Ever wondered whether you had completely missed some of the most important papers in your discipline? Or whether you just read enough? Well, now you can’t stop wondering, since the answer is right here in this new post. About our latest paper, a paper that recommends to read recommended papers.

In ecology. Yeah, I know, the title doesn’t specify “in ecology”. And it should, since a list of ecology papers is going to be of no interest whatsoever for you guys in astrophysics or neurobiology. Plus, the Sheldon Coopers and Amy Farrah Fowlers among you are now probably going to smirk about our classics. My official excuse is that you should always try to have as short a title as possible, in order to be attractive (after all, we are living in an era of unsalvageable lazy millennials). But the real reason is that I wanted to give my blog a little boost, after months of abstinence, so that was on purpose. But instead of frowning with your judgmental scorn, please consider that I didn’t put sex, GoT or Trump in the title, be merciful, and go forward to all your friends.

Now that you’ve made a healthy re-acquaintance with my annoying habit to not-cut-to-the-chase, I should probably start. After all, rule#1 for a successful blog: short posts (see one of my first entries).

For a few years, I’ve been wondering whether I was missing the important papers, and more worryingly, if my students were. There are now so many papers to read, and so little time to do it, it’s easy to stay confined within a small niche of papers – your area of expertise – and miss the big picture, those papers that made your field, and from which the wise professors probably get part of their wisdom.

So, I have been thinking for quite some time of the best way to come up with such a list. It was not easy, because important papers are a very subjective thing to select, let alone rank. But I came up with a simple solution: ask the wise professors. Or more exactly, ask the 665 experts in the Editorial Board of the highest ranking, generalist journals in ecology, who probably are the best suited to evaluate the worth of papers regardless of their field. After receiving all their nominees, an internet vote and clever statistical analyses by my brilliant co-author and good friend Corey Bradshaw, at the time in sabbatical in my group, we came up with …

(hint: click on the image to get the list – I really must tell you everything…)

This came up with a few surprises, such as the discrepancy between the articles that experts recommend to students and those they have actually read themselves, the fact that the average scientist reads ~40 papers per month (if you thought that maybe you were lazy, now you know for sure), or the huge gender bias in authors of said articles, but, damned, I don’t have any space left (nor you any patience left) to discuss that. I really should learn to focus on the important stuff. Well, this said, for those you interested in the full story, it is now published in Nature Ecology & Evolution. As for the pdfs of those articles, I’m sure they somehow will be found on SciHub…

Ok, remember, you’re supposed to read at least 40 papers per month, so the 100 papers’ list is not going to be a huge additional load in your PhD. So, don’t blame us and go start reading your share. And no, this post doesn’t count as a reading.


Oh, and if you find one or several such papers were utterly useless to you, don’t blame me for choosing them, I didn’t. Don’t even blame me for making you read them, I didn’t either…


The 100 selected articles:

  1. Darwin, C.R.; Wallace, A.R. 1858. On the tendency of species to form varieties; and on the perpetuation of varieties and species by natural means of selection. Zoological Journal of the Linnean Society 3:45-62
  2. Hardin, G. 1960. The competitive exclusion principle. Science 131:1292-1297
  3. Paine, R.T. 1966. Food Web Complexity and Species Diversity. The American Naturalist 100:65-75
  1. Hutchinson, G.E. 1961. The Paradox of the Plankton. The American Naturalist 95:137-145
  2. Hutchinson, G.E. 1959. Homage to Santa Rosalia or Why Are There So Many Kinds of Animals? The American Naturalist 93:145
  3. MacArthur, R.H.; Wilson, E.O. 1963. An Equilibrium Theory of Insular Zoogeography. Evolution 17:373-387
  1. Hutchinson, G.E. 1957. Concluding Remarks. Cold Spring Harbor Symposia on Quantitative Biology 22:415-427
  2. Hairston, N.G.; Smith, F.; Slobodkin, L. 1960. Community structure, population control, and competition. The American Naturalist 94:421-425
  1. Connell, J.H. 1978. Diversity in tropical rain forests and coral reefs. Science 199:1302-1310
  2. Janzen, D.H. 1970. Herbivores and the Number of Tree Species in Tropical Forests. The American Naturalist 104:501
  3. May R.M. 1974. Biological populations with non-overlapping generations: stable points, stable cycles, and chaos. Science 186:645-647
  4. Gause, G.F. 1934. Experimental Analysis of Vito Volterra’S Mathematical Theory of the Struggle for Existence. Science 79:16-17
  5. Chesson, P. 2000. Mechanisms of Maintenance of Species Diversity. Annual Review of Ecology and Systematics 31:343-366
  1. Carpenter, S.R.; Kitchell, J.F.; Hodgson, J.R. 1985. Cascading trophic interactions and lake productivity. BioScience 35:634-639
  2. Levin, S.A. 1992. The problem of pattern and scale in ecology: the Robert H. MacArthur Award lecture. Ecology 73:1943-1967
  3. Hanski, I. 1998. Metapopulation dynamics. Nature 396:41-49
  4. MacArthur, R.; Levins, R. 1967. The Limiting Similarity, Convergence, and Divergence of Coexisting Species. The American Naturalist 101:377-385
  5. Tilman, D. 1977. Resource Competition Between Plankton Algae: An Experimental and Theoritical Approach. Ecology 58:338-348
  6. Hamilton, W.D. 1964a. The genetical evolution of social behaviour. I. Journal of Theoretical Biology 7:42370
  7. Charnov, E.L. 1976. Optimal foraging, the marginal value theorem. Theoretical Population Biology 9:129-136
  8. Tilman, D. 1996a. Biodiversity: Population versus ecosystem stability. Ecology 77:350-363
  9. Rosenzweig, M. 1971. Paradox of enrichment: destabilization of exploitation ecosystems in ecological time. Science 171:385-387
  10. Connell, J.H. 1961. The Influence of Interspecific Competition and Other Factors on the Distribution of the Barnacle Chthamalus Stellatus. Ecology 42:710-743
  11. MacArthur, R.; Levins, R. 1964. Competition, habitat selection, and character displacement in a patchy environment. Proceedings of the National Academy of Sciences of the United States of America 51:1207-1210
  12. Hardin, G.J. 1968. The tragedy of the commons. Science 162:1243-1248
  13. Levin, S.A. & Paine, R.T. 1974. Disturbance, patch formation, and community structure. Proceedings of the National Academy of Sciences of the United States of America 71:2744-2747
  14. Felsenstein, J. 1981. Skepticism towards Santa Rosalia, or why are there so few kinds of animals? Evolution 35:124-138
  15. Tilman, D. 1994a. Competition and biodiversity in spatially structured habitats. Ecology 75:42401
  16. Holling, C.S. 1973. Resilience and Stability of Ecological Systems. Annual Review of Ecology and Systematics 4:44927
  17. Hurlbert, S.H. 1984. Pseudoreplication and the Design of Ecological Field Experiments. Ecological Monographs 54:187
  18. Vitousek, P.M. et al. 1997b. Human Domination of Earth’s Ecosystems. Science 277:494-499
  19. May R.M. 1972. Will a large complex system be stable? Nature 238:413-414
  20. Pianka, E.R. 1970. On r- and K-selection. American Naturalist 104:592-597
  21. Brown, J.H. et al. 2004. Toward a metabolic theory of ecology. Ecology 85:1771-1789
  22. Ehrlich, P.R.; Raven, P.H. 1964. Butterflies and plants: a study in coevolution. Evolution 18:586-608
  23. MacArthur, R.H.; McArthur, J. 1961. On bird species diversity. Ecology 42:594-598
  24. Simberloff, D.S. et al. 1969. Experimental Zoogeography of Islands: The Colonization of Empty Islands. Ecology 50:278-296
  25. Grime, J.P. 1977. Evidence for the existence of three primary strategies in plants and its relevance to ecological and evolutionary theory. The American Naturalist 111:1169-1194
  26. Brown, J.H. 1984. On the Relationship between Abundance and Distribution of Species. The American Naturalist 124:255
  27. Connell, J.H. 1961a. Effects of competition, predation by Thais lapillus, and other factors on natural populations of the barnacle Balanus balanoides. Ecological Monographs 31:61-104
  28. Holt, R.D. 1977. Predation, apparent competition, and the structure of prey communities. Theoretical Population Biology 12:197-229
  29. Anderson, R.M; May, R.M. 1979. Population biology of infectious diseases: Part I. Nature 280:361-367
  30. Huffaker, C.B. 1958. Experimental studies on predation: dispersion factors and predator-prey oscillations. Hilgardia 27:343-383
  31. Clements, F.E. 1936. Nature and structure of the climax. Journal of Ecology 24:252-284
  32. Pulliam, D.W. 1988. Sources, Sinks, and Population Regulation. The American Naturalist 132:652-661
  33. Lawton, J.H. 1999. Are there general laws in ecology? Oikos 84:177-192
  34. Lindeman, R.L. 1942. The trophic-dynamic aspect of ecology. Ecology 23:399-418
  35. Kimura, M. 1968. Evolutionary Rate at the Molecular Level. Nature 217:624-626
  36. May R.M. 1976. Simple mathematical models with very complicated dynamics. Nature 261:459-467
  37. Trivers, R.L. 1974 Parent-Offspring Conflict. American Zoologist 14:249-264
  38. Paine, R.T. 1980. Food Webs: Linkage, Interaction Strength and Community Infrastructure. Journal of Animal Ecology 49:666-685
  39. Tilman, D.; Wedin, D.; Knops, J. 1996. Productivity and sustainability influenced by biodiversity in grassland ecosystems. Nature 379:718-720
  40. MacArthur, R.H. 1958. Population ecology of some warblers of northeastern coniferous forests. Ecology 39:599-619
  41. May R.M. 1977. Thresholds and breakpoints in ecosystms with a multiplicity of stable states. Nature 260:471-477
  42. Simberloff, D. 1976. Experimental Zoogeography of Islands : Effects of Island Size. Ecology 57:629-648
  43. Schindler, D.W. 1977. Evolution of phosphorus limitation in lakes. Science 195:260-262
  44. Kunin, W.E.; Gaston, K.J. 1993. The biology of rarity: Patterns, causes and consequences. Trends in Ecology & Evolution 8:298-301
  45. Vitousek, P. M.; Reiners W.A. 1975. Ecosystem succession and nutrient retention: a hypothesis. BioScience 25:376-381
  46. Tilman, D. 1980. Resources: a Graphical-Mechanistic Approach To Competition and Predation. The American Naturalist 116:362-393
  47. Lande, R. 1980. Sexual dimorphism, sexual selection, and adaptation in polygenic characters. Evolution 34:292-305
  48. Tilman, D. et al. 1994. Habitat destruction and the extinction debt. Nature 371:65-66
  49. Fretwell S.D. & Lucas H.L. 1970. On territorial behavior and others factors influencing habitat distribution in birds. I. Theoretical development. Acta Biothereotica 19:16-36
  50. May R.M. 1973a. Qualitative stability in model ecosystems. Ecology 54:638-641
  51. Redfield, A.C. 1958. The biological control of chemical factors in the environment. American Scientist 46:205-221
  52. Tilman, D. et al. 1997. The Influence of Functional Diversity and Composition on Ecosystem Processes. Science 277:1300-1302
  53. Hamilton, W.D. 1967. Extraordinary Sex Ratios. Science 156:477-488
  54. Schluter, D. & McPhail, J.D. 1992. Ecological character displacement and speciation in sticklebacks. The American Naturalist 140:85-108
  55. Hanski, I. 1994. A practical model of metapopulation dynamics. Journal of Animal Ecology. 63:151–162
  56. Hamilton, W.D. 1964b. The genetical evolution of social behaviour. II. Journal of Theoretical Biology 7:17-52
  57. Likens, G.E. et al. 1970. Effects of Forest Cutting and Herbicide Treatment on Nutrient Budgets in the Hubbard Brook Watershed-Ecosystem. Ecological Monographs 40:23-47
  58. Odum, E.P. 1969. The strategy of ecosystem development. Science 164:262-270
  59. Hubbell, S.P. 1979. Tree dispersion, abundance, and diversity in a tropical dry forest. Science 203:1299-1309
  60. Grinnell, B.Y. 1917. The niche-relationships of the california thrasher. The Auk 34:427-433
  61. MacArthur, R.H.; Pianka, E. R. 1966. On optimal use of a patchy environment. American Naturalist 100:603-609
  62. Tilman, D.; Forest, I.; Cowles, J.M. 2014. Biodiversity and ecosystem functioning. Annual Review of Ecology, Evolution, and Systematics 45:471-493
  63. May, R.M. & MacArthur, R.H. 1972a. Niche overlap as a function of environmental variability. Proceedings of the National Academy of Sciences of the United States of America 69:1109-1113
  64. Leibold, M.A. et al. 2004. The metacommunity concept: a framework for multi-scale community ecology. Ecology Letters 7:601-613
  65. Axelrod, R.; Hamilton, W. D. 1981. The Evolution of Cooperation. Science 211:1390-1396
  66. Gleason, H.A. 1926. The Individualistic Concept of the Plant Association. Bulletin of the Torrey Botanical Club 53:46204
  67. Grime, J.P. 1998. Benefits of plant diversity to ecosystems: immediate, filter and founder effects. Journal of Ecology 86:902-910
  68. Gould S.J.; Lewontin R.C. 1979. The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptionist programme. Proceedings of the Royal Society B: Biological Sciences 205:581-5981017
  69. Grant, P.R; Grant, B.R. 1995. The Founding of a New Population of Darwin’s Finches. Evolution 49:229-240
  70. Stearns, S.C. 1976. Life-history tactics: a review of the ideas. The Quarterly Review of Biology 51:3
  71. Vitousek, P.M. 1994. Beyond global warming: ecology and global change. Ecology 75:1861-1876
  72. Janzen D.H. 1967. Why mountain passes are higher in the tropics. The American Naturalist 101:233
  73. Carpenter, S.R. et al. 1987. Regulation of lake primary productivity by food web structure. Ecology 68:1863-1876
  74. Stenseth, N.C. 1997. Population regulation in snowshoe hare and Canadian lynx: asymmetric food web configurations between hare and lynx. Proceedings of the National Academy of Sciences of the United States of America 94:5147-5152
  75. Anderson, R.M; May, R.M. 1978. Regulation and Stability of Host-Parasite Population Interactions. Journal of Animal Ecology 47:219-247
  76. Krebs, C.J. et al. 1995. Impact of Food and Predation on the Snowshoe Hare Cycle. Science 269:1112-1115
  77. Ginzburg, L.R.; Jensen, C.X.J. 2004. Rules of thumb for judging ecological theories. Trends in Ecology and Evolution 19:121-126
  78. Chave,J. 2013. The problem of pattern and scale in ecology: what have we learned in 20 years? Ecology Letters 16:42461
  79. MacArthur, R. 1955. Fluctuations of Animal Populations and a Measure of Community Stability. Ecology 36:533
  80. Ricklefs, R.E. 1987. Community diversity: relative roles of local and regional processes. Science 235:167-171
  81. Levins, R. 1966. The strategy of model building in population biology. American Scientist 54:421-431
  82. Anderson, R.M; May, R.M. 1981. The Population Dynamics of Microparasites and Their Invertebrate Hosts. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 291:451-524.
  83. Brown, W.L.; Wilson, E.O. 1986. Character displacement. Systematic Zoology 5:49-64
  84. Lande, R. 1993. Risks of Population Extinction from Demographic and Environmental Stochasticity and Random Catastrophes. The American Naturalist 142:911-927
  85. May R.M. & Anderson, R.M. 1979. Population biology of infectious diseases: Part II. Nature 280:455-461
  86. Parmesan, C.; Yohe, G. 2003. A globally coherent fingerprint of climate change impacts across natural systems. Nature 421:37-42
  87. Power, M.E. 1990. Effects of fish in river food webs. Science 250:811-81



PS: if you want the pdf of the 545 nominated articles – including the 100 – you may find them here.



  1. Alexandre Terrigeol says:

    Thank you for this interesting list, even if I personally feel bad to see that I only read 2 or 3 of the articles listed. This will ne longer be the case as I plan to read them now. Even if a post had been created asking people how they were reading articles, it would be interesting to know how you do it, as i expect people to perform very differently!


  2. […] If you can access the article directly, then the 100 top-ranked list is available in a table (Box 1), and Franck Courchamp has provided the full, ranked list on his blog. […]


  3. These papers are only representative of ecology to the extent that you believe the handling editors of some of the more prominent ecological journals are representative of ecology. I’ve not done the study, but I strongly suspect the population of handling editors that was surveyed to identify papers was heavily biased towards white males in North America and Europe. This suspicion is reinforced by a quick look at the final list of papers, which generally lack representation by females, minorities, and researchers from entire geographic regions of the world.

    Would it be any surprise that when you survey white males from rich countries that they respond by saying their own work is the most important?

    Frankly, I won’t be having my students waste their time on this list, as I don’t find it to be a good representation of the field. It comes across to me as a bit self-serving.


    • You say that on the postulate that white males from rich countries have a sense of belonging and corporatism that would make them select as reading commendation only papers from people who look like them (and who were active 40 years ago, the average age of the 100 papers…). This is nonsensical. I don’t look at the gender of first authors (or their skin color) when I read a paper. Classical (often older) papers are indeed often authored by white males in North America and Europe because most papers at the time were, period. If the list had ended up with a perfect gender+racial+country blend, I would find it highly suspicious.
      Plus, I strongly doubt that your students would waste their time browsing through this list. It is not my opinion, it is the consensual opinion of the 665 editorial members asked for recommendations. You are of course free to ignore it; hopefully your students won’t 😉


      • Bradley J Cardinale says:

        Dear Franck,

        In your article, you wrote “Our aim was to collate a list of objectively chosen and ranked
        seminal papers deemed to be of major importance in ecology …” This statement is demonstrably false. What you produced was a sample that was a biased representation of viewpoints from handling editors of ecologically-oriented journals. It is well-established that this population is not representative of the discipline – therefore, you did not produce an objectively chosen selection of papers from the field as claimed in the article.

        You now have a lot of people, including an entire movement of women in science #WomenInSTEM, who are offended by your lack of attention to these biases in your paper, and upset by your attempts to sell the list as if it’s representative of ecology. In addition, we recently learned that you were well aware of these biases in your original paper, because you and co-author have acknowledged that you have a second paper coming out that explicitly details some of these biases. So you published the original paper knowing very well that it was a non-representative sample of the discipline, and then you justified the publication of a non-representative sample with a second paper acknowledging the biases. That has come across to some as unethical.

        Your responses to these concerns have been nothing but defensive. My suggestion is that you and co-author should own up to the problems, acknowledge the concerns of your critics, and work with them to fix this and produce a better list. Everyone makes mistakes. But reputable scientists fix them.



      • Brad,
        I hesitated before approving your response, because I think it puts a dirty smear on my very pretty blog. But I don’t want to be accused of censorship, and frankly your blind accusations and patronizing tone got the better of me…

        You are right that no such list can be entirely objective. We tried our best, however, to limit subjectivity; and that is why we asked these experienced experts to try to avoid personal biases.

        You are free to believe that what was produced was a “biased representation of viewpoints from handling editors of ecologically-oriented journals”. Perhaps most students will be happy to have a list of recommended papers from these. I know I would have, because as biased as you say they are, they remain arguably among the best experienced to judge importance of papers. You are free to think that they are not, but you are yet to come with a better alternative.

        But that’s about the only extent were you are right.
        A lot of people are offended? They shouldn’t be. They lacked the wisdom of taking the time to mull a bit over the results and intended outcome of the paper before over-reacting. In addition, an overwhelming proportion of the well-over-1000 reactions were positive. The majority of the (minority) criticisms were unfounded, just people blindly repeating what they have read somewhere else without checking the fact themselves, like yourself here saying that we have biased and knew it, and let them be publish and published an additional paper to acknowledge the bias. You say this is unethical. Please check your facts before making such strong accusations.

        Just so that you have not completely lost your time (and mine), please note that the 665 editors we contacted contained 22% women, and this is about the proportion of women first authors in ecology these days. So, in this regard our sample was no more gender biased than the scientist population. Which arguably is what we would seek in such a survey (if that was a survey, which this is not). From that sample, fewer women responded (10% vs 26% for men), making the final sample indeed more biased. Not because we made a biased sample, but because fewer women responded. We did not anticipate this, and did not notice it until after the submission of the first ms, and therefore made a more complete analyses of this interesting fact in a separate ms (which incidentally showed that not only men but also women are men-biased, so having more women would not have changed the result that much). But that does not change the fact that the initial list of respondent remains more adequate than any alternative I can think of to propose to students a long list of recommended articles by experienced ecologists.

        We are not selling a list, as you say, we are not selling anything. We are working our asses off to benefit the younger generation of scientists, and you if are not a buyer, I won’t lose my sleep over it. Reputable scientists don’t write this type of comments, being unfair and mean, while trying to appear polite and constructive. My response is still defensive, yes, but that is because you are offensive.

        Liked by 2 people

      • Emilio Bruna says:

        I appreciate the discussion the paper has generated.

        I wouldn’t be so quick to dismiss Cardinale’s hypothesis (which you paraphrase as “white males from rich countries have a sense of belonging and corporatism that would make them select as reading commendation only papers from people who look like them”). Men self-cite their articles more than women, and there is ample evidence that where articles are published and how much they are cited – both signals of “importance” – are biased against authors from the global south. And since the editorial boards of these journals are dominated by men from US and a few Western European countries, it seems like a reasonable question to ask.

        As an aside, there was a rich tradition of ecological research outside of Western Europe and the USA 40 years ago. Just because you (or the editors surveyed) didn’t read papers by, e.g., Aziz Ab’Sáber or Arturo Gomez-Pompa doesn’t mean they didn’t exist or werent weren’t important contributions.

        I look forward to the continued discussion about your work.


      • This was not really presented as an hypothesis by Cardinale, but rather as a patronizing accusation, which was following some rather unpleasant tweets of his that got me indeed defensive.
        True, the journals I selected are representative of “the western world” (2 journals from Scandinavia, one from France, one from the UK and three from the US, plus Faculty of 1000), but the editorial members from the most renown journals in general ecology is the best “pool” of experts I could think of to conduct the survey.
        And true their editorial board is also gender biased (22% women), but that is the % of articles nowadays first-authored by women, so the bias is in our population, and with regards to our population, our pool is not further biased.
        And yes, many, many important contributions were not proposed, or voted for. By women, by scientists from the South and by white male scientists as well. Not being in the list doesn’t mean students shouldn’t read them.
        But our literature being heavily biased towards white males, it shouldn’t come as a surprise (nor as an injustice) that the most recommended of them are also biased this way.


    • Harko Werkman says:

      It appears that there’s something in the offing already, but there’s a very simple way to test the assertion that the original sampling/analysis for this paper was biased, and that is to look at the gender/location profiles in this sample compared to the profiles of the available corpus of literature in the disciplines covered. Of course one would need to account for confounding cofactors, but that shouldn’t be onerous for ecologists…

      Alternatively one could sample the papers of female and non-Anglophone authors, and assess the relative ‘importance’ of these papers compared to the current list.

      Indeed, when it comes down to it, there are any number of ways to test the validly of BJC’s grievance. I’m not sure though that conspicuous fulminations on a blog post serve to advance the work described in what is an interesting and useful paper, regardless of the opinions of some.

      Whilst I’m here, the ordinate labels for figures 1e and 1f should probably be labelled ‘Article age’ rather than ‘Log10 article age’, given the numbering of the y axis. It appears that someone has confused the logarithmic scaling of the axis with the numerical values depicted on it.


  4. […] a beer with him, I mean we had separate bottles. We didn’t actually share the same beer.)  Here is Franck’s post and here is Corey’s post. Franck’s blog post briefly identified this bias an […]


  5. […] most seminal ecology papers. They show the ranked top 100 list. The authors comment on their paper here and here. The paper predictably lit up the ecology intertubes, which is why I’m reluctant to […]


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