The ID Cookie Crumbles...


During this year of celebration of Darwin and evolutionary biology, Intelligent Design (ID) supporters are fond of asserting that a branch of the biological sciences that currently accounts for over 100 regularly published journals (containing over 1000 peer-reviewed scientific reports) per year, over 1000 books published by reputable scientific publishers per year, and involving grant and foundation support amounting to several billion dollars per year is "crumbling", while ID, which accounts for not one peer-reviewed scientific journal and one peer-reviewed book (published over a decade ago) is replacing it.

I can go to Mann Library here at Cornell (the second largest library of biology in the world, comprising over a million books and bound periodicals) and find the equivalent of an entire floor devoted to evolutionary biology. I couldn't carry this month's issues of the various journals devoted to evolutionary biology to the loan desk, even if I used a large laundry basket and made several trips. I have a paltry selection of the most current books on the subject of evolution in my personal library: only 1000+ volumes published in the past ten years or so. If I had unlimited funds, I could buy ten times as many, and still could not keep up with the field.

Virtually every large university in the world has a department of ecology and evolutionary biology. Here at Cornell we have such a department, with almost two dozen professors and dozens of graduate students, and there are at least five other departments at Cornell who number evolutionary biologists among their members. There are almost half a dozen undergraduate and graduate organizations devoted to the scientific aspects of evolutionary biology at Cornell; branches of such societies are found worldwide.

By contrast, there are two tenured professors in the entire world who explicitly support ID, only one of whom is in a department devoted to an empirical science (the other teaches at a theological seminary). Neither of them is currently engaged in empirical research intended to validate ID.

Of the 35+ undergraduate IDEA clubs (a very liberal estimate) that were founded during the heyday of ID (the late 1990s and early 2000s), not one is currently maintaining a website or apparently meeting regularly. And according to Google Trends, interest by the news media in ID has fallen almost to zero since the Kitzmiller v. Dover trial in late 2005, while interest in evolutionary biology is at an all-time high and still increasing with no end in sight.

So, based on the empirical evidence, which is "crumbling", evolutionary biology or ID?

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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen

Books in Celebration of the 150th Anniversary of the Origin of Species


2009 marks several important anniversaries in the science of evolutionary biology. Perhaps the two most important are the 200th anniversary of the birth of Charles Darwin and the 150th anniversary of the publication of his most important book, On the Origin of Species, originally published in November of 1859. Scientific societies around the world are celebrating these two events, with “Darwin Day” observances at hundreds of colleges, universities, and museums, including Cornell University and the Museum of the Earth at the Paleontological Research Institution in Ithaca, New York.

Publishers are also celebrating these two anniversaries, bringing out a flood of books on Darwin and evolution. There isn’t space in this blogpost to note all of these publications, but we can mention some of the most noteworthy and relevant of these publications. Here is an annotated list of some of the best and most comprehensive publications that have come out recently, celebrating the life and work of Charles Darwin and his theory of evolution by natural selection. The full publication information for each work is included in this list, followed by a brief paragraph describing its contents.

DARWIN'S BOOKS (in print)
Darwin, Charles (Edward O. Wilson, editor) (2005) From So Simple a Beginning: Darwin's Four Great Books. W.W. Norton & Co.: New York, NY, ISBN 0393061345 (hardcover, $39.95), 1706 pages. Available here.
Perhaps the best way to celebrate Darwin’s life and work is to read his most important and influential books. In preparation for the Darwin bicentennial several publishers have brought out various versions of his books. This one is the best, not only because it includes his four most influential publications, but also because the editor, evolutionary biologist Edward O. Wilson, provides a brief but illuminating introduction to each one. Wilson places Darwin’s work in its historical and scientific context, and shows how each of his four great books laid the groundwork for the modern science of biology. If you only have time for one of these books, let this compendium be the one you read this year.


Darwin, Charles (James T. Costa, editor) (2009) The Annotated Origin: A Facsimile of the First Edition of On the Origin of Species. Belknap Press of Harvard University Press: Cambridge, MA, ISBN 0674032810 (hardcover, $35.00), 576 pages. Available here.
Darwin didn’t originally intend to publish the book we now know as the Origin of Species. He was working on a much longer multi-volume work that he intended to call Natural Selection, and rushed to publish an “abstract” of this work in 1859 to forestall losing his priority for the idea to Alfred Russell Wallace. As a consequence, the Origin of Species has no footnotes, endnotes, nor bibliography. This version of the Origin makes up for that lack. Ably edited by James Costa, The Annotated Origin contains many of the annotations that the original Origin of Species lacked, and provides the reader with a comprehensive grounding in the natural history that Darwin marshaled in support of his revolutionary theory.


Darwin, Charles (David Quammen, editor) (2008) On the Origin of Species: The Illustrated Edition. Sterling Publishers: New York, NY, ISBN 1402756399 (hardcover, $35.00), 560 pages. Available here.
Darwin rushed to publish the Origin of Species, and included only one technical illustration in the first edition. This lack of illustrations has been rectified in David Quammen’s beautifully illustrated version of the Origin. Perhaps best known for his masterful book on island biogeography and extinction, The Song of the Dodo, Quammen has chosen a huge selection of images that illuminate Darwin’s theory in ways that Darwin himself would have found both fascinating and extremely valuable for a deeper understanding of his theory.

DARWIN'S BOOKS (online)

The Internet has provided readers and scholars today with an unparalleled opportunity to study the works of Darwin online. By far the best of these electronic resources is a website initiated and maintained by John Van Whye and his colleagues in England. Available online at http://darwin-online.org.uk/, The Complete Works of Charles Darwin includes not only all of Darwin’s books (available in all of their various editions), but also photographic facsimiles of these works side-by-side with searchable full text versions, plus all of Darwin’s scientific publications and a massive and growing collection of his voluminous correspondence. Here is a sampling:

Darwin, Charles (1845) Journal of Researches into the Natural History and Geology of the Countries Visited During the Voyage of H.M.S. Beagle Round the World, Under the Command of Capt. Fitz Roy, R.N. 2d edition. John Murray: London, UK, 536 pages. Text and images available here.
This was Darwin’s first and most popular book for the general public, establishing him as the premier naturalist in England at the time of its publication. Continuously in print since 1845, the Voyage of the Beagle (as it is most often referred to) is both a marvelous compendium of natural history and a fascinating journal of a voyage of discovery almost unparalleled in the literature of science.


Darwin, Charles (1859) On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. John Murray: London, UK, 522 pages. Text and images available here.
Darwin’s most important book, and perhaps the most important book ever published in the science of biology. Indeed, a strong argument could be made that this book single-handedly founded the modern science of biology. This online edition juxtaposes the text of the Origin with photographs of the corresponding pages from the first edition.


Darwin, Charles (1871) The Descent of Man, and Selection in Relation to Sex. John Murray: London, UK, 424 pages. Text and images available here.
Darwin’s second most important book, and perhaps the most controversial of his published works, the Descent of Man (as it is most often referred to) contains Darwin’s proposal that humans have evolved from “lower” primates, and a detailed exploration of what Darwin believed to be the most important mechanism in the evolution of humans: sexual selection. Darwin also speculates on the evolutionary origin of such uniquely human traits as art, language, and morality.


Darwin, Charles (1871) The Expression of the Emotions in Man and Animals. John Murray: London, UK, 374 pages. Text and images available here.
One of Darwin’s lesser-known books, but no less important in its own way, The Expression of the Emotions in Man and Animals is often credited with founding the modern science of animal behavior. In it, Darwin explored the biological basis for the expression of emotions in humans and other animals, with detailed examples (many with accompanying photographs) that show how human emotions are most probably derived from the emotions of other “lower” animals, including dogs as well as monkeys and other primates.

DARWIN'S LIFE AND WORK (adults/general)

Many people (including many evolutionary biologists) not only have never read Darwin's books, but also do not know how Darwin came to write them. Furthermore, opponents of Darwin's theories often portray Darwin's life and personal beliefs in as negative a light as possible, in an attempt to discredit his theories. Perhaps the best antidotes to both of these deficiencies are the books listed here, which together provide a comprehensive view of Darwin's life and work. Most of them are written for non-scientists, and all of them provide a fascinating glimpse into the work of the founder of the science of biology.

Browne, Janet (1996) Charles Darwin: A Biography, Volume 1: Voyaging. Princeton University Press: Princeton, NJ, ISBN 0691026068 (paperback, $25.95), 622 pages. Available here.
Although not explicitly published in celebration of the Darwin bicentennial, Janet Browne’s two-volume biography of Darwin is widely recognized as the premier biography of the founder of evolutionary biology. This first volume covers Darwin’s youth and early career, up to the completion of the voyage of HMS Beagle and the germination of his theory of evolution by natural selection. Using the extensive collection of Darwin’s correspondence along with personal papers and those of his family and scientific associates, Browne draws a detailed and sometimes surprising portrait of the founder of the most comprehensive and controversial theory in the natural sciences.


Browne, Janet (2003) Charles Darwin: A Biography, Volume 2: The Power of Place. Princeton University Press: Princeton, NJ, ISBN 0691114390 (paperback, $25.95), 600 pages. Available here.
In this second volume of Janet Browne’s two-volume biography of Darwin, Janet Browne explores the origin of the Origin of Species and Darwin’s other ground breaking and revolutionary works. Unlike some of Darwin’s other biographers, Browne gets most of the science right in her biography, and illuminates Darwin’s ideas in light of his detailed work in natural history. She also provides insights into Darwin’s personality, including his much-noted reclusive nature, and shows how his scientific work and ideas were influenced by events in his private life.


Darwin, Charles (Nora Barlow, editor) (1993) The Autobiography of Charles Darwin: 1809-1882. W.W. Norton & Co.: New York, NY, ISBN 0393310698 (paperback, $14.95), 224 pages. Available here.
Although Darwin wrote an autobiography, he never intended it to be a comprehensive history of his life or his ideas. Rather, he did it almost on a lark, as a kind of “gift” to the members of his family and closest friends. Despite this, his autobiography provides some fascinating insights into his personality, and especially his views on art, literature, religion, ethics, and philosophy. This edition includes autobiographical material that was suppressed by Darwin’s widow and son, and should be considered to be the definitive edition.


Darwin, Charles (Gordon Chancellor & John Van Wyhe, editors) (2009) Charles Darwin’s Notebooks from the Voyage of the Beagle. Cambridge University Press: New York, NY, ISBN 0521517575 (hardcover, $150.00), 650 pages. Available here.
This is an expensive, but absolutely invaluable publication, brought out by Cambridge University Press as part of this year’s bicentennial celebration and edited by Gordon Chancellor and John Van Whye (the same John Whye who maintains the complete works of Darwin online). It contains the entirety of Darwin’s handwritten journals of the voyage of HMS Beagle, written during the voyage. In it one can trace the development of Darwin’s revolutionary ideas as they first occurred to him, and see how the events of the voyage laid the groundwork for his revolutionary theories. This is a must-have edition for anyone who wants to understand how and when the Darwinian revolution began.


Desmond, Adrian & Moore, James (2009) Darwin's Sacred Cause: How a Hatred of Slavery Shaped Darwin's Views on Human Evolution. Houghton Mifflin Harcourt: New York, NY, ISBN 0547055269 (hardcover, $14.95), 448 pages. Available here.
Charles Darwin was born on February 12, 1809, the same day as Abraham Lincoln. This propinquity of origins is all the more notable when one realizes that Darwin and his family were staunch abolitionists. Darwin himself tells how he became horrified by the institution of slavery while on the voyage of HMS Beagle.

Desmond and Moore, whose 1994 biography of Darwin (Darwin: The Life of a Tormented Evolutionist, available here) is widely recognized as one of the best biographies of Darwin, have in this volume dug deeper into Darwin’s personal history, showing how his views on slavery and its abolition helped shape his views on the evolution of humans, as outlined in his Descent of Man.


McCalman, Iain (2009) Darwin's Armada: Four Voyages and the Battle for the Theory of Evolution. W.W. Norton & Co.: New York, NY, ISBN 0393068145 (hardcover, $29.95), 432 pages. Available here.
Although perhaps the best known, Darwin’s round-the-world voyage aboard HMS Beagle was not by any means the only such voyage of discovery in the 19th century. Three of Darwin’s closest friends and collaborators in evolutionary biology — botanist Joseph Hooker, zoologist Thomas Henry Huxley, and professional collector and naturalist Alfred Russell Wallace — all participated in similar voyages, and all contributed to Darwin’s theories. McCalman describes all four voyages, and shows how the insights and information about natural history gained by these four great naturalists provided the foundation for the science of evolutionary biology in the 19th and early 20th centuries.


Quammen, David) (2007) The Reluctant Mr. Darwin: An Intimate Portrait of Charles Darwin and the Making of His Theory of Evolution. Atlas Books: New York, NY, ISBN 039332995X (paperback, $35.00), 304 pages. Available here.
Quammen’s biography of Darwin’s personal origin of the Origin is a perfect companion to Janet Browne’s two-volume biography of Darwin. Focusing on the period in Darwin’s life leading up to the original publication of the Origin, Quammen shows how Darwin’s personality and early education predisposed him to the intellectual and emotional labor that resulted in the publication of the Origin and its reception by the scientific community.

DARWIN'S LIFE AND WORK (children/school)

All too often people think that the work of great scientists such as Charles Darwin is only accessible to adults, especially scientists. Here is a brief selection of books for children, all published during this bicentennial year, which present Darwin and his work in a way that anyone, including young children, can understand and appreciate.

Heiligman, Deborah (2009) Charles and Emma: The Darwins' Leap of Faith. Henry Holt and Co.: New York, NY, ISBN 0805087214 (hardcover, $18.95), 272 pages. Available here.
Heiligman’s biography of Charles and Emma Darwin begins with Charles drawing up a list of the pros and cons of marriage to his first cousin, Emma. His decision to marry her set in motion an emotional odyssey for Charles and Emma, who had very different views of religion. Charles’ theory of evolution threatened Emma’s deep religious belief, and forced Darwin to thoroughly document his views before publishing them. This sympathetic rendering of their happy, though sometimes turbulent, marriage is a wonderful introduction to the real life of a famous scientist, one that young adults will find particularly relevant to their own lives.


Lasky, Kathryn (2009) One Beetle Too Many: The Extraordinary Adventures of Charles Darwin. Candlewick Press: Somerville, MA, ISBN 076361436X (hardcover, $17.99), 48 pages. Available here.
Lasky’s copiously illustrated account of Darwin’s life and work, focusing especially on his adventures while on the voyage of HMS Beagle, gives younger readers an engaging look at the work of the founder of the theory of evolution by natural selection. Lasky does not shy away from Darwin’s views on evolution and religion, presenting them in Darwin’s own words. The illustrations are particularly charming, rendered in various media in a quirky and engaging style.

McGinty, Alice B. & Azarian, Mary (2009) Darwin, with Glimpses into his Private Journal and Letters. Houghton Mifflin Books for Children: New York, NY, ISBN 0618995315 (hardcover, $18.00), 48 pages. Available here.

Another illustrated account of Darwin’s life and work, this time covering his childhood, his adventures while on the voyage of HMS Beagle, and the subsequent development of his theory of evolution by natural selection. gives younger readers an engaging look at the work of the founder of the theory of evolution by natural selection. Illustrated with woodcuts and watercolor, McGinty and Azarian’s biographical history also touches on Darwin’s views on religion and philosophy in a way that is accessible to younger readers.


Schanzer, Rosalyn (2009) What Darwin Saw: The Journey That Changed the World. National Geographic Children's Books: Des Moines, IA, ISBN 1426303963 (hardcover, $18.00), 48 pages. Available here.
Schanzer’s illustrated account of Darwin’s voyage of HMS Beagle is more limited in scope than the others in this list, but succeeds in presenting Darwin’s natural history investigations in an engaging (and sometimes humorous) way. More like a graphic novel than a standard biography (complete with cartoon-like illustrations and speech balloons), Schanzer presents selected observations that led Darwin to his theory of evolution. Unlike the other three books for younger readers in this list, Schanzer avoids discussion of Darwin’s views on religion and philosophy, sticky strictly to the natural history of his historic voyage and the science that it inspired.

In addition to the books listed and reviewed here, those who are interested in the life and works of Charles Darwin can search Amazon.com, where most of the books listed above can be browsed, along with many others published in connection with the Darwin bicentennial. Books on the subject of evolution are listed here, including books both supportive of Darwin’s theory and critical of it.

I hope this list provides you with some ideas of how to celebrate Darwin's life and work by learning more about it.

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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen

Macroevolution: What Were the Evolutionary Ancestors of Whales?


ID supporters often assert that, although there is abundant empirical evidence for microevolution, there is no such evidence for macroevolution. One of the macroevolutionary transitions that they often cite is the evolution of whales from a land-dwelling ancestor. Which leads me to ask the following question:

What empirical evidence would verify (i.e. support) or falsify (i.e. undermine) the hypothesis that whales had evolved from a land-dwelling mammal? Please note that this is a hypothesis about macroevolution, not microevolution.

A basic principle of hypothesis validation in the natural sciences is that if one can find multiple lines of evidence, all of which support the hypothesis, then such evidence is much stronger than if there were only a single line of evidence. This is especially the case if the different lines of evidence are from very widely separated fields.

Until recently the main line of evidence for the evolution of whales (i.e. members of the mammalian order Cetacea) from even-toed ungulates (i.e. members of the mammalian order Artiodactyla) was anatomical. This anatomical evidence was derived from two sources:

1) similarities between the anatomy (especially skeletal anatomy) of living (i.e. "extant") Artiodactyls and Cetacea, and

2) an evolutionary phylogeny of the transition from terrestrial Artiodactyls to aquatic Cetacea, based on fossils.

Rather than summarize this comparative anatomical evidence here, I recommend that interested readers follow this link. What you will find is a fairly detailed summary of the evidence from comparative anatomy, all of it pointing to the conclusion that whales (i.e. Cetaceans) evolved from even-toed ungulates (i.e. Artiodactyls). It is this evidence that most evolutionary biologists have until recently cited as support for the macroevolutionary derivation of Cetaceans from Artiodactyl ancestors.

However, one can also ask the question Does the comparative genomics of Artiodactyls and Cetaceans support the same hypothesis? That is, are there observable DNA sequence similarities and differences that are similar in both scope and timing to the similarities and differences in the fossil record (and as reflected in the comparative anatomy of Artiodactyls and Cetaceans)?

This is an easily falsified hypothesis: If the genomic evidence does not support the Artiodactyl into Cetacean hypothesis — e.g. that Cetaceans evolved from some other clade, or that they had not evolved at all, but rather sprang into existence fully-formed and without genetic evidence of a macroevolutionary transition — then this evidence would not support the evidence from comparative anatomy and the macroevolutionary hypothesis based on comparative anatomy would be called into question.

So, what does the comparative genomic evidence indicate about the macroevolutionary relationships between the Artiodactyla and the Cetacea? Here’s a summary of the findings from comparative genomics:

The idea that whales evolved from within the Artiodactyla was based on analysis of DNA sequences. In the initial molecular analyses, whales were shown to be more closely related to ruminants (such as cattle and deer) than ruminants are to pigs. In order for the order name to reflect a real evolutionary unit, the term Cetartiodactyla was coined.

Later molecular analyses included a wider sampling of artiodactyls and produced a more complete tale. Hippos were determined to be the closest relative of whales, ruminants were related to a whale/hippo clade, and pigs were more distant. In addition to producing the controversial whale/hippo clade, these analyses debunked the idea that hippos and pigs are closely related. This had been a popular taxonomic hypothesis (Suiformes) based on similarities in morphological (physical) characteristics.

In addition to DNA and protein sequences, researchers tracked the movement of transposons called SINEs in the genome. A transposon is a DNA sequence that will occasionally make a copy of itself and insert that copy into another part of the genome. It is considered highly unlikely that SINEs will insert themselves into the exact same part of a genome by chance. The data indicate that several transposons inserted themselves at the same point in the genomes of whales, ruminants and hippos (sometimes referred to as "pseudoruminants" because although they have four-chambered stomachs like true ruminants, they do not chew the cud). This insertion point is not shared with camels and pigs.

This hypothesis has been tested with DNA sequences from a host of genes: the complete mitochondrial genome (as well as several of its genes independently), beta-casein, kappa-casein, von Willebrand factor, breast cancer 1, recombination activating genes 1 and 2, cannabinoid receptor 1, and several others. These sequence data and the transposons converge on the same conclusion that hippos and whales are more closely related to one another than either is to other artiodactyls.

Sequences analyzed in combined analyses with morphological characters have also produced the same results as sequences alone. Some have argued that the sheer number of characters (one for each nucleotide) in sequences swamps out the effects of morphology. There have been a few morphology-based studies that have suggested (weakly) the same results as the molecular results, but overall most morphological studies have conflicted with the whale/hippo hypothesis of Cetartiodactyla.

An important exception is a recent conducted by Boisserie et al. (2005). They examined 80 hard morphological characters of fossil and extant cetartiodactylan taxa. Their results suggest that hippopotamids evolved from within a clade of anthracotheres. That anthracothere/hippopotamid clade appears to be sister to the Cetacea and supports the molecular results.

[sources: http://en.wikipedia.org/wiki/Cetartiodactyla (summary article), where you can find links to many primary references]


Note that much of the genomic data (especially from transposon sequences) supporting the macroevolutionary hypothesis is based on non-adaptive DNA sequences. That is, DNA sequences that do not code for adaptive characteristics, and in many cases that do not code for anything at all. This is like figuring out which students have been copying the answers to test questions from other students by comparing their wrong answers. The right answers are the same for everybody, but wrong answers vary from student to student in virtually random ways. If two students have the same wrong answers, you can be reasonably confident that one of them copied the wrong answers from the other. You can then test this hypothesis by looking at seating charts, past test performance (cheaters are often identified by sudden increases in test scores without apparent increases in effort), and – often the last resort – asking them if they copied answers.

Conclusion: The empirical evidence from comparative genomics closely matches the empirical evidence from (both extant and fossil) comparative anatomy.

Is that all, or is there yet another line of evidence that might be pursued to verify or falsify the Artiodactyl into Cetacean hypothesis? Yes, there is. Consider the observable fact that whales reproduce much more slowly than even-toed ungulates (such as deer and hippos). Indeed, there is a general principle in zoology that the larger the members of a species are (on the average) the fewer offspring they have, the more widely spaced those offspring are in time, the fewer offspring they can have over their lifetime, and the longer the average lifespan of individuals.

For example, deer can have offspring every year, and under good conditions can sometimes have twins or even triplets in one reproductive cycle. By comparison, baleen whales can only have offspring every few years (it can take up to two years for one pregnancy in large baleen whales), they virtually never have more than one calf at a time, they have only a few reproductive life cycles per lifetime, and they have much longer lifespans than deer.

This means that, if Cetaceans evolved from Artiodactyls, one might be able to find empirical evidence that the rate of the macroevolutionary transition from Artiodactyl ancestors into Cetacean descendants had slowed down as the result of the increase in size, decrease in number of offspring per reproductive cycle, decrease in total number of offspring per lifetime, and increase in average lifespan. In brief, there might be evidence that the macroevolutionary “clock” slowed down as Cetaceans evolved larger and larger size.

Here’s the latest genomic evidence vis-a-vis this hypothesis:

Big and slow: phylogenetic estimates of molecular evolution in baleen whales (suborder mysticeti).
Molecular Biology and Evolution. 2009 Nov;26(11):2427-40. Epub 2009 Jul 31.
Jackson JA, Baker CS, Vant M, Steel DJ, Medrano-González L, Palumbi SR.
Marine Mammal Institute, Hatfield Marine Science Center, Oregon State University, OR, USA.

ABSTRACT: Baleen whales are the largest animals that have ever lived. To develop an improved estimation of substitution rate for nuclear and mitochondrial DNA for this taxon, we implemented a relaxed-clock phylogenetic approach using three fossil calibration dates: the divergence between odontocetes and mysticetes approximately 34 million years ago (Ma), between the balaenids and balaenopterids approximately 28 Ma, and the time to most recent common ancestor within the Balaenopteridae approximately 12 Ma. We examined seven mitochondrial genomes, a large number of mitochondrial control region sequences (219 haplotypes for 465 bp) and nine nuclear introns representing five species of whales, within which multiple species-specific alleles were sequenced to account for within-species diversity (1-15 for each locus). The total data set represents >1.65 Mbp of mitogenome and nuclear genomic sequence. The estimated substitution rate for the humpback whale control region (3.9%/million years, My) was higher than previous estimates for baleen whales but slow relative to other mammal species with similar generation times (e.g., human-chimp mean rate > 20%/My). The mitogenomic third codon position rate was also slow relative to other mammals (mean estimate 1%/My compared with a mammalian average of 9.8%/My for the cytochrome b gene). The mean nuclear genomic substitution rate (0.05%/My) was substantially slower than average synonymous estimates for other mammals (0.21-0.37%/My across a range of studies).

CONCLUSION: The nuclear and mitogenome rate estimates for baleen whales were thus roughly consistent with an 8- to 10-fold slowing due to a combination of large body size and long generation times. Surprisingly, despite the large data set of nuclear intron sequences, there was only weak and conflicting support for alternate hypotheses about the phylogeny of balaenopterid whales, suggesting that interspecies introgressions or a rapid radiation has obscured species relationships in the nuclear genome. [emphasis added]


So, there are indeed empirically falsifiable hypotheses for the macroevolution of whales from land-dwelling ancestors. If whales (Cetacea) evolved from even-toed ungulates (Artiodactyla), then the following predictions should be supported by the observable data:
• that there should be anatomical similarities between extant Artiodactyls and Cetaceans,
• that there should also be anatomical similarities between fossil Artiodactyls and Cetaceans,
• that there should be shared similarities and differences between the genomes of extant clades of Artiodactyls and Cetaceans, and that the overwhelming majority of these similarities and differences would mirror the comparative anatomical evidence for the macroevolutionary origin of the various clades of the Cetartiodactyla, and
• that the inferred slowing of macroevolutionary change during the transition from Artiodactyl ancestors to Cetacean descendants should also be consistent with the hypothesis that the rate of this transition would have slowed as the result of increasing body size, increasing reproductive spacing, decreasing numbers of offspring per life cycle, and increasing longevity.

And it is.

Clearly, an ID supporter might then ask for specific empirical evidence on how the various transitions occurred at the genetic and developmental level, and if these details could unambiguously distinguish between natural and supernatural causes for such genetic mechanisms. Evolutionary developmental biologists are currently working on answers to the first part, but I personally cannot imagine how one could empirically test the second part. Furthermore, it seems to me that invoking a supernatural cause for the macroevolutionary transition from Artiodactyls to Cetaceans would be unnecessary, and would add nothing whatsoever to our understanding of the mechanisms by which this transition occurred.

Ergo, if I were doing this research and publishing my results I wouldn’t mention it, as it would be completely unnecessary for a scientific explanation of this phenomenon.

Just out of curiosity, ask yourself how one might use any of the foregoing as positive or negative empirical evidence for the existence of God. I mention this because some evolutionary biologists believe they can use the data of evolutionary biology to disprove the existence of God, and some ID supporters believe they can use the data of evolutionary biology to prove the existence of God. Personally, I believe both attempts are misguided, pointless, and ultimately futile. That’s why I don’t make such attempts, and wonder why anyone would.

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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen

The Darwinian Revolutions Video Series


The Darwinian Revolutions

An online video lecture series
in honor of the 150th anniversary
of the original publication of
Charles Darwin's Origin of Species


Produced by:
The Cybertower at Cornell University

Written, directed and narrated by:
Allen MacNeill, Senior Lecturer
The Biology Learning Skills Center
Ecology & Evolutionary Biology

Videography by:
Dina Banning

Sound Engineering by:
Colbert McClennan

Technical Direction by:
Becky Lane

Videotaped at:
The Museum of the Earth
The Paleontological Research Institution
Ithaca, New York

Voiceover Narration Recorded at:
Fall Creek Studios
1201 North Tioga Street
Ithaca, New York

Images Obtained at:
WikiMedia Commons
Stebbins/Simpson/Dobzhanky photo credit: Martin Tracey

Galapagos Video Credit:
Prof. William Provine

It's finally done! After more than a year of meetings, writing, image acquisition, videotaping, sound recording, editing, revising, captioning, and (most of all) thinking, our video series on the Darwinian revolutions is now online!

This series of six online videos is a brief introduction to Darwin's theory of evolution by natural selection and its implications. Here is a brief synopsis of the six episodes (click on each episode title to go to the linked video):


Episode One: Darwinian Revolutions
We begin with an overview of the series, which has been released to coincide with the 150th anniversary of the publication of Darwin's Origin of Species. Darwin's theory of evolution by natural selection revolutionized both the biological sciences and our understanding of ourselves and the world around us. In this episode we learn that Darwin's theory has itself evolved in the 150 years since it was published. We also learn that Darwin actually presented two theories:
• a theory of descent with modification from common ancestors, and
• the theory of natural selection, Darwin's mechanism for evolution.


Episode Two: Evolutionary Ancestors
Beginning with an overview of Darwin's predecessors, we learn how the idea of evolution by natural means alone goes back more than two thousand years, to ancient Greece and Rome. Democritus of Abdera first proposed the "ground rules" for naturalist evolution, which were later extended by the Roman poet and philosopher, Lucretius. However, these early naturalistic theories were eclipsed for almost two millennia by the ideas of Plato and Aristotle.


Episode Three: Lamarck's Theory
In the 19th century, Jean Baptiste Lamarck set the stage for Darwin's monumental achievement with his Philosophie Zoologique (published in 1809), which advanced a theory of evolution by means of the inheritance of acquired characteristics. Lamarck's theory was the first theory of evolution to include a testable mechanism for evolutionary change — the inheritance of acquired characteristics — and provides a useful comparison with Darwin's theory.


Episode Four: One Long Argument
Darwin, whose academic training at Cambridge University was in Anglican theology, became an acclaimed naturalist and science writer following the five-year voyage of HMS Beagle. Using the notes and specimens that he had collected during the voyage, Darwin spent twenty years refining his theory, first published in 1859, of evolution by natural selection.


Episode Five: Mendel and the Eclipse of Darwin
Darwin's theory of descent with modification was accepted by most scientists worldwide within ten years of its publication in 1859. However, his theory of natural selection was widely criticized, and by the turn of the 20th century was widely considered to be dead. However, the work of Gregor Mendel, who discovered the foundations of what we now call genetics, provided a mechanism by which Darwin's theory could be revived and expanded.


Episode Six: The Evolving Synthesis
In the final segment of this series, we visit the The Museum of the Earth in Ithaca, New York, whose director, Dr. Warren Allman, discusses the importance of such museums to the science of evolutionary biology. We also hear from Cornell professor William Provine, who discusses Darwin's work and its importance to the history and philosophy of biology. He tells us how Darwin's original theory of natural selection was integrated into the sciences of population genetics, ecology, physiology, paleontology, embryology, and botany, to produce a "modern synthesis" of evolutionary theory. Prof. Provine also tells us how the "modern synthesis" has continued to evolve, and that today is the most exciting time yet in the history of Darwin's scientific revolution.

This has been an exciting year: the 200th anniversary of the publication of Lamarck's Philosophy Zoologique, the 200th anniversary of the birth of Charles Darwin, and the 150th anniversary of the publication of Darwin's Origin of Species. There have been many events marking these anniversaries, and there will be many more. As Will Provine says, the theory of evolution is more dynamic, more exciting, more widely accepted, and more widely applied than at any time in the past century and a half. With the accelerating pace of discoveries in evolutionary biology and their applications in biology, medicine, psychology, economics, and even literature and art, the 21st century shows all indications of being what the founders of the "modern synthesis" called it back in 1959: the "century of Darwin" and his theory of evolution by natural selection.

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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen

Why I Post Comments on Creationist and ID Blogs


A fellow blogger asked me recently "Why waste your time posting [at creationist and ID blogs] at all? You do not seem to have a receptive audience." Other people ask me why I generally treat creationists and ID supporters with respect, rather than taking every opportunity to heap scorn and ridicule upon them. Here's what I hope is an adequate explanation to both of these questions.

I post comments at creationist and ID blogs (when I have the time, which is definitely not all the time) because I expect that there are a lot of people who read these comments without participating directly in such debates. This is why I try to keep as civil a tone as possible, especially when my opponents use ad hominem arguments, character assassination, insults, and ridicule. The contrast between their tactics and mine undermines their credibility (and, by extension, that of their soi dissant “science”). Indeed, some commentator’s comments are so insulting that I refuse to respond to them, and I believe that this does not pass unnoticed by readers who are not yet irrationally committed to one side or the other.

Also, as a teacher (currently in my 35th year of teaching at Cornell University), I feel a professional responsibility to correct some of the more egregious misrepresentations and misunderstandings of the science of evolutionary biology which are promulgated at creationist and ID blogs. Some of these misrepresentations clearly stem from ignorance, and in a gratifying number of cases some the commentators whom I have corrected have thanked me for the information and references I have provided.

However, other misrepresentations are apparently part of a deliberate and ongoing effort to distort the public record and deliberately misrepresent the relevant scientific information for political and religious purposes. I strongly believe that this kind of mendacity should be exposed for what it is, and for what it isn’t (i.e. it isn’t science).

Furthermore, I have always tried to emulate the long-standing Quaker tradition of “speaking truth to power”, which is the opposite of “preaching to the choir”. It means confronting directly what I perceive to be misunderstanding (and what I perceive to be deliberate mendacity), rather than limiting my interactions to people with whom I already agree. I agree with Charles Darwin, who said that he paid much more attention to the criticisms of people who disagreed with him than the praise of people who agreed with him. Like Darwin, I find that debating with people with whom I disagree helps me greatly to clarify my own position on the relevant issues, and to help my opponents clarify theirs. I believe that it would be a terribly boring (and non-progressive) world in which everyone agreed upon every subject, and so I am grateful to some of the commentators here for helping me improve my understanding of the relevant issues and my ability to argue persuasively for what I perceive to be the best supported position.

Like Hegel, I believe that genuine synthesis usually arises out of the clash between thesis and antithesis, and that progress in human understanding is almost always gained at the price of diligence, honesty, and honor. As my fencing master often says, “a gentleman is always gracious and dignified in defeat, humble and gentle in victory”. I have to the best of my ability tried to account myself according to this standard of conduct, and believe that the world would be a better place if everyone tried to do so.

Finally, to the best of my ability, I try to “fight the good fight” in defense of what I understand to be an accurate description of reality. I expect my opponents to do the same. When they do, I tell them so. Indeed, if they do a good job, I congratulate them, especially if they persuade me to change my mind, as the result of sufficiently convincing arguments. However, when my opponents depart from honorable and honest argumentation and stoop to ad hominem attacks, character assassination, insults, and ridicule, I call them on it and I inform them in no uncertain terms that I will no longer respond to them.

The same principles apply to this blog, and can be read here.

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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen

Rescuing Darwin from His Detractors (and Supporters)


TITLE: Rescuing Darwin

AUTHOR: Paul Fayter

SOURCE: The United Church Observer

SUMMARY: Ever since the Origin of Species was first published 150 years ago, scientists and creationists have tried to spin the religious views of Charles Darwin their way.

COMMENTARY: This article by Paul Fayter (appearing on a website sponsored by The United Church Observer, a Christian organization) directly contradicts many of the assertions made by some evolutionary biologists and most ID supporters, including most of the egregious distortions in Benjamin Wiker's recent book,
The Darwin Myth: The Life and Lies of Charles Darwin.


Unlike Wiker and most ID supporters, Paul Fayter has spent a significant fraction of his professional life studying Charles Darwin and his work. Here's what he found:

- Darwin was not an atheist when he wrote the Origin of Species and therefore could not possibly have written it as an "apology" for his atheism;

- the most important factors in his eventual agnosticism were 1) the death of his daughter, Annie, and 2) his view that a benevolent deity would not have created a world with such horrors as parasites (and the indiscriminate death of innocents);

- that despite his agnosticism, Darwin remained a "practicing" member of both the Church of England and his local parish church;

- that despite his embrace of the science of evolutionary theory, he did not descend into a life of libertinism and immorality, nor did he distort his theory to support any political position (including eugenics or "social darwinism");

- he remained a dedicated scientist, a loving husband, a doting father, a devoted member of his parish, and an unwavering opponent of slavery and its concomitant evils; and

- he also remained what would now be considered to be a "theistic evolutionist" until his death.

Here is Faytor's conclusion:

Darwin was able to reconcile the power and glory of a good and loving God with nature's cold indifference and manifest cruelty - the infamous and pitiless "survival of the fittest" - by viewing struggle, pain, suffering and death not as the direct will of God but as the result of the impersonal operation of universal laws. The process of evolution by means of natural selection was deadly and wasteful, and yet, as Darwin concluded in The Origin, it had a higher, nobler purpose. Higher species would evolve. The Creator - the God of scientific theism - lawfully drew good out of evil and progress out of pain.

Near the end of his life, Darwin thought it impossible to conceive that "this immense and wonderful universe" was "the result of blind chance or necessity." No, it still seemed that the world had been willed into being. "I feel compelled to look to a First Cause having an intelligent mind in some degree analogous to that of man," he wrote in his autobiography, "and I deserve to be called a Theist." At the same time, Darwin believed that "the mystery of the beginning of all things" was simply unsolvable; and so he also declared, "I for one must be content to remain an agnostic."

All of this directly contradicts Wiker's biography of Darwin and the opinions of most ID supporters. Demonizing Darwin, especially when there are very good (and unbiased) biographies of Darwin (not to mention his own, very candid autobiography) does nothing to harm Darwin's scientific reputation. By contrast, the fundamental misunderstanding among many ID supporters of how real science is done does quite a bit to harm ID's scientific reputation.

Neither evolutionary biology nor ID will be advanced or forestalled as scientific enterprises by pro- or anti-hagiographies of Charles Darwin (or any other individual scientist or ID supporter). This will happen only when sufficient field and laboratory work has been done and the results published in reputable scientific journals to decide between them. So far ID supporters have chosen to pursue a political program and the vilification of world-renowned scientists, rather than do the requisite science. Until they decide to abandon ad hominem arguments and actually focus on the science, they will remain (like "scientific creationism") a cautionary footnote to the history of the triumph of evolutionary biology.

In closing, I find it quite significant that only ID supporters refer to Darwin as a "little God". Like all evolutionary biologists, I consider him to have been a dedicated and talented observer of nature and a genuinely good person, but (like all of us) a plain and simple (and, of course, fallible) human.

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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen

Evolution, Intelligent Design, and the Banana Diet


There has been a lot of speculation lately about the status of "intelligent design". Last December, I posted an essay on the demise of the Intelligent Design and Evolution Awareness clubs. At that time it was clear that the IDEA club movement at American colleges and universities was dead. Despite protestations to the contrary, the available data indicate that this is still the case.

Which leads to a more interesting question: what is the current status of the intelligent design "movement"? This question is even more significant this month, as November 24th marks the 150th anniversary of the original publication of Charles Darwin's Origin of Species, the founding document of the science of biology and the most influential book ever published in the natural sciences. Although some ID supporters assert that ID is compatible with evolutionary theory, it is clear from even a cursory examination of their views that the majority of ID supporters are opposed to the idea of evolution in virtually all of its versions.

Which brings me to the point of this blogpost: what impact has ID had on mainstream evolutionary theory, and which of these alternative explanations for the origin of adaptations and the diversity of life is gaining in veracity and which is losing? One way to answer this question is to analyze the frequency with which the terms "evolution" and "intelligent design" appear in internet searches and in news stories in the mainstream media. To assess this, Google has a marvelous tool: Google Trends. The following two graphs were produced using Google Trends. I have copied the graphs directly from the relevant web pages, adding only the red line at 1.5 (on the Y axis) to make possible comparison of relative values.


The first graph (above) shows the frequency of the term "evolution" as it appeared in Google searches and in news articles on the web over the past six years. What the upper graph shows is that Google search volume for the term "evolution" has fluctuated over the past six years, but held relatively steady. The lower graph shows that there has been a steady increase in news articles about evolution over the past six years.


The second graph (above) shows the frequency of the term "intelligent design" as it appeared in Google searches and in news articles on the web over the same six years. As you can see, these graphs are markedly different than those for "evolution". These graphs show that prior to 2005 there were very few searches and almost no news stories for "intelligent design". In 2005 this pattern changed abruptly: by the end of the year, there were over ten times as many Google searches for the term "intelligent design", and a concomitant spike in news articles on the same subject. This spike corresponds to the Kitzmiller vs. Dover Area School District trialin Dover, Pennsylvania, in the course of which the ID movement gained national attention. The decision in that trial, issued by Judge John E. Jones (a Republican, appointed by President George W. Bush), was widely hailed as a massive defeat for the ID movement.

This assessment is borne out by the remainder of the graph. As you can see, both internet searches and news articles on ID have declined precipitously since December of 2005, falling to almost unmeasurably low levels.


This graph (above), comparing the search volume and news article frequency of the terms "evolution" (in blue) and "intelligent design" (in red) is even more revealing. Here it can easily be seen that both search volume and news articles on "evolution" have consistently dwarfed those for "intelligent design", even during the period covering the Kitzmiller v. Dover trial. Furthermore, there is a steady upward trend in the frequency of news articles on evolution, while the frequency of news articles on intelligent design has fallen to almost unmeasurable levels.

So, what are we to make of these trends? Clearly, interest in evolution is on a steady upward course, which just as clearly reflects its importance in the sciences. As I have written elsewhere, the fundamental concepts of the theory of evolution by natural selection have been steadily spreading into all of the branches of biology, and have recently begun to transform such disparate fields as psychology, literature, and even diets and nutrition and art.

By contrast, interest in intelligent design exploded in 2005 and has crashed since then. What other phenomena exhibit this same "boom and bust" pattern? One such transitory phenomenon is the explosion in fad diets. For example, this graph shows the Google search volume and news article frequency for the term "banana diet" (the choice of this term was at least partly in honor of young Earth creationist and ID supporter, Ray Comfort):


The similarity between this graph (above) and the graph for "intelligent design" is striking, and is probably no accident. As many critics of ID have asserted, the ID movement has essentially consisted of a public relations campaign, rather than a scientific research program. Public relations campaigns, like advertising campaigns in general, often show the "boom and bust" pattern shown in the "intelligent design" and "banana diet" graphs. This is because public relations campaigns do not depend on veracity, but rather on appearance. The supporters of ID (such as the Discovery Institute in Seattle, WA) have consistently promoted ID as a scientific research program, but even a cursory examination shows that, while there is abundant evidence that ID is a quasi-religious movement, there is virtually no empirical research being conducted by its supporters.

And so, as we approach the 150th anniversary of the founding of the rapidly expanding science of evolutionary biology, we may note in passing that this anniversary can also be used to mark the demise of the ID movement, a phenomenon with all of the earmarks of a passing fad and all of the scientific content of the banana diet.

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As always, comments, criticisms, and suggestions are warmly welcomed!

--Allen