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
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leads a study abroad program in India. His research evaluates
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A. Malcolm Campbellteaches biology at Davidson College,
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A. Malcolm Campbell
Evolutionary History
Evolutionary History
10 9 8 7 6 5 4 3 2 1
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
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.
35
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
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
b c
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
12 EVOLUTIONARY HISTORY
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).
DESCENT WITH MODIFICATION AND ADAPTIVE RADIATIONS 13
= 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
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
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.
DESCENT WITH MODIFICATION AND ADAPTIVE RADIATIONS 15
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
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
18 EVOLUTIONARY HISTORY
A B C
(bryophytes)
nonvascular
plants
1 hornworts
land plants
mosses
other vp
seedless
vascular
plants
vascular plants
2 ferns
conifers
plants
seed
3
flowering plants
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
A B
C D
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
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
28 EVOLUTIONARY HISTORY
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)
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.
Humans Evolved From Hominid Ancestors in Africa 29
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,
30 EVOLUTIONARY HISTORY
(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
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
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
Humans Evolved From Hominid Ancestors in Africa 33
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
34 EVOLUTIONARY HISTORY
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.
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.
36 EVOLUTIONARY HISTORY
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
Humans Evolved From Hominid Ancestors in Africa 37
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.
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evolution (translated by Gomme R), Malden, MA, 2006, Blackwell
Publishing.
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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,
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Lieberman DE, McBratney BM, Krovitz G: The evolution and devel-
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99(3):11341139, 2002.
Pickford M: Orientation of the foramen magnum in late Miocene to
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of the Upper Miocene Toros-Menalla hominid locality, Chad, Nature
418(6894):152155, 2002.
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dimorphism, Current Anthropology 17(4):579, 1976
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2002.
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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
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
42 EVOLUTIONARY HISTORY
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
5,000,000
4,000,000
3,000,000
2,000,000
1,000,000
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
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
44 EVOLUTIONARY HISTORY
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
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
46 EVOLUTIONARY HISTORY
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).
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.
EVOLUTION CAN OCCUR QUICKLY 49
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)
5 resistant
populations
5 resistant
populations
Culex torrentium
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.
54 EVOLUTIONARY HISTORY
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
EVOLUTION CAN OCCUR QUICKLY 57
Bibliography
Breiman RF, Butler JC, Tenover FC, et al.: Emergence of drug-
resistant pneumococcal infections in the United States, J Am Med
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Casida JE, Quistad GB: Golden age of insecticide research: past, present,
or future? Annu Rev Entomol 43:116, 1998.
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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
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1982.
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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
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A. Malcolm Campbell