Classic Works in Evolutionary Biology

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

What is This Page?

This page provides a list of classic works in evolutionary biology that I (Kurt Schwenk) compiled and partially annotated. It grew out of a graduate seminar I taught in EEB (the Dept. of Ecology and Evolutionary Biology, University of Connecticut) during the spring semester, 2008. 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 Diego's presentation 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 we read during the semester, as well as additional papers included in the 'master list' of classic works in evolutionary biology that I compiled along the way. Many of these are available as pdfs on the list page.

The list 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 was never 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 in which I can hardly claim expertise. 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 (e.g., vertebrates; 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) (to pull an example out of the air...) 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 not 'classics', themselves, but are helpful in appreciating the context or significance of the listed work. This is a type of emendation/addition/annotation that would be particularly useful (see suggestions, below, about annnotations).

Who Can Use These Pages?

EEBians can download all papers on the list where available (LINK HERE) simply by logging in using your UConn Net ID (click on link at upper right corner of the page OR simply click on a pdf icon and you will be directed to log in). EEBians are also encouraged to annotate the list according to the guidelines noted below. 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 have a UConn Net ID, but are not a member of EEB, and you wish to have full access to the list (i.e., the ability to download everything), you can make your case to me ( or Paul ( for the addition of your Net ID to the user list. Permission can be granted in special cases.

Obviously, you are welcome to read this page, peruse the LIST and download a copy of the list using the link on this page, below, but you will be unable to download most of the pdfs. This is for reasons of copyright. However, a number of items on the list have links to open sources or full text documents on the web. If you have any questions or comments about the list, or wish to make annotations or suggestions for additions/deletions, please contact me directly (

Classic Works in Evolutionary Biology With Annotations and Links to PDFs

Kurt contemplates the mystery of his existence and concludes, "I didn't evolve from no stinkin' monkeys!"

Please note that the downloadable pdf version of the classics list may not be up-to-date. For the most recent version of the list and live links, see the web page.



Changing or Annotating the List

I want you.jpg
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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

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 other than JSTOR or other standard electronic databases; or in some cases there are entire books online to which 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 the list. 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.

Create your own specialty/subdiscipline list! For example, you might want to create a 'classics' list for particular topics, such as phylogenetics, behavioral evolution, or form and function. If you do this, please provide a link to this page and a link on this page to yours.


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 ( or EEB Webmaster, Paul Lewis (


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