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Wednesday, February 15th, 2012 | ISSN 1556-5696

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Photograph of Charles Darwin taken around 1874 by Leonard Darwin (source: http://commons.wikimedia.org/wiki/File:1878_Darwin_photo_by_Leonard_from_Woodall_1884.jpg)
About this week’s eSkeptic

In this week’s eSkeptic, Donald R. Prothero remembers Charles Darwin (on the occasion of what would have been his 203rd birthday this past Sunday). Prothero reminds us that it was 40 years ago this year that the most frequently cited paper in the history of paleontology was published: none other than the legendary 1972 article by Niles Eldredge and Stephen Jay Gould which proposed the “punctuated equilibrium” hypothesis. Prothero also shares some insights from his own research.
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Darwin’s Legacy

by Donald R. Prothero

Niles Eldredge

Niles Eldredge as he looks today, Curator Emeritus at the American Museum of Natural History

Last Sunday, February 12, we celebrated the 203rd birthday of two of the most important figures in world history, Abraham Lincoln—and Charles Darwin. To mark the occasion properly, I spent part of my weekend visiting the Creation Museum in Santee, California, with Carrie Poppy and Ross Blocher of the podcast “Oh no, Ross and Carrie!” (more on that trip in my March 7 Skepticblog post). But I thought I’d mark this anniversary with a discussion of another important anniversary in the history of evolutionary science.

It was 40 years ago this year that the most frequently cited paper in the history of paleontology was published. That was none other than the legendary 1972 article by Niles Eldredge and Stephen Jay Gould which proposed the “punctuated equilibrium” hypothesis. (Full disclosure: I took seminars from Niles while I was a student at the American Museum of Natural History, and Steve Gould was very interested in and supportive of my research even though I was not his student in a formal sense). At the time the paper came out, the dominant concept about speciation was the allopatric speciation model. In a nutshell, good biological evidence showed that new species arise not in the large mainland populations (with their extensive gene mixing) but in small isolated populations with unusual gene frequencies (peripheral isolates), usually living separate (allopatric) from the mainland population. Once these allopatric populations were no longer isolated but remixed with the mainland population, they would be genetically and behaviorally distinct from their parent species. Thus, they would be no longer capable of interbreeding, which is part of the definition of a biological species.

Experimental biology … may reveal what happens to a hundred rats in the course of ten years under fixed and simple conditions, but not what happened to a billion rats in the course of ten million years under the fluctuating conditions of earth history. Obviously the latter problem is more important.

—George Gaylord Simpson, 1944, Tempo and Mode in Evolution

Even though the allopatric speciation model was accepted by biologists as early as 1942, it took paleontologists 30 years to recognize its implications. In their historic 1972 paper, Niles and Steve pointed out that if you took Ernst Mayr’s allopatric speciation model seriously, it would predict that species should arise in a normal biological time frame: a few years to a few hundred years at most. That’s a geologic instant, the difference between one bedding plane and the next in strata that span millions of years. The allopatric speciation model also predicted that species should arise in small, peripherally isolated areas, so they were unlikely to be fossilized in the few places for which we have a good fossil record. Rather than slow gradual change through millions of years of strata (the “phyletic gradualism” model), the allopatric speciation model accepted by biologists should give a fossil record where species seem to appear suddenly without any gradual transition preserved (“punctuation”), and then persist for long periods of time without change (“equilibrium”).

Stephen Jay Gould, Michael Shermer, and Donald Prothero

(left–right) Stephen Jay Gould, Michael Shermer, and yours truly, Mt. Wilson Observatory, Pasadena,CA (2001)

The “punctuated equilibrium” paper is a masterpiece of writing and incisive thinking, which poses a number of interesting issues. The first part is a general discourse on the philosophy of science, which argues that all scientists are products of their time and culture and tend to see what they expect to see. In this context, Darwin led paleontologists to expect phyletic gradualism, which they vainly tried to document for over a century before the allopatric speciation model came along. Then Eldredge and Gould introduced the details of the allopatric model, described punctuated equilibrium, and give examples from their own research (phacopid trilobites from Eldredge, Bahamian land snails from Gould). Every time I teach a paleontology class, I always assign the original paper as required reading, and then lead a class discussion section teasing it apart. Like fine wine, the paper gets better every time I reread it. I’m always amazed at what insights it contains, what future debates it triggered or foreshadowed, and how different students pick up different elements when they read it for the first time.

For the first decade after the paper was published, it was the most controversial and hotly argued idea in all of paleontology. Soon the great debate among paleontologists boiled down to just a few central points, which Gould and Eldredge (1977) nicely summarized on the fifth anniversary of the paper’s release. The first major discovery was that stasis was much more prevalent in the fossil record than had been previously supposed. Many paleontologists came forward and pointed out that the geological literature was one vast monument to stasis, with relatively few cases where anyone had observed gradual evolution. If species didn’t appear suddenly in the fossil record and remain relatively unchanged, then biostratigraphy would never work—and yet almost two centuries of successful biostratigraphic correlations was evidence of just this kind of pattern. As Gould put it, it was the “dirty little secret” hidden in the paleontological closet. Most paleontologists were trained to focus on gradual evolution as the only pattern of interest, and ignored stasis as “not evolutionary change” and therefore uninteresting, to be overlooked or minimized. Once Eldredge and Gould had pointed out that stasis was equally important (“stasis is data” in Gould’s words), paleontologists all over the world saw that stasis was the general pattern, and that gradualism was rare—and that is still the consensus 40 years later.

The debate was less than a decade old when I was wrapping up my dissertation work in 1981. In my dissertation on the incredibly abundant and well preserved fossil mammals of the Big Badlands of the High Plains, I had over 160 well-dated, well-sampled lineages of mammals, so I could evaluate the relative frequency of gradualism versus stasis in an entire regional fauna. I also had a wide geographic spread (from Montana and Saskatchewan to Texas, but mostly in the Dakotas, Nebraska, Wyoming, and Colorado). I had large fossil samples of many species, with dozens at each level, and excellent stratigraphic data. When I finally plunged in and plotted and analyzed my data carefully, it was clear that nearly every lineage showed stasis, with one minor example of gradual size reduction in the little oreodont Miniochoerus. I could point to this data set and make the case for the prevalence of stasis without any criticism of bias in my sampling. More importantly, the fossil mammals showed no sign of responding to the biggest climate change of the past 50 million years (the Eocene-Oligocene transition, when glaciers appeared in Antarctica after 200 million years). In North America, dense forests gave way to open scrublands, crocodiles and pond turtles were replaced by land tortoises, and the snails changed from those typical of Nicaragua to those of Baja California. Yet out of all the 160 lineages of mammals in this time interval, there was virtually no response. To paraphrase Rhett Butler, “Frankly, my dear, the mammals don’t give a damn.”

This result intrigued me, so I began to re-examine the uncritical acceptance of the notion that fossil mammals track environmental changes. It occurred to me that our excellent database of North American fossil mammals and global climate change might be a good place to test this hypothesis. In a 1999 paper, I argued that for the four biggest independently documented periods of climatic change in the past 50 million years, the mammals either do not respond at all, or show much less speciation and extinction than they do at times when there is no evidence of climatic change. One interval included the middle-late Eocene climate change at 37 million years ago, when turnover was merely at background level, despite evidence of floral change elsewhere in North America, and a big climatic cooling event in the global oceans. The second was the Eocene-Oligocene transition just discussed. The third was the great expansion of modern grasslands at 7.5 million years ago, long after mammals with high-crowned grazing teeth appeared at 15 million years ago. In fact, there is almost no significant faunal change at 7.5 million years ago. The final example is the last 2 million years of ice ages, when climate changed dramatically, but speciation did not occur in response to climate. Instead, most ice age mammals simply migrated north and south in response to the movements of the ice sheets.

About six years ago, I decided to test this last idea. Back in 2006, I had a bunch of excellent students in my paleontology classes at both Occidental and Caltech, and several wanted to do research with me. I was at a loss over how to supervise so many undergraduates with limited backgrounds, until I realized that we could do a group of related projects practically in our back yard: the Page Museum at La Brea tar pits in Los Angeles. I equipped each one of them with calipers and gave them a measuring protocol, and got them access to the La Brea fossils through the collections managers. Then they each measured a different Ice Age mammal or bird, collecting data on hundreds of bones from all the tar pits with good radiocarbon dates: saber-toothed cats, dire wolves, giant lions, bison, horses, camels, ground sloths, plus the five most common birds: golden and bald eagles, condors, caracaras, and turkeys. After six years of work and publication, the conclusion is clear: none of the common Ice Age mammals and birds responded to any of the climate changes at La Brea in the last 35,000 years, even though the region went from dry chaparral to snowy piñon-juniper forests during the peak glacial 20,000 years ago, and then back to the modern chaparral again.

In four of the biggest climatic-vegetational events of the last 50 million years, the mammals and birds show no noticeable change in response to changing climates. No matter how many presentations I give where I show these data, no one (including myself) has a good explanation yet for such widespread stasis despite the obvious selective pressures of changing climate. Rather than answers, we have more questions—and that’s a good thing! Science advances when we discover what we don’t know, or we discover that simple answers we’d been following for years no longer work.

Clearly, it is an interesting time to be a paleontologist. As Gould (1983) put it, from the “irrelevance” of the early 1900s to the “subservient” role that George Gaylord Simpson placed us in 1944, paleontology is now in the driver’s seat, and trying to reach the “High Table” where the “High Priests” of evolution and genetics have long ruled unchallenged. Who knows where this will all end? Whatever the result, we’re clearly making advances by challenging the accepted dogmas of the time and finding their faults, and hopefully discovering something new and interesting. As Gould and Eldredge (1977) put it, “Why be a paleontologist if we are condemned only to verify what students of living organisms can propose directly?”

References
  • Eldredge, N. 1985. Time Frames: The Rethinking of Darwinian Evolution and the Theory of Punctuated Equilibria. Simon & Schuster, New York.
  • Eldredge, N. 1995. Reinventing Darwin: The Great Debate at the High Table of Evolutionary Theory. John Wiley & Sons, New York.
  • Eldredge, N., and S. J. Gould, 1972. Punctuated equilibria: an alternative to phyletic gradualism, pp. 82-115, in Schoft, T.J.M., Models in Paleobiology. Freeman Cooper, San Francisco.
  • Gould, S.J. 1980. Is a new and more general theory of evolution emerging? Paleobiology6: 119–130.
  • Gould, S.J. 1982. Darwinism and the expansion of evolutionary theory. Science 216:380–387.
  • Gould, S.J. 1983. Irrelevance, submission, and partnership: the changing roles of paleontology in Darwin’s three centennials, and a modest proposal for macroevolution, pp. 347-366, in Bendall, D.S. (ed.), Evolution from Molecules to Men. Cambridge University Press, Cambridge.
  • Gould, S.J. 2002. The Structure of Evolutionary Theory. Harvard University Press, Cambridge, Mass.
  • Gould, S.J., and Eldredge, N. 1977. Punctuated equilibria: the tempo and mode of evolution reconsidered. Paleobiology 3:115–151.
  • Prothero, D.R., 1992.  Punctuated equilibrium at twenty: a paleontological perspective.  Skeptic 1(3): 38–47.
  • Prothero, D.R. 1999. Does climatic change drive mammalian evolution? GSA Today   9(9):1–5.
  • Prothero, D.R., Syverson, V., Raymond, K.R., Madan, M., Molina, S., Fragomeni, A., DeSantis, S., Sutyagina, A., and Gage, G.L. (in press) Size and Shape Stasis in Late Pleistocene Mammals and Birds from Rancho La Brea during the last Glacial-Interglacial Cycle. Quaternary Science Reviews(in press)
  • Prothero, D.R., and T.H. Heaton, 1996, Faunal stability during the early Oligocene climatic crash. Palaeogeography, Palaeoclimatology, Palaeoecology 127:239–256.
About the Author
photo

DR. DONALD R. PROTHERO was Professor of Geology at Occidental College in Los Angeles, and Lecturer in Geobiology at the California Institute of Technology in Pasadena. He earned M.A., M.Phil., and Ph.D. degrees in geological sciences from Columbia University in 1982, and a B.A. in geology and biology (highest honors, Phi Beta Kappa) from the University of California, Riverside. He is currently the author, co-author, editor, or co-editor of 32 books and over 250 scientific papers, including five leading geology textbooks and five trade books as well as edited symposium volumes and other technical works. He is on the editorial board of Skeptic magazine, and in the past has served as an associate or technical editor for Geology, Paleobiology and Journal of Paleontology. He is a Fellow of the Geological Society of America, the Paleontological Society, and the Linnaean Society of London, and has also received fellowships from the Guggenheim Foundation and the National Science Foundation. He has served as the President and Vice President of the Pacific Section of SEPM (Society of Sedimentary Geology), and five years as the Program Chair for the Society of Vertebrate Paleontology. In 1991, he received the Schuchert Award of the Paleontological Society for the outstanding paleontologist under the age of 40. He has also been featured on several television documentaries, including episodes of Paleoworld (BBC), Prehistoric Monsters Revealed (History Channel), Entelodon and Hyaenodon (National Geographic Channel) and Walking with Prehistoric Beasts (BBC). His website is: www.donaldprothero.com. Check out Donald Prothero’s page at Shop Skeptic.

Skeptical perspectives on Darwin, Gould and punctuated equilibrium…
cover Skeptic magazine 1.3:
Revolution in Evolution?

In this issue: Darwin & Gould; A Skeptical Investigation at Edgar Cayce’s Association for Research and Enlightenment; 20 years of Punctuated Equilibrium: A Paleontological Perspective; Historic 1842 Sketch on Evolution; the Nature of Change; Darwin, Darwinism & the Darwinian Culture, and more… GET THIS ISSUE or SUBSCRIBE TO SKEPTIC MAGAZINE.

cover A Darwin Day 200th Birthday Celebration
with Dr. Donald Prothero, Dr. Michael Shermer, and Dr. Joel Smith

2009 marked the 200th anniversary of Charles Darwin’s birth on February 12, 1809 — the same day as Abraham Lincoln — and the 150th anniversary of the publication of On the Origin of Species on November 24, 1859. This event was a special celebration of the life and science of one of the greatest scientists in history. GET THE DVD.

cover Evolution Revolution: Paleontology, History, Biography (Festschrift 2000 for Stephen Jay Gould, Part I)
with Dr. Donald Prothero, Dr. Michael Shermer,
and Dr. Frank J. Sulloway

Dr. Donald Prothero discusses how Gould helped launch a revolution in paleontology that continues to this day. Dr. Michael Shermer presents a slide show of an exhaustive literary taxonomy and content analysis of all 300 of Gould’s essays. Dr. Frank J. Sulloway focuses on the science of biography, using a dual metaphor found in much of Gould’s writing… GET THE DVD.

cover Evolution Revolution: Skepticism, Education, Environment (Festschrift 2000 for Stephen Jay Gould, Part II)
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James “The Amazing” Randi, the world’s foremost spokesman against pseudoscience, discussed Gould’s role in skepticism. Dr. Carol Tavris delivered one of the most popular lectures, discussing her fascinating and important work in gender studies, showing how pervasive the male bias is in all aspects of research. Bill Nye, “The Science Guy” mixed humor with important moral homilies on science, education, and on instilling kids with a sense of wonder and creativity. Dr. Paul MacCready emphasized the importance of education, particularly teaching kids how to think “out of the box” to find creative new solutions to old problems such as overpopulation, global warming, and pollution. GET THE DVD.

cover Evolution Revolution: Keynote Address by Gould (Festschrift 2000 for Stephen Jay Gould, Part III)
with Dr. Stephen Jay Gould

Dr. Stephen Jay Gould is one of the best known and most highly decorated scientists of our age. Gould delivers a remarkable lecture filled with wit, charm, and historical anecdote. He traced the history of Western culture’s uneasy relationship with the pedestal-shattering discoveries of science. The single best lecture in the history of the Skeptics Society. Don’t miss this one!
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6 Comments »

6 Comments

  1. Jim Hull says:

    A wonderfully interesting article by Dr. Prothero. It sums up and showcases some fascinating developments in paleontology — a field we lay readers have found irresistible since we were kids playing with dinosaur toys — and gives a sense of the adventure of doing this kind of science. It also presents a demonstration of the imaginative and critical thinking scientists (and skeptics) must employ to unravel the complex mysteries of nature. Instructive!

  2. BaronPike says:

    And yet we have the average human getting taller in some areas of Europe an getting shorter in the US. So does stasis represent the most successful adaptation (and perhaps in the end the least malleable), or the most successful use of strategies to avoid adapting physically, etc.

    • Dr. S says:

      immigration???
      bogus statistic

      rjp

      • Donald Prothero says:

        As paleontologists use the term, “stasis” is over millions of years. What is happening to humans in a few generations is ecophenotypic change, not genetic change, so it is not the realm of evolution nor paleontology.

        • Dr. S says:

          thank you
          I might suggest using the term “eCONOphenotypic as well.
          or Politicopyenotypic
          or ethnophenotypic?

          Maybe people in the U.S are getting shorter is that the tall guys play basketball in high school.
          This takes away from homework time, and as a result they can’t get in college, so they become drunk and end up in lower economic class and produce shorter offspring because of poor nutrition in childhood!!!

          This might be called spheroprojectilotypic.

          Others??

          Dr. Sidethink Hp.D

  3. Josh says:

    Very enjoyable article, Don. Thank you for the history lesson as well as a great breakdown of punc-eq. And, as one of those “high-priests” to which you refer, I’ll go on record as saying I think paleontology is the bee’s knees.

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