Difference between revisions of "Classic Works in Evolutionary Biology"

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[homeostasis of populations and individuals (CS)]<br/><br/>
[homeostasis of populations and individuals (CS)]<br/><br/>
'''Lewontin, R. C. 1978.  Adaptation. Scientific American 239:212-228.''' (Adams; Schwenk)<br/>
'''Lewontin, R. C. 1978.  Adaptation. Scientific American 239:212-228.'''<br/>
[strangely enough, this article published in the popular press Scientific American is one of the best single treatments of this critical, yet slippery, concept (KS)]<br/><br/>
[strangely enough, this article published in the popular press Scientific American is one of the best single treatments of this critical, yet slippery, concept (KS)]<br/>
'''Lewontin R. C. and L. C. Birch. 1966. Hybridization as a source of variation for adaptation to new environments. Evolution 20: 315-336.'''<br/>
'''Lewontin R. C. and L. C. Birch. 1966. Hybridization as a source of variation for adaptation to new environments. Evolution 20: 315-336.'''<br/>

Revision as of 04:29, 17 May 2008

Charles Darwin in later years
"Darwin's finches" (Geospiza) from the Galapagos Islands

What is This Page?

This page grew out of a graduate seminar taught in EEB during the spring semester, 2008, by Kurt Schwenk. The seminar was very successful—so successful, in fact, that one student was motivated to prepare a multi-media summary of what he learned—you can download it HERE (Diego is appropriately modest about his achievement). There was a general consensus that it would be useful to graduate students and the EEB community at large to have continuing access to the papers and book selections that were read during the semester, as well as additional papers included in the master list of 'classic' works in evolutionary biology. Many of these pdfs are available in the list below.

Who Can Access and Use This Page?

Anyone can access this page, read its contents and even learn something. However, only members of the EEB community can edit the page and download pdfs for personal use. If you are a member of EEB, all you need to do is log in using your University Net ID and password. You can log in by clicking the tiny 'log in' link at the upper right corner of the page. Alternatively, you can click on a pdf icon or an 'edit' link and you will be directed to log in before proceeding. If you are a member of the EEB community and haven't done so already, please send Paul Lewis (Mr. EEBedia) your net ID (but not your password). Once you are on the 'acceptable' list, you have full access. Paul has created a system for this page and future course pages whereby it is not necessary to remember a username and password to access pdfs - it is all accomplished by logging in with your Net ID.

If you are a part of the UConn community with a Net ID, but not a member of EEB, and you wish to have full access to the list, you can make your case to Paul (paul.lewis@uconn.edu) for the addition of your Net ID to the user list.

What is 'The Master List'?

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Kurt contemplates the mystery of his existence and concludes that he "didn't evolve from no stinkin' monkeys"

The 'master list' is a list of classic works in evolutionary biology that I (Kurt Schwenk) compiled and partially annotated during the course of the semester. It was created based on my own choices, as well as suggestions from EEB colleagues and various web sources. Obviously there are potentially as many lists of 'classic works' as there are evolutionary biologists - this particular list is not intended to be definitive. It reflects, to some degree, my own personal interests and biases. However, an attempt was made to be inclusive; for example, there are a number of very important botanical references (thanks largely to Carl Schlichting), as well as obviously important works in population genetics, molecular biology and other areas I can hardly claim expertise in. As much as possible, I included only works that are generally important to evolutionary biology, as a whole, rather than the many papers that are of more circumscribed, if nonetheless critical, importance to a particular subdisciplines (e.g., phylogenetic analysis) or within a particular taxon (although I admit that a couple of important vertebrate papers somehow wandered into the list...). I excluded books and papers that are too recent. Thus, while there is no doubt that Schlichting and Pigliucci's Phenotypic Evolution (1998) (for example) will be considered a classic in the future, it is simply too recently published to have achieved this status now (I have added a separate section for such works, with the oxymoronic title, 'Modern Classics'). Finally, I note that several of the listed works are followed by a sublist of related papers that are, themselves, not necessarily 'classics', but are nevertheless helpful in appreciating the context or significance of the listed work. This is a type of emmendation/addition/annotation that would be particularly useful (see note below about annnotations).

Pdficon small.gifEvolutionary Classics Master List (version 5/15/08)

A Comment on Modifying the Master List

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Kurt reacts to someone f***ing with the Master List pdf.

A great advantage of using an EEBedia page for this purpose is that it can be edited and added to by the EEB community (thank you Paul Lewis!). However, I have a request - as much as I would like to see additions, changes, annotations, etc., I ask that a copy of the 'original' master list (i.e., the pdf linked above) be preserved. Thus, please modify the list posted below. Alternatively, you can create your own separate list - for example, you might want to create a 'classics' list for a particular subdiscipline, such as phylogenetics or behavioral evolution - and create a link to a separate pdf or a new EEBedia page. As such, the list posted below should not be modified for 'specialty' purposes. Additions should be carefully considered and restricted to books and papers the contributor feels are of genuine, general importance to evolutionary biology sensu lato.

The pdf version of the Master List will periodically be revised and the updated version uploaded to the link above. However, there will be periods of time during which the pdf version of the list does not correspond exactly to the 'editable' version below.

I request that you do not make deletions from the posted list without consulting me (Schwenk) first. While I accept that it might be a good idea to trim the list in some cases, I would prefer to err on the side of conservatism and make the decision to delete through consultation with colleagues. If there is a consensus that a particular work should be stricken fron the list, then it will be stricken! Conversely, if an addition is made to the list that causes widespread grumbling, it might be deleted.

If you make additions to the list, I strongly encourage you to upload pdfs of the new papers whenever possible. One of the principle purposes of this page, obviously, is to educate and one specific way to do this is to help graduate students preparing for their oral qualifying exams. Making sources easy to obtain would be a great service to students and other users. Obvioulsly not everything is available in electronic format, but often more is available than one might think and can be located by doing a Google search on the paper title (i.e., pdfs are sometimes available through web sources outside of JSTOR or other standard electronic databases; or in some cases there are entire books online so that a link can be provided) . Scanned documents are good, but watch out for memory hogs - there is a limit of 4 MB on documents uploaded to this page. If you have a document larger than that, or if you don't want to take the time to upload the paper yourself, feel free to send it to Schwenk for posting.

Changing or Annotating the List

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Useful ways to modify the list:

  • add an annotation - a summary or short note explaining the significance or context of the listed work (add your initials here and to the 'key')
  • correct or add to an existing annotation
  • add subsidiary/supplemental citations below a listed work if it helps inform about the work's importance (see examples)
  • add a link to a site with a copy of the text or with supplementary information related to the listed work
  • upload a pdf of the listed paper using the form link at the top of the list
  • add a book or paper to the list (please do not do this casually - give it careful thought with regard to the comments above about the spirit of the list)
  • create your own 'specialty list' of classics in a particular subdiscipine (e.g., vertebrate form and function) - create a new, linked EEBedia page or upload a pdf
  • if you own a hard copy of a listed book or paper that is not available electronically and you are willing to loan it, indicate this with your last name in parentheses after the reference


Classic Works in Evolutionary Biology With Annotations and Links to PDFs


annotation initials key:

KS = Kurt Schwenk
CS = Carl Schlichting

Note 1: a name in parentheses after a citation indicates a faculty member who has a hard copy of the listed work she/he is willing to loan
Note 2: PDFs of many of the listed papers are being uploaded on a regular basis (it takes awhile) - keep checking back!


Baldwin, J. M. 1902. Development and Evolution. MacMillan, London.

Bonner, J. T. 1958. The Evolution of Development. Cambridge Univ. Press, Cambridge.

Bonner, J. T. 1988. The Evolution of Complexity by Means of Natural Selection. Princeton Univ. Press, Princeton. (Schwenk)

Darwin, C. 1859. On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. 1st ed. John Murray, London.
[see http://darwin-online.org.uk/ for Darwin’s collected works online (KS)]
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Darwin, C. 1872. On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. 6th ed. with additions and corrections. John Murray, London.
[see http://darwin-online.org.uk/ for Darwin’s collected works online (KS)]

Darwin, C. 1871. The Descent of Man, and Selection in Relation to Sex. Vols. 1 & 2. 1st ed. John Murray, London.
[see http://darwin-online.org.uk/ for Darwin’s collected works online (KS)]

Dawkins, R. 1976. The Selfish Gene. Oxford Univ. Press, Oxford. (Schwenk)
[phenotypes as contrivances of genes to replicate themselves (KS)]

de Beer, G. R. (editor).1938. Evolution. Essays on Aspects of Evolutionary Biology Presented to Professor E. S. Goodrich on His Birthday. Oxford/Clarendon Press, Oxford. (Schwenk)

de Beer, G. R. 1958. Embryos and Ancestors. 3rd ed. Oxford/Clarendon Press, Oxford. (Schwenk)
[1st edition published in 1940 (KS)]

Dobzhansky, T. 1937. Genetics and the Origin of Species. Columbia Univ. Press, New York. (Schwenk)
[+2nd and 3rd editions in 1941 and 1951, respectively (KS)]

Dobzhansky, T. 1970. Genetics of the Evolutionary Process. Columbia Univ. Press, New York. (Schwenk)

Endler, J. A. 1986. Natural Selection in the Wild. Mongraphs in Population Biology No. 21. Princeton Univ. Press, Princeton, NJ. (Schwenk)
[in addition to technical material on measuring selection, etc., Endler has an excellent and thoughtful general/philosophical discussion of natural selection - a very good introduction to the concept (KS)]

Fisher, R. A. 1930. The Genetical Theory of Natural Selection. Clarendon Press, Oxford.

Frazzetta, T. H. 1975. Complex Adaptations in Evolving Populations. Sinauer Assoc., Sunderland, MA. (Schwenk)
[a quirky and often overlooked little book on the evolution of 'complex' adaptations, character complexes, etc. (KS)]

Goldschmidt, R. 1940. The Material Basis of Evolution. Yale Univ. Press, New Haven. (Schwenk)
[ever wonder where the term "hopeful monster" came from? (KS)]

Gould, S. J. 1977. Ontogeny and Phylogeny. Belknap/Harvard Univ. Press, Cambridge. (Schwenk)
[read this book for the first half - a fantastic history of developmental morphology - Haeckel, von Baer - those guys (KS)]

Grant, V. 1963. The Origin of Adaptations. Columbia Univ. Press, New York. (Schwenk)

Gregory, W. K. 1951. Evolution Emerging. A Survey of Changing Patterns From Primeval Life to Man. Vols. 1 & 2. MacMillan, New York. (Schwenk)
[The first volume is a history of life and was probably a bit dated even when it was published; the second volume is an amazing collection of figures of organisms, skulls, fossils, etc. Together they represent an amazing post-war treatise on biodiversity and the physical evidence for evolution (KS)]

Haldane, J. B. S. 1932. The Causes of Evolution. Longmans, Green and Co., London.
[Birth of quantitative genetics; appendix has summary of his papers on selection intensity, etc. (CS)]

Hennig, W. 1966. Phylogenetic Systematics. Univ. of Illinois Press, Urbana. (Schwenk)
[translated by D. Dwight Davis and R. Zangerl; this book represents a revised and expanded version of Hennig’s Grundzüge einer Theorie der phylogenetischen Systematik (1950) and is therefore a new book rather than a simple translation. This is the bible of cladistics that when introduced in this country caused a paradigm shift in systematics (KS)]

Huxley, J. S. 1942. Evolution, the Modern Synthesis. Harper, New York. (Schwenk)
[new edition in 1963, Allen and Unwin, London, with new Introduction by Huxley (KS)]

Jepsen, G. L., G. G. Simpson and E. Mayr (editors). 1949. Genetics, Paleontology and Evolution. Princeton Univ. Press, Princeton, NJ. (Schwenk)
[the short Foreward by Jepsen is a nice capsule summary of the aims and scope of the Synthesis (KS)]

Malthus, T. 1798. An Essay on the Principle of Population, as it Affects the Future Improvement of Society with Remarks on the Speculations of Mr. Godwin, M. Condorcet, and Other Writers. Printed for J. Johnson, in St. Paul's Church-yard, London.
[not a work on evolution, but this famous essay was instrumental to Darwin in formulating the notion of the 'struggle for existence', which played a critical part in his theory of natural selection (KS)]
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Mayr, E. 1942. Systematics and the Origin of Species. Columbia Univ. Press, New York. (Schwenk)

Mayr, E. 1963. Animal Species and Evolution. Belknap/Harvard Univ. Press, Cambridge, MA. (Schwenk)
[synthesis of the Synthesis from the man who gave us the Synthesis; explicates Mayr’s view on geographic/allopatric speciation, among other things (KS)]

Rensch, B. 1959. Evolution Above the Species Level. Columbia Univ. Press, New York. (Schwenk)
[originally published in 1954 in German; Rensch was a double PhD ("Herr Prof. Dr. Dr" - I'm not kidding) in philosophy and zoology; his philosophical bent is obvious in his writings. Interestingly, his student, Gerhard Roth - who works on evolutionary neuroanatomy of the brain and especially sensory systems in amphibians - is also a Herr Prof. Dr. Dr. in philosophy and zoology...) (KS)]

Riedl, R. 1978. Order in Living Organisms. John Wiley and Sons, New York.
[fascinating, but rather opaque; read his 1977 paper (below) for the essentials; Gunter Wagner at Yale was a Riedl student (KS)]

Schmalhausen, I. I. 1949. Factors of Evolution. The Theory of Stabilizing Selection. The Blakiston Co., Philadelphia. (Schwenk)
[reprinted 1986 by the Univ. of Chicago Press. Schmalhausen worked in isolation in Stalinist Russia and his accomplishments are all the more remarkable because of this (KS)]

Simpson, G. G. 1944. Tempo and Mode in Evolution. Columbia Univ. Press, New York. (Schwenk)

Simpson, G. G. 1949. The Meaning of Evolution. Yale Univ. Press, New Haven. (Schwenk)
[more of a popular book, but influential (KS)]

Simpson, G. G. 1953. The Major Features of Evolution. Columbia Univ. Press, New York. (Schwenk)
[a complete reworking of Tempo and Mode; virtually a new, more synthetic book. If you can only read on Simpson book, read this one. The job of relating the population/genetic/population microevolutionary phenomena of concern to most of the ‘synthesists’ to macroevolutionary/deep time patterns evident in the fossil record fell to Simpson. He makes a heroic effort here and was way ahead of his time. Originates the notion of the 'adaptive zone' and discusses the relation between adaptive zones and adaptive readiations at length (KS)]
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Stebbins, G. L., Jr. 1950. Variation and Evolution in Plants. Columbia Univ. Press, New York.
[Stebbins was the one botanist ‘officially’ welcomed into the Synthesis fold (KS)}

Thompson, D. W. 1942. On Growth and Form: A New Edition. Cambridge Univ. Press, Cambridge. (Schwenk)
[reprinted unabridged by Dover Press, 1992. An amazing book dealing with allometry, morphological transformation and quantification, among other things. Introduces the application of Cartesian coordinates to examine 2-D shape change (KS)]

Waddington, C. H. 1957. The Strategy of the Genes. A Discussion of Some Aspects of Theoretical Biology. Macmillan, New York. (Schwenk)
[the dawning of the modern evo-devo movement; a critically important, but often neglected book - a 'must read' for people interested in development and phenotypic evolution. Explicates notion of 'canalization' among other important concepts (KS)]

White, M. J. D. 1978. Modes of Speciation. W. H. Freeman, San Francisco. (Schwenk)
[the view from cytogenetics, which at the time was very big (cytogenetics has been largely supplanted by molecular-genetic approaches (KS)]

Whyte, L. L. 1965. Internal Factors in Evolution. George Braziller, New York. (Schwenk)
[more philosophical than biological, this book, virtually ignored at the time, is becoming increasingly influential; deals with internal selection and organismal ‘homeostasis’. Introduces the important concept of 'internal selection' (KS)]

Williams, G. C. 1966. Adaptation and Natural Selection. Princeton University Press, Princeton NJ. (Schwenk)

Williams, G. C. 1992. Natual Selection. Domains, Levels, and Challenges. Oxford Univ. Press, Oxford. (Schwenk)

Wright, S. 1968. Evolution and the Genetics of Populations. Vol. 1. Genetics and Biometric Foundations. Univ. of Chicago Press, Chicago.

Wright, S. 1969. Evolution and the Genetics of Populations. Vol. 2. The Theory of Gene Frequencies. Univ. of Chicago Press, Chicago.
Wright, S. 1977. Evolution and the Genetics of Populations. Vol. 3. Experimental Results and Evolutionary Deductions. Univ. of Chicago Press, Chicago.
Wright, S. 1978. Evolution and the Genetics of Populations. Vol. 4. Variability Within and Among Natural Populations. Univ. of Chicago Press, Chicago. (Schwenk)


Alberch, P., S. J. Gould, G. F. Oster and D. B. Wake. 1979. Size and shape in ontogeny and phylogeny. Paleobiology 5:296-317.
[rejects Gould’s (1977) ‘clock model’ of heterochrony and formalizes the notion of ‘ontogenetic trajectories’; proposes a formal lexicon of heterochrony terms (KS)]
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Arnold, S. J. 1983. Morphology, performance and fitness. American Zoologist 23:347-361.
[a critical paper for anyone interested in functional biology and evolution (KS)]
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Antonovics, J. 1976. The nature of limits to natural selection. Annals of the Missouri Botanical Garden 63:224-247.
[insufficient genetic variability and the swamping effects of gene flow are inadequate explanations of limits to natural selection. Comparison of evolutionary responses in different populations subjected to similar selective forces, comparison of rare and widespread species, and comparison of marginal and central populations are all neglected research areas that bear on the nature of limits to natural selection. Plant populations provide us with well-defined, operationally viable systems for addressing these comparisons. Several possible constraints on range extension of ecologically marginal populations are considered in detail. Selection on fitness components that are themselves negatively correlated will be ineffective: such negative correlations are to be expected in natural populations. Small size of marginal populations will reduce severely the probability of obtaining appropriate character combinations; it will increase the swamping effects of gene flow; and it may lead to inbreeding depression effects. Gene flow will have different effects depending on whether the genes concerned are effectively neutral, advantageous, or deleterious in the population into which they migrate. Gene flow will spread beneficial genes rapidly, but may retard divergence if density of marginal populations is low and swamping effects are high. Finally a population entering a new habitat is likely to meet new competitors and predators: the coevolutionary responses of the latter may counteract adaptive responses by the species undergoing range extension. All these factors are likely to interact in important ways in marginal populations. The study of limits to natural selection is likely to be a fruitful future research area, and one in which the detailed documentation of the systematist will provide invaluable baseline information (CS)]

Baldwin, J. M. 1896. A new factor in evolution. Amer. Nat. 30:441-451, 536-553.
[for some reason this has been called 'the Baldwin Effect'; see http://en.wikipedia.org/wiki/Baldwin_effect for more information. Also note that Baldwin's paper was divided into two portions in Am. Nat., hence the two pdfs. The 'Baldwin Effect' remains contentious (KS)]
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Boag, P. T., and P. R. Grant. 1981. Intense natural selection in a population of Darwin’s finches in the Galapagos. Science 214: 82-85.
[see Grant & Grant paper below (KS)]
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Bock, W. J., and G. von Wahlert. 1965. Adaptation and the form-function complex. Evolution 19:269-299.
[distinguishes the concepts of structure, function and biological role; extremely useful for those who think in terms of 'the target of selection' , i.e., what is actually being selected for? (also relevant to ideas about 'levels' of selection) (KS)]
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Brown, W. L., Jr., and E. O. Wilson. 1956. Character displacement. Systematic Zoology 5:49-64.
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Clausen, J., and W. M. Hiesey. 1958. Experimental studies on the nature of species. IV. Genetic structure of ecological races. Publication 615, Carnegie Institution of Washington, Washington, DC.

Conway Morris, S. 1989. Burgess shale faunas and the Cambrian explosion. Science 246: 339-346.
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Coyne, J. A. and H. A. Orr. 1989. Patterns of speciation in Drosophila. Evoluion 43:362-381.
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Coyne, J. A. and H. A. Orr. 1997. “Patterns of speciation in Drosophila” revisited. Evolution 51:295-303. Pdficon small.gif

Dobzhansky, T. and O. Pavlovsky. 1957. An experimental study of interaction between genetic drift and natural selection. Evolution 11:311-319.
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Ehrlich, P. R. and P. H. Raven. 1964. Butterflies and plants: a study in coevolution. Evolution 18:586-608.
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Ehrlich, P. R. and P. H. Raven. 1969. Differentiation of populations. Science 165:1228-1232.
[seminal paper for our understanding of gene flow and species coherence (KS)]
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Eldredge, N. and S. J. Gould. 1972. Punctuated equilibria: An alternative to phyletic gradualism. In T. J. M. Schopf (ed.), Models in Paleobiology, pp. 82-115. Freeman, Cooper and Company, San Francisco.
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Epling, C. and T. Dobzhansky. 1942. Genetics of natural populations. VI. Microgeographic races in Linanthus parryae. Genetics 27:317-332.
[populations seem to represent the action of genetic drift: fixation of one or the other of two alleles (CS)]

Fisher, R. A. 1932. The evolutionary modification of genetic phenomena. Pp. 165-172. In: Proceedings of the Sixth International Congress of Genetics, Vol. 1. D. F. Jones (ed.).
[go to http://www.esp.org/books/6th-congress/facsimile/title3.html (KS)]

Frazzetta, T. H. 1970. From hopeful monsters to bolyerine snakes? Amer. Nat. 104:55-72.
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Gottlieb, L. D. 1984. Genetics and morphological evolution in plants. American Naturalist 123:681-709.
[The genetic basis of differences in morphology within and between flowering plant species is reviewed in order to elucidate how many genetic changes are responsible for the evolution of new characters. Two broad morphological categories are evident. Differences in structure, shape, orientation, and presence versus absence are frequently discrete and appear to be governed by one or two genes. Differences in dimensions, weight, and number usually exhibit continuous variation and are influenced by numerous genes, though many of them probably act only indirectly via general effects at the whole organ or whole plant levels. Although it is difficult to specify the relative contributions of the two morphological categories during evolutionary divergence, it is clear that discrete character differences are more common in plants than in animals. I propose that their prevalence in plants is a direct consequence of the open, less integrative, and plastic patterns of plant morphogenesis which permit large changes in morphology on the basis of relatively few genetic changes. Morphological divergence among genera or families of flowering plants may reflect many fewer genetic changes than is the case for similar taxonomic levels of higher animals. Accurate estimates of the number of genes responsible for character divergence require knowledge of the ontogenetic and anatomical details of character development and these must be coordinated with genetic analyses. Until this knowledge becomes available, general conclusions about the number of genetic changes responsible for morphological diversity are premature (CS)]

Gould, S. J. and N. Eldredge. 1977. Punctuated equilibria: the tempo and mode of evolution reconsidered. Paleobiology 3:115-151.
[A formalization and expansion of the concept introduced by Eldredge and Gould in 1972 as Gould gradually makes the concept his own. A valiant attempt to find a raprochement between microevolution/population-level phenomena and macroevolutionary patterns revealed in the fossil record. The concept remains contentious - for example, see Levinton and Simon critique below (KS)]
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Levinton, J. S., and C. M. Simon. 1980. A Critique of the punctuated equilibria model and implications for the detection of speciation in the fossil record. Systematic Zoology 29:130-142. PDF
[our own Chris Simon critiqued the punctuated equilibria model on several levels, including its restriction of speciation models to peripheral isolates, the confounding of species identification in the fossil record with 'stasis' and the assertion that species selection is random with respect to phenotypic trends (KS)]

Gould, S. J. and R. C. Lewontin. 1979. The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme. Proceedings of the Royal Society of London. Series B. 205:581-598.
[Whatever you think of this paper, there is no denying that it was absolutely seminal; probably the most critiqued paper ever published in terms of its cross-disciplinary appeal - including an entire edited volume analyzing it from a rhetorical perspective! It marks the beginning of the modern era of constraint theory (KS)
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Queller, D. C. 1995. The spaniels of St. Marx and the Panglossian paradox: A critique of a rhetorical programme. Quart. Rev. Biology 70:485-489. PDF
[one of many critiques published about G & L – an indication of its seminal influence (KS)]
Cain, A. J. 1964. The perfection of animals. Pp. 36-63. In: Viewpoints in Biology. Carthy, J. D. and C. L. Duddington (eds.). Butterworths, London. PDF
[reprinted in Biol. J. Linn. Soc. 36:3-29 (1989); G & L are often criticized for creating a ‘straw man’ in the ‘adaptationist programme’, but this paper exemplifies some of the extreme adaptationist thinking of the time (KS)]
Du Brul, E. L. and H. Sicher. 1954. The Adaptive Chin. American Lecture Series, Publication No. 180. Charles C Thomas, Springfield. (Schwenk)
[G & L refer to the human chin as their “favorite example” of incorrect identification of an atomized character. This monograph provides the foundation for their views (the chin as an integrated field, not a ‘thing’). Gould cites it in his 1977 book, but G & L do not, despite referring to it implicitly (KS)]

Gould, S. J. and E. S. Vrba. 1982. Exaptation—a missing term in the science of form. Paleobiology 8:4-15.
[turns out it really wasn’t missing – it was just called ‘preadaptation’, which, for reasons they never fully justify, they eschew. They build a new vocabulary around the term ‘aptation’ to distinguish past vs. present function and selection. Although their terminology is sometimes applied, it is often done so self-consciously, i.e., a term such as "exaptation" is used in a sentence, but it is alwayts followed by the citation, as such: (sensu Gould and Verba, 1982) - a sign that the language hasn't really caught on and become part of our evolutionary vernacular (KS)]
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Grant, P. R. and B. R. Grant. 2002. Unpredictable evolution in a 30-year study of Darwin’s Finches. Science 296:707-711.
[the one exception to the ‘too recent to qualify as a classic rule’ – the summation of 30 years work on the action of natural selection is simply too incredible and important not to include. The Grants demonstrate remarkable phenotypic lability in the beak related to climate change and its effect on food availability. Can 30 years of data be generalized to macroevolutionary patterns? —the big question (KS)]
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Haffer, J. 1969. Speciation in Amazonian forest birds. Science 165:131-137.
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Haldane, J. B. S. 1924. A mathematical theory of natural and artificial selection. Part I. Trans. Cambridge Phil. Soc. 23:19-41.
[In a series of 10 papers from 1924-1934, Haldane outlines the first mathematical models for many cases of evolution due to selection, an important concept in the modern evolutionary synthesis]
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Haldane, J. B. S. 1924. A mathematical theory of natural and artificial selection. Part II. The influence of partial self-fertilisation, inbreeding, assortative mating and selective fertilisation on the composition of Mendelian populations and on natural selection. Proc. Cambridge Phil. Soc. 1:158-163.
Haldane, J. B. S. 1926. A mathematical theory of natural and artificial selection. Part III. Proc. Cambridge Phil. Soc. 23:363-372.
Haldane, J. B. S. 1927. A mathematical theory of natural and artificial selection. Part IV. Proc. Cambridge Phil. Soc. 23:607-615.
Haldane, J. B. S. 1927. A mathematical theory of natural and artificial selection. Part V. Selection and mutation. Proc. Cambridge Phil. Soc. 23:838-844.
Haldane, J. B. S. 1930. A mathematical theory of natural and artificial selection. Part VI. Isolation. Proc. Cambridge Phil. Soc. 26:220-230.
Haldane, J. B. S. 1931. A mathematical theory of natural and artificial selection. Part VII. Selection intensity as a function of mortality rate. Proc. Cambridge Phil. Soc. 27:131-136.
Haldane, J. B. S. 1932. A mathematical theory of natural and artificial selection. Part VIII. Metastable populations. Proc. Cambridge Phil. Soc. 26:220-230.
Haldane, J. B. S. 1932. A mathematical theory of natural and artificial selection. Part IX. Rapid selection. Proc. Cambridge Phil. Soc. 28:244-248.
Haldane, J. B. S. 1934. A mathematical theory of natural and artificial selection. Part X. Some theorems in artificial selection. Genetics 19:412-429. PDF

Haldane, J. B. S. 1932. The time of action of genes, and its bearing on some evolutionary problems. American Naturalist 66:5-24.
[points out the importance of knowledge about the age/stage of gene expression - gametophytes and gametes to zygotes, embryos and immature and mature organisms (CS)]
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Haldane, J. B. S. 1932. Can evolution be explained in terms of known genetical facts? Pp. 185-189. In: Proceedings of the Sixth International Congress of Genetics, Vol. 1. D. F. Jones (ed.).
[see http://www.esp.org/books/6th-congress/facsimile/title3.html (KS)]

Haldane, J. B. S. 1957. The cost of natural selection. Journal of Genetics 55:511-524.
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Hamilton, W. D. 1964. The genetical evolution of social behavior. 1. J. Theor. Biol. 7:1-16.

Hamilton, W. D. 1964. The genetical evolution of social behavior. 2. J. Theor. Biol. 7:17-52.

Hamilton W. D. and M. Zuk. 1982. Heritable true fitness and bright birds: a role for parasites? Science 218:384-387.
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Hardy, G. H. 1908. Mendelian proportions in a mixed population. Science 28:49-50.
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Huxley, J. S. 1924. Constant differential growth-ratios and their significance. Nature 114:895-896.

Jacob, F. 1977. Evolution and tinkering. Science 196:1161-1166.
[A wonderful 'perspective' essay that puts together many modern themes around the notion of "hierarchy" - ahead of its time (KS)]
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Kimura, M. 1968. Evolutionary rate at the molecular level. Nature 217:624-626.

King, J. L. & Jukes, T. H. 1969. Non-Darwinian evolution. Science 164:788-798.
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King, M.-C. & Wilson, A. C. 1975. Evolution at two levels in humans and chimpanzees. Science 188: 107-116.
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Lande, R. 1982. A quantitative genetic theory of life history evolution. Ecology 63:607-615.
[Dynamic models of quantitative (polygenic) characters are more generally applicable in the analysis of life history evolution than are static optimization methods or one and two locus genetic models. A dynamic theory of life history evolution is derived by synthesizing population demography with quantitative genetics. In a population under weak selection with a nearly stable age distribution, the relative fitness of individuals with a particular life history phenotype can be approximated as an average of age-specific relative fecundity and mortality rates, weighted respectively by the present productivity and future reproductive value of each age-class. An adaptive topography is constructed showing that, with phenotype- and age-specific fecundity and mortality rates constant in time, evolution of the mean life history maximizes the intrinsic rate of increase of a population. However, the rate and direction of evolution in response to selection are strongly influenced by genetic correlations among characters. Negative genetic correlations among major components of fitness are often obscured phenotypically by positive environmental correlations, but commonly constitute the ultimate constraint on life history evolution, as illustrated by artificial selection experiments. Methods are suggested for measuring selective forces and evolutionary constraints that effect life history characters in natural populations (CS)]
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Lande, R. and S. J. Arnold. 1983. The measurement of selection on correlated characters. Evolution 37:1210-1226.
[reanalysis of Bumpus' data (CS)]
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Lewontin, R. C. 1957. The adaptations of populations to varying environments. Cold Spring Harbor Symposium on Quantitative Biology 22:395-408.
[homeostasis of populations and individuals (CS)]

Lewontin, R. C. 1978. Adaptation. Scientific American 239:212-228.
[strangely enough, this article published in the popular press Scientific American is one of the best single treatments of this critical, yet slippery, concept (KS)]
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Lewontin R. C. and L. C. Birch. 1966. Hybridization as a source of variation for adaptation to new environments. Evolution 20: 315-336.
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Lewontin, R. C. and J. L. Hubby. 1966. A molecular approach to the study of genic heterozygosity in natural populations. II. Amount of variation and degree of heterozygosity in natural populations of Drosophia pseudoobscura. Genetics 54:595-609.
[a pretty boring and seemingly unremarkable paper to read now, but it marks the beginning of molecular population genetics (KS)]

Maynard Smith, J. 1966. Sympatric speciation. American Naturalist 100:637-650.
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Maynard Smith, J., R. Burian, S. Kauffman, P. Alberch, J. Campbell, B. Goodwin, R. Lande, D. Raup and L. Wolpert. 1985. Developmental constraints and evolution: a perspective from the Mountain Lake Conference on Development and Evolution. Quart. Rev. Biol. 60:265-287.
[an important and oft-cited work on evolutionary/developmental constraint marred by internal contradictions reflecting its having been ‘written by committee’ (KS)]
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Mayr, E. 1940. Speciation phenomena in birds. American Naturalist 74: 249-278.
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Mayr, E. 1949. Speciation and selection. Proc. Amer. Phil. Soc. 93:514-519.

Mayr, E. 1949. Speciation and systematics. Proc. Pp. 281-298, In: Jepsen, G. L., G. G. Simpson and E. Mayr (eds.), Genetics, Paleontology and Evolution. Princeton Univ. Press, Princeton, NJ. (Schwenk)
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Mayr, E. 1954. Change of genetic environment and evolution. Pp. 157-180. In: Evolution as a Process. J. Huxley, A. C. Hardy & E. B. Ford (eds.). Allen and Unwin, London.
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Mayr, E. 1981. Biological classification: toward a synthesis of opposing methodologies. Science 214:510-516.
[written at the height of the 'classification wars', this paper makes the case for 'evolutionary taxonomy' - a hybrid approach that allows for cladistic methods in phylogeny construction, but which aims for classifications to emphasize morphological differences by permitting paraphyletic groups, e.g., a 'Reptilia' that excludes birds (KS)]
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Northcutt, R. G., and C. Gans. 1983. The genesis of neural crest and epidermal placodes: a reinterpretation of vertebrate origins. Quarterly Review of Biology 58:1-28.
[okay, this one is really of most interest to vertebrate biologists, but it is a fantastic story about the origin of a novel tissue type/germ layer - the neural crest- and how it is almost single-handedly responsible for the craniate head - a true novelty and complex structure if ever there was one (KS)]
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Orr, H. A., and J. A. Coyne. 1992. The genetics of adaptation: a reassessment. American Naturalist 140:725-742.
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Raup, D. M. 1961. The geometry of coiling in gastropods. Proceedings of the National Academy of Sciences USA 47:602-609.
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Raup, D. M. 1966. Geometric analysis of shell coiling: general problems. Journal of Paleontology 40:1178-1190.
[critically important paper for constraint theory—although ironically, Raup does not invoke constraint himself; Raup formalizes the notion of potential morphospace as a way of assessing observed biodiversity (KS)]
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Riedl, R. 1977. A systems-analytical approach to macro-evolutionary phenomena. Quart. Rev. Biol. 52:351-370.
[a must read for complex systems/phenotypes/constraint freaks (KS)]
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Roth, V. L. 1984. On homology. Biol. J. Linn. Soc. 22:13-29. (Schwenk)
[the best synthesis of the homology concept to that time; the starting point for many subsequent treatments (KS)]

Stebbins, G. L., Jr. 1949. Reality and efficacy of selection in plants. Proc. Amer. Phil. Soc. 93:501-513.

Stern, C. 1943. The Hardy-Weinberg law. Science 97:137-138.
[have you ever wondered why it's called the 'Hardy-Weinberg' principle? (KS)]
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Trivers, R. L. 1971. The evolution of reciprocal altruism. Quarterly Review of Biology 46:35-57.
[have you ever wondered why it's called the 'Hardy-Weinberg' principle? (KS)]
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Trivers, R. L. 1974. Parent-offspring conflict. American Zoologist 14:249-264.

Turesson, G. 1922. The genotypical response of the plant species to the habitat. Hereditas 3:211-350.

Van Valen, L. 1973. A new evolutionary law. Evolutionary Theory 1:1-30.
[the ‘red queen hypothesis’- the idea that you must evolve just to stay in place ("for an evolutionary system, continuing development is needed just in order to maintain its fitness relative to the systems it is co-evolving with"); predator-prey arms races are a specific example of the red queen phenomenon (KS)]

Vermeij, G. 1974. Adaptation, versatility, and evolution. Systematic Zoology 22:466-477.

Weldon, W. F. R. 1901/1902. A first study of natural selection in Clausilla laminata (Montagu). Biometrika 1:109-115.

Williams, G. C. 1957. Pleiotropy, natural selection and the evolution of senescence. Evolution 11:398-411.
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Wright, S. 1931. Evolution in Mendelian populations. Genetics 16:97-159.
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Wright. S. 1932. The roles of mutation, inbreeding, crossbreeding and selection in evolution. Pp. 356-366. In: Proceedings of the Sixth International Congress of Genetics, Vol. 1. D. F. Jones (ed.).
[A terrific paper that introduces several fundamental concepts including Wright's famous 'adaptive landscape' metaphor; you can get open access HERE (KS)]

Wright, S. 1951. The genetical structure of populations. Annals of Eugenics 15: 323-354.


Comment: The list above is reserved for actual scientific contributions in the history of post-Darwinian evolutionary biology—it does not contain secondary sources, i.e., books and papers primarily about the history and philosophy of evolution. There are many such contributions, particularly histories of Darwinism and neo-Darwinism (the Synthesis). Some of the latter have been written by historians and philosophers of biology, some by biologists—notably Ernst Mayr’s various historical treatments of the Synthesis. William Provine is another important author in this area who is much less dogmatic. However, we are presently in the midst of what might eventually be interpreted as a Kuhnian ‘paradigm shift’ in evolutionary biology, with the roles of development, ontogeny and organismal phenotype playing an increasingly important part in our views about evolutionary mechanisms and patterns. The modern evo-devo movement, though often thought of as ‘new’, is actually based on a very old tradition originating in the early 19th century, primarily from the German school of evolutionary and developmental morphology ('transcendental morphology', e.g., Haeckel, von Baer), carried into the 20th century by Baldwin, de Beer and Waddington, to name a few. There is ongoing debate about the importance of evo-devo in a general theory of evolution—one extreme views it as virtually irrelevant and that the genetic-population-selection principles of the neo-Darwinian Synthesis are both necessary and sufficient to account for phenotypic—'macroevoluion' is simply the extrapolation of microevolutionary processes over deep time. The opposite extreme views selection as a minor player in phenotypic evolution and posits that developmental ‘rules of form’ have primacy. It is likely, as usual, that the truth lies somewhere in between. The particular books listed below ae notable because they deal with the history of modern evolutionary theory explicitly from the vantage point of evo-devo, and as such, provide novel perspectives and analyses. They also discuss historical figures usually ignored or even denigrated in ‘traditional’ histories. Beware historical revisionism in Science, as well as politics—it is up to us to sift through the various views to see which is consonant with the primary literature. Of course, this is not always easy to do—for example, not everyone can read through von Baer’s (1828) 300+ page Über Entwickelungsgeschichte der Thiere: Beobachtung und Reflexion (“On the Developmental History of Animals: Observations and Reflection”).

Amundson, R. 2005. The Changing Role of the Embryo in Evolutionary Thought. Roots of Evo-Devo. Cambridge Studies in Philosophy and Biology. Cambridge Univ. Press, Cambridge (Schwenk)
[Ron Amundson is an extraordinarily thoughtful and insightful historian/philosophers of biology. His essays on adaptation and constraint are top notch. I expect this book is the same (KS)].

Richards, R. J. 1992. The Meaning of Evolution. The Morphological Construction and Ideological Reconstruction of Darwin’s Theory. Univ. of Chicago Press, Chicago. (Schwenk)
[Richards provides an excellent short history of the role and importance of the (mostly) German school of morphology and evolutionary morphology/development and its role in the formation of evolutionary theory—a perspective that is not easy to come by and which speaks to the modern evo-devo movement. An alternative to the Mayrian neoDarwinian, revisionist view of evolutionary history.]

Richards, R. J. 2002. The Romantic Conception of Life. Science and Philosphy in the Age of Goethe. Univ. of Chicago Press, Chicago.

Modern Classics?

Schlichting: A classic in the making?

Gould, S. J. 2002. The Structure of Evolutionary Theory. Belknap/Harvard Univ. Press, Cambridge, MA.
[so massive it's got to be a classic... synthesizes all of Gould's favorite themes—theory, history, hierarchy, punc. equil., macroevolution, constraint, adaptation]

Schlichting, C. and M. Pigliucci (1998) Phenotypic Evolution. A Reaction Norm Perspective. Sinauer, Sunderland, MA.
[Carl intends to hang on long enough to see the day this is moved up to the main list... of course, he might take advantage of EEBedia's editing capacity to slip it up there late one night when no one is looking. Go HERE to buy it now! (KS)]

Günter Wagner in Vienna, 2005, with tiny, glaring, out-of-focus Kurt in the background...

Schwenk, K., and G. P. Wagner (2004) The relativism of constraints on phenotypic evolution. Pp. 390-408. In: Phenotypic Integration: Studying the Ecology and Evolution of Complex Phenotypes. M. Pigliucci & K. Preston (eds.). Oxford Univ. Press, Oxford. PDF
[the absolute final word on evolutionary constraint—you can take my word on this (KS)]

Massimo Pigliucci, EEB PhD - classy if not yet classic.

Wagner, G. P., and L. Altenberg (1996) Complex adaptations and the evolution of evolvability. Evolution 50:967-976. PDF
[The merging of evolutionary biology and computer science. An extremely important paper on the evolution of evolvability through modularity at different hierarchical levels; introduces the terms genotype-phenotype map and variability, meaning 'the tendency to vary (as in 'solubility'). An absolute must-read for grad students with an evolutionary or evo-devo bent (KS)]

Questions About This Page?

Contact Kurt Schwenk (kurt.schwenk@uconn.edu) or EEB Webmaster, Paul Lewis (paul.lewis@uconn.edu)


This page would not be possible without the hard work of Paul Lewis who created and maintains EEBedia. Paul devoted considerable additional time to helping me learn the ins-and-outs of EEBedia and even modifying it to make the page do exactly what I wanted it to do. I also thank my colleagues who made suggestions of books and papers for inclusion in the Master List (even if, in the end, their nominations didn't make the final cut): Eldridge Adams, Andy Bush, John Cooley, Charlie Henry, Carl Schlichting, Chris Simon and Gregor Yanega. Finally, I thank the students in EEB 485, spring 2008, for their enthusiastic participation, advocacy of this web page and for teaching me a lot: Dave Hudson, Diego Sustaita, Rogerta Engel, Florian Reyda, Frank Smith, Jim Freyermuth, Jason Hill, Kat Shaw, Nic Tippery, Maria Pickering, Thiago Rangel, Tobias Landberg and Trina Schneider. --KS