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“We see in these facts some deep organic bond”
:
The Origin
, 350.

“On the theory of descent with modification”
: Ibid., 340.

“the animal in its less modified state”
: Ibid., 449.

the “very soul” of natural history
: Ibid., 434.

And the “same great law” of homologous parts
: Ibid.

“that it has so pleased the Creator”
: Ibid.

but somehow “natural” and objective
: Ibid., 413.

simply “reveals the plan of the Creator”
: Ibid.

“all true classification is genealogical”
: Ibid., 420.

“the hidden bond which naturalists”
: Ibid.

he also calls them “atrophied” or “aborted”
: Ibid., 450.

the entire book is essentially “one long argument”
: Ibid., 459.

“a considerable revolution in natural history”
: Ibid., 484.

“In the distant future”
: Ibid., 488.

“Light will be thrown on the origin of man”
: Ibid.

“To my mind it accords better”
: Ibid.

“There is grandeur,” he says finally
: Ibid., 490.

“This Abstract, which I now publish”
: Ibid., 2.

“If I had
space,”
he claims later
: Ibid., 31.

“…but I have not space here”
: Ibid., 89.

“I could show by a long catalogue of facts”
: Ibid., 45.

“I shall reserve for my future work”
: Ibid., 53.

“if I had space, I could show that they are conformable”
: Ibid., 230.

“This view may be true, and yet it may never be capable”
: Ibid., 338.

“the assertion is quite incapable of proof”
: Ibid., 468.

“I think it highly probable that” and “I am convinced that”
: Ibid., 18, 43.

The adjective “random” appears nowhere
: Attested by Barrett, et al. (1981), 606.

to say that variations are “due to chance”
:
The Origin
, 131.

sounds concrete and sensible: the “effects of use and disuse”
: Ibid., section heading, 134.

“I think there can be little doubt that use”
: Ibid., 134.

“I believe that the nearly wingless condition”
: Ibid.

“The real triumph of Darwin's book”
: Peckham (1959), 25.

“If I lived twenty more years”
: F. Darwin, ed. (1903), 2: 379.

The Fittest Idea

variations occur in response to “conditions of life”
:
The Origin
, 131.

“Our ignorance of the laws of variation”
: Ibid., 167.

variations occur “in no determinate way”
: F. Darwin, ed. (1909), 85.

he had described them as “accidents”
:
CD's Notebooks
, 633.

to say they are “due to chance”
:
The Origin
, 131.

“a dish of rank materialism cleverly cooked”
: Quoted in Hull (1973), 169.

suggesting that “man might be a transmuted ape”
: Quoted in ibid., 177.

positing instead some “internal innate force”
: Quoted in ibid., 409.

“The ‘Doctrine of Uniformity' in Geology Briefly Refuted”
:
Proceedings of the Royal Society of Edinburgh
, vol. 5, 1866, cited in Gould (2002), 492–93.

far less than the “incomprehensibly vast”
:
The Origin
, 282.

“sufficient to disprove the doctrine that transmutation”
: Quoted in Gould (2002), 497.

about the “odious spectre” of Thomson
: Marchant (1975), 220.

cut “incomprehensibly vast” to merely “vast”
: Peckham (1959), 478.

“we have no means of determining how long”
: Ibid., 486.

“Fleeming Jenkin has given me much trouble”
: F. Darwin, ed. (1903), 2: 379.

“the tendency to revert to parent forms”
:
CD's Notebooks
, 248.

“elaborated by Mr. Darwin in his celebrated”
: Wallace (1962), 102.

“moral and higher intellectual nature of man”
: From “Sir Charles Lyell on Geological Climates and the Origin of Species,”
The Quarterly Review
, 1869, vol. 126, excerpted in Smith, ed. (1991), 31.

“that an Overruling Intelligence has watched”
: Ibid., 33–34.

“I shall be intensely curious to read”
: F. Darwin, ed. (1903), 39.

Darwin scratched “No!!!”
: Browne (2002), 318.

“Lamarckism in a modern form”
: Packard (1980), 393, quoting himself from an earlier paper.

“nearer the truth than Darwinism proper”
: Ibid.

became known as “the law of acceleration”
: Bowler (1992), 128; Gould (2002), 367.

a book about the “self-adaptation” of plants
: Quoted in Bowler (1992), 86.

a claim that internal “laws of growth” dictate
: Quoted in ibid., 90.

“that the actual course of orthogenetic evolution”
: Ibid., 151.

“must advance by the shortest and slowest steps”
:
The Origin
, 194.

“I know that you are studying hybrids”
: Quoted in Gould (2002), 419.

whereas others are
recessive
(Mendel's terminology)
: Mendel (1965), 8.

He argued that the
germ plasm
: Mayr (1982), 700.

“based on logic and on interpretation of many kinds”
: Futuyma (1998), 11.

“fully vindicate his hypothesis”
: Ibid., 12.

“about 30,000 genes, with 99%”
:
Nature
, vol. 420, December 5, 2002, 509.

The resemblance between our 30,000 human genes
: Dr. Futuyma was rounding off, based on the best gene counts available at that time. Since my conversation with him, on January 21, 2004, further work in genomics has led to a slight revision: Scientists now believe there are only about 20,000 to 25,000 human genes, rather than 26,000 to 31,000, as thought when the hurried results of the Human Genome Project were first announced. The mouse genome also continues to be studied; the overwhelming degree of mouse-human similarity hasn't been challenged; and the downward revision in gene number doesn't affect the validity of Futuyma's point.

“Comparative genome analysis is perhaps”
:
Nature
, vol. 420, December 5, 2002, 557.

The Last Beetle

with salutations from a “sincere admirer”
: Quoted in Desmond and Moore (1991), 601.

“interesting letter” about hairiness on the ears
: F. Darwin, ed. (1903), 2: 53–54.

“It has always pleased me to exalt”
:
Autobiography
, 135.

Bessy was “not good at practical things”
: Raverat (1952), 146.

“Don't think that it is not my affair”
: Reprinted in
Autobiography
, 237.

“When I am dead, know that many”
: Ibid.

“might possibly interest my children”
:
Autobiography
, 21.

“it is odd that I can remember hardly anything”
: Ibid., 22.

“You care for nothing but shooting”
: Ibid., 28.

“But no pursuit at Cambridge was followed”
: Ibid., 62.

“a third and new kind, which I could not bear to lose”
: Ibid.

is titled “Religious Belief”
: Ibid., 85, but absent from the table of contents, 19.

“gradually came to disbelieve in Christianity”
: Ibid., 86.

“Thus disbelief crept over me”
: Ibid., 87.

“I can indeed hardly see how anyone”
: Ibid.

“it revolts our understanding”
: Ibid., 90.

“I cannot pretend to throw the least light”
: Ibid., 94.

the alternative, “atheist,” was too aggressive
: Desmond and Moore (1991), 657.

“now that the law of natural selection”
:
Autobiography
, 87.

“a prize article” with a sort of Buddha calm
: Quoted in Browne (2002), 438.

“He has a pretty mouth and expression”
: Quoted in ibid., 438.

grandpa was “Baba” and Bernard
: Ibid., 491.

“Worms do not possess any sense of hearing”
: CD,
The Formation of Vegetable Mould
, 29.

“I am not the least afraid to die”
: Desmond and Moore (1991), 662.

“My love, my precious love”
: Ibid., 661.

“If I could but die,” and repeated the phrase
: Ibid., 662.

“I have placed the shell in fresh-water”
: F. Darwin, ed. (1903), 2:29.

“As the wretched beetle was still”
: Ibid. 2: 29.

Bibliography

Charles Darwin was a prolific writer. His lifetime output includes the published books and articles, the private notebooks, and a formidable abundance of personal letters. The best evidence for who he was, and for what he thought, is what he wrote. Close attention to the text of the first edition of
The Origin of Species
, for instance, and to the changes he made in later editions, is the right substitute and remedy for blurred, secondary notions of what is or isn't “Darwinian.” Darwin's letters are very illuminating, especially as edited and annotated by Frederick Burkhardt and a team of other scholars in the ongoing multi-volume series from Cambridge University Press. The transmutation notebooks, transcribed and edited by Paul H. Barrett and some colleagues, reveal much about how Darwin pieced his ideas together. His
Autobiography,
restored to completeness (after earlier suppression of some passages, in deference to Emma Darwin) and edited by his granddaughter, Nora Barlow, is also a telling document. These have been my primary sources. Confession: I haven't read every word that Darwin put into print. My selective list of his books, below, records only those that have been most influential scientifically and that were crucial or useful to me.

The secondary literature on Darwin and his work (sometimes called the “Darwin Industry” of scholarship and commentary) is huge and still growing. Every week, it seems, someone publishes another erudite research article or argumentative book with his name in the title. The list below is, again, partial and personal. Think of it as merely a small sample, purposeful but somewhat subjective on my part, of what's out there on the subject of Charles Darwin.

In case you find yourself hungry for further reading—of Darwin's own writings, or about him—I'll offer a few suggestions. For a start that takes you straight to the core of the man and his work, read
The Origin of Species
, preferably in a reprint of the first edition.
The Voyage of the Beagle
(as his
Journal of Researches
has been titled in later editions) is also a wonderful book in its way—not so important as
The Origin of Species
(how many books are?) but more relaxed and amusing, a travel narrative filled with acutely observed natural history and told in the voice of a likable, unpretentious young Englishman.
The Autobiography of Charles Darwin
, first published in 1887 as part of
The Life and Letters of Charles Darwin
, and later available in the Nora Barlow edition, is warm and personable (mainly because he wrote it for his children, not for distant readers such as us), graced by his gentle spirit and his extraordinary honesty. For those who enjoy reading letters, you don't need to plunge into the oceanic thoroughness of that series from Cambridge; Frederick Burkhardt has also edited a nice little book,
Charles Darwin's Letters: A Selection
. Better still, find a reprint of Francis Darwin's original 1887 compilation,
The Life and Letters of Charles Darwin
, in two modest-sized volumes. Volume I of that set gives you, besides letters and the autobiography, a long chapter of reminiscences by Francis about his father's character, working habits, and everyday life.

Among the many Darwin biographies, the two most impressive are also the two most appealing: Janet Browne's and the one by Adrian Desmond and James Moore. Both works are enriched by formidable scholarship and enlivened by sharp insights and good writing. Desmond and Moore are especially strong on the political context surrounding Darwin and his thought. Browne's two-volume life is particularly good on the social milieu in which Darwin lived, and on the high-minded ruthlessness with which he presumed upon the females of his own family, as well as friends and minions around the world, to indulge his disposition and serve his intellectual demands. The biographical portrait by Darwin's great-great-grandson, Randal Keynes, published in the United States as
Darwin, His Daughter, and Human Evolution
(originally in the UK as
Annie's Box
) is also a valuable book, poignant but levelheaded, and informed by insider material.

For Darwin's own works, what I cite below are the original bibliographical data—details of the first editions—so you can see at a glance how his publishing career unfolded. In my Source Notes, on the other hand, I place quotations from Darwin within the editions that were available to me. Sorry for this inconsistency, but on the whole I think it keeps things more clear. In the list of secondary material, I cite the editions that came into my hands. Some of those books and articles represent milestones in the development of evolutionary theory, published long ago and since reprinted. In such cases (for instance, Mendel's paper), where the year of first publication helps clarify the historical significance, I supply the original year in parentheses.

1. Works published by Charles Darwin

1839.
Journal of Researches into the Geology and Natural History of the Various Countries Visited by H.M.S. Beagle, Under the Command of Captain FitzRoy, R.N. from 1832 to 1836
. London: Henry Colburn.

1839. “Observations on the Parallel Roads of Glen Roy,”
Philosophical Transactions of the Royal Society of London
, 39–81.

1842.
The Structure and Distribution of Coral Reefs. Being the first part of the geology of the voyage of the Beagle, under the command of Capt. FitzRoy, R.N. during the years 1832 to 1836
. London: Smith, Elder.

1845.
Journal of Researches into the Natural History and Geology of Countries Visited during the Voyage of H.M.S. Beagle round the World, Under the Command of Capt. FitzRoy, R.N.
London: John Murray.

1851–54a.
A Monograph of the Sub-Class Cirripedia
. Vol. I:
The Lepadidae, or Pedunculated Cirripedes
, 1851. Vol. II:
The Balanidae, (or Sessile Cirripedes); the Verrucidae, etc.
, 1854. London: The Ray Society.

1851–54b.
A Monograph of the Fossil Lepadidae, or Pedunculated Cirripedes of Great Britain
, 1851.
A Monograph of the Fossil Balanidae and Verrucidae of Great Britain
, 1854. London: Palaeontographical Society.

1859.
On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life
. London: John Murray.

1862.
On the Various Contrivances by which British and Foreign Orchids Are Fertilised by Insects, and on the Good Effects of Intercrossing.
London: John Murray.

1868.
The Variation of Animals and Plants Under Domestication.
London: John Murray.

1871.
The Descent of Man, and Selection in Relation to Sex
. London: John Murray.

1872.
The Expression of the Emotions in Man and Animals
. London: John Murray.

1875.
Insectivorous Plants
. London: John Murray.

1875.
The Movements and Habits of Climbing Plants
. London: John Murray.

1877.
The Different Forms of Flowers on Plants of the Same Species
. London: John Murray.

1881.
The Formation of Vegetable Mould, through the Action of Worms, with Observations on Their Habits.
London: John Murray.

1882. “On the Dispersal of Freshwater Bivalves,”
Nature
, vol. 25, April 6, 1882.

Darwin, Charles, and A. R. Wallace. 1858. “On the Tendency of Species to Form Varieties; and on the Perpetuation of Varieties & Species by Means of Natural Selection.” Read on July 1, 1858, and first published in the
Journal of the Linnean Society of London
, vol. 3, 1858.

2. Other writings by Charles Darwin, unpublished in his lifetime

Barlow, Nora, ed. 1969.
The Autobiography of Charles Darwin, 1809–1882. With original omissions restored
. New York: W. W. Norton.

Barrett, Paul H., Peter J. Gautrey, Sandra Herbert, David Kohn, and Sydney Smith, eds. 1987.
Charles Darwin's Notebooks, 1836–1844.
Ithaca, NY: Cornell University Press.

Burkhardt, Frederick, ed. 1996.
Charles Darwin's Letters: A Selection
. Cambridge: Cambridge University Press.

———, Sydney Smith, et al., eds. 1985–93.
The Correspondence of Charles
Darwin
, Vols. 1–8, covering 1821–60. Cambridge: Cambridge University Press.

Darwin, Francis, ed. 1887.
The Life and Letters of Charles Darwin, including an autobiographical chapter
. 2 vols. London: John Murray.

———, ed. 1909.
The Foundations of the “Origin of Species”: Two Essays Written in 1842 and 1844 by Charles Darwin
. Cambridge: Cambridge University Press (also available on the Web, along with many of Darwin's other writings, edited by John van Wyhe, at http://pages.britishlibrary.net/charles.darwin).

———, and A. C. Seward, eds. 1903.
More Letters of Charles Darwin: A Record of his Work in a Series of Hitherto Unpublished Letters
. 2 vols. London: John Murray.

Keynes, Richard Darwin, ed. 1988.
Charles Darwin's Beagle Diary
. Cambridge: Cambridge University Press.

———, ed.
Charles Darwin's Zoology Notes & Specimen Lists from H.M.S. Beagle
. Cambridge: Cambridge University Press.

Stauffer, R. C., ed. 1987.
Charles Darwin's Natural Selection: Being the Second Part of his Big Species Book Written from 1856 to 1858
. Cambridge: Cambridge University Press.

3. Additional sources

Appleman, Philip, ed. 2001.
Darwin: A Norton Critical Edition
. New York: W. W. Norton.

Ayala, Francisco J. 1982.
Population and Evolutionary Genetics: A Primer.
Menlo Park, CA: Benjamin/Cummings.

Baker, Allan J., C. H. Daughterty, Rogan Colbourne, and J. L. McLennan. 1995. “Flightless Brown Kiwis of New Zealand Possess Extremely Subdivided Population Structure and Cryptic Species Like Small Mammals,”
Proceedings of the National Academy of Science
, vol. 92.

Barrett, Paul H., Donald J. Weinshank, and Timothy T. Gottleber. 1981.
A Concordance to Darwin's “Origin of Species,” First Edition
. Ithaca, NY: Cornell University Press.

Bowlby, John. 1992.
Charles Darwin: A New Life
. New York: W. W. Norton.

Bowler, Peter J. 1989.
Evolution: The History of an Idea
. Berkeley: University of California Press.

———. 1992.
The Eclipse of Darwinism: Anti-Darwinian Evolutionary Theories in the Decades Around 1900.
Baltimore: Johns Hopkins University Press.

———. 1992.
The Non-Darwinian Revolution: Reinterpreting a Historical Myth
. Baltimore: Johns Hopkins University Press.

Brent, Peter. 1983.
Charles Darwin: A Man of Enlarged Curiosity
. New York: W. W. Norton.

Brooks, John Langdon. 1984.
Just Before the Origin: Alfred Russel Wallace's Theory of Evolution
. New York: Columbia University Press.

Browne, Janet. 1983.
The Secular Ark: Studies in the History of Biogeography
. New Haven: Yale University Press.

————. 1996.
Charles Darwin: Voyaging
. Princeton: Princeton University Press.

____. 2002.
Charles Darwin: The Power of Place
. New York: Alfred A. Knopf.

Burkhardt, Richard W., Jr., ed. 1977.
The Spirit of System: Lamarck and Evolutionary Biology
. Cambridge, MA: Harvard University Press.

Calder, William A. III. 1978. “The Kiwi,”
Scientific American
, vol. 239.

————. 1979. “The Kiwi and Egg Design: Evolution as a Package Deal,”
Bio-Science
, vol. 29, no. 8.

Camerini, Jane R., ed. 2002.
The Alfred Russel Wallace Reader: A Selection of Writings from the Field
. Baltimore: Johns Hopkins University Press.

Chambers, Robert. 1994 (1844).
Vestiges of the Natural History of Creation, and Other Evolutionary Writings
, ed. James Secord. Chicago: University of Chicago Press.

Cracraft, Joel. 1974. “Phylogeny and Evolution of the Ratite Birds,”
Ibis,
vol. 116, no. 4.

Dawkins, Richard. 1991.
The Blind Watchmaker
. London: Penguin Books.

Dennett, Daniel C. 1995.
Darwin's Dangerous Idea: Evolution and the Meanings of Life
. New York: Simon & Schuster.

Depew, David J., and Bruce H. Weber. 1996.
Darwinism Evolving: Systems Dynamics and the Genealogy of Natural Selection
. Cambridge, MA: MIT Press.

———, eds. 1985.
Evolution at a Crossroads: The New Biology and the New Philosophy of Science
. Cambridge, MA: MIT Press.

Desmond, Adrian. 1992.
The Politics of Evolution: Morphology, Medicine, and Reform in Radical London.
Chicago: University of Chicago Press.

———, and James Moore. 1991.
Charles Darwin: The Life of a Tormented Evolutionist
. New York: Warner Books.

Dobzhansky, Theodosius. 1982 (1937).
Genetics and the Origin of Species
. New York: Columbia University Press.

Eldredge, Niles. 1986.
Time Frames: The Rethinking of Darwinian Evolution and the Theory of Punctuated Equilibria
. New York: Touchstone.

———. 1995.
Reinventing Darwin: The Great Debate at the High Table of Evolutionary Theory
. New York: John Wiley & Sons.

———. 2000.
The Triumph of Evolution, and the Failure of Creationism.
New York: W. H. Freeman.

———. 2005.
Darwin: Discovering the Tree of Life
. New York: W.W. Norton.

Ellegård, Alvar. 1990 (1958).
Darwin and the General Reader: The Reception of Darwin's Theory of Evolution in the British Periodical Press, 1859–1872
. Chicago: University of Chicago Press.

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