Open Questions: Evolutionary Theory
Molecular evolution --
Evolutionary psychology --
Evolutionary history of animals --
Evolutionary milestones --
Nothing in biology makes sense except in the light of evolution.
There is not a single Why? question in biology that can be answered
adequately without a consideration of evolution.
Ever since Darwin's Origin of Species was published in 1859, his
theory of evolution has generated vigorous controversy -- not because it
was obviously wrong, but, certainly, because for all of its originality
and power, it left so many tantalizing questions unanswered.
The theory of evolution is a prime example of an "emergent" sort of theory,
in which the phenomena it describes and explains are treated on the
basis of principles which are formulated directly in terms of the phenomena
which are to be explained, in this case, the historical origins of living
things on Earth.
Although this seems like a reasonable and proper way
to proceed, and the theory of evolution itself was a brilliant and
successful development, it has suffered from one nagging weakness:
Insofar as the theory is founded on observation of the available
evidence, many key facts will be forever outside our reach, simply
because they relate to events in the distant past which we can never
actually observe. All we have to work with are the living things which
we can observe today, and a few haphazard fossil remains of living things
which flourished millions or billions of years ago.
In short, although we can often make shrewd guesses as to why things
have turned out the way they seem to have, our conclusions often remain
just guesses. We frequently find ourselves wanting key pieces of evidence
which would confirm or disconfirm the guesses.
As a result, controversies remain, for lack of the evidence which could
settle them conclusively beyond a reasonable doubt.
For example: Is evolution generally a smooth and continuous process, with
one animal species changing slowly and almost imperceptibly into one or
more descendent species? Or does it, instead, more often involve sudden,
discontinuous changes -- perhaps provoked by radical environmental changes --
with species otherwise remaining essentially unchanged for millions of
years? It's usually difficult to determine this from the fossil record,
given the few samples we frequently must deal with, where the distances
between their origins in time and space are both large and not very certain.
Given only a few
bones, how much can we really say about the relationship between the
animals they were once a part of?
It would be a tremendous help in resolving such questions if we had
available entirely different sorts of evidence to work with. And now,
half a century after the determination of the structure and function of
DNA in cells, we do.
It should be cause for serious reflection that Darwin was as successful
as he was in formulating his theory, given that he knew almost nothing
of the actual science of genetics which controls how living creatures
pass on their characteristics from one generation to the next. And yet,
the fact that this inheritance takes place is absolutely key to the
theory. Astonishingly enough, Darwin knew nothing of Mendelian genetics,
and in fact he believed in the Lamarckian idea of the inheritance of
This is a tribute to what can be done with a non-reductionistic theory
in the hands of someone as brilliant as Darwin. And yet, there is so
much more that can be done, so many unresolved puzzles which can be
solved, once we have learned enough to reduce many of the known facts of
evolution to the more fundamental insights painstakingly acquired
through the science of molecular biology.
Open Directory Project: Evolution
- Categorized and annotated links. A version of this
list is at
Google, with entries sorted in "page rank" order. May also be
Unoficial Stephen Jay Gould Archive: Links
- Relatively short but carefully selected list of links
related to Gould and evolution.
Evolution and Behavior
- Many external links on evolutionary theory in general, by
Sites of Interest to Botanists, Ecologists, and Evolutionary
- External links collected by Kent Holsinger.
Biology Links: Evolution
- From the Harvard
Department of Molecular and Cellular Biology.
Evolutionary Biology Resources
- Good list, orgainized into ten categories.
- Categorized site directory. Entries usually include
Sites with general resources
UCMP Exhibit Halls: Evolution
- Information on the
theory of evolution and
history of evolutionary thought. Part of the University of
California at Berkeley's
Museum of Paleontology
The Talk.Origins Archive
- Excellent general collection of resources on evolutionary
New Scientist Special Report on Evolution
- Primarily offers links to many news articles from the past
several years of New Scientist magazine. But there are
also other features, including facts and figures, external links,
frequently-asked questions, and a short bibliography.
- Also known as the Evolution Education Wiki -- "a free, reader-built
encyclopedia of evolution, biology, and origins. It is inspired
Talk.Origins Archive and the
Evolution Resources From the National Academies
- "This Web page is designed to provide easy access to books,
position statements, and additional resources on evolution
education and research."
The World of Richard Dawkins
- An unofficial Web site dealing with the work of evolutionary
theorist Richard Dawkins. It contains a great deal of information
and external links for many controversial topics in evolutionary
The Unofficial Stephen Jay Gould Archive
- Site contains a wide variety of material related to Gould.
This includes an extensive online
library of articles and papers on evolution by Gould
and others, a
bibliography of books and papers by Gould, and a very good
selected external links on evolution.
Niles Eldredge Evolutionist
- Site contains a wide variety of material related to Eldredge.
This includes a
"Library of Evolution", consisting of many of Eldredge's
papers on evolution, and
information on books by Eldredge.
Evolution and firstname.lastname@example.org
- A portal to relevant Nature Publishing Group resouces in the
field of evolution and ecology.
BBC Evolution Website
- Large collection of educational material related to
the history and content of evolutionary theory.
Includes a very good
bibliography of both print and online reference material.
- The site is intended for science teachers, to explain
some of the common misconceptions about evolution in order to
help students understand it more easily. Provides a good basic
introduction to the subject, including good external
readings and resources.
- Material from a PBS series. Includes a large number of
articles, interviews, videos, recommended readings, and links.
Best accessed through the
site map or the
frequently asked questions.
Surveys, overviews, tutorials
Category: Evolutionary biology
- Topic category from
- Article from
Modern evolutionary synthesis,
List of evolutionary biology topics.
- Article from
Nothing in Biology Makes Sense Except in the Light of Evolution
- An important and well-known article published in 1973 by
Theodosius Dobzhansky. It provides both a concise overview of
evolutionary theory, and the facts and reasoning which make it
Genetics and Evolution Tutorial
- Good tutotrial, covering genetics as well as evolutionary
theory, located at the
Biology Online site.
- Good overview lecture by Eugenie Scott, at the
- Single-page overview from
Kimball's Biology Pages.
Evolution: Fact AND Theory
- General page on evidence for evolution. Includes
Notes on Evolution -- class notes from a course.
- April 2006 online article in Seed Magazine - an interview
with Ricardo Azevedo, who has developed computer models of
evolution, about the evolutionary importance of sex.
The Woodstock of Evolution
- June 2005 Scientific American In Focus article, subtitled
"The World Summit on Evolution, held in the Galapagos Islands,
revealed a science rich in history and tradition, data and theory,
as well as controversy and debate."
- October 2000 Scientific American news article about
an example of very rapid speciation.
Sex and Speciation
- September 2000 Scientific American news article about
how different mating systems in insects affect the rate of
Score One for Punk Eek
- July 1996 Scientific American Explore article about
an experiment involving the evolution of bacteria which claims
to provide support for the theory of punctuated equilibrium.
Sexual conflict pushes species making
- September 2000 news article in Science News, about
speciation in inssects due to sexual conflict.
Should Evolutionary Theory Evolve?
The Scientist, January 2010
- Some biologists are calling for a rethink of the rules of
A Most Private Evolution
Science News, February 14, 2009
- Dumb designs for sex: Evolutionary biology walks on the weird side.
Science News, January 16, 2009
- Scientists sift through genetic data sets to better map
twisting branches in the tree of life.
- Darwin's Living Legacy--Evolutionary Theory 150 Years Later
Scientific American, January 2009
- Evolutionary theory has broadened and changed as Darwin's
ideas have been melded with genetics. Evolutionary biology still
must contend with some of the same questions that preoccupied
Darwin: What, for one, is a species?
- Testing Natural Selection with Genetics
H. Allen Orr
Scientific American, January 2009
- Biologists working with the most sophisticated genetic
tools are demonstrating that natural selection plays a greater
role in the evolution of genes than even most evolutionists
- What Is a Species?
Scientific American, June 2008
- Formal taxonomic systems first identified species based
on visual traits such as fins or fur. Later, the species
concept changed, specifying that two organisms should be
capable of breeding.
- Founder Mutations
Scientific American, October 2005
- Was Darwin Wrong?
National Geographic, November 2004, pp. 2-35
- Shows and explains, with many illustrations, that the evidence
for evolution is overwhelming.
- Evolutionary Shocker?
Science News, June 22, 2002, pp. 394-396
- The protein known as heat shock protein 90 may assist
evolution by allowing genetic changes to remain unexpressed
until an organism experieces stress, such as elevated temperature.
Alarming Butterflies and Go-Getter Fish
Science News, July 21, 2001, pp. 42-44
- Additional conditions that lead to the emergence of new species
continue to be discovered.
- Evolution: A Lizard's Tale
Jonathan B. Losos
Scientific American, March 2001
- The Mother of Mass Extinctions
Douglas H. Erwin
Scientific American, July 1996, pp. 72-78
- Catastrophic changes in the Earth's environment 250 million
years ago caused the largest episode of mass biological extinction
in history. About 90% of ocean-dwelling species, for example,
disappeared, but the exact evolutionary factors that account for
which species survived and which didn't is not clear.
- Ernst Mayr -- What Evolution Is
Basic Books, 2001
- Mayr is one of the 20th century's
most outstanding contributors to evolutionary theory and
began publishing (on ornithology) in 1923. The present book is
a detailed introduction to evolution for general readers. Topics
include how evolutionary change and adaptation occur,
speciation, and macroevolution. The final part of the book
deals with human evolution.
- Menno Schilthuizen -- Frogs, Flies, and Danelions: The Making
Oxford University Press, 2001
- Although Darwin called his book The Origin of Species,
and his ideas were a giant leap forward,
he never really answered the question of what causes
new species to appear, or even exactly what a "species" is.
Biologists today are still very actively investigating these
difficult questions. Schilthuizen's relatively brief book provides a
good snapshot of where things stand today.
- Kim Sterelny -- Dawkins vs. Gould: Survival of the Fittest
Icon Books, Ltd., 2001
- The author, a professor of philosophy, presents key points
of controversy in the modern theory of evolution, especially
with respect to the well-publicized diasgreements between
Richard Dawkins and Stephen Jay Gould. The book provides a good
introduction to modern evolutionary theory and its main issues in a
- Richard Morris -- The Evolutionists: The Struggle for Darwin's Soul
Henry Holt and Company, 2001
- The theme is the ongoing controversy within evolutionary
theory between "orthodox" interpreations and dissenting opinions
offered by scientists such as Stephen Gould. In the course of
an even-handed and relatively brief discussion of the controversy,
Morris is able to impart a useful understanding of topics such
as the content of Darwin's theory, the evidence for it, complexity
theory, and evolutionary psychology.
- Steve Jones -- Darwin's Ghost: The Origin of Species
Ballantine Books, 2001
- Jones, a distinguished geneticist, rewrites Darwin's classic
to reflect the vast quantity of findings in biology since 1859.
The outline of Darwin's work, and some of the original passages,
are retained. The result is a very good introduction to modern
- Jeffrey H. Schwartz -- Sudden Origins: Fossils, Genes, and the
Emergence of Species
John Wiley and Sons, 1999
- Schwartz tackles head-on the problem of speciation in
evolution. There is a long review of the history of evolutionary
theory, followed by arguments regarding the importance for speciation
of recent findings in developmental biology -- especially the
role of gene regulatory mechanisms and the homeobox genes.
- Henry Gee -- In Search of Deep Time: Beyond the Fossil Record
to a New History of Life
Cornell University Press, 1999
- "Deep time" refers to the geological time scale, measured in
millions of years. The author presents a view of evolutionary
theory for general readers which is based on the idea that in
"deep time" it is not possible to make testable hypotheses
about ancestry relationships between species, but only more
general statements about degree of relatedness. He explains this
using the tools of "cladistics", for which this is a good
- Laurent Keller, ed. -- Levels of Selection in Evolution
Princeton University Press, 1999
- This is a volume of contributed papers addressing many issues
in the controversies over "levels of selection". The goal is to
synthesize divergent views and move beyond old arguments. Many of
the papers are technical, but well worth reading to understand
the complexities of the overall problem.
- Michael Ruse -- Mystery of Mysteries: Is Evolution a Social
Harvard University Press, 1999
- The author has held professorships in both philosophy and
biology. Although this book has the explicit purpose of
enquiring into the nature of science using evolutionary theory
as a case study, it is as valuable for the presentation of the
theory as for the philosophy. It examines the work
of a number of the leading thinkers about evolution, from Darwin (both
Erasmus and Charles) to contemporary scietists like Edward Wilson
and Stephen Jay Gould. It's a good introduction to many of
the current controversies in the field.
- Michael R. Rose -- Darwin's Spectre: Evolutionary Biology in the
Princeton University Press, 1998
- Rose, a professor of evolutionary biology, presents a history
of the theory of evolution from its origins to the present. The
history includes a brief explanation of the theory, how it has
itself evolved over time, and some of its effects on society and
the way we view ourselves and the world. This short book makes a
good introduction for general readers to many aspects of the theory.
- John Maynard Smith -- Evolutionary Genetics
Oxford University Press, 1998
- Here is an understandable textbook by a master of the subject,
on the genetic mechanisms that underlie evolutionary theory.
Simple mathematical models help illuminate otherwise difficult
issues such as natural selection, population biology,
genetic drift, evolutionary strategies,
sexual reproduction, speciation, and macroevolution.
- Richard Dawkins -- Climbing Mount Improbable
W. W. Norton & Company, 1996
- Dawkins shows, through many examples, the ways in which long
evolutionary processes have succeeded in producing very complex
and seemingly improbable biological structures. An especially
interesting example is the numerous independent evolutions of
"eyes" (light-collecting devices) in various species.
- Richard Dawkins -- The Blind Watchmaker
W. W. Norton & Company, 1996
- The author provides an account of the theory of evolution for
a general audience, with special emphasis on how is is possible
for complex biological forms to arise without the need for a
- Daniel C. Dennett -- Darwin's Dangerous Idea: Evolution and the
Meanings of Life
Simon and Schuster, 1995
- This is a superb work by a philosopher who is scientifically
informed. It considers the arguments that have been made, pro and
con, about evolution over the years, and it show how the general
model of biological evolution applies in other areas of scientific
interest as well.
- Richard Dawkins -- River Out of Eden: A Darwinian View of
Basic Books, 1995
- A short overview of the theory of evolution from the author's
customary perspective. Dawkins focuses on "utility functions" --
the maximization of which is in some sense the "purpose" of
evolution, another way of referring to the "good" that evolution
is metaphorically directed towards.
- Richard Dawkins -- The Selfish Gene
Oxford University Press, 1989
- This is a revised edition of the first of the author's series
of books on evolutionary theory for a general audience. It explains
the theory in the light of Dawkins' emphasis on the central role
individual genes play in the unending marathon race for survival.
- Richard Dawkins -- The Extended Phenotype
Oxford University Press, 1983
- In this volume, Dawkins provides a sequel to The Selfish
Gene aimed at an audience with more background in biology.
The theme remains not a new variation of the theory of evolution
so much as a new way of looking at it where the focus is on
"replicators" such as the individual gene, instead of organisms,
which are merely vehicles for the replicators.
Copyright © 2002 by Charles Daney, All Rights Reserved