Web PDF

EBOOKS Evolutionary History
FOR THE Christopher J. Paradise A. Malcolm Campbell

APPLIED BIOLOGY COLLECTION
This book describes how evolutionary history is studied using
SCIENCES
several well-known examples and also using evolutionary trees.
LIBRARY Evolutionary trees are analyzed and used to explain adaptive
Create your own radiations of orchids and the diversification of bats over geo-
Customized Content logic time. Evolutionary trees and genetic evidence is used to
infer when and from what ancestors terrestrial plants evolved
Evolutionary
Bundlethe more
and invaded land. Specific adaptations of early land plants led
books you buy,
to the evolution of terrestrial plants and their success on land.
the greater your
History
Evidence about the ancestors and habitats of humans is used
discount!
to infer and analyze the evolution of the human family tree,
whose populations were subject to the same forces of evolution
THE CONTENT to which other species are subject. Human evolution was not
Energy Physics linear, involved offshoot species that did not survive, and took
Engineering many thousands of years. In contrast, evolution can be seen in
just a few years or less in other examples, and analysis of the
Biotechnology
evolution of mechanisms of pesticide resistance in insects will
Biology
be used to illustrate this rapid evolution.
Mathematics
Chemistry Christopher J. Paradiseis professor of biology and environ-
mental studies at Davidson College. He teaches introductory
biology, ecology, entomology, and topical seminars on ecotoxi-
THE TERMS
cology and renewable natural resources. He also occasionally
Perpetual access
leads a study abroad program in India. His research evaluates
for a one time fee anthropogenic factors that influence insect biodiversity at a
No subscriptions or variety of scales. His current research interests include effects of
access fees land use patterns on pollinator communities in parks.
Unlimited
A. Malcolm Campbellteaches biology at Davidson College,
concurrent usage
NC. He received national and international education awards:
Downloadable PDFs
Genetics Society of America (2013); American Association for the
Free MARC records
Advancement of Science (2012); and American Society for Cell
Biology (2006). He was the founding co-editor in chief of CBE Life
For further information,
Sciences Education; founding director of Genome Consortium
a free trial, or to order,
contact: for Active Teaching (GCAT); and member of the American Soci- Christopher J. Paradise
sales@momentumpress.net ety for Cell Biology governing council (20122014).
A. Malcolm Campbell
Evolutionary History
Christopher J. Paradise, PhD

A. Malcolm Campbell, PhD
Copyright Christopher J. Paradise and A. Malcolm Campbell. 2016.
All rights reserved. No part of this publication may be reproduced, stored

in a retrieval system, or transmitted in any form or by any means
electronic, mechanical, photocopy, recording, or any other except for
brief quotations, not to exceed 250 words, without the prior permission
of the publisher.
First published in 2016 by

Momentum Press, LLC
222 East 46th Street, New York, NY 10017
www.momentumpress.net
ISBN-13: 978-1-60650-965-4 (print)

ISBN-13: 978-1-60650-966-1 (e-book)
Momentum Press Biology Collection
Cover and interior design by S4Carlisle Publishing Services Private Ltd.,

Chennai, India
10 9 8 7 6 5 4 3 2 1
Printed in the United States of America

Abstract
This book describes how evolutionary history is studied using several
well-known examples and also using evolutionary trees. Evolutionary
trees are analyzed and used to explain adaptive radiations of orchids and
the diversification of bats over geologic time. Evolutionary trees and
genetic evidence is used to infer when and from what ancestors terres-
trial plants evolved and invaded land. Specific adaptations of early land
plants led to the evolution of terrestrial plants and their success on land.
Evidence about the ancestors and habitats of humans is used to infer
and analyze the evolution of the human family tree, whose populations
were subject to the same forces of evolution to which other species are
subject. Human evolution was not linear, involved offshoot species that
did not survive, and took many thousands of years. In contrast, evolu-
tion can be seen in just a few years or less in other examples, and analysis
of the evolution of mechanisms of pesticide resistance in insects will be
used to illustrate this rapid evolution.
Keywords
species, extinction, evolution, population, speciation, adaptive radiations,
natural selection, ancestors, common ancestor, lineage, descendants,
evolutionary tree, phenotype, hominids, radioisotope dating, relative
dating, population genetics, heterozygous, antibiotic resistance, homozy-
gous, pesticide resistance, adaptation
Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 Descent with Modification and Adaptive Radiations
can be Observed.................................................................1
Adaptive Radiation of Orchids from a Common
Ancestor.........................................................................3
Rapid Diversification in Bats..............................................9
Chapter 2 Terrestrial Plants Evolved from Aquatic Ancestors
Millions of Years Ago.......................................................17
Chapter 3 Humans Evolved From Hominid Ancestors in Africa.......25
Ethical, Legal, Social Implications: Eugenics
Yesterday and Today......................................................34
Ethical, Legal, Social Implications: Evolution has not
Reached its Peak; Humans are Still Evolving.................37
Chapter 4 Evolution can Occur Quickly in Response to Strong
Selection..........................................................................41
Ethical, Legal, Social Implications: Overuse of
Chemicals like Pesticides and Antibiotics can Have
Detrimental Effects.......................................................54
Conclusion............................................................................................59
Glossary................................................................................................61
Index....................................................................................................63
Preface
This book about evolutionary history and the analysis of evolutionary
history is part of a thirty book series that collectively surveys all of the
major themes in biology. Rather than just present information as a col-
lection 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 impor-
tant information from a single book with the potential to see how the
major themes span all size scales: molecular, cellular, organismal, popula-
tion and ecologic systems. The major themes of biology encapsulate the
entire discipline: information, evolution, cells, homeostasis and emer-
gent 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 evolutionary history 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 evolutionary history the way professional scientists
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 gener-
ous 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
administrators 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
Charles Darwin wrote in The Origin of Species that the affinities, or
relationships, of all the beings of the same class can be represented by
a great tree, wherein the twigs represent existing species. He was in
effect describing the radiation of a successful group of species and the
extinction of less successful species. As organisms produce successive
generations, individuals in one generation are linked to their ancestors
by lines of descent. And yet the descendants may not look exactly like
the ancestors. More recently, the evolutionary biologist, Sewall Wright,
wrote a treatise on evolution toward the end of his 70-year career.
Regarding the population concept of a species, he wrote that prior to
Darwin a species was thought of as a type of organism; a more or less
static entity. After Darwin, the concept of a species began to shift to
one of an interbreeding population of organisms. If this population was
distributed over a very large geographic range, the individuals at one end
of the range may not look exactly like individuals at the other end of the
range, but throughout, the different variations merged gradually with
one another through a continuous series of intermediate forms. Species
thus began to be conceived of as dynamic entitiesthe populations, which
varied through time and space. The study of evolution at the popula-
tion level progressed significantly with Wrights insights. Speciation will
be explored, but in this book it will be examined in relation to change
over evolutionary time and adaptive radiations. The main themes of
evolution will be evident throughout the book. Natural selection is the
mechanism that accounts for adaptation to selective pressures. The evo-
lution of plants that can live on land led to entirely new communities
on the planet, and those communities are the ancestors of modern day
terrestrial plant communities. Life continues to evolve as the environment
changes.
CHAPTER 1
Descent with Modification

and Adaptive Radiations
can be Observed
Although we cannot go back in time to observe an ancestral species,

scientists can use modern techniques to infer historical events in the
evolution of any group of species. Darwin argued two theses in The
Origin of Species. First, that all organisms have descended with modifi-
cation from a common ancestor, which is one from which a group of
related species has evolved. Second, that the chief agent of modification
is natural selection. All descendants of a common ancestor belong to the
same lineage, that is, a line of descent. Variation arises through muta-
tion, recombination, and environmental factors, and natural selection
operates on that variation within a population. These concepts provide
a clue as to how descendants can be modified from their ancestors over
long periods of time. Although descendants are modified from their
ancestors, there are still common characteristics that can be used to
determine evolutionary relationships.
Common characteristics are examined in living and fossil specimens
and in anatomical and molecular studies. Scientists infer evolution-
ary history from these data. An evolutionary tree can be constructed
to show the hypothesized relationships among groups of organisms
(Figure 1). An evolutionary tree is a diagram that shows the evolutionary
relationships among various species. Evolutionary trees show the evo-
lutionary relationships among species hypothesized to have a common
ancestor. In the tree, each branch point with descendants arising from
it represents the most recent common ancestor of the descendants. The
ends of the branches represent the species or groups of species being
2 EVOLUTIONARY HISTORY
mosses ferns pines, spruce, fir flowering plants
plant common ancestor
Figure 1 Evolutionary tree of major groups of plant species.

The base of the tree trunk represents the common ancestor
of plants. Flowering plants include orchids.
Source: See text for explanation of a, b, and c.
studied. For instance, in Figure 1, all mosses are grouped together at

the end of the leftmost branch to indicate that they are all more closely
related to each other than any moss is related to any species on another
branch. The base of the tree represents the common ancestor of all plants.
The common ancestor of all mosses evolved along branch a. There is
great diversity within the flowering plants; orchids are one diverse group
among many.
Knowing how to read evolutionary trees is a useful skill for a biologist.
Following the logic in Figure 1 allows scientists to construct knowledge
that will allow them to interpret more complicated trees. The species that
existed at branch point b was the common ancestor of ferns, the pines,
spruce, and fir (the conifers), and the flowering plants. This common an-
cestor was a species that existed in the distant past, and science may never
know exactly what this species looked like, but because of our knowledge
of the relationship of all the descendant species, scientists know this com-
mon ancestor existed. All species of flowering plants had a single common
ancestor, which existed on that branch just after point c where the co-
nifers split off from the flowering plants. Although the evolutionary tree
DESCENT WITH MODIFICATION AND ADAPTIVE RADIATIONS 3
shown in Figure 1 shows only one branch leading to the flowering plants,
keep in mind that this branch represents over 250,000 living species of
plants, including the orchids.
Adaptive Radiation of Orchids

from a Common Ancestor
Orchids are flowering plants, often with highly modified flowers. As with
other organisms, they continue to evolve and possess adaptations that
have led to their success in terms of numbers of orchid species. Biologists
sometimes measure the success of a group of species by how many species
there are. Because there are about 30,000 species of orchidsmore than
any other plant familyorchids are considered quite successful. Adapta-
tions possessed by individuals have led to the success of species. We can
examine orchid evolutionary history by studying evidence of their de-
scent from a common ancestor. Scientists want to piece together orchid
evolutionary history to understand the characteristics that made them so
successful.
To determine the evolutionary history of orchids, including to what
other plants they are related, when the common ancestor of orchids lived,
and when orchids diversified, Naomi Pierce and her colleagues studied fos-
silized pollen and DNA sequences of orchids (Ramirez et al., 2007). The
orchid pollen was found on a bee trapped in amber. The amber was embed-
ded in rock estimated to be between 15 and 20 million years old. Attached
to the back of the bee, between its wings, were two packets of pollen from
a single flower with each packet containing many pollen grains.
Because there are very few fossilized orchids or parts of orchids, this
was a new species, named Meliorchis caribea. The phenotype of the fossil-
ized pollen packets is found only in one subfamily of the orchid family.
A subfamily is a taxonomic group of related organisms ranking between
a family and a genus, while a family is a taxonomic group of related
organisms that includes all subfamilies, genera, and species that evolved
from a common ancestor. All orchids are in the same family, and they are
subdivided, based on evolutionary relationships, into five subfamilies. All
orchids have one common ancestor, but each subfamily has a more recent
common ancestor within the orchid family.
Based on characteristics of the pollen and how it was attached to the

back of the bee, the scientists hypothesized what the flowers of this orchid
species looked like. Because other orchids also have flowers that place
pollen packets on the backs of their pollinators, the scientists conjectured
that the extinct orchid had flowers with a lip, a specialized petal, upon
which pollinators land. The pollen packet lays at the end of the erect
column, which contains both the anther with the pollen, and the
stigma, which receives pollen from a pollinator. This column projects
out over the top of the lip. In order to collect nectar, the pollinator must
crawl on the lip, under the column, at which time the sticky pollen packet
becomes stuck to its back.
In living orchids thought to be related to M. caribea, the placement
of pollen packets results in attachment on the pollinators mouthparts.
Attachment of the pollen to the dorsal surface of a bee is only possible
with the column arranged such that it is projected over the lip, where the
insect crawls into the flower and the pollen packet is stuck to the bee as
it backs out. The scientists compared the ancient species to living species
and constructed a table of phenotypes with the particular shape or struc-
ture of each phenotypic character (Table 1). For each phenotype, coded as
a letter, the particular form of that phenotype observed in each species is
coded as a number. For instance, orientation of the flower column and lip
is one phenotypic characteristic (a in Table 1). Parallel flower orienta-
tion was coded as 0 and perpendicular flower orientation coded as 1.
From just the position of the pollen packet on the back of the fossil-
ized bee, scientists inferred many things about a plant they knew existed
Table 1 Phenotypic characters for several orchids listed by genus.

Only a subset of the species and phenotypes used by Pierce and her
colleagues is shown here. Meliorchis refers to M. caribea and is the
fossilized orchid. Each phenotype is denoted by a letter.
specimen a b c d e f g h i j k l m n
Altensteinia 0 0 0 0 1 0 0 1 0 1 0 0 2 1
Gomphichis 1 0 0 0 0 0 0 0 0 1 0 0 1 1
Goodyera 0 1 0 0 1 0 0 1 0 1 1 1 0 1
Microchilus 0 1 0 0 1 0 0 1 0 0 1 1 0 1
Meliorchis 0 1 1 0 1 0 0 1 0 0 1 1 0 1
Source: Data from Ramirez et al., 2007, supplemental materials.

15 to 20 million years ago (MYA). Once they constructed phenotypes of

this orchid species, they could compare it to living species. From Table 1
and data from the other species that were analyzed, it was determined that
the phenotypes of some orchids are similar to the fossil species and the
phenotypes of others are dissimilar. If all species have the same number in
a column, then the phenotype exhibits no variation and all species exhibit
the same phenotype. If two species have the same variation for each phe-
notype (that is, two rows are identical), then the two species are similar
to each other for all phenotypes examined. That is not the same as stating
that the two species are identical, because variation exists among other
phenotypes not listed in Table 1.
DNA changes over time as mutations accumulate, recombination
occurs, and other mechanisms of evolution act on populations, lead-
ing to variation in characteristics. If these changes are great enough,
then some individuals may no longer be able to interbreed. This hap-
pened many times in the evolutionary history of orchids, leading to an
adaptive radiation, caused by isolation of populations and adaptation
to local conditions and resulting in the high species diversity of orchids.
An adaptive radiation is a rapid evolutionary diversification characterized
by an increase in the number of species in a lineage. Phenotypes that
changed over time and were selected for could be maintained in isolated
populations that eventually became new species.
Pierce and her colleagues used the data, including the data in Table 1,
to place Meliorchis caribea into a particular orchid subfamily, the Or-
chidoideae. They then compared orchid DNA sequences for 55 species
of orchids, distributed among all five orchid subfamilies, and used the
similarities and differences in the sequences to construct an evolutionary
tree. Each nucleotide position in the sequence is akin to a phenotype
a mutation adds variation to that position, and large differences in the
sequences indicate a possible speciation event. Using knowledge of how
old the fossil M. caribea was and how similar it was to other members
of its subfamily (from the full set of data on which Table 1 is based), the
scientists placed a timeline on the evolutionary tree. Species that are not
very similar split a long time ago and similar species split more recently.
Scientists can use the information in tables such as Table 1 to quantify the
similarity between species and construct evolutionary trees.
In their evolutionary analysis, the scientists also included DNA

sequences from plants thought to be closely related to orchids. This
allowed determination of the dates when the ancestor of all orchids lived
and when particular groups split. Two sets of dates were used to calculate
orchid divergence times: the oldest and youngest estimates of the ages of
the orchid fossil, as well as several related plant fossils. The age boundaries
of M. caribea (15 to 20 million years old) were then used to construct two
separate scales, based on the range of the age estimate.
Pierce and her colleagues estimated that orchids evolved and began
to form new species before dinosaurs went extinct 65 MYA (the K/T
boundary, now known as the K-PG boundary). They concluded that the
most recent common ancestor lived 76.5 to 84.6 MYA. Their evolution-
ary tree also suggested that all or most of the subfamilies had evolved
prior to 65 MYA. This indicates that an adaptive radiation occurred early
in the evolution of orchids.
There are several characteristics associated with the high species di-
versity of orchids. The most diverse subfamily, Epidendroideae, contains
over 15,000 epiphyte species that live almost exclusively on trees and
are not rooted in the soil. Well over half of all orchids belong to this
subfamily. The high diversity has been hypothesized to be due to liv-
ing in a highly specialized habitat, attached to trees in the canopy of
forests. The forest and trees can be broken up into several specialized
habitats. Epiphytes can attach themselves to the main trunk of a host
tree, to secondary trunks higher up in the tree, or out closer to the tips of
branches. Most epiphyte species occur in the secondary branches, with
very few out at the very tips of host branches and in the lower parts of
main trunks.
Possessing adaptations that allow small plants to live in trees provides
them with high light conditions, as opposed to being small and living
in the shade on the ground. Orchids and other plants with this adapta-
tion are able to compete successfully for light, which may be limiting in
forests. Although living at the very tops of trees might provide orchids
with the highest light conditions, very few species live there. The dis-
advantages caused by high wind conditions, extreme daily temperature
variations, and lack of support from weak branches may outweigh the
advantages of high light. However, living high enough to gather light but
gaining protection through a strong attachment to branches or trunks

appears to be of benefit to individuals and their descendants. Once it
evolved, it led to many species of orchids with the epiphytic lifestyle.
If specific adaptations are required to survive in these habitats, indi-
viduals that evolve adaptations will be successful in these areas, and this
can lead to isolation of populations, which can result in speciation. This
hypothesis can be tested by comparing species diversity in orchids and
non-orchid plants living in trees to related plants living on the ground.
Barbara Gravendeel and her colleagues tested the hypotheses that
living aboveground promotes species diversity. They randomly selected
100 genera of orchids that live in trees and 100 genera of orchids that
live on the ground (Figure 2A). A genera, or genus (singular), refers to the
taxonomic group above the species level.
Each orchid genus in Gravendeel and colleagues database contained
at least one orchid species. They repeated their selection procedure for
non-orchid plants living in trees and on the ground (Figure 2B). The
scientists then determined how many genera had a certain range of
species. For instance, there are 33 genera of tree-dwelling orchids that
have between two and five species each.
Although there are many orchid genera with only one species per
genus, there are more single-species genera that live on the ground. There
are five genera of orchids living on trees that have over 300 species each
in them. No ground-dwelling orchid genera have that many species. For
non-orchids, there are no genera with more than 500 species in them, and
there are two genera with 301 to 500 species that are tree-dwelling and
only one that is ground-dwelling. These trends support the hypothesis
that living in trees fosters high species diversity.
A second hypothesis is related to pollination specialization. Orchids
are typically pollinated by only one or several species of pollinator. If
pollinators specialize on orchids, then high species diversity of orchids
could result from isolation of populations visited by only one species of
pollinator. Pollen from individuals in one population would not be trans-
ferred to another population, leading to isolation of the two populations.
One mutation that spreads through a population may be all it takes to
isolate that population from other populations. This hypothesis can also
be tested by comparing subfamilies of orchids and their pollinators.
35
# of genera having x species 30

= in trees
25
= on ground
20
15
10
0
ly
10
00
00
2-
-2
-5
20
30
50
on
A
6-
-1
10
11
21
1-
1-
1-
51
1
1-
10
20
30
50
35
# of genera having x species
30
25
20
15
10
0
ly
10
00
00
2-
-2
-5
20
30
50
on
B
6-
-1
10
11
21
1-
1-
1-
51
1
1-
10
20
30
50
# of species in genus
Figure 2 Frequency distributions of orchids (A) and non-orchid

plants living on trees aboveground (dark gray bars) and terrestrially,
on the ground (light gray bars). The x-axis shows the range of species
in a genus, and the y-axis represents the number of genera having
that range of species.
Source: From Gravendeel et al., 2004, Table 1.
From a study of orchids and their pollinators, the scientists exam-

ined the numbers of pollinators per species of orchid across the five or-
chid subfamilies. With the exception of subFamily Epidendroideae, the
other four subfamilies primarily contain species that live on the ground.
The researchers calculated the mean number of pollinators per species
for each subfamily, and compared species diversity of the subfamilies
with the mean number of pollinator species per orchid species. The least
diverse subfamily, Apostasioideae, has 17 known species and an average

of 1.5 ( 0.5 s.e.) pollinator species per orchid species. Cypripedioideae
has about 175 known species and an average of 3.3 ( 1.3 s.e.) pollina-
tor species per orchid species. Vanilloideae, the subfamily that contains
the commonly used vanilla plant, from which vanilla flavoring is
derived, has about 235 known species and 4.7 ( 1.8 s.e.) pollinator
species per orchid species. Orchidoideae, the subfamily that contains
the fossil orchid discussed earlier, is an order of magnitude more diverse,
with 4,600 known species. Orchids in this subfamily have on average
4.6 ( 1.2 s.e.) pollinator species per orchid species. Finally, the most
diverse subfamily, Epidendroideae, has on average 3.3 ( 0.3 s.e.) pol-
linator species per orchid species.
Pollinator specialization, unlike the epiphyte hypothesis, does not
seem to be related to high species diversity. The subfamily with the
least number of species (Apostasioideae) has just over one pollinator per
species, on average. The subfamily with the largest number of species
(Epidendroideae) has over three pollinators per species. This led Graven-
deel and her colleagues to reject the pollinator hypothesis, even though
Epidendroideae, with the most species, has more specialized pollinators
than Orchidoideae, with the second most species.
Among scientists studying orchids, there is still a debate about what
adaptations led to the extreme species diversity of orchids; the data
are not conclusive. For other groups of organisms, the major successful
adaptation leading to high species diversity is known. For instance, all
flying animals are relatively more diverse than related non-flying animals.
However, how wings and flight evolved is not clearly known. As with
the evolutionary history of orchids, scientists can only infer evolution by
examination of fossil evidence of individuals and patterns of relationships
and adaptations among living individuals.
Rapid Diversification in Bats

We often cannot directly observe evolutionary changes occurring, because
they either have occurred too long ago, or they take too long for humans
to observe in a lifetime. But we can infer that these changes occurred
from a number of types of evidence. Consider the evolution of a complex
structure, such as wings. The wings themselves are not necessarily com-
plex, but if they are used for flight, then there are many associated struc-
tures and functions associated with flight, which all evolved together to
lead to a descendant capable of flight.
The evolution of wings capable of flight appears to have occurred at
least three different times in evolutionary history. We can infer this from
the fossil record, anatomical studies, and our understanding of the evolu-
tionary relationships among birds, bats, and insects, which are the three
groups of animals capable of true flight. An evolutionary tree shows rela-
tionships among these animals (Figure 3).
All insects are more closely related to each other than any insect is
to either birds or bats. The arrow labeled a in Figure 3 indicates the
evolution of the common ancestor of all insects. Likewise, birds and bats
are more closely related to each other than either is to insects. The arrow
labeled b indicates the position of the common ancestor of all birds,
which is thought to be a dinosaur, and c indicates the ancestor of all
mammals. Despite the close relationship between bats and birds, they
still have distinct evolutionary histories, with bats evolving flight from
within the mammals and birds evolving flight separately, from a reptilian
ancestor. In other words, while both groups are vertebrates and are more
closely related to each other than either is to insects, the ancestors of
each that evolved flight had already separated evolutionarily prior to the
insects birds bats
b c
common animal ancestor
Figure 3 Evolutionary relationships among birds, bats, and insects.

The evolutionary tree for all animals is much more complex.
evolution of flight. Because bats are mammals, bats and mice are more
closely related to each other than either is to any bird. The common an-
cestor of all bats was some ancient mammal, incapable of flight.
Studying bats can help explain the adaptive value of wings and the
evolution of flight. The oldest known fossil bat belonged to a species
named Icaronycteris index and is estimated to be about 50 million years
old. The wings of fossil bats and living bats are basically a membrane of
skin stretched between several elongated fingers of the forelimb, and the
fossil bat I. index, had a forelimb skeletal structure that is remarkably
similar to that of living bats. In other words, bat forelimbs do not seem to
have changed much structurally in the past 50 million years, at least based
on this superficial analysis.
The bat species that is ancestral to the many species of living bats
evolved from non-flying mammals, and the transition from non-winged
to winged is hypothesized to be in the fossil record. In the absence of
that piece of evidence, some scientists have used other approaches to un-
derstand the evolutionary success of bats. For instance, to determine if
the skeletons of bats had changed much over 50 million years, as would
be true if the fossil bat was not capable of flight, Lee Niswander and her
colleagues measured the lengths of digits of fossil bats and living bats.
Specifically, the scientists measured the length of the fifth metacarpal
bone, which is one of the bones of the pinky digit, and they compared
that to an index of body size determined from measurements of other
bones.
Inspection of the skeletons of fossil bats and living bats, along with
the analysis of digit length versus body size, indicates that fossil bats look
very much like modern bats. The relationship between the log of digit
length and an index of body size was a strong linear fit, revealing that
all bats, whether living or extinct, all had fifth metacarpals that were
the same length relative to body size. As body size changed, so did digit
length, but in a predictable manner, whether bats were small or large,
living or extinct. Niswander and her colleagues concluded that a bat that
lived 50 MYA had fully formed wings and was capable of flight. Studies
of DNA sequences in mammals suggest that bats evolved from a small
four-legged mammal with legs similar to those of mice. The lack of fos-
sil bats with shorter digits and, presumably, smaller wings may indicate
that the evolution of wings in bats occurred over a relatively short time
span, geologically speaking. If so, there would be limited fossils of those
individuals in ancient rocks.
There are two hypotheses regarding the selection for longer digits. The
first hypothesis states that bat ancestors evolved as gliding animals. That
is, the long digits and stretched skin would not have worked as wings for
true flight early on but could have aided them if they lived in trees and
glided from branch to branch. The second hypothesis states that bat an-
cestors lived on the ground and used the flaps of skin to maintain some
lift while jumping, perhaps off a small hill. In neither case did these ani-
mals have the necessary musculature or behavior for true flight. A mouse-
like animal with long digits existed more than 50 MYA, and selection
favored it in a particular environment. Scientists can use clues from living
species to understand the evolution of long digits.
Niswander and her colleagues studied the development of fore-
limbs in the short-tailed fruit bat Carollia perspicillata and the mouse
Mus musculus. Using a comparative approach, the scientists compared
the development of bats and mice. During development, bones grow
through proliferation, or rapid cell division, of cartilage cells, cartilage
being a tough, elastic tissue found in vertebrate joints and embryonic
skeletons. Growth is controlled such that proliferation occurs more in
some groups of cells than in others, leading to changes in relative lengths
and shapes of bones. In the long bones of the digits, the cartilage cells
go through a series of steps as they divide and mature, including pro-
liferation and differentiation, which is development of a cell into a
particular cell type.
Niswander and colleagues examined the proliferation and differentia-
tion of bat and mouse forelimbs during equivalent developmental stages
of digit bone elongation (Figure 4). Even though mice mature faster
than bats, stages that are equivalent are given the same number. Stage 18
occurs earlier in mice than in bats in terms of actual time, but stage 18
is comparable for both mice and bats in terms of growth of particu-
lar structures. Next, Niswander and colleagues compared the length of
the fifth metacarpal to the percentage of cells in the differentiation/
elongation step in developing bats during those same developmental
stages (Figure 5).
= mouse differentiation
40 = mouse proliferation
= bat differentiation
= bat proliferation
35
region of 3rd-5th forelimb digits
percentage of cells of growing
30
25
20
15
10
0
18 19 20 21 22
developmental stage equilavents
Figure 4 The percentage of growing third to fifth forelimb

digits composed of proliferating (dark gray) and differentiating/
elongating (light gray) cells in mice (triangles) and bats
(circles) at numbered developmental stages. Stages are
numbered sothat animals in the same stage can be compared.
Source: Modified from Sears et al., 2006, Figure 2c.
70
60
length of 5th metacarpal (m)
50
40
30
20
10
0
10 15 20 25 30 35 40
percent of cells of 5th digit in differentiation/elongation
Figure 5 Percentage of cells in the elongation and

differentiation zone of the fifth metacarpal and length of
the fifth metacarpal as development proceeds.
Source: From Sears et al., 2006, Figure 2d.
There are many genes that affect cartilage cell growth during devel-
opment. Genes that code for proteins called bone morphogenetic proteins
(BMPs) are important in cartilage cell growth. The researchers tested
the response of mouse and bat limbs to bone morphogenetic protein 2
(BMP2) or a BMP2 inhibitor, and they also determined what develop-
mental genes were expressed in bats and mice during different stages of
digit development. Embryonic bat forelimb cells cultured with BMP2,
its inhibitor, or a control, revealed that BMP2 led to an average 239 m
increase in metacarpal length compared with the forelimbs cultured in
control media. In contrast, the inhibition treatment resulted in metacar-
pals that were on average 183 m shorter than controls. They also found
that both bats and mice express BMP2, but that expression is 31% higher
in bat than mice forelimb metacarpals.
Examination of the developing cartilage at the cellular level revealed
that the major difference in the growing portion of the bat digit cartilage
is in the differentiation/elongation phase of cartilage cell maturation and
that this difference appeared in the third, fourth, and fifth digits of the
forelimb, corresponding to our own middle finger, ring finger, and pinky
(see Figure 4). As the percentage of cells that were in differentiation/
elongation increased, so did the absolute length of the digits.
Flight in bats was made possible, in part, by the elongation of the
third to fifth forelimb digits. The analysis of Niswander and her col-
leagues revealed a single gene whose expression is enhanced in bats during
development of the forelimb. BMP2 increases the length of mouse and
rat limbs grown in cell cultures, whereas the presence of a chemical that
inhibits BMP2 causes the limbs to be stunted. This suggests that BMP2
plays a role in the elongation of bat wing digits during development. This
one small evolutionary change led to the emergent property of wings and
may help explain the evolution of bats.
Many other evolutionary changes had to occur in the nervous sys-
tem, muscles, and behavior before powered flight occurred. Once an in-
dividual gained an advantage over other individuals, gliding would have
spread quickly. Other changes that increased the benefit of this character-
istic would have been selected for and also spread. Flying bats could have
evolved over the relatively short geological time frame of a few million
years from non-flying mouse-like ancestors.
In this section, evolution of individuals over the course of millions of

years was examined, as was the linkages between genes, cells, individuals,
populations, and ecological systems in the evolution of two very diverse
groups of organisms, orchids and bats. In neither case were we able to
observe the actual evolution of the phenotypes that made these groups so
successful, but careful observation and experimentation allowed scientists
to draw conclusions about how individuals evolved phenotypes that led
to multiple speciation events over great spans of time, an adaptive radia-
tion. Evolutionary changes that shaped entire ecological systems can also
be examined using similar strategies and techniques, and the evolution
of terrestrial plant communities will be examined in the next chapter.
Bibliography
Cameron KM: Age and beauty: kin to both irises and onions, orchids
have a long history and a large repertoire of enticing tricks, Natural
History June:2632, 2004.
Cooper KL, Tabin CJ: Understanding of bat wing evolution takes flight,
Genes Dev 22(2):121124, 2008.
Gravendeel B, Smithson A, Slik FJW, et al.: Epiphytism and pollinator
specialization: drivers for orchid diversity? Philos Trans R Soc Lond B
Biol Sci 359(1450):15231535, 2004.
Ramirez SR, Gravendeel B, Singer RB, et al.: Dating the origin of
the Orchidaceae from a fossil orchid with its pollinator, Nature
448(7157):10421045, 2007.
Sears KE, Behringer RR, Rasweiler JJ 4th, et al.: Development of bat
flight: morphologic and molecular evolution of bat wing digits, Proc
Natl Acad Sci USA 103(17):65816586, 2006.
Teeling EC, Springer MC, Madsen O, et al.: A molecular phylogeny
for bats illuminates biogeography and the fossil record, Science
307(5709):580584, 2005.
CHAPTER 2
Terrestrial Plants Evolved

from Aquatic Ancestors
Millions of Years Ago
Coevolution of plants and animals has been partly responsible for the
incredible diversity observed in terrestrial communities. Both diffuse
and pairwise coevolution have been important in the evolutionary his-
tories of herbivores and the plants on which they feed, fruit-producing
plants and fruit-eating animals, flowering plants and their pollinators,
and plants and their pathogens. Plants have not always lived on land,
and we have good evidence that terrestrial plants evolved from aquatic
plants sometime around 400 MYA and maybe as early as 475 MYA. The
evolution of a novel way of living often leads to a burst of speciation, and
the move from aquatic to terrestrial existence was such an evolutionary
novelty. In addition, this adaptive radiation led to the evolution of entire
terrestrial ecological systems. As with any evolutionary change, moving
from an aquatic to a terrestrial existence required specific adaptations
and many millions of years of gradual change.
Both plants and animals faced major challenges as they moved from
an aquatic to a terrestrial existence. As anyone knows from swimming,
water is much more supportive than air. Water is more buoyant than
air, and living a terrestrial existence requires a sturdier support system,
such as a tree trunk or a skeleton. All organisms exchange materials with
their surrounding environment. Plants take in carbon dioxide (CO2) for
photosynthesis and either give off or take in oxygen (O2) depending on
their metabolic needs. In aquatic environments, CO2 is plentiful, and O2
may be limited, whereas in terrestrial environments, O2 makes up almost
21% of the atmosphere, and CO2 makes up only about 0.038%so the
latter could be limited to terrestrial plants. All organisms require water,

and living on land means working hard to obtain enough water to sur-
vive. Animals drink, and plants take up water through their roots. In
order for plants to evolve to live on land, they must adapt to the very dif-
ferent conditions and evolve support, gas exchange, and water acquisition
mechanisms.
The evolution of terrestrial plants and animals did not occur over-
night. Wellman and his colleagues examined fossils from the Arabian
Peninsula that offered clues to the evolution of terrestrial plants. The
rock from which the fossils were found was dated to somewhere between
488 and 443 MYA. From other studies, they knew that the region from
which the fossils came was a shore habitat. The rock layers that contained
the fossils were made of shale and sandstone.
Shale is made of tiny particles of clay that are formed into a layer
of rock when exposed to heat and pressure. Clay particles are depos-
ited by movement of water, implying that the region was underwater at
the time. Layers higher up formed later, and these were made of sand-
stone and interpreted as rock formed when the ocean receded. The sand-
stone is indicative of wave action depositing sand onto a shore, and it
implies that the area was on the shore at that time. So this region was first
below and then above the surface of the ocean, which occurs as oceans
and land masses rise or fall.
The fossils were found in rock that was dated accurately, because it
was sandwiched between layers of shale that contained fossils of marine
animals with hard shells, such as clams and snails, which have been well
studied. Scientists know when species of common fossil clams and snails
lived, and presence of these species was used to assign dates to the layers.
Wellman and his colleagues extracted fossils from the rocks. The tiny
fossils in question were found in the sandstone, meaning that the organ-
isms lived in a near shore habitat, possibly near a marsh, estuary, or other
shallow water habitat. Many of the fossils were indicative of non-marine
species. They found large plant fragments and spores, which are single-
celled structures produced by multicellular organisms that can grow into
a new organism without fusing with another cell.
Although spores had been found from fossils dated to that period by
other scientists, this was the first instance of fossil fragments of the plants
TERRESTRIAL PLANTS EVOLVED FROM AQUATIC ANCESTORS 19
that actually produced the spores. The fragments consisted of sections of

the plant that produce, store, and shed spores. The covering of this struc-
ture contained swirls of material observed in electron micrographs, which
Wellman and his colleagues interpreted to be made up of sporopollenin
released from spores as they degenerated. Sporopollenin is a stable chemi-
cal that surrounds spores and pollen grains of terrestrial plants. As spores
degrade in a dead plant, the sporopollenin, which is resistant to degrada-
tion, is released from the spores and ends up in the covering material. The
scientists came to this conclusion based on other research on degradation
of spores and the parts of plants that produce spores.
There is good evidence that the fossils discovered by Wellman and his
colleagues were among the first plants to have colonized land. Although
they lived on land, at that point they still lived close to water. This may
mean that they still did not possess all the adaptations needed to live com-
pletely free from standing water. But finding the fossils in rocks formed
from non-marine sediments is the evidence that they had evolved to live
on land.
The shape and size of the fossil spores were similar to spores of
other land plants. In particular, the fossil spores resemble the spores of
living liverworts, small plants that have no vascular tissue belonging
to a group of plants called bryophytes that include liverworts, mosses,
and hornworts (Figure 6). The spores found here were identical to other
spores found in other rocks of similar age. The key difference here is that
these spores are still attached to fragments of the plants from which they
came, giving scientists their first clues as to what the plants were like, and
the small size of these plants is suggestive of the bryophytes. The large
numbers of spores produced and stored in special structures is further
evidence that these fossils are from terrestrial plants. A key adaptation for
living on land is the presence of sporopollenin, which helps protect spores
from the dry terrestrial environment during the dispersal phase by provid-
ing a stable coat around the spore.
This fossil discovery is valuable data used to piece together the evolu-
tionary history of land plants. Techniques in molecular biology can also
be used to assess relationships among and origins of living plants. The
speculation by Wellman and his colleagues that liverworts were the first
land plants can be used as a hypothesis in genetic studies. Land plants are
A B C
Figure 6 Bryophytes. (A) The liverwort is approximately 3 to 4 cm

across and 1 cm high. (B) The moss mat is 10 to 15 cm across and
1 to 2 cm high, and it includes many individual mosses. (C) The
hornwort is 4 to 5 cm across and 2 to 4 cm high.
Source: A, From http://upload.wikimedia.org/wikipedia/commons/4/41/Liverwort.jpg
Author: Eric Guinther (en:User:Marshman). This file is licensed under the Creative
Commons Attribution-Share Alike 3.0 Unported license. B, http://commons.wikimedia.org/
wiki/Moss#mediaviewer/File:Moos_5769.jpg Manfred Morgner (ka-em-zwei-ein) - selbst.
Moos 5769CC BY-SA 3.0. C, From http://www.botany.hawaii.edu/faculty/webb/BOT311/
Cyanobacteria/AnthoGametoSporoOver500.jpg, Photo by David Webb Author: Jason Hollinger,
2007. This file is licensed under the Creative Commons Attribution 2.0 Generic license.
very diverse and the hundreds of thousands of species are hypothesized to

have all evolved from marine photosynthesizing algae.
If the colonization of land occurred once, then all land plants would
be descendants of one ancestral species. There might be some existing
species, such as the bryophytes, that still retain ancestral characteristics.
By studying many plants from a variety of taxonomic groups, scientists
can test the hypotheses that all land plants evolved from green algae and
that the bryophytes retain ancestral characters and are the most likely
ancestors of all subsequent land plants.
There are several types of algae, but the green algae are thought to
be most closely related to land plants because of several shared charac-
ters. Both groups store carbohydrates as starch, have cell walls made of
cellulose, and possess the photosynthetic pigments chlorophyll a and b
and the accessory pigment -carotene. Other types of algae have different
combinations of these and other pigments.
Bryophytes (liverworts, mosses, and hornworts) have no vascular
tissue, which includes xylem and phloem. Vascular tissue is the supportive
and conductive tissue in plants. Xylem is supporting and water-conducting
tissue, which brings water and nutrients from the roots to the rest of the
evolutionary taxonomic typical intron

tree group pattern
1 2 3
red algae aquatic
algae
green algae
liverworts
(bryophytes)
nonvascular
plants
1 hornworts
land plants
mosses
other vp
seedless
vascular
plants
vascular plants
2 ferns
conifers
plants
seed
3
flowering plants
1 = original of land plants (~475 mya)

2 = original of vascular plants (~420 mya)
3 = original of seed plants (~305 mya)
Figure 7 Evolutionary tree of plants and presence or absence of three

mitochondrial introns, indicated by black or white boxes, respectively
(The gray box indicates presence in 50% of species tested.), among
land plants and two types of algae. vp = vascular plants.
Source: Intron data from Qui et al., 1998; figure modified from http://bio1151.nicerweb.com/
Locked/media/ch29/PlantPhyla.html.
plant. Phloem is tissue that transports organic compounds from photosyn-

thesizing tissues to rest of the plant. These tissues evolved in large terrestrial
plants. Bryophytes thus lack this derived character and retain more ancestral
characters. They are mostly restricted to shady, moist habitats. Bryophytes
reproduce via spores and do not produce flowers, seeds, or fruits. Because
they are limited by not having vascular tissue, they are typically very small,
because vascular tissue is a necessary adaptation for growing large.
There are similarities among the three types of bryophytes, and
Wellman and colleagues suggested that the first land plants were most
like todays liverworts. Qui and colleagues tested the relationships of a
diversity of algae and land plants using the presence of three introns in
two mitochondrial genes, called cox2 and nad1. The pattern in presence
or absence of introns is based on other DNA sequences (Figure 7).
Science will probably never know exactly what the first land plant
looked like, but from the fossil and evolutionary evidence, scientists can
piece together a fairly good, if a bit speculative, picture of the attributes

of that plant. We know the environmental challenges faced by adapting
to life on land and that early land plants had evolved at least one trait,
sporopollenin, which allowed them to disperse spores on land.
Early plants were not free from living in close proximity to water,
and they were small and without vascular or support tissue. These traits
are similar to bryophytes that exist today. Over the 475-million-year
history of land plants, some lineages evolved new traits that allowed
them to expand into new habitats and grow to larger sizes than bryo-
phytes. Bryophytes possess fewer of these derived traits. Terrestrial eco-
logical systems began to form, composed of a growing variety of land
plants as new species formed during the adaptive radiation that followed
colonization of land.
The evolutionary tree in Figure 7 shows that green algae are more
closely related to all land plants than are red algae; the red algae lineage
splits off from all other plants before the green algae lineage does. Qui
and colleagues found that the introns they studied did not exist in
any red algae, green algae, or liverworts; and they concluded that the
introns evolved after a speciation event that separated the liverwort
ancestor from what would become the ancestor of all other land plants.
The lack of the introns in liverworts and algae suggests that liverworts
are more closely related to algae than other land plants. The introns were
found in hornworts and mosses. It is unlikely that these two groups of
bryophytes evolved the introns independently of more derived land
plants, and that event occurred after colonization of land. Mosses and
hornworts are thus more closely related to other land plants than the
liverworts. The data from fossils and the evolutionary tree analysis both
point to liverworts being most closely related to the ancestors of all
land plants.
The evidence put together by these researchers point to the coloniza-
tion of land by plants occurring once, with all subsequent land plants
having evolved from one common ancestor. That ancestor appears to
have attributes similar to living liverworts, which have few adaptations
for terrestrial existence. As other adaptations (such as, vascular tissue,
seeds instead of spores, and flowers) evolved, adaptive radiations occurred
that led to new ecological systems made up of many interacting species.
As animals evolved to live on land, they found habitats filled with rich
resources that spurred an adaptive radiation of land animals.
Bibliography
Qiu YL, Cho Y, Cox JC, et al.: The gain of three mitochondrial introns
identifies liverworts as the earliest land plants, Nature 394(6694):
671674, 1998.
Wellman CH, Osterloff PL, Mohiuddin U: Fragments of the earliest land
plants, Nature 425(6955):282285, 2003.
CHAPTER 3
Humans Evolved From

Hominid Ancestors in Africa
The major mechanisms of evolution, mutation, natural selection, gene

flow, and genetic drift, can all be used to examine long-term patterns
of evolutionary change. While learning about these mechanisms, earlier
chapters in this book explored the time course of evolution in diverse
groups of species. This knowledge of evolution will be applied to humans.
Humans, Homo sapiens, evolved into the species we are today and were
subject to all the mechanisms of evolution to which any other species has
been subjected. Humans have even attempted to direct their own evolu-
tion through eugenics programs (see the Ethical, Legal, Social Implica-
tions section Eugenics yesterday and today). Most people have a curiosity
about where they come from and how they came to be. From a scientific
perspective, the past few decades have revealed great insights into the
evolution of humans.
The evidence for where and when humans evolved is buried in the
fossil record from which scientists have reconstructed an evolution-
ary tree of hominids. A hominid is any member of the human fam-
ily, including extinct human ancestors and living humans. The tree has
changed dramatically during the past few decades as new evidence and
reinterpretations of old evidence have altered our view of our own evolu-
tionary history. New fossils are discovered all the time, and the evidence
is strong that humans evolved in Africa. The discovery of fossils dated
from 6 to 7 MYA have extended the human family tree beyond what
scientists thought was possible just several decades ago. In 2001 and
2002, a team of researchers led by Michel Brunet discovered several
fossils in Chad in central Africa. Brunet and his team found an almost
A B
C D
Figure 8 Skull of Sahelanthropus tchadensis discovered in Chad.

(A) View from the front; note that skull has been compressed on
one side. (B) Side view. (C) View from above. (D) View from below.
(E) Actual size replica in a human hand to show scale.
Source: Photos courtesy of Chris Paradise.
complete skull (Figure 8) along with several other bone fragments. They
estimated the measurements of the skull size and other key features that
could be compared to other ancient and modern species (Table 2). This
helped determine whether the fossils were from a previously undiscov-
ered species. The distance from the base of the nasal openings to the
base of the upper teeth, which is a measure of how far forward the face
extends, was measured and compared to several other specimens, living
and extinct (Table 2).
Humans Evolved From Hominid Ancestors in Africa 27
Table 2 Skull measurements for unknown hominid fossil and several

known species. If the range from multiple specimens is known, that is
shown. Empty cells represent missing data. The upper lip length is
the distance from the base of the nasal cavity to the base of the upper
teeth. Canines are teeth. *Estimates based on reconstruction of the
compressed skull. #canines of P. boisei described as relatively small
compared to Australopithecus. A. = Australopithecus,
P. = Paranthropus, Pan troglodytes is the chimpanzee.
upper lip upper canine lower canine lower canine browridge
length (mm) thickness (mm) width (mm) thickness (mm) thickness (mm)
Chad fossil* 22 10.2 11.0 8.5 18.2
A. afarensis 3033 9.312.5 7.511.7 8.812.4
A. africanus 21.130 610
P. boisei 42.2 # # #
Homo habilis 2531
Homo sapiens short 6.57.7 6.510.4 1.810.1 0~.5
Pan troglodytes 9.511.8 7.017.9 11.4 5.211.8
Gorilla (gorilla) 11.316.8 8.020.9 7.317.5
Source: Data from Wolpoff, 1976, Tables 5, 6, and 7; Brunet et al., 2002, Table 3; Lieberman
etal., 2002, Table 2; and http://www.talkorigins.org/faqs/homs/species.html.
The face of the Chad fossil has a large browridge, from which the
authors concluded that the specimen was a male. The middle of the face
is rather short and does not jut forward as much as it does in chimpan-
zees or other ancient hominids. Other measurements, from this study
and other research, show differences among the different species (Table 2;
Figure 9).
The brain capacity of the Chad fossil is comparable to that of living
chimpanzees and is smaller than a gorilla. The browridge was thicker
than even a large male gorilla, yet the face did not extend forward as it
does with many great apes and human ancestors. The smallish canines
were human-like and unlike great apes (chimpanzees, gorillas, and orang-
utans). Although the size of the canines was within the range shown for
some other species, the smaller values for other species in Table 2 usu-
ally are from females, and Brunet and his colleagues concluded that the
Chad fossil was a male, based on phenotypes such as the large browridge.
The small canines are a derived phenotype of early human ancestors,
and the possession of these traits suggests that the Chad fossil is an early
hominid species in the human family tree. A derived phenotype is one
that has undergone change from the ancestral condition. At the same
time, the small brain size and large browridge, being a primitive trait,
suggests a more apelike species. Brunet and his colleagues concluded that
1600
H. sap.
1400
range of brain volumes (cm3)
1200
H. ere.
H. erg.
1000
800
H. hab.
600 P. rob. Gor.
Au. afar.
P. bos.
400 Au. afr.
Pan
Chad fossil
200
7 6 5 4 3 2 1 0
estimated dates of existence (millions of years)
Figure 9 Estimates of brain volume ranges from fossil evidence of a

variety of hominid species and three living species. The Chad fossil is
at the lower left. Other extinct hominids are shown (Australopithecus
afarensis [Au. afar.], Australopithecus africanus [Au. afr.],
Paranthropus boisei [P. bos.], Paranthropus robustus [P. rob.], Homo
habilis [H. hab.], Homo ergaster [H. erg.], Homo erectus [H. ere.]).
Modern humans, Homo sapiens (H. sap.), chimpanzees (Pan), and
gorillas (Gor.) are shown for comparison.
Source: Data from Wood, 2002, Figure 2 for dates and other sources for brain capacity.
this species was sufficiently different from any other species known to
science and gave the species a new name, Sahelanthropus tchadensis. The
mixture of both primitive and derived traits led the researchers to further
conclude that S. tchadensis is close to the base of the human family tree,
possibly being close to the last common ancestor of hominids and apes.
Prior to the discovery of this fossil, it was thought (based on molecu-
lar evidence) that the last common ancestor of hominids and apes existed
about 5 MYA. The analysis of the Chad fossil, which extended the date
back by 1 to 2 million years, was radical and controversial. In addition
to examination of the skull itself, the fossils and rocks in which it was
discovered must also be examined in order for Brunet and his colleagues
to accurately date the fossil.
One of Brunets colleagues, Vignaud, was the lead scientist for this
analysis. The area where the S. tchadensis fossils were found was rich with
other fossils, and the team found over 700 identifiable mammal fossils
from 24 taxonomic groups just from the locality where the S. tchadensis
skull was found. The research team also examined the geologic forma-
tions of the area. The area is currently a desert, but it was under a lake in
the past and was a desert prior to the lake forming. This means that the
rock layers were formed from sediments deposited at the bottom of the
lake, as well as from sediments deposited from winds and episodic flash
floods that occurred during times when the area was not under water.
Because the rocks are not volcanic in origin, they could not be dated using
radioisotope dating, so the researchers used a relative dating method.
Radioisotope dating is a technique that compares the observed abun-
dance of a naturally occurring radioactive isotope and its decay products,
using known decay rates. Relative dating is a technique that uses layers of
rocks and the fossils contained within to estimate the order of prehistoric
events.
There were several layers of sedimentary rock, but most of the mam-
mal fossils they found were within one particular layer. Wave ripples in
the rock layers, formed from water running over sediments, were run-
ning in many different directions. This led Vignaud and his colleagues
to conclude that this particular layer experienced episodic flooding and
draining, as well as gradation of a lake ecological system into the des-
ert environment that had prevailed in earlier geologic times. A shallow,
semiaquatic area would have provided many different habitats, including
aquatic, shoreline, and from grassland to forest to desert.
The many freshwater fish fossils are known from other fossil depos-
its to have been present in waters of central Africa since about 8 MYA,
and most of them have living descendants still in the region. Terrestrial
mammals include extinct species of hyenas, cats, an otter, a monkey, vari-
ous rodents, several very common hooved mammals, several elephants,
and other species. Some species that were amphibious, like turtles, were
also found. Based on a comparison of other sites of known age that
either contained or did not contain these species, the researchers came
to the conclusion that the site containing S. tchadensis was between
6 and 7 million years old. Based on the geologic and biological evidence,
they also concluded that S. tchadensis lived in an area that contained

aquatic habitats, including swamps and a lake, quite likely bounded by
forest close to shore with open grassland being the dominant habitat away
from the shore.
Many of the fossils found alongside the Chad fossil have descendants
alive today. Hyenas, many hooved mammals, and elephants are all com-
mon in grasslands, although they may also be found in forests. Otters and
amphibious species are found in or near bodies of water. In addition to
using animals as evidence for habitat, it is known that trees often grow
close to the edge of water, even in habitats dominated by grasses, so it may
be reasonable to conclude that both forest and grassland existed near the
lake with the forest being closer to the shore. A careful analysis of data
from ecological systems allows reconstruction of past habitat. Knowing
the past environment is critical when considering under what environ-
mental conditions adaptations evolved. In retracing hominid evolution,
humans are interested in whether bipedalism and large brains evolved
in forest or grassland. We are also interested in when those adaptations
evolved and in what ancestral species.
The Chad fossil, dated beyond the previously accepted divergence
point of the human lineage and the great ape lineage (5 MYA), indicates
that the human family tree began to evolve 1 to 2 million years before
the prior estimate. Features uniquely human were evident even back then,
including the small canines and the more vertically oriented face. Since
that time, other derived traits have evolved, including the large brain
(see Figure 9). The human family tree is more like a bush with multiple
branches, and the data in the figures and Table 2 allow scientists to place
the species into multiple groups. The similarities between hominid ances-
tors and great apes living today are unmistakable, but these groups do
have differences, too.
Hominids are distinguished from great apes by having smaller teeth
as well as other skull features, such as a small browridge. Skeletal fea-
tures that facilitated bipedal locomotion, such as the way pelvic bones
and legs were put together, differ between apes and hominids. However,
not all hominids were similar in every way. Some species possessed a small
brain, large teeth and were facultatively bipedal, meaning they could
walk upright on their hind legs, but they were not obligated to do so
(those species are placed in the Australopithecus genus, shown in the pink
box in Figure 10). Other species in the Paranthropus genus also had a
small brain and were facultatively bipedal, but had much larger teeth
(the light blue box in Figure 10). Finally, species in the genus Homo,
including modern humans (the yellow box in Figure 10), had a large
brain, small teeth, and were/are obligately bipedal, meaning that they
only walk upright on their legsthey dont, as adults, use their fore limbs
(arms) to locomote.
Figure 10 reveals that there were actually many species of hominids,
and we can construct groups that share particular features. In addition,
this evolutionary tree, based on available fossil evidence of known homi-
nids, suggests several conclusions about the evolution from S. tchadensis
to H. sapiens.
The data reveal evolutionary lineages that went extinct with species
that died out without giving rise to descendant species. Brain size, teeth
Sahelanthropus tchadensis
Orrorin tugenensis
Ardipithecus ramidus
Ardipithecus
kadabba Au. anamensis Au. garhi
Au. afarensis Au. africanus
P. boisei
= Homo P. aethiopicus
= Paranthropus P. robustus
= Australopithecus
= early hominids
H. ergaster
= age of fossils H. floresiensis
H. habilis
H. erectus
= inferred relationships between
hominid species
H. neanderthalensis
= relationships of major groups
of hominids H. heidelbergensis H. sapiens
miocene pliocene pleistocene halocene

7 6 5 4 3 2 1 0
millions of years ago
Figure 10 Evolutionary reconstruction of hominids; the human

family tree. Age ranges of fossils are shown by solid gray lines, and
dashed gray lines represent inferred relationships based on fossil
evidence. Major groups are contained within shaded boxes.
Au. = Australopithecus; P. = Paranthropus; H. = Homo.
Source: From http://farm1.static.flickr.com/193/ 463256571_931cf92975.jpg?v=0, Zimmer, and
http://www.efossils.org/ page/phylogeny.
size, and bipedalism all varied, even among species that lived at the same
time. Some species existed for longer periods of time than others, at least
as evidenced by the fossil record. That may indicate their success, but it
may simply be that either no fossils exist for a species at other times, or
they have just not been found yet. Finally, the evidence and analysis by
experts of all these fossils suggests that the evolution from S. tchadensis to
H. sapiens was not linear, and the evolutionary tree of hominids is more
like a bush than a tree.
Many hominid species existed between 3.6 and 1.1 MYA. To further
explore hominid evolution of those species that existed more recently
than S. tchadensis, and to determine the kind of habitats in which the
different major groups of hominids existed at that time, Kaye Reed used
fossil evidence of the mammalian communities in eastern and south-
ern Africa, much as Vignaud and his colleagues did to reconstruct the
environment of S. tchadensis. These regions are areas where major fossil
finds of many hominids have been found. Reeds analysis showed that
the community of mammals in these major regions changed over time.
Scientists can use such data to infer the environmental changes under
which a species evolved. Reed summarized the analysis by plotting the
percentages of grazing mammals, because that indicates open habitats and
grasslands, and the percentages of tree-dwelling and fruit-eating mam-
mals, because those indicate shrubby or forested habitat. If the percentage
of grazing mammals increases over time, this suggests that the grassland
in that area is expanding.
Reeds analysis compares habitats over large spatial and time scales,
and there is a lot of variation from one fossil locality to another.
Australopithecus species, which existed between 4.5 and 2.5 MYA, seemed
to have lived in woodland or shrubby habitat with some grassland.
They may have even existed in flood plains near rivers. Australopithecus
afarensis has generally been found in localities that contained water and
trees. Anatomical studies indicate this species possessed adaptations for
moving in and through trees. Australopithecus africanus probably lived
in the same type of habitat, although possibly somewhat drier, accord-
ing to Reeds analysis. With the exception of some wide variations in the
percentages of tree-dwelling mammals at the sites, percentages of other
mammal groups remained fairly steady prior to 2.5 MYA. This suggested
to Reed that these hominids were constrained in where they lived by the
presence of tree cover and water, either in rivers, lakes, or plentiful rainfall.
Around 2.5 MYA there was a change in the environment, which
caused selection for individuals that could survive drier conditions in
habitats that were more open and less forested. Tree-dwelling or climbing
mammals declined sharply in both southern and eastern Africa at around
the same time, which corresponded to an increase in grazing mammals
between 2.3 and 1.8 MYA. Fruit eating mammals declined between 2.5
and 1.8 MYA in eastern Africa. Australopithecus species evolved into other
species or went extinct.
The hominids that arose after the demise of Australopithecus species
have been placed in the genus Paranthropus. The corresponding habitats
in which these species evolved have been classified as shrubby habitat
to open woodland regions with grasslands as a major component of the
habitat. Some of the fossil localities where Paranthropus fossils have been
found are thought to have contained river deltas and floodplains, indicat-
ing that these species lived fairly close to water, as Australopithecus prob-
ably did. Paranthropus has not been found in habitats thought to be arid
grassland environments, suggesting that as the environment became more
arid and open, these hominids faced a changing environment, which
could have led to their extinction.
About 2 MYA, the first Homo species appear in the fossil record in
Africa. This corresponds to a time period when the environment in east-
ern and southern Africa was becoming drier with more open habitats
that were either partially wooded or complete grassland. Some of the
localities where Homo has been found have been classified by Reed and
other researchers as arid plains and shrub land with possible river-edge
woodland. There also appears to be a time period where Homo overlapped
with Paranthropus. With the appearance of Homo erectus about 1.8 MYA,
the reconstructed environment in localities where this hominid was found
was mostly open grasslands, or savanna.
We now have a rough sketch of the changes that occurred in the
landscape where human ancestors evolved and what some of the selec-
tive factors might have been that led to evolution or extinction of several
groups of hominid species. The genus Homo arose about 2 MYA in an
environment that was becoming more open and arid, possibly selecting
for individuals that had to walk upright to see far distances and scan
for predators. Not only did the environment change, but the available
resources changed, forcing them to adapt to new food sources or face
extinction. Over the 2-million-year history of Homo, brain size continued
to increase, the ability to walk upright became obligate, tool use devel-
oped, and complex social interactions evolved. All of these evolutionary
changes were selected for in a changing environment, and all but one
species of Homo, the one to which humans belong, went extinct.
Ethical, Legal, Social Implications:

Eugenics Yesterday and Today
There are differences between individuals of any species, even humans.
Differences arise from the influence of genes and environments. Chances
are pretty good that any two randomly selected people do not look alike.
The variation among humans is a result of the evolutionary history of
populations of humans living in distinct locations subject to the forces
of mutation, natural selection, gene flow, and genetic drift. Today, those
distinctly variable humans often live side-by-side, in the same coun-
try, city, or neighborhood. The superficial, physical differences between
groups of people are often at the heart of conflicts and prejudices.
To be prejudiced is to have a preconceived judgment of someone
based on appearance or some other characteristic. Racial prejudice is
often associated with an irrational hostility toward a race or ethnic group
different from a persons own. Prejudice may also be associated with a
sense of superiority of ones own race, and it may be used to justify geno-
cide, as it was by Nazi Germany in an attempt to exterminate the Jewish
race. The sense of superiority may also lead to the view that certain traits
are genetically controlled and that some traits are less desirable and some
more beneficial than others.
After the publication of Charles Darwins The Origin of Species and
the rediscovery of Gregor Mendels publications on inheritance, some
scientists sought to apply principles of genetics to improving humanity.
If plant and animal breeders could select for particular traits in agricul-
tural crops and domesticated animals by selective mating, why not do the
same with humans? Eugenics, the science that deals with improving the
human race through selective breeding, had both positive and negative
approaches. Positive eugenics sought to use voluntary breeding programs
to improve the human race and held the view that humans could self-
direct their own evolution through breeding. In Britain, this movement
sought ways to perpetuate the ruling class.
Negative eugenics sought to prevent unfit people from breeding.
People were judged to be unfit based on possession of a variety of traits.
Proponents of this view used degeneracy theory as a guiding principle.
According to this theory, unfit people came from bad environments,
which damaged genes and led to offspring that were also degenerate.
Faulty genes caused degeneracy, and genes could deteriorate over time. If
people that possessed those genes were prevented from breeding, bad
genes would be eliminated from the population. Degeneracy theory was
based on a flawed understanding of genes and heredity. In the same way
that degeneracy theory attempted to explain the heritability of degenerate
behavior, it was also used to predict, incorrectly, that degenerate behavior
could be removed from a family within three generations by moving the
family to a good social environment.
When intelligence quotient (IQ) testing developed, researchers could
quantify intelligence. Low IQs were associated with abnormal behavior,
poverty, and crime. There was a fear among eugenics researchers that
these feebleminded people might appear normal and possibly repro-
duce with intelligent people, mixing bad genes with good genes. There
was also widespread acceptance that genes alone explained all these attri-
butes and that the traits were recessive. A misconception was that people
with low IQs were more likely to exhibit abnormal behavior and be
criminals. This was degeneracy theory in action. Another misconception
was that prevention of homozygous recessive individuals from breeding
would rapidly reduce the occurrence of the recessive trait. Population
genetics and the Hardy-Weinberg equilibrium revealed that this was
not so because of heterozygous carriers.
Eugenics researchers used pedigrees (family trees) to trace traits with
the assumption that traits such as alcoholism, feeblemindedness, lack of
moral control, and shiftlessness were caused by defects in single genes
that were passed on to offspring. The misconception here was that com-
plex behavioral traits were thought to be determined by a single gene.
Today, we know two things about genes and heredity that directly con-
tradict this flawed view: complex, continuous traits are very rarely, if ever,
controlled by a single gene, and although traits like alcoholism may have a
genetic component, there is a substantial environmental component that
determines whether an individual will become an alcoholic.
The eugenics movement got underway in the United States during
the early part of the twentieth century and focused more on negative
eugenics, although there was a popular movement toward better breeding
of the upper class. Soon after eugenics research got under way, eugenics
began to be used to spout racist and anti-immigration propaganda. In the
areas of immigration restrictions, interracial marriages, and forced steril-
ization of unfit individuals, eugenics had significant effects on US history.
Consider the forced sterilization of sex offenders, epileptics, and the
mentally retarded. Laws were passed in about 30 US states that allowed
prisons and mental institutions to sterilize inmates without their consent.
These laws were advocated by eugenics researchers and physicians who
maintained that these people must be prevented from breeding, because
this might contaminate the good gene pool.
The downfall of eugenics began at the end of World War II, as
knowledge of the Holocaust, the attempt to achieve racial purity in Nazi
Germany, came to light. Some US laws remained in force until the 1970s,
and over 60,000 people were sterilized until the laws were repealed and
the practice was discredited as bad science. Some US states have officially
apologized to individuals and the families of sterilized individuals.
Eugenics made a lasting mark on US history, but prejudice and the
potential misuse of genetics still looms large. As we learn more about
the human genome through the Human Genome Project, we learn
more about the influence of genes on traits. In recent years, studies have
shown correlations between possession of a certain allele and a particular
trait. News stories in the media have been written about the gay gene
or the alcoholic gene. Of course, the scientists performing these studies
have shown only a correlation between possession of an allele and a trait,
yet the news stories are often skewed to suggest that possession of that
gay allele will determine whether someone becomes homosexual.
Around the world, there have been recent attempts of one popula-
tion to completely exterminate individuals of another race or ethnicity. In
Cambodia, Sudan, Somalia, Rwanda, and the Balkans, there have been
recent acts of genocide. Most of these examples, when analyzed closely,
are about access to and control of resources, but consider whether it be-
comes easier for one person to kill another if he or she is taught that
people of another race or ethnic group are inferior. Propaganda is put
forth by one group to promote the view that another group is inferior and
that they do not deserve access to resources. In order for the first group
to survive and have a higher quality of life, they must eliminate the other
group. The differences between groups are the excuse used by some to
gain power over others.
Ethical, Legal, Social Implications: Evolution has not

Reached its Peak; Humans are Still Evolving
Hominids evolved over a very long time, often with several coexisting
hominid species living in the same regions at the same time. There is only
one hominid species left on this planet, Homo sapiens. Questions that
arise from this knowledge are whether humans are the peak of evolution
and whether humans are still evolving.
Humans are considered by many to be the pinnacle and endpoint of
evolution. If we have arrived at that pinnacle, there would be no reason to
continue evolving. Over the course of evolutionary history, there is a ten-
dency toward more complex and intelligent life forms, which has, accord-
ing to some, culminated in our species. This view presupposes that natural
selection will lead to more complexity over time, but this is not necessarily
true. Evolution is not directed and does not have a goal, so the conclu-
sion that humanity is the endpoint of evolution is a misunderstanding
of biological evolution. The mechanisms of evolution, including natural
selection, mutation, gene flow, and genetic drift, are not goal-directed, and
they will continue to occur in every species as long as DNA is replicated.
Natural selection, which works with existing structures and modifies
them over many generations, can lead to complexity, but there is no dic-
tate from natural selection that complexity must increase. Environmental
factors, such as living in social groups, an altered diet, and coordinated
effort needed for parental care and food acquisition, contributed to the
evolution of a large, complex, intelligent brain. During the course of
hominid evolution, selection could just as easily favored a solitary exis-

tence if, for instance, human ancestors ate a diet that could be obtained
individually and they lived free from major predators. If that occurred
after a complex brain had begun to evolve, the large brain, which is costly
to maintain, may not have been necessary. It might have shrunk over
time, because a complex brain was not needed to survive in the particular
environment in which hominids found themselves and the energy needed
to maintain the brain was not readily available.
Yet here we are. As discussed in the previous Ethical, Legal, Social Impli-
cations Eugenics yesterday and today, humans have attempted to direct their
own evolution through breeding programs. There may be other ways that
we direct our own evolution, speeding it up or slowing it down, through
the development of technology. We are on the cusp of being able to select
the genes or traits that our offspring will have. The genomic revolution may
soon make it possible for individuals to be able to direct the evolution of
their own families. Alternatively, technology may make evolution obsolete.
Perhaps we have achieved so much technologically that we do not
need to evolve further. Humans may not be the pinnacle of evolution,
but our current level of technological sophistication may negate natural
selection. There can be no doubt that great strides have been made in
medicine, allowing many people to live longer, healthier lives. Natural
selection may no longer be acting on human populations that have access
to the medicine and technology that will keep people alive even in the
face of debilitating or deadly diseases or injury.
Technology has also made the world smaller. Distances of many thou-
sands of kilometers can be travelled in hours, instead of days and months.
One consequence of this is increased gene flow, increasing the genetic
connections among populations. The result of a high level of gene flow
would be homogenization of the gene pool. Mutations could spread from
one to other populations, and separate populations would not be likely
to diverge. Speciation of Homo sapiens is unlikely in this scenario. Some
populations, such as some aboriginal tribes, however, are still relatively
isolated and often exhibit differences in genetic traits. For instance, the
frequencies of two alleles in the MN blood group vary widely among pop-
ulations of Australian Aborigines, Greenland Eskimos, and United States
Caucasians. This suggests very little gene flow among these populations.
If all human populations were connected genetically and diseases

were eliminated, then there would be little natural selection or genetic
drift occurring. Without these two factors occurring and with the preva-
lence of gene flow, there may be little to push human populations geneti-
cally in any one direction, randomly or non-randomly. Mutations would
still occur, of course. Despite these considerations, whether we can
choose our offsprings genes, whether were the pinnacle of evolution, or
whether our technology has stopped our evolution, we can still find evi-
dence that humans are evolving. For instance, in over 75% of the human
population lactase (the enzyme that breaks down lactose, the sugar
in breast milk) declines in activity by 95% at birth. However, this condi-
tion does not affect populations equally. Adult lactase activity is preva-
lent in 95% of European-derived populations but in only about 10%
of Asian and African populations. The rise of lactase persistence seems
to coincide with the rise of dairy farming during the past 10,000 years.
In regions of Africa, selective pressures still exist due, in part, to
inadequate access to healthcare, and this suggests that evolution is
occurring. Malaria affects millions of people annually. Malaria resistance
has been linked to several genes, one of which codes for a metabolic en-
zyme. A mutation in the gene coding for this enzyme causes a deficiency,
which leads to a condition affecting over 400 million people. Although
a deficiency causes problems in the blood, it also improves resistance to
malaria. The mutation has high frequencies in areas that have malaria
outbreaks, suggesting selection. The mutation arose relatively recently;
evolution of humans is still occurring.
If malaria becomes more prevalent in new areas around the world,
the environment has changed. This changes selection pressures, and genes
that confer resistance to malaria may now be favored in this altered en-
vironment. They may spread through gene flow and natural selection.
However, modern medicine may lessen the selection for that gene if drugs
to combat malaria are readily available.
It is likely that we are neither the pinnacle of evolution nor have we
stopped evolving. Knowledge of the mechanisms of evolution would lead
us to suspect that Homo sapiens is subject to these mechanisms in the same
fashion as other species. The evidence that we are still evolving suggests
that we are not the pinnacle of evolution.
Bibliography
Arsuaga JL, Martinez I: The chosen species: the long march of human
evolution (translated by Gomme R), Malden, MA, 2006, Blackwell
Publishing.
Brunet M, Guy F, Pilbeam D, et al.: A new hominid from the Upper
Miocene of Chad, Central Africa, Nature 418(6894):145151, 2002.
Bulhes AC, Goldani HAS, Oliveira FS, et al.: Correlation between lac-
tose absorption and the C/T-12910 and G/A-22018 mutations of the
lactase-phlorizin hydrolase (LCT) gene in adult-type hypolactasia,
Braz J Med Biol Res 40:14411446, 2007.
Carlson EA: The unfit: history of a bad idea, Cold Spring Harbor, NY,
2001, Cold Spring Harbor Laboratory Press.
Lieberman DE, McBratney BM, Krovitz G: The evolution and devel-
opment of cranial form in Homo sapiens, Proc Natl Acad Sci USA
99(3):11341139, 2002.
Pickford M: Orientation of the foramen magnum in late Miocene to
extant African apes and hominids, Anthropologie XLIII(2-3):
191198, 2005.
Micklos D, Carlson E: Engineering American society: the lesson of
eugenics, Nat Rev Genet 1(2):153158, 2000.
Reed KE: Early hominid evolution and ecological change through the
African Plio-Pleistocene, J Hum Evol 32(23):289322, 1997.
Vignaud P, Duringer P, Mackaye HT, et al.: Geology and palaeontology
of the Upper Miocene Toros-Menalla hominid locality, Chad, Nature
418(6894):152155, 2002.
Wolpoff MH: Some aspects of the evolution of early hominid sexual
dimorphism, Current Anthropology 17(4):579, 1976
Wood B: Hominid revelations from Chad, Nature 418(6894):133135,
2002.
Zimmer C: Smithsonian intimate guide to human origins, New York, 2007,
Smithsonian Books/HarperCollins.
Tishkoff SA, Varkonyi R, Cahinhinan N, et al.: Haplotype diversity and
linkage disequilibrium at human G6PD: recent origin of alleles that
confer malarial resistance, Science 293(5529):455462, 2001.
CHAPTER 4
Evolution can Occur

Quickly in Response to
Strong Selection
Since the early part of the 1900s, new synthetic chemicals have been used
by humans to control pests of agricultural crops and pests that transmit
diseases to humans. As the effectiveness of these chemicals was realized,
there was an exponential increase in the number of chemicals developed
by the chemical industry and used by humanity. Some of the earliest
discoveries are still in use today, although their effectiveness varies.
Dichloro-diphenyl-trichloroethane, or DDT by its more common
name, is a synthetic chlorinated hydrocarbon. The insecticidal proper-
ties of DDT were discovered in 1939, and because it was inexpensive to
manufacture, it was quickly adopted as a weapon of choice against mos-
quitoes. Female mosquitoes of the genus Anopheles can carry the parasite
that causes malaria. Female mosquitoes suck blood from other animals.
The parasite is injected with mosquito saliva. Mosquitoes that do not
carry the parasite pick it up when they suck blood from an infected
person and can later pass it on. Malaria is a common and debilitating
infectious disease and a global public health problem. The World Health
Organization (WHO) estimates there are between 300 million and
500 million cases of malaria every year. Anywhere from 1 to 2 million
deaths result from these infections annually with about 90% occurring
in Africa. Malaria is caused by a protozoan (a unicellular eukaryotic
microorganism) parasite (genus Plasmodium) and leads to anemia, fever,
nausea, flulike symptoms, coma, and death.
The incidence of malaria in many countries followed the pattern
observed in Italy after the introduction of DDT with incidences often
dropping from the hundreds of millions to zero or near zero. The United
Nations and WHO used DDT to attempt to eradicate malaria from the
planet by eliminating the mosquito vectors. Samples of mosquito larval
habitat (such as swamps, ponds, and stagnant bodies of water) initially
revealed few or no surviving mosquitoes after an area was first sprayed
with DDT. In some areas where success was thought to have occurred,
biologists began to find larval and adult mosquitoes in areas where they
were thought to have been eradicated. More aggressive spraying of DDT
followed, and in many countries, such as India, public health authorities
tracked the incidence of malaria and DDT usage, where they observed
incidence actually increases with DDT usage (Figure 11). Public health
officials worldwide studied the crisis with concern.
DDT acts by opening sodium ion channels in neurons, causing
them to fire spontaneously. This leads to spasms and eventual death.
DDT is known to have other effects on animals. In any region where
DDT was initially sprayed, the reduction in mosquitoes and malaria
cases followed the pattern observed in Italy. In India and other areas,
mosquitoes evolved resistance, an evolved ability to withstand toxin
exposure, to DDT and became unaffected by the pesticide. Their popu-
lations resurged and often malaria returned with them. Disease-carrying
animals, mainly insects, that evolve resistance to insecticides continue
to cause human suffering. Global resurgence of mosquitoes and malaria
occurred, despite continued and aggressive spraying as illustrated for
India in Figure 11. Control of mosquitoes required larger doses of DDT,
which became less effective over time, eventually becoming totally inef-
fective in some regions. Humans are forced to continually develop new
ways to control these pests.
The resistance adaptation was sought to determine how these insects
evolved resistance so quickly. Scientists looked at several detoxification
enzymes, which break down toxic chemicals that enter the body. Some of
these enzymes are specific and others are general, with the general ones
being able to use many chemicals as substrates.
Glutathione S-transferases (GSTs) are a family of related enzymes that
perform several functions. One function is to detoxify fat soluble chemi-
cals by breaking them apart and making them more water soluble. This
inactivates the toxin and makes it easier for the animal to excrete it. Each
EVOLUTION CAN OCCUR QUICKLY 43
7,000,000
malaria incidence (number of new cases) 6,000,000
5,000,000
4,000,000
3,000,000
2,000,000
1,000,000
A 1964 1966 1968 1970 1972 1974 1976 1978 1980

year
12,000
11,000
DDT usage in public health
and agriculture (tons)
10,000
9,000
8,000
7,000
6,000
5,000
4,000
B 1964 1966 1968 1970 1972 1974 1976 1978 1980
year
Figure 11 Malaria incidence (number of new cases) and DDT use in

India from 19651978.
Source: From Sharma and Mehrotra, 1982, Table 1.
of the genes that codes for one of the GSTs is slightly different, coding for
variable versions of GST.
DDT was sprayed in Tanzania from 1961 to 1967 and then again
from 1983 to 1988. It was primarily sprayed on walls of houses. The
major transmitter of the malaria parasite in Tanzania is Anopheles gambiae,
the females of which rest on walls after they take a blood meal. Resting on
walls sprayed with DDT exposes them to the chemical, and susceptible
individuals die in minutes to hours after exposure. La-aied Prapanthadara

and colleagues examined the multiple forms of GST in populations of
A. gambiae that were resistant and susceptible to DDT. Resistant indi-
viduals were collected from Tanzania, and susceptible individuals were
collected from Gambia.
Adults were allowed to reproduce, and bioassays, experiments designed
to measure the effects of a substance on a living organism, were performed
on resulting mosquito larvae. Groups of larvae were exposed to a series of
DDT concentrations for 24 hours, at which point the number of dead
and live larvae were counted. The scientists determined the DDT concen-
tration at which 50% and 90% of each test population died (Table 3).
Low concentrations mean the population is more susceptible. Much lower
concentrations of DDT killed 50% or 90% of the Gambian mosquito
population than the Tanzanian mosquito population, meaning it is more
toxic to the Gambian mosquitoes. Because the mosquitoes used were off-
spring of individuals collected in the field, there was no prior exposure of
the test population, and it was concluded that the Tanzanian population
had evolved resistance. The adaptation was passed on to the offspring.
GST was extracted from mosquitoes and pooled together. Seven
variants of GST were separated using chemical techniques. The mass
of each variant per 30 grams of pooled mosquito larvae was determined
for each population (Figure 12A). Prapanthadara and colleagues then
determined the rate at which each variant in each population detoxified
DDT (Figure 12B).
Scientists discovered that the GST family of genes was responsible for
conferring resistance to this population of mosquitoes. In the Tanzanian
population, there was a greater amount of almost every GST enzyme, and
some of those enzymes had much higher activity in the resistant than the
susceptible population. Exposure to toxic chemicals can lead to increased
Table 3 Concentration of DDT (mg/L) that causes 50%

and 90% mortality of two populations of A. gambiae.
mortality rate Gambian population Tanzanian population
50% 0.017 0.339
90% 0.040 1.160
Source: Data obtained from Prapanthadara et al., 1995, Figure 2.

25
= Gambia
20 = Tanzania
mass (mg)
15
10
0
1 2 3 4 5 6 7
A GST variant
12,000
activity of GST on DDT (nmole/min)
10,000
8,000
6,000
4,000
2,000
0
1 2 3 4 5 6 7
B GST variant
Figure 12 Results from extraction of GST from two populations of

A. gambiae, one from Gambia and one from Tanzania. (A) Mass of
each variant of GST. (B) Ability of each variant to break down DDT.
Activity for variants 1, 2, and 3 were not determined for the Gambia
population due to low levels of the enzyme.
Source: From Prapanthadara et al., 1995, Table 2.
expression of genes, as is evident for several variants, but the higher activ-
ity for the same variant in the Tanzanian population than the Gambian
population indicates a mutation changed the allele to one that detoxifies
DDT faster.
Different populations of pest insects exposed to the same pesticide
do not necessarily evolve the same mechanism of resistance. In the pre-
vious example, a change in the rate of pesticide detoxification confers
resistance, because the pesticide resides in the mosquito for a shorter pe-
riod of time. Other ways to become resistant are to adapt to detect and
avoid the chemical or to evolve an altered target site at which the pesticide
acts. A target site can be an organ, tissue, cell, or molecule. DDT binds to
the sodium channel in insect neurons. The sodium channel is the target
site and is a large multi-subunit protein that spans the cell membrane and
through which sodium ions flow to initiate a nerve impulse. When DDT
binds to this protein, the channel is held open, causing spontaneous and
inappropriate nerve impulses. Spasms occur first, and death follows if the
concentration is high enough to compromise many nerves.
Frank Collins and his colleagues used A. gambiae mosquitoes from
several populations to identify mechanisms of resistance to DDT and
another pesticide, permethrin (Ranson et al., 2000). Permethrin acts in
a manner similar to DDT, targeting the sodium channel in nerves, but
it is detoxified in insects by a different kind of enzyme. One A. gambiae
population from Kenya had been exposed to bed nets impregnated with
permethrin, another was the DDT-resistant population from Tanzania,
and a third population was a susceptible strain that had been maintained
in the laboratory through several generations. Collins and colleagues ex-
posed adults from each population to either 0.5% permethrin or 4%
DDT. One other treatment included piperonyl butoxide (PBO), which
inhibits the enzyme known to detoxify permethrin and related pesti-
cides. The time to reach 50% or 90% mortality was determined for each
population and condition (Table 4).
Table 4 Time in minutes to 50% and 90% mortality of A. gambiae

populations when exposed to a 0.5% solution of permethrin or a 4%
DDT solution. pop. = population; nd = not determined.
pesticide/ permethrin- permethrin- DDT-exposed laboratory
mortality rate exposed Kenyan exposed Kenyan Tanzanian control
population population w/ PBO population population
permethrin
50% 32 25 nd 17
90% 98 79 nd 36
DDT
50% 57 nd 145 <15
90% 115 nd >250 41
Source: From Ranson et al., 2000, Figure 1.

Given that PBO inhibits the detoxification enzyme, if PBO is added

to a treatment with permethrin, and permethrin resistance is conferred
only by this detox enzyme, then time to death should be the same as
in the control population. However, that is not what was observed by
the scientists. There is some reduction in time to death, but it is not as
short as in the control, suggesting that there is also another mechanism of
resistance at work.
In many insects, evolution of resistance to one type of pesticide con-
fers resistance to other pesticides. Pesticides that have the same target
site in the body, or are detoxified by the same enzyme, would be inef-
fective against individuals that had a mutated gene that led to an altered
target site or detoxification enzyme, respectively. It is important to know
the mechanism of resistance in resistant populations to make wise deci-
sions regarding use of pesticides. Using permethrin in Tanzania would
be advisable, because it is known from the previous study that DDT
resistance in those mosquitoes is caused by alteration of GSTases. These
are not the main enzymes that detoxify permethrin. Alternatively, use of
DDT against the permethrin-resistant mosquitoes in Kenya would be
inadvisable. Collins and his colleagues found that these mosquitoes had
two mechanisms of resistance. There was evidence that PBO inhibition
led to an increased rate of mortality and shorter time for 50% or 90%
mortality, which suggested that the detoxification enzyme was being
inhibited. But the major effect was not due to detoxification; rather it
was in an altered target site. The evidence for this was that permethrin-
resistant mosquitoes were also resistant to DDT and in fact survived
longer in presence of DDT than presence of permethrin.
Collins and colleagues went on to sequence regions of the sodium
channel gene and found that there was one point mutation that led to
permethrin-resistance in A. gambiae from Kenya. In another case of
pesticide resistance a team of French researchers documented a similar
situation where one nucleotide change in the gene coding for acetylcho-
linesterase led to the evolution of resistance (Weill et al., 2003).
Acetylcholinesterase (ACHe) is an enzyme that breaks down acetyl-
choline, a neurotransmitter primarily found in the junctions between
neurons and muscle cells. These motor neurons release acetylcholine to
activate muscles. Acetylcholine is released at the axon terminal end of a
neuron into the junction between the neuron and the muscle. It then
binds to a receptor on the muscle cell, which triggers a muscle contrac-
tion. Acetylcholine is then released and broken down by acetylcholines-
terase, which is found in the junction. After it is broken down into acetate
and choline, those two components are taken up by the axon terminal of
the neuron that secreted the acetylcholine and they are used to remake
acetylcholine. This all occurs very rapidly, as is obvious when considering
how fast muscles can contract in response to a stimulus.
Acetylcholine is also used as a neurotransmitter in the autonomic ner-
vous system, the part of the nervous system that affects the functioning of
internal organs, and regulates bodily functions such as heart rate, diges-
tion, respiratory rate, urination, and sexual arousal. As a neurotransmitter
between two neurons, it acts in much the same way as it does between
a neuron and a muscle cell, except in this case it causes generation of a
nerve impulse in the receiving neuron, as opposed to a muscle cell con-
traction. In digestion, acetylcholine helps to activate parietal cells.
Chemicals that affect parts of this system, either the acetylcholine,
the receptor, or the acetylcholinesterase, can have very damaging effects
ranging from paralysis to convulsions. For example, two classes of pesti-
cides, organophosphates and carbamates (the latter being organic com-
pounds derived from carbamic acid (NH2COOH)), block the active site
of acetylcholinesterase, inhibiting its action. The end result of this is that
acetylcholine does not get broken down in the synapse and the action of
acetylcholine is increased because degradation is delayed. Other chemicals
that do this are some nerve agents, such as Sarin. When degradation of
acetylcholine is inhibited, the resulting accumulation of acetylcholine in
the nerve synapses causes continuous stimulation of the muscles, glands,
and central nervous system, which can result in fatal convulsions if the
dose is high.
Organophosphates and carbamates are frequently used as pesticides
because of the effects they have on insect nervous systems, and while they
can be more toxic to insects, these chemicals also have potential for high
toxicity in birds and mammals, although that varies depending upon the
exact chemical formula. Because carbamates, for instance, have been used
in the environment for decades to control mosquitoes and other insect
pests, the possibility of resistance evolution is high.
The French researchers began their research by documenting popu-

lations of mosquitoes resistant to carbamates. They chose one resistant
population of Culex pipiens and one susceptible population of the same
species that had never been exposed to carbamates. Culex pipiens is called
the house mosquito because it is the most common mosquito in urban
settings in many parts of the world. It is vector of multiple diseases includ-
ing various types of encephalitis, as well as meningitis and West Nile virus.
In these two mosquito populations the researchers determined the
complete messenger RNA coding sequence for acetylcholinesterase, which
codes for the 702 amino acid enzyme. From the mRNA, the research-
ers created complementary DNA and found 27 nucleotide differences
between them, but only one of which generates a difference in the amino
acid sequence of the acetylcholinesterase protein produced by the two
populations, a change at amino acid number 119 from glycine to serine.
The researchers inserted the gene for acetylcholinesterase from both
resistant and susceptible mosquitoes into different cell lines derived from
a fruit fly (called S2 cells), resulting in two cell lines, one that produced
the resistant form of acetylcholinesterase and one that produced the sus-
ceptible form. They also took many individual mosquitoes from each
population and ground them up to create a homogenate, which is a sus-
pension of cell fragments and cell constituents obtained when tissue is
homogenized.
The researchers then exposed the homogenates (which contained
acetylcholinesterase, of course) from resistant and susceptible mosquito
populations and the cells with the acetylcholinesterase gene from the
two mosquito populations inserted into them, as well as a cell line with
no acetylcholinesterase (S2 cells) to varying concentrations of a carbamate
pesticide (Figure 13). At each concentration, the researchers measured the
ability of either the homogenate or the cell line to break down acetylcho-
line, which is a measure of acetylcholinesterase activity.
The data clearly show that acetylcholinesterase activity drops very
quickly in both the susceptible mosquito homogenate and the S2 cells
that had the susceptible acetylcholinesterase gene inserted into its genome
as the carbamate concentration increases. In fact, the responses are very
similar suggesting that it is fact the acetylcholinesterase activity that is
being affected by the carbamate. It was important to test the homogenate,
120 = R-mosq ACHe

acetylcholinesterase activity (%)
= S-mosq ACHe
100
80
60
40
20
0
A 0 1e-8 1e-7 1e-6 1e-5 1e-4 1e-3 0.01
= R-ACHe in S2
120 = S-ACHe in S2
acetylcholinesterase activity (%)
= S2 cells
100
80
60
40
20
0
0 1e-8 1e-7 1e-6 1e-5 1e-4 1e-3 0.01
B carbamate pesticide concentration (M)
Figure 13 Acetylcholinesterase activity of A, homogenates of resistant

(dark gray bars) and susceptible (gray bars) mosquitoes and B, lysates
from fruit fly cells with either the gene for resistant acetylcholinesterase
(dark gray bars), susceptible acetylcholinesterase (gray bars), or
unmanipulated fruit fly cells (light gray bars) after exposure for
15minutes to the specified concentration of the carbamate insecticide.
Source: From Weill et al., 2003, Figure 1.
which contains acetylcholinesterase from the actual mosquitoes, to show

that those mosquitoes are actually susceptible. It was also important to
isolate the acetylcholinesterase gene and insert it into the cell line to show
that it is the acetylcholinesterase that is being acted upon by the car
bamate. The no acetylcholinesterase S2 control was needed to show that
there was no unknown acetylcholinesterase or acetylcholinesterase-like

enzyme in the S2 cell line.
In comparison to the susceptible population, the data from the
resistant populations homogenate and its gene inserted into the S2 cells
clearly show that this population has a version of acetylcholinesterase
that is not susceptible to even high concentrations of carbamate. Ac-
tivity remains high across all concentrations and only drops to about
80% when carbamate concentrations are extremely high, at 0.01 M.
The homogenate and the S2 cells with the acetylcholinesterase gene
have similar responses.
The scientists next performed an evolutionary tree analysis on vari-
ous populations of C. pipiens from around the world, as well as one
population of a closely related species, C. torrentium (Figure 14). Culex
pipiens exists as different subspecies, C. pipiens quinquefasciatus, which is
found in more tropical locations, and C. pipiens pipiens, which is found
in more temperate, northern locations. Multiple populations from each
from around the world were examined for their acetylcholinesterase
gene sequence. Those sequences were used to build the tree shown in
Figure 14. The researchers knew for each population which were resis-
tant to carbamates and which were susceptible.
The researchers found many differences in the acetylcholinesterase
gene sequence among the many populations, both resistant and suscepti-
ble, but one important aspect to note is that in every resistant population
the DNA sequences had mutated to produce acetylcholinesterase with
serine instead of glycine at position 119.
What the researchers found was that the particular mutation that
leads to resistance arose independently at least twice among the C. pipiens
populations. This is evident from inspection of the evolutionary tree. This
tree shows that the resistant populations of C. pipiens pipiens all group
together on the tree, as do the susceptible ones. In addition, the resis-
tant populations of C. pipiens quinquefasciatus are all more closely re-
lated to each other than any of them are to either susceptible C. pipiens
quinquefasciatus populations or to any population of C. pipiens pipiens.
In addition, the resistant populations of C. pipiens pipiens are not more
closely related to the resistant populations of C. pipiens quinquefasciatus
than the latter populations are to susceptible populations of C. pipiens
5 resistant
populations
Culex pipiens quinquefasciatus

14 susceptible
populations
5 resistant
populations
Culex pipiens pipiens

5 susceptible
populations
Culex torrentium
Figure 14 Evolutionary tree of C. pipiens populations from around

the world belonging to two subspecies, using the acetylcholinesterase
DNA sequence. Both resistant and susceptible populations from each
subspecies were sequenced. The acetylcholinesterase sequence from
one C. torrentium population was used for comparison.
Source: Data from Weill et al., 2003, phylogenetic analysis courtesy of M. Franzen.
quinquefasciatus. Because C. pipiens pipiens and C. pipiens quinquefascia-

tus belong to the same species (just different subspecies), these mosquitoes
can interbreed. The resistant acetylcholinesterase gene could have spread
throughout populations of both subspecies after it originated the first
time, but it did not. If it had, then the pattern on the tree would look dif-
ferent. For instance, all the resistant populations, regardless of subspecies,
would have clumped together on the tree.
The resistance gene might have spread through the resistant popula-
tions of each subspecies, and that is quite possible given that they clump
together on the tree, but we do not have evidence for that either way.
Examination of the evolutionary tree should lead to the conclusion that
the resistant acetylcholinesterase evolved at least twice, and possibly as
many as 10 times in this species, meaning that the same exact mutation
evolved multiple times. The scientists who performed this research also
documented one population of Anopheles gambiae that also had the same
exact mutation in its acetylcholinesterase.
The evolution of the same mutation occurring in different popula-
tions as a response to the evolutionary selective factor of a pesticide
is quite amazing. Scientists could not predict that evolution would
work in that exact manner, given what is known about the processes of
mutation and natural selection. For this to happen, the exact same
random mutation of the acetylcholinesterase gene had to occur in each
population. Many other mutations also occurred, but that is the one
that conferred resistance to the population when exposed to carbamate
pesticides. Given enough time, large mosquito populations, and high
exposure to carbamates, an understanding of evolutionary processes
should allow one to conclude that this can, and does, happen. It is still
quite amazing, though.
The mutations examined in this chapter all led to variation among
individuals in their respective populations. When those individuals
were exposed to DDT, permethrin, or carbamates, those without the
mutation were killed. Individuals with the mutation passed it on to
their offspring, increasing the proportion of individuals in the popula-
tion with the mutation. This is natural selection in action, occurring
in a period of time short enough for us to observe. Further evidence
of populations of A. gambiae in Africa indicates enough isolation
such that mating of individuals from different populations does not
occur. Scientists studying these populations suspect that these popula-
tions are diverging into new species in front of our eyes. Evolution of
species in the context of their environment, including in the context
of other species, is the subject of another book in this series, Evolution
of Interactions in Communities.
Ethical, Legal, Social Implications:

Overuse of Chemicals like Pesticides and
Antibiotics can Have Detrimental Effects
Discoveries of new antibiotics, substances or compounds that kill or
inhibit the growth of bacteria, and pesticides have saved millions of lives.
Antibiotics have cured people with deadly bacterial infections, and pesti-
cides have prevented transmission of deadly diseases. Decreasing preva-
lence of disease has increased the health and quality of life for many
people.
Uses of both kinds of chemicals have increased in recent decades, but
too much of a good thing can have adverse effects. Pests resistant to pesti-
cides can re-emerge as an economic threat to a crop. In the United States,
insecticide resistance forces farmers to spend an additional $40 million
annually to apply additional controls. The annual costs of crop losses may
be in the tens of millions of dollars and have caused complete collapses
of some crops and the economies that sustain them. These problems arise
due to pesticide misuse and overuse.
The introduction of inorganic pesticides after World War II
increased crop yields dramatically. The pesticides were cheap and effec-
tive and were considered the solution to feed the worlds growing human
population. Humans would be able to rid themselves of crop pests and
disease-carrying insects. However, within a short time, dozens of insect
species had evolved resistance to pesticides. It is now estimated that over
1,000 species of insects and hundreds of weed species are resistant to at
least one type of pesticide.
The scientists who developed these chemicals probably looked
upon themselves as saviors of a sort. They were almost certainly act-
ing with good intentions to solve world hunger and raise people out
of poverty. They might argue that millions of lives were saved by these
chemicals. Yet, someone else might argue that overuse of chemicals
designed to kill has had adverse effects. There are documented cases
of population decline, local extinction, and evolutionary changes in
non-target species. Low concentrations of some pesticide residues
have been shown to mimic hormones, affecting animal and human
health.
Just as insects have evolved resistance to insecticides, bacterial patho-

gens have evolved resistance to antibiotics. Worldwide use of penicillin
began in the 1940s, and more than 150 other antibiotics have been dis-
covered. It is hard to estimate the countless lives saved by the use of these
drugs against diseases like tuberculosis, tetanus, typhoid fever, diphtheria,
and cholera. As with pesticides, antibiotics have been misused and over-
used. The evolution of resistance to penicillin was observed only a few
short years after mass production began. The first bacterium to resist pen-
icillin was Staphylococcus aureus, which is often harmless, but can cause
pneumonia. Today, antibiotic resistance is a global health crisis. Strains of
nearly all important bacteria in the world are resistant to commonly used
antibiotics.
As with any population, individual bacteria exhibit variability. Some
individuals, even at low frequency, are able to withstand an antibiotic.
When a patient takes an antibiotic, the drug selects for individuals that
are mildly affected or unaffected by the antibiotic. These bacteria in-
crease in proportion relative to susceptible individuals, and this can
occur quickly. The penicillin-resistant strain of Pneumococcus increased
from 0.02% in the early 1980s to 6.6% of the population by 1994. Bac-
teria may acquire resistance through mutation or exchanges of genes. A
resistance gene that evolves in one species can even transfer to another
species!
The biology of bacteria facilitates the evolution of resistance but so
can misuse of antibiotics. One misuse is not taking them as prescribed.
Patients prescribed an antibiotic for a legitimate reason must take the
entire course of medication to kill all the harmful bacteria. A patient that
discontinues use when he begins to feel better often wipes out only the
most susceptible bacteria, while allowing survival of mildly resistant bac-
teria. Resistant bacteria then become widespread, leading to infections
that are more difficult to treat. This is a problem of individual underuse,
but overuse and over-prescribing are also problems.
Antibiotics are often prescribed by doctors for illnesses that are viral.
Antibiotics are ineffective against viruses, and they should not be used for
colds or the flu. Patients often pressure doctors for antibiotics and may
not understand that they will not work for these diseases, but doctors
who relent to patients demands perpetuate antibiotic misuse. Antibiotic
resistance may be selected for in other bacteria in the patients body, lead-
ing to trouble in the future, as when a beneficial bacteria exchanges genes
with a pathogenic bacteria and provides it with a resistance gene.
Superbugs have emerged due to the continued patterns of misuse
and overuse of antibiotics. When faced with a bacterium resistant to a
first choice antibiotic, doctors often prescribe a stronger antibiotic, but
the bacteria may evolve resistance to the more potent drug as well, perpet-
uating a cycle in which increasingly powerful drugs are required to treat
infections. Today, one in seven new tuberculosis cases are resistant to the
two drugs most commonly used to treat it, and 5% of these patients die.
For years, the potent antibiotic, vancomycin, was a reliable last line
of defense against certain severe infections, notably those caused by
Staphylococcus. But in recent years, some superbugs have become resis-
tant to even vancomycin, including a bacterium known as Enterococcus.
The fear now is that the gene conferring resistance to vancomycin will
spread to other strains and species of bacteria. This resistant strain of
Enterococcus has already appeared in hospitals in several countries.
Patients with resistant bacterial infections may be ill for longer
periods of time with increased risk of complications and death. Costs
to individuals include increased hospital stays and more expensive tests
and treatments. People die each year of infections that they contract in
hospitals from antibiotic-resistant bacteria. The failure of first line anti-
biotics means that doctors must resort to other medications. The antibi-
otics needed to treat multidrug-resistant forms of tuberculosis are very
expensive, have toxic side effects, and may need to be taken for months.
Some patients may be denied the medicine that they need because of
their socioeconomic status; they may be too poor to pay for it or have no
health insurance.
Other costs go beyond the individual. Costs to society include
higher health insurance premiums, the need for society to treat unin-
sured or underinsured individuals, loss of income and worker productiv-
ity, and allocation of government resources to monitor the problems.
Although experts are working to develop new antibiotics to keep pace
with antibiotic-resistant strains of bacteria, infectious organisms adapt
quickly. Some strategies that have been advocated to slow the evolu-
tion of resistance include better hygiene and education of the healthcare
community and faster identification and isolation of patients with

antibiotic-resistant infections. Antibiotic-resistant bacteria will continue
to be a global health concern, and using antibiotics wisely is an impor-
tant part of preventing their spread.
Bibliography
Breiman RF, Butler JC, Tenover FC, et al.: Emergence of drug-
resistant pneumococcal infections in the United States, J Am Med
Assoc 271(23):18311835, 1994.
Casida JE, Quistad GB: Golden age of insecticide research: past, present,
or future? Annu Rev Entomol 43:116, 1998.
Lewis R: The rise of antibiotic-resistant infections, FDA (website): http://
www.fda.gov/Fdac/features/795_antibio.html. Accessed June 18, 2014.
Prapanthadara L, Hemingway J, Ketterman AJ: DDT-resistance in
Anopheles gambiae (Diptera: Culicidae) from Zanzibar, Tanzania,
based on increased DDT-dehydrochlorinase activity of glutathione
S-transferases, Bull Entomol Res 85:267274, 1995.
Ranson H, Jensen B, Vulule JM, et al.: Identification of a point muta-
tion in the voltage-gated sodium channel gene of Kenyan Anopheles
gambiae associated with resistance to DDT and pyrethroids, Insect
Mol Bio 9(5):491497, 2000.
Reimer L, Fondjo E, Patchok S, et al.: Relationship between kdr mutation
and resistance to pyrethroid and DDT insecticides in natural popu
lations of Anopheles gambiae, J Med Entomol 45(2):260266, 2008.
Sharma VP, Mehrotra KN: Return of malaria, Nature 298(5870):210,
1982.
Unicef/World Health Organization: 2005 WHO world malaria report
(PDF online): http://apps.who.int/iris/bitstream/10665/43213/1/
9241593199_eng.pdf. Accessed September 25, 2015.
Conclusion
Genetic changes over time create lines of descent among lineages of
organisms. Variation among individuals that occurs at the genetic level
leads to descent with modification and speciation. Natural selection
acts on individuals and leads to changes in populations. Those changes
in populations can lead to speciation events if populations are isolated
from one another and enough genetic changes accumulate. Species are
dynamic entities as Sewall Wright envisioned many years ago. Wrights
critical insights into the concept of a population allowed scientists to con-
ceive of theories and experiments to illuminate how evolutionary change
is brought about.
Many speciation events over a short period of evolutionary time
within a lineage constitute an adaptive radiation, and these radiations
begin with variation among individuals. The evolution of plants that can
live on land led to entirely new communities on the planet, and those
communities are the ancestors of modern day terrestrial plant commu-
nities. These organisms are linked by lines of descent. Life continues to
evolve as the environment changes, and humans are a major contributor
to changes in our twenty-first century environment.
Glossary
adaptive radiation. An adaptive radiation is a rapid evolutionary diversification
characterized by an increase in the number of species in a lineage.
ancestors. Early type of organism from which other species have evolved.
anther. the part of the flower that produces pollen.
antibiotics. Antibiotics are substances or compounds that kill or inhibit the
growth of bacteria.
bioassay. A bioassay is an experiment designed to measure the effects of a sub-
stance on a living organism.
bryophytes. A group of small flowerless plants that include liverworts, mosses,
and hornworts.
coevolution. Coevolution is reciprocal natural selection of two or more species
where each species acts as a selective agent on at least one other species for traits
specific to their interaction.
column. In orchids, the column contains both the anther with the pollen, and
the stigma.
common ancestor. A common ancestor is one from which a group of related
species has evolved.
derived. A derived phenotype is one that has undergone change from the ances-
tral condition.
descendants. Species or organisms descended from a particular ancestral species
or organism.
differentiation. Differentiation is development of a cell into a particular cell type.
eugenics. The study or practice that deals with improving the human race
through selective breeding.
evolution. the scientific explanation for the origin of life and its continual change
over time.
evolutionary tree. An evolutionary tree is a diagram that shows the evolutionary
relationships among various species.
extinction. the permanent loss of a species from the planet when no individuals
of that species are left living.
family. A family is a taxonomic group of related organisms that includes all sub-
families, genera, and species that evolved from a common ancestor.
genera. Genera, or genus (singular), refers to the taxonomic group above the
species level.
genes. A distinct sequence of nucleotides forming part of a chromosome, which
determines the order of monomers in a polypeptide or nucleic acid molecule
which a cell may synthesize.
62 GLOSSARY
Hardy-Weinberg equilibrium. Principle that states that allele and genotype fre-
quencies in a population will remain constant from generation to generation in
the absence of evolutionary mechanisms acting upon the population.
heterozygous. Heterozygous individuals have two different alleles of a gene.
hominids. Animals, specifically primates of Family Hominidae that includes hu-
mans and their fossil ancestors.
hornworts. A small semi-aquatic plant with narrow forked leaves that become
translucent and horny as they age.
lineage. A lineage is a line of descent.
liverworts. A small flowerless green plant with leaf-like stems or lobed leaves, oc-
curring in moist habitats and lacking true roots.
mosses. A small flowerless plant that lacks true roots, grows in low carpets in
damp habitats and reproduces using spores released from stalked capsules.
natural selection. Adaptive mechanism by which evolution takes place and is
often summarized as differential survival and reproduction.
pedigrees. The recorded ancestry of an organism.
phenotype. Phenotype is the way an organism appears or behaves due to its
genetic makeup.
phloem. Phloem is tissue that transports organic compounds from photosynthe-
sizing tissues to rest of the plant.
pollen. A particle or grain that contains a male gamete discharged from the male
part of a flower or a male cone.
population. A population is a group of individuals of the same species living in
the same place at the same time
population genetics. Study of the distribution and change in frequency of alleles
and genotypes within populations.
proliferation. Proliferation is rapid cell division.
protozoan. A protozoan is a unicellular eukaryotic microorganism.
radioisotope dating. Radioisotope dating is a technique that compares the
observed abundance of a naturally occurring radioactive isotope and its decay
products, using known decay rates.
relative dating. Relative dating is a technique that uses layers of rocks and the
fossils contained within to estimate the order of prehistoric events.
resistance. Resistance is an evolved ability to withstand exposure to a toxin.
speciation. The formation of new species through any evolutionary mechanism.
species. A group of organisms consisting of similar individuals capable of
exchanging genes or interbreeding; the natural taxonomic unit, ranking below a
genus and denoted by a Latin binomial.
stigma. The part of the pistil upon which pollen lands and germinates.
subfamily. A subfamily is a taxonomic group of related organisms ranking
between a family and a genus.
vascular tissue. Vascular tissue is the supportive and conductive tissue in plants.
xylem. Xylem is supporting and water-conducting tissue, which brings water and
nutrients from the roots to the rest of the plant.
Index
Acetylcholine, 4748 Collins, Frank, 46
Acetylcholinesterase (ACHe), 4748 Common ancestors, 1
important aspect of, 51 Cox2, 21
RNA coding sequence for, 4950 Culex pipiens, 49
Adaptive radiation, 2223 as different subspecies, 51
definition of, 5 mutation of, 5152
of orchids, 39 Cypripedioideae, 9
Adult lactase activity, 39
Algae, types of, 20 Darwin, Charles, 34
Anopheles gambiae, 43 Degeneracy theory, 35
Antibiotic-resistant bacteria, 5657 Descendants
Antibiotics, over use of, 5457 with modification and adaptive
Ape lineage, 30 radiations, 13
Apostasioideae, 9 orchids, adaptive radiation of, 39
Aquatic ancestors, terrestrial plants rapid diversification in bats, 915
from, 1723 Detoxification enzymes, 42
Australopithecus africanus, 31, 3233 Dichloro-diphenyl-trichloroethane, 41
as opening sodium ion channels, 42
Bats in Tanzania, 4344
developing cartilage at the cellular DNA, sequences of orchids, 3
level, 14
proliferation and differentiation Embryonic bat forelimb cells, 14
of, 1213 Enterococcus, 56
rapid diversification in, 915 Epidendroideae, 6, 9
Bioassays, 44 Epiphytes, 6
Bipedal locomotion, 3031 Episodic flooding, 29
Bone morphogenetic proteins Eugenics, 3435
(BMPs), 14 downfall of, 36
Brunet, Michel, 2526 recent acts of genocide, 37
Bryophytes, 19, 20 in United States, 36
types of, 21 Evolutionary tree, 13
analysis on populations of C.
Canines, 2728 pipiens, 51, 52
Carbamates, 48 green algae, 22
Carollia perspicillata, 12 of plants, 21
Chad fossil relationships among birds, bats, and
analysis of, 28 insects, 1011
brain capacity of, 27
face of, 27 Female mosquitoes, 41
Climbing mammals, 33 Fifth metacarpal bone, 1112
Coevolution, of plants and Fossils
animals,17 bats, skeletons of, 1112
64 INDEX
brain volume ranges from, 28 Malaria

discovery of, 25 incidence of, 4142
fragments of plants, 1819 resistance, 39
freshwater fish, 2930 Meliorchis caribea, 3, 5
in rock, 18 Mendel, Gregor, 34
as shore habitat, 18 Metacarpal bone, fifth, 1112
skull measurements for unknown Mosquitoes, global resurgence of, 42
hominid, 27 Mosses, 19, 20
spores, shape and size of, 19 Mus musculus, 12
Fruit fly. See S2 cells Mutations
of Culex pipiens, 5152
Genetic drift, 39 individuals with, 53
Glutathione S-transferases
(GSTs),4243 Nad1, 21
exposure to toxic chemicals, 4445 Natural selection, 37
seven variants of, 44 Negative eugenics, 35
Gravendeel, Barbara, 7 Neurotransmitter
Green algae, evolutionary tree, 22 in autonomic nervous system, 48
Niswander, Lee, 11
Hardy-Weinberg equilibrium, 35
Hominid ancestors, humans Orchidoideae, 9
from, 2534 Orchids, adaptive radiation
ethical, legal, social of, 39
implications,3437 Organophosphates, 48
pinnacle and endpoint of The Origin of Species, 34
evolution,3739
Homo erectus, 33 Paranthropus genus, 31, 33
Homo sapiens, 25 Parasite, 41
Hornworts, 19, 20 Penicillin, use of, 55
House mosquito. See Culex pipiens Permethrin, 46
Human Genome Project, 36 in Tanzania, 47
Human lineage, 30 Pesticides
Humans detoxification of, 4546
from hominid ancestors evolution of resistance to, 47
inAfrica,2534 organophosphates and carbamates
ethical, legal, social implications, as, 48
3437 two classes of, 48
pinnacle and endpoint of over use of, 5457
evolution, 3739 Phenotypic characters, for
variation among, 34 orchids,45
Phloem, 21
Icaronycteris index, 11 Pierce, Naomi, 3
Intelligence quotient (IQ), 35 Piperonyl butoxide (PBO), 4647
Introns, in liverworts, 22 Pneumococcus, 55
Pollens, 7
Land plants, 1920 Pollination specialization, 7
Liverworts, 19, 20 Population genetics, 35
introns in, 22 Positive eugenics, 35
Longer digits, hypotheses regarding Proliferation, 12
selection for, 12 Protozoan, 41
INDEX
65
Racial prejudice, 34 Shale, 18

Radioisotope dating, 29 Sporopollenin, 19
Rapid diversification Staphylococcus aureus, 55
in bats, 915 Subfamily, 3
Red algae, 22 Superbugs, 56
Reed, Kaye, 32
Relative dating method, 29 Terrestrial plants, from aquatic
Resistance gene, 53 ancestors, 1723
Tree-dwelling mammals, 33
S2 cells, 4951
Sahelanthropus tchadensis Vancomycin, 56
to H. sapiens, 32 Vanilloideae, 9
skull of, 26, 28 Vascular tissue, 20
Sandstone, 18
Sarin, 48 Xylem, 2021
OTHER TITLES IN OUR BIOLOGY
COLLECTION
Behavior and Information Exchangeby Christopher J. Paradise and A. Malcolm Campbell

Cells in Tissuesby Christopher J. Paradise and A. Malcolm Campbell
Ecological Dynamicsby Christopher J. Paradise and A. Malcolm Campbell
Evolution of Interactions in Communitiesby Christopher J. Paradise and
A.Malcolm Campbell
Effects of Genetic and Pathogenic Diseases on Cellsby Christopher J.Paradise and
A.Malcolm Campbell
Information in the Environmentby Christopher J. Paradise and A. Malcolm Campbell
Mechanisms of Evolutionby Christopher J. Paradise and A. Malcolm Campbell
Properties in and of Populationsby Christopher J. Paradise and A. Malcolm Campbell
Variation and Population Geneticsby Christopher J. Paradise and A. Malcolm Campbell
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
Announcing Digital Content Crafted by Librarians

Momentum Press offers digital content as authoritative treatments of advanced engineering
topics by leaders in their field. Hosted on ebrary, MP provides practitioners, researchers, faculty,
and students in engineering, science, and industry with innovative electronic content in sensors
and controls engineering, advanced energy engineering, manufacturing, and materials science.
Momentum Press offers library-friendly terms:

perpetual access for a one-time fee
no subscriptions or access fees required
unlimited concurrent usage permitted
downloadable PDFs provided
free MARC records included
free trials
The Momentum Press digital library is very affordable, with no obligation to buy in future
years.
For more information, please visit www.momentumpress.net/library or to set up a trial in the
US, please contact mpsales@globalepress.com.
EBOOKS Evolutionary History
FOR THE Christopher J. Paradise A. Malcolm Campbell
APPLIED BIOLOGY COLLECTION
This book describes how evolutionary history is studied using
SCIENCES
several well-known examples and also using evolutionary trees.
LIBRARY Evolutionary trees are analyzed and used to explain adaptive
Create your own radiations of orchids and the diversification of bats over geo-
Customized Content logic time. Evolutionary trees and genetic evidence is used to
infer when and from what ancestors terrestrial plants evolved
Evolutionary
Bundlethe more
and invaded land. Specific adaptations of early land plants led
books you buy,
to the evolution of terrestrial plants and their success on land.
the greater your
History
Evidence about the ancestors and habitats of humans is used
discount!
to infer and analyze the evolution of the human family tree,
whose populations were subject to the same forces of evolution
THE CONTENT to which other species are subject. Human evolution was not
Energy Physics linear, involved offshoot species that did not survive, and took
Engineering many thousands of years. In contrast, evolution can be seen in
just a few years or less in other examples, and analysis of the
Biotechnology
evolution of mechanisms of pesticide resistance in insects will
Biology
be used to illustrate this rapid evolution.
Mathematics
Chemistry Christopher J. Paradiseis professor of biology and environ-
mental studies at Davidson College. He teaches introductory
biology, ecology, entomology, and topical seminars on ecotoxi-
THE TERMS
cology and renewable natural resources. He also occasionally
Perpetual access
leads a study abroad program in India. His research evaluates
for a one time fee anthropogenic factors that influence insect biodiversity at a
No subscriptions or variety of scales. His current research interests include effects of
access fees land use patterns on pollinator communities in parks.
Unlimited
A. Malcolm Campbellteaches biology at Davidson College,
concurrent usage
NC. He received national and international education awards:
Downloadable PDFs
Genetics Society of America (2013); American Association for the
Free MARC records
Advancement of Science (2012); and American Society for Cell
Biology (2006). He was the founding co-editor in chief of CBE Life
For further information,
Sciences Education; founding director of Genome Consortium
a free trial, or to order,
contact: for Active Teaching (GCAT); and member of the American Soci- Christopher J. Paradise
sales@momentumpress.net ety for Cell Biology governing council (20122014).
A. Malcolm Campbell

Web PDF

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Web PDF

Uploaded by

Copyright:

Available Formats

EBOOKS Evolutionary History

FOR THE Christopher J. Paradise A. Malcolm Campbell

Christopher J. Paradise, PhD

All rights reserved. No part of this publication may be reproduced, stored

First published in 2016 by

ISBN-13: 978-1-60650-965-4 (print)

Momentum Press Biology Collection

Cover and interior design by S4Carlisle Publishing Services Private Ltd.,

Printed in the United States of America

Descent with Modification

Although we cannot go back in time to observe an ancestral species,

mosses ferns pines, spruce, fir flowering plants

plant common ancestor

Figure 1 Evolutionary tree of major groups of plant species.

studied. For instance, in Figure 1, all mosses are grouped together at

Adaptive Radiation of Orchids

Based on characteristics of the pollen and how it was attached to the

Table 1 Phenotypic characters for several orchids listed by genus.

Source: Data from Ramirez et al., 2007, supplemental materials.

15 to 20 million years ago (MYA). Once they constructed phenotypes of

In their evolutionary analysis, the scientists also included DNA

gaining protection through a strong attachment to branches or trunks

# of genera having x species 30

Figure 2 Frequency distributions of orchids (A) and non-orchid

From a study of orchids and their pollinators, the scientists exam-

diverse subfamily, Apostasioideae, has 17 known species and an average

Rapid Diversification in Bats

insects birds bats

common animal ancestor

Figure 3 Evolutionary relationships among birds, bats, and insects.

Figure 4 The percentage of growing third to fifth forelimb

Figure 5 Percentage of cells in the elongation and

In this section, evolution of individuals over the course of millions of

Terrestrial Plants Evolved

latter could be limited to terrestrial plants. All organisms require water,

that actually produced the spores. The fragments consisted of sections of

Figure 6 Bryophytes. (A) The liverwort is approximately 3 to 4 cm

very diverse and the hundreds of thousands of species are hypothesized to

evolutionary taxonomic typical intron

1 = original of land plants (~475 mya)

Figure 7 Evolutionary tree of plants and presence or absence of three

plant. Phloem is tissue that transports organic compounds from photosyn-

piece together a fairly good, if a bit speculative, picture of the attributes

Humans Evolved From

The major mechanisms of evolution, mutation, natural selection, gene

Figure 8 Skull of Sahelanthropus tchadensis discovered in Chad.

Table 2 Skull measurements for unknown hominid fossil and several

Figure 9 Estimates of brain volume ranges from fossil evidence of a

they also concluded that S. tchadensis lived in an area that contained

Au. afarensis Au. africanus

miocene pliocene pleistocene halocene

Figure 10 Evolutionary reconstruction of hominids; the human

Ethical, Legal, Social Implications:

Ethical, Legal, Social Implications: Evolution has not

hominid evolution, selection could just as easily favored a solitary exis-

If all human populations were connected genetically and diseases

Evolution can Occur

malaria incidence (number of new cases) 6,000,000

A 1964 1966 1968 1970 1972 1974 1976 1978 1980

Figure 11 Malaria incidence (number of new cases) and DDT use in

individuals die in minutes to hours after exposure. La-aied Prapanthadara

Table 3 Concentration of DDT (mg/L) that causes 50%

Source: Data obtained from Prapanthadara et al., 1995, Figure 2.