guppies Pollinators feeding strategy, 26
brightness of color, 34 Population
classification of, 2 bottleneck, 3740
in predator-containing aquarium, 16 isolation, 3337
total survival of, 23 structure, 2
variation in behavior of, 1 Precautionary principle, 1920
Predation, 23
Heritability, 1718
Heterozygosity, 3435, 38 Rainfall, evolutionary response to
Heterozygous, 8 changing, 1417
Homozygous, 8 Random Amplification of
Human-induced climate change, 18 Polymorphic DNA
Hyphae, 24 (RAPD), 34
Recessive allele, 8
Inbreeding, 39 Restriction enzymes, 28
Intergovernmental Panel on Climate Rose, species of, 3437
Change (IPCC), 11, 17
Schemske, Doug, 69
Kay, Eric, 2325 Schmitt, Je, 2527
Selective agent, 23
Linanthus parryae. See Desert snow Silene vulgaris. See Bladder
campion
McCauley, D. E., 2829 Solar activity, temperature increase
Mushroom, 2425 and, 11
genetic similarity of individuals, 24 Spearman rank correlation
limited dispersal on, 25 test, 4
movement and genetic relatedness Species evolution, in response to
of, 24 climate change
rainfall, 1417
Natural selection range expansion, 1213
act on behaviors, 16 Squirrels, 23
through agent of predation, 3 Sternus vulgaris. See European
description of, 1 starlings
on discrete trait, 69 Sunflower-like plants, 2527
populations evolution in absence,
3340 Tetrao tetrix. See Black grouse
Non-adaptive evolution, 33
United Nations Environment
Obama, Barack, 18 Programme (UNEP), 18
Olson, M. S., 2829 US Fish and Wildlife Service
Oyster mushroom. See Mushroom (USFWS), 30
 INDEX
47

Vilgalys, Rytas, 2425 genetic differences in, 16

Volcanoes, temperature increases
and, 11
Weight-of-evidence approach, 20
Wild mustard, 1417
flowering time
distribution of, 15
heritability of, 1617
mortality differences in, 16
survival of, 1516
World Meteorological Organization
(WMO), 17
Yellow bellflower, 3437
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Ecological Homeostasisby Christopher J. Paradise and A. Malcolm Campbell
Ecological Interactionsby Christopher J. Paradise and A. Malcolm Campbell
Emergent Properties of Individual Organismsby Christopher J. Paradise and
A. Malcolm Campbell
Organismal Homeostasisby Christopher J. Paradise and A. Malcolm Campbell
Population Homeostasisby Christopher J. Paradise and A. Malcolm Campbell

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