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February 05, 2005

Eulogy: Ernst Mayr 1904-2005

Ernst Mayr has died at the very old age of one hundred.  It is hard to
imagine that he is gone.  I remember back in 1970 at a conference in
Indiana, where Mayr rather dominated the discussion.  I thought at the time:
"Well, we won't have to put up with this much longer."  It was not the last
time or the only way in which I was very much mistaken about Ernst Mayr.

Ernst Mayr was born in Germany, in Bavaria.  His father was a lawyer, but
the family tradition was medicine and it was that profession towards which
young Ernst was directed.  He was a keen bird watcher and, through this, was
diverted to a lifetime career as a professional biologist.  German
universities when Mayr was a young man were incredibly hierarchical.  His
professor (Erwin Stresemann) told him that, because there was another bright
young scholar (Bernhard Rensch) ahead of him, there was little chance of the
coveted professorship, and that hence he should seek other pastures.  Mayr
went west, and for the next two decades worked at the American Museum of
Natural History in New York.

For many reasons, this was an incredibly fortunate move.  First, he was now
working with one of the greatest collections of bird skins in the world.  He
was able to study variation in detail and range in a way that was impossible
for virtually anyone else.  He met and became close friends with a group of
men who were determined to put evolutionary studies on a new and proper
foundation.  Most important of all was the Russian-born geneticist
Theodosius Dobzhansky, closely followed by the brilliant paleontologist
George Gaylord Simpson.  He met Greta -- a fellow German - who was to become
his wife.  And, more than anything, he missed the rise of the Nazis, and the
corruption that that brought to his homeland.

As a student in Germany, Mayr had been rather disdainful of evolution.  I do
not mean that he was not an evolutionist - he never had any doubts on that
score.  It was just that he did not really think it a proper subject for a
professional biologist.  This may seem strange, and even stranger is the
fact that the man who was to become one of the dominating figures of
neo-Darwinism was at this time a committed Lamarckian - he believed that the
main causal force was the inherited of acquired characteristics.  In fact,
the odd thing would have been if Mayr had thought evolution worthy of study
by an ambitious young biologist, or had he been other than a Lamarckian.
For all that Darwin, back in the mid-nineteenth century, had discovered the
mechanism of natural selection - the survival of the fittest - evolution had
been captured by those (Herbert Spencer, Ernst Haeckel, Thomas Henry and
grandson Julian Huxley) who were determined to make evolution into a kind of
secular alternative to the dominant Christianity.  It was at best an
inferior science and at worst a vehicle on which any enthusiast could hang
wild metaphysical theories and hypotheses.  Selection was thought a minor
factor in significant change, and Lamarckism was one of the favored

Theodosius Dobzhansky changed all of that.  He was captivated by the
adaptive landscape metaphor of Sewall Wright, and used it as the basis of
Genetics and the Origin of Species, his paradigm-making survey of the forces
of organic change, first given as lectures at Columbia University in 1936
and then published as a book the year following.  As everyone knows,
Dobzhansky was an enthusiast, gathering up co-workers and students.  Mayr
was brought into the circle, and at Dobzhansky's urgings gave his own
lectures, and published the result as Systematics and the Origin of Species
in 1942.

By this time, everyone was starting to see selection as the most important
force for change, and Mayr's book reflected that.  His main contribution was
to demonstrate beyond doubt the variation that exists in nature, and how
everything points to gradual change - a key plank in the Darwinian program.
Particularly important was the evidence of groups that range all the way
from good species (interbreeding populations, isolated from others), to
species-in the-making, with subgroups starting to break apart, and on to
fully defined separate species.  Although Mayr's actual examples have been
challenged by later biologists, really significant were so-called "rings of
races" where on has a chain of groups (sub species) touching and
interbreeding, except at the ends that come together (completing the circle)
where the touching groups behave as distinct species.

The work of Dobzhansky and Mayr and Simpson (who published the
paleontological contribution, Tempo and Mode in Evolution, in 1944)
collectively founded what came to be known in America as the "Synthetic
Theory of Evolution" and in Britain as "Neo-Darwinism."  (The botanical
contribution was delayed.  Edgar Anderson was supposed to write it - he gave
lectures at Columbia at the same time as Mayr - but proved unable to produce
the book.  Dobzhansky then recruited G. Ledyard Stebbins, who published
Variation and Evolution in Plants in 1950.)

One must take some care however to understand the synthetic theory and
Mayr's contribution to it.  Most obviously the theory is an updating of
Darwin's theory in the Origin.  There, Darwin offered what his mentor, the
great historian and philosopher of science William Whewell, called a
"consilience of inductions."  First Darwin presented and justified his
causal heart, evolution through natural selection.  Then he used this to
explain biological phenomena across a wide range of subjects - instinct,
paleontology, geographical distribution, morphology, embryology, and more.
He argued that the success in explanation in turn provides confirmation of
the unifying mechanism.

Following Sewall Wright's work in population genetics, showing how Mendelism
blends with selection, apparently Dobzhansky and the others worked in the
tradition of Darwin, using population genetics (especially as incorporated
in the adaptive landscape) as the updated version of Darwin's presentation
of natural selection.  Then, Dobzhansky having put empirical flesh on Sewall
Wright's theoretical skeleton, the other synthesizers worked on the range -
Mayr on biogeographical distribution, Simpson on paleontology, and so forth.

There is a lot of truth in this picture, although one should not
over-emphasize the extent to which Wright and the early Dobzhansky were
enthused by natural selection.  Sewall Wright was (following both his own
father and his Harvard teacher, L. J. Henderson) an ardent Spencerian, and
for him upward progress and the balance between homogeneity and
heterogeneity was what really counted.  Selection was secondary, drift was
highlighted, and the same was true for Dobzhansky in the first edition of
Genetics and the Origin of Species.  It was around 1940 that everything
became a lot more selectionist, when Dobzhansky discovered synchronic
chromosome changes in separate populations of fruitflies.  Selection can
explain these and drift cannot.

Mayr fits into this picture, but rather uncomfortably.  The adaptive
landscape model is there, but it is not dominant.  Mayr used to deny (to me,
very often) that he really cared about the model at all - he said that he
just dropped it in to keep others happy.  This is not quite true, but it is
true that Mayr was not doing what Simpson was doing, namely trying to
complete the neo-Darwinian picture as sketched above.  Mayr was still
fighting the battles of the 1920s in Germany.  There, the big enemies of
naturalists like himself were the geneticists.  They were arguing that
really significant change is caused by macromutations, jumps from one form
to another.  Mayr and his fellows knew that this could not be true - nature
showed them that there was gradual change.

This was the battle that Mayr was still fighting in Systematics and the
Origin of Species.  In fact, Germany had come to New York.  One of the
leading scientific refugees from the Nazi oppression was Richard
Goldschmidt, who was arguing strongly for macromutations and jumps - what
are known in the trade as "saltations."  Mayr took him on and wrestled him
to the ground.  In fact, there was a satisfying personal element here.
Goldschmidt was a distinguished Herr Professor.  Mayr was an upstart, a
museum worker.  (Hard to imagine him in this role, but it is true.)
Personally they got on well - Mayr loved to tell the tale of taking
Goldschmidt home one Saturday lunch for soup prepared by Greta - but
fighting this battle was for Mayr very much (to change metaphors) very much
a matter of winning his spurs.

The battle also helps to explain what always struck me as a rather curious
feature of Mayr's attitude to his science.  For all that genetics was
fundamental to the synthetic theory, and for all that in the early 1950s
Mayr himself extended Sewall Wright's thinking - Sewall Wright said that
Mayr pinched Sewall Wright's thinking - about the significance for evolution
of isolated groups that undergo genetic revolutions and rapidly become new
species (the so-called "founder principle"), Mayr was forever rude about and
belittling of genetics.  Not molecular genetics, so much, as Mendelian
genetics.  He spoke of it contemptuously as "beanbag genetics" (causing J.
B. S. Haldane to speak in favour of beanbag genetics), he excoriated its
reductionistic nature, and somewhat pettily when in the early 1970s he
organized two major conferences on the founding of the synthetic theory, he
did not invite Sewall Wright.

It would be easy to say simply that it was all simply personal - for the
record, I think the founder principle does come out of Sewall Wright's work,
but that Mayr deserves full credit for its development and application (and,
for all that today evolutionists are questioning its significance, that the
principle has proved itself a very important guide to productive research).
It would be truer to say that the roots of Mayr's hostility lies back in the
battles when he was a student and genetics was the enemy of naturalists,
rather than the foundation of modern evolutionary theory.

It would be unfair to say that after Systematics and the Origin of Species,
the rest of Mayr's career was simply footnotes and appendices.  Apart from
the founder principle, he was a very important systematist, working on the
theory of classification.   It is true that around him grew up the new
taxonomic approach of phylogenetic systematics or cladism, and Mayr could
never buy into this.  In fact, in respects he became the Goldschmidt for the
new generation of Young Turks.  But without genetics, Mayr could not have
been what he is, and I suspect that without Mayr modern taxonomy could not
be what it is.  However, one does sense that after Systematics and the
Origin of Species, to use yet another metaphor Mayr was skiing down the
other side.   In 1963 he published a major synthesis, Animal Species and
Evolution, but it was more a synthesis of what had been done than a work to
inspire for the future.

Conceptually that is.  Because Mayr was now about to make an even more
significant contribution to twentieth-century, evolutionary biology.  The
1940s saw major moves by the evolutionists to bring workers together and to
make a formal discipline from and for evolutionary studies - a journal, a
society, grants, students, jobs, and the rest.  Mayr was in the thick of
things.  He was the first editor of the new journal Evolution, and showing
that he was German in more things than one, whether accepting papers or
rejecting them, he never wrote a sentence when a paragraph would do.  He ran
the society.  Once, somewhat sardonically he said to me:  "Unlike my good
friends Dobzhansky and Simpson, I never thought that the only position I
could take in an organization was that of president."  He moved to Harvard -
the immigrant boy made good, as Dobzhansky once said of himself and Mayr -
and worked unceasingly to promote organismic biology at that institution, at
a time when the molecular biologists were all conquering, and not very nice
about it either.

To quote Ed Wilson:  "Jim Watson?  The most unpleasant man I ever knew."
(Actually, one of Mayr's daughters dated Watson for a while.  Although more
on this topic in a moment.)  The fact is that without Ernst Mayr we would
not have the professional discipline of evolutionary biology that we have
today.  I do not mean that nothing would exist - others were organizing,
including Dobzhansky with his many students and (over in England) E B
"Henry" Ford was also showing himlsef a master of organization and funding
finding.  But without Mayr, things would simply not be as well developed as
they are now.

But Mayr had many more years of active life.  Even last year he was
scrounging one of my books from our shared publisher, Harvard University
Press, so that he could put the boot into me one more time before he was
done.  By about 1960, Mayr was starting to move interests from pure science
to its history and its philosophy.  I know only second-hand about the
history, although I do know that he was much involved in the development of
the History of Science department at Harvard, he worked hard to promote the
careers of young scholars (Frank Sulloway, notably), he himself organized
the above mentioned conferences on the history of the synthetic theory (and
then co-edited with Will Provine the collection of the proceedings)  and
Mayr was one of those who worked hard to get the Journal of the History of
Biology off the ground.

Mayr himself was much interested in the history of biology and eventually
was to write a massive work on the history of evolutionary biology, the only
volume of what had been intended to be a complete history of the life
sciences.  When he had finished the evolutionary side, he realized that he
had said what he wanted to say.  For one should never think of Mayr as
interested in history for its own sake.  He was always writing for a purpose
- a purpose that I would describe as philosophical.

As a student in Germany, Mayr had studied the classics of philosophy - and
he came from the cultured middle-classes, where discussion of philosophical
issues would have been normal rather than a sign of pretension - and so, at
one level, philosophy was part of his very being.  But, by about 1960, it
became much more.  It became a tool in his fight for his vision of the life
sciences, where evolution took an honored and central place.  (When
Dobzhansky said that nothing in biology makes sense except in the light of
evolution, he was not just making an epistemological claim.  He was making a
political statement.  A war cry to rally the troops.)

I have said that I never sensed in Mayr's attitude towards molecular
biology, the deep hostility he felt to the genetics of the 1920s.   Early
fights and hatred run more deeply and last forever.  I have disliked people
in my life, but never with the visceral hatred that I still feel towards my
headmaster.  (Mark you, he was late middle-aged, and I suspect his feelings
about me prove the exception to what I have just said about early
experiences causing the really deep antagonism.)  However, whatever the
ordering of the threats that Mayr felt, it is undoubtedly true that with
good reason he could see that by 1959 - the year of the centenary of the
Origin and of the celebration of the triumph of neo-Darwinism - the
molecular biologists were grabbing all of the goodies.

Not just the Nobel prizes, but the students and grants and research space
and much more.  The biology of whole organisms was being pushed aside.
(Read Jim Watson's Double Helix if you want more proof.)  Mayr was ever a
fighter.  He was not about to sit by and let this happen.  He turned to
philosophy to prove that organismic biology will always have a place in the
sciences, that it cannot simply be subsumed under - absorbed into - the
molecular sciences, of physics and chemistry.  In other words, he started a
fight against what he saw as unwarranted reductionism.  Not because he was a
Marxist (like Steve Gould and Dick Lewontin in the decades later), but
because he wanted to preserve the autonomy of those parts of the life
sciences that had evolutionary thought as central.  If he could make his
case, then biology as he knew it could continue to survive and flourish.

And so from Mayr, as from others working alongside him (notably Simpson), we
got a stream of articles and writings and conference interjections promoting
the autonomy of whole-organism biology.  We learnt that (unlike the physical
sciences) it has a differentiating historical dimensions, that it deals with
unique phenomena, and that above all it is teleological - forward-looking.
Although, in a clever move, realizing that teleology has a bad name in
biology - Mayr himself battered would-be contributors to Evolution for
teleological tendencies - he took up the name of "teleonomy."  This is
sanitized teleology - forward-looking, dealing with functions, without the
unneeded vital forces and so forth.

Above all, Mayr was a holist, meaning that he thought breaking things down
to small components is not only not necessarily the right way to go in
biological science, it is often positively exactly the wrong way to go.  For
this reason, when Michael Ghiselin and David Hull in the 1970s began arguing
that species are not classes - groups of member organisms - but individuals
- integrated wholes - Mayr embraced their thesis with enthusiasm.  And it
was for this reason that Mayr wrote a lot of his history.

It is true that old battles were not forgotten.  Those dreadful geneticists
had ignored variation, the basis of Mayr's gradualism, and so a lot of the
history was devoted to showing that that rotter Plato had illicitly
introduced essentialism - the idea that groups have no variation - into
biological thought.  Only slowly and gradually, thanks primarily to Darwin
and to a certain immigrant to the United States of America, had population
thinking finally triumphed.  But this history telling was only part of the
story, for the important underlying message was that whole-organism thinking
has a grand tradition and, as one recognizes this, one recognizes that such
thinking has its own autonomous problems and ways of solution.  Molecular
biology is important, but only a molecular biologist would think it all
important.  And that in itself tells you something about their limitations.

Was Mayr successful in his efforts?  Well, yes I think he was, although how
much was due to Mayr himself and how much to others, and what the nature of
that success are perhaps questions that yield answers Mayr would not
entirely have liked.  On the one hand, Mayr's labours in the history and
philosophy of science - always as much organizational as conceptual - paid
off greatly in the development of both the history and philosophy of biology
as thriving enterprises as we have them today.

Mayr encouraged many of us, and helped in so many ways.  He may have
irritated me - I have never been much of a holist - but he encouraged me and
said very kind things when my first book appeared.  When I founded Biology
and Philosophy, I knew that Ernst Mayr (and David Hull) would always write a
report on a paper.  I generally knew what he would say before I sent it to
him, but it always came back with lots of useful comments.  And I know that
others have had the same experiences.

At the same time however, in history particularly, my suspicion is that
generally the (justified) feeling is that Mayr's extra-historical motives
distort his thinking to the point where what he wrote is of minimal value.
Certainly the feeling among historians is that not only is the essentialism
thesis just plain false, but that we have wasted a lot of valuable time
getting over it.  Perhaps this is just a reaction of the present, but I
suspect there is some truth to it.

On the other hand, today organismic biology does have a full and honored
place in the biological spectrum.  Far from the molecular biologists
swallowing up evolutionary biology, the molecular biologists have been
turned on by the problems of evolution.  Witness the thriving success of the
field of evolutionary development (evo-devo).  But this in a way is the very
point.  Evolutionary biology today is so exciting because of molecular
biology, not in the face of it.  Bring on the molecules, we all say.  A
rather different cry from that of the Ernst Mayr of the 1960s.

What sort of man was Ernst Mayr?  I trust that Ed Wilson of Harvard and Guy
Bush of Michigan State, both men who encountered Mayr at Harvard when they
were very junior, will not think I am betraying confidences when I say that
he was not always much fun for the young and less established.  He could be
formidable to the point of - beyond the point of - bullying.  As is often
the case, one suspects that this concealed certain insecurities - the
foreigner in the new world, in a field that is under siege.

But it does not make for easiness, whatever.  At times, Mayr was desperately
- almost pathetically - eager to make his own place in history.  He truly
loved Dobzhansky and admired him greatly, but worried that he (Mayr) would
always have second place.  It is easy to joke about this, as I did above,
but it has an edge nevertheless.  He was deeply hurt that, at the Origin
centenary celebration in Chicago in 1959, Dobzhansky, Simpson, and Julian
Huxley were given honorary degrees and not him.

However, insecure or not, no one could deny that Ernst Mayr had a deep love
of the living world.  To walk with him around his cottage in New Hampshire
was to see a man who simply loved animals and plants - loved them for their
own sake, loved them because he was trying to understand them.  Mayr was a
man who could think of no higher calling.  He told me that reading Ernst
Haeckel as a child had turned him away from Christianity.  He joked, when I
admired a photo of him in his twenties in New Guinea, where he was wearing a
beard, that people took him for a missionary and he shaved at once!  He was
much influenced by Julian Huxley's Religion without Revelation.  He told me
that he felt with Huxley and with others (including, very much, Wilson) that
it is possible to be a deeply religious person in the complete absence of
theology.  He meant it and proved it.

It is certainly true that, although Mayr may have been rough on people like
Wilson and Bush, he respected them and other biologists in a way that he
never felt about others, including historians and philosophers.  I always
felt a bit like a court eunuch when I was around him.  Useful and
interesting, but ultimately not a real man.  Or perhaps court jester.  Mayr
was much easer on us than on the biologists, but that was because we existed
only to help biology.  This did not preclude real friendship.  Over the
years he and I built up a close relationship.  I did earn some respect.  At
a discussion group in the 1980s at Harvard, I told him to his face that I
thought the species-as-individuals thesis is nonsense.  He came at me across
the room, shaking his finger, and yelling.  I stood my gourd, and got
invited to lunch the next day!

I never sensed that Mayr was a man deeply engaged in politics.  He told me
once that as a young man, like everyone in his class, he was mildly
anti-Semitic, and then mentioned the Holocaust and remarked how far we had
moved since then.  After Mayr retired, one of his secretaries was a gay man,
who died of AIDS.  I saw genuine concern and affection.  He liked kids - my
children thought he was a terrific old man - and he (unlike a lot of
academics) always treated my wife Lizzie with affection.  He remembered her
name without prompting.  Some of my fondest memories are of lunchtime soup
with the Mayrs and of his wife pushing him around and making him go for an
after-lunch walk.

My favorite Mayr story is of an event about twenty years ago.  A French film
company decided to do a one-hour biography of Mayr.  When finished, flyers
were sent far and wide, advertising this film.  Memorably, it described Mayr
as a man of intelligence, a man of humility, a man of humor.  I wrote to
Ernst saying that I certainly thought of him as a man of intelligence, and
that although he was a German I was sure that at least once in his life he
had cracked funny joke, but that - think goodness - he was not a man of
humility.  He was a man of pride and determination and no nonsense about
falsely putting himself down.

Mayr was absolutely livid.  What did I mean by saying that he had no sense
of humor?  He was the funniest man that he knew.  The next time we met, he
would order me to listen to his jokes and to laugh.  (Actually, his favorite
joke with me was to call me at about ten in the morning and apologize for
getting me out of bed.  Mayr was famously an early riser - four in the
morning, every day.  He told me that at the beginning of the War, when he
was an enemy alien - he had not then taken out citizenship - and fearful for
his job at the Museum, he had decided to be a milkman if all else failed.
He liked getting up early in the morning, and he liked the outdoors life.)

My final memory is four years old.  I had just retired from thirty-five
years of teaching in Canada, and moved to Florida State University here in
Tallahassee.  With my professorship comes money for (and expectations of)
putting on conferences.  So for the first, in 2001, I organized a conference
on the philosophy of biology.  Mayr was still in the habit of spending the
winter at a small college in central Florida, and Betty Smocovitis at the
University of Florida - an old friend of both Mayr and me - arranged to
bring Ernst along.  All sorts of prior warnings were sent that Ernst was old
and frail and would need lots of time out.  Although the conference was from
Friday to Sunday, he would need to leave on Saturday afternoon.
Unfortunately he would not be able to go on the field trip to Wakulla
Springs, a nature reserves full of birds and alligators (and the site of the
original Tarzan movies as well as of The Creature from the Black Lagoon and
Airport 77.)

Along came Betty and Ernst.  Inevitably as soon as the paper was over, Ernst
was on his feet.  This continued for the rest of Friday and all day
Saturday.  Who, on Saturday at Wakulla, was at the front of the boat?  You
guessed it!  Ernst Mayr having a wonderful time, telling us that this was
his first visit to Wakulla since 1931.  And who saw the Saturday barbecue
out to the bitter end, until finally I said: "Ernst go to bed."  None other
than our frail old friend and evolutionist.

Mayr once told me that the greatest influence on his life had been his
mother, who had impressed upon him the obligation of people with his talents
and background and education to serve others.  I wish I could tell his
mother how proud she could be of him.  I wish I could tell his father, a
deeply educated and cultured man, how proud he could be of his son.  The
truly great person is not he or she who simply does things easily, without
strain.  The truly great person triumphs above his or her limitations, and
makes them seem trivial and unimportant.  Ernst Mayr was a truly great man.
What a life.  What vitality.  What service.  What a privilege to have known

Ernst Mayr at FSU, 2001

Posted by Michael Ruse on February 05, 2005 at 06:24 PM | Permalink


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A very nice piece. Even though you are British, you deserve a nice meal for it, so get somebody else to cook it. I gladly volunteer.

I think you are right on the mark about Mayr and his contributions to the history of biology. Making history and writing it are indeed two things, as even Churchill knew.

I also agree with you--how scary is that—about your characterization of the role of the 1920's fight between naturalists and geneticists in Germany. I just want to offer one caveat. All those geneticists were great naturalists themselves, so it is more a fight about methods and the choice of the right question, which brings us to the role of experiments and into the best of all possible periods, the 19th century and the role of Helmholtz et al for setting the standards for science. Maybe you want to clarify this somewhat. The way you generalize about genetics of the 1920s is not quite right, I think.



Posted by: Manfred Laubichler | February 5, 2005 06:37 PM

I briefly met John Maynard Smith - another lamented departure - a few years ago and the talk turned to Mayr. "Oh, Uncle Ernie is one of my favorite people" he said, nearly causing me to choke on the sandwich he had given me. I never thought of Mayr as an "Uncle Ernie"...

Posted by: John Wilkins | February 5, 2005 08:13 PM

Michael--it is indeed a nice piece. You neglected to add that he did a great deal of this on one kidney. The other one was removed in the early 1950s when it was found to have a tumor.

He and Ledyard Stebbins disliked each other immensely to the end. Ledyard--a life-long liberal-- would spit things out like "I bet Ernst is a Republican" when he got angry (he wasn't of course). Ernst would spit things out like "Ledyard is such a child."

Posted by: Vassiliki Betty Smocovitis | February 5, 2005 08:49 PM

Here is a little story some of you have not heard. Ernst was a participant at the first IUBS symposium "Towards a Theoretical Biology" in 1966. There is a picture of Ernst at that symposium on the inside of the front cover of the published volume "Towards a Theoretical Biology 1. Prolegomena." He is wearing a checked short-sleeved shirt. Fast forward 27 years to the summer 1993 meeting of ISHPSSB at Brandeis. Ernst was there wearing a light-green and white checked short-sleeved shirt that looked suspiciously like the one in the 1966 picture. So I asked him about it, and he said it was indeed the very same shirt. He said something about giving it to me someday, but that never happened.

Malcolm Kottler

Posted by: Malcolm Kottler | February 6, 2005 02:31 PM

Michael mentiones in his splendid piece that Ernst "was not always much fun for the young and less established". I cannot judge the situations he is alluding to but I do have very different experiences with Ernst. When I met Ernst in 1982 for the first time at the Darwin centennial conference in Cambridge, I was really everything else but established -- AND I was a physicist turned philosopher. But I was always treated by Ernst in an absolutely decent way, he always took me seriously, he never made me feel the incredibly huge gap in our scientific status, and he was always supportive. (Of course, discussions about reductionism were not always easy with him, and after a while, I gave them up.) Sometime during the late 80s or early 90s, he took me along to the Harvard Faculty Club as his guest when Quine's guest was Follesdal (surely I was flattered!).

Here is my favourite Ernst story. Ernst was 42 years senior to me. He once introduced me to someone with the words: "This is Paul. He is an old friend of mine."

Posted by: Paul Hoyningen-Huene | February 7, 2005 04:15 AM

When a small group of titanic individuals re-make something - the US Founding Fathers and the United States, the neo-Darwinians and the evolutionary synthesis - historians have work for decades trying to figure out which are the contributions of each titan and how the product might have looked had Titan One or Titan Two been absent. Perhaps historians can also be characterized as holists or reductionists, those who might see the product as completely different had Titan One been absent, and those who might see the product only measurably distinct.

From my perspective, without Mayr the entire product of the evolutionary synthesis would have been different. It seems to me, as a practitioner, with an acknowledged concomitant limit to my historical perspective, that Mayr contributed in four ways that influenced beyond easy description the structure of modern evolutionary biology. Michael Ruse mentions all of them, so perhaps our interpretations differ only in nuance. First, Mayr drove home the problem of continuous variation. Mayr's work hammered home the fact that macromutations, as hypotheses about interesting, evolutionarily relevant variation, are few in number and rare in consequence. Elaborate studies of mutational effects confirmed Mayr's claims, largely based on measuring and observing extant variation. Second, Mayr almost created the viewpoint of "population thinking," although we could give credit to Ford as well. While everyone used the word "population," Mayr seemed to be the foremost proponent of using the word in an operational sense. Third, Mayr connected the process of speciation to genetic mechanisms in a way that no one else was doing. While his verbal model of founder effect speciation (peripatric in today's nomenclature) was, ahem, inconsistent and perhaps at times not perfectly in synch with mathematical genetic logic, he is the founder of an intellectual lineage that would include Carson, Templeton, Bush, Orr, and even Gavrilets (even if many of them might not think so). Fourth, Mayr played a critical role in what we would now call the infrastructure of science. His strong hand as editor of EVOLUTION, the influence of his books (which were used as textbooks for a few generations of students, including mine), and his unending arguments over nearly every important subject in evolutionary biology molded, eventually, the discipline as it exists today, at least to a very significant extent.

Historians have a rich bed of material to mine to decide which of these four might be ranked most highly, and I have no doubt that some of my colleagues, historians or biologists, might delete one of those four and add another. That we can have this argument speaks to Mayr's power as an intellect and his intellectual omnipresence.

Two aspects of this legacy strike me as worthy of thought. First, we often glance sideways at ourselves when contemplating building infrastructure - as in, the best thinkers keep thinking and writing and those who get tired start doing service. Those who get very tired take up outreach. Let me make full disclosure: I have spent a lot of time in service, as editor of THE AMERICAN NATURALIST and in university administration, so if the reader thinks I'm being defensive, so be it. I also point out I have kept thinking, writing, and turning out research, for what that might be worth. Now, disclosure made, I suggest that at least in Mayr's case, there is much to be said for building infrastructure. In many ways, his influence on what we do, what we teach, and how we think about evolution might be much the greater for that fourth contribution than the others. Anyone who has been an editor of a major journal, and a heavy-handed one at that, knows the power one wields and the ability one has to influence a discipline. After all, an editor molds hundreds or thousands of manuscripts, depending upon his or her stamina, while authors mold, well, dozens, maybe.

The second point that strikes me about Mayr is that he was so influential even though he seems to have been wrong in his thinking about some issues. I was never struck by the command of population genetics I saw in his writing nor by the grasp of the logic of thematical genetic arguments that he was criticizing and I never found his attack on beanbag genetics to be compelling or even strong (although I much enjoyed the lecture he offered at the Werkmeister Conference that Michael describes). I don't think he fully grasped Wright's arguments (which, contrary to how I read Michael's comments, did not emphasize drift over selection, but a role for drift in creating a lot of genetic options in different local populations among which selection might choose) and the logic of his hypothesis about founder effect speciation seems a bit slippery. But stepping back, I see these mis-steps as rather minor scrapes on a rather imposing and impressive suit of intellectual armor. That Mayr got everyone thinking about these arguments, and perhaps did his part to inspire serious research on the genetics of speciation, seems more than enough to compensate for whatever skewness characterized his opinions.

I had heard the stories of his imperious nature and argumentative disposition but had never met him until the Werkmeister Conference that Michael describes. My favorite moment in that weekend was at Wakulla Springs, where I was leading the group outing, describing natural history and the local culture, mindful that Mayr himself had visited there in the 1930's as a working ornithologist. A graduate student who did not know his entire history engaged him in conversation, and Mayr and the student were discussing birds that we were seeing. The student said, innocently, and here I paraphrase, you seem to know a lot about birds, are you very interested in them, to which Mayr responded, with graciousness and warmth, why, yes, I have long been interested in birds. The student and Mayr continued in this vein for several minutes, through which Mayr gave no hint that, should he choose, he could talk rings around all of us about birds.

I would like to be that gracious when I grow up.

Posted by: Joe Travis | February 7, 2005 02:49 PM

Actually, one of Mayr's daughters dated Watson for a while. Although more on this topic in a moment.

Unless I missed it somehow, we're still waiting for the "more". At least, I am. Waiting, I mean. With bated breath.

(That was a wonderful piece. We should all be remembered with such warmth and eloquence.)

Posted by: sennoma | February 9, 2005 05:11 PM

A wonderful essay – I sent it to all of my graduate students. Something I always thought was curious about Mayr was his dismissal of sexual selection as a force in evolution. I don’t know whether his attitude had anything to do with the sociobiology controversy of the 70s and beyond; certainly he was a mentor to Bob Trivers, among others. But I was interested to note that several years ago, in a review that Mayr wrote of Janet Browne’s biography of Darwin, he persisted in this belief. When he mentioned Darwin’s Descent of Man and Selection in Relation to Sex, he referred to it only in passing, and as a book only concerned with humans. Of course, this is not the case – Darwin was extremely interested in the evolution of differences between males and females, and thought sexual selection was very important.

This is also interesting in light of Mayr’s feeling slighted by not receiving the Nobel Prize, but consoling himself with the thought that Darwin would not have received it either. Of course, a Nobel Prize was indeed awarded to scientists in his field, at least if you interpret it broadly: the 1973 Prize in Physiology and Medicine went jointly to Karl von Frisch, Niko Tinbergen, and Konrad Lorenz. Although they studied behavior, they were forerunners of modern behavioral ecology, which has a firm connection to evolution in the best Mayr tradition.

Posted by: Marlene Zuk | February 10, 2005 08:04 PM

I very much appreciate Michael's eulogy of Ernst, which I have shared with family, friends and students. I appreciate it all the more for my memories of Ernst and Michael jousting and joking about this and that.

What follows is a late addition, in the sense that I wrote it last week. There are a few other reminiscences and stories about Ernst that I would also like to share. I will post them asap.


5 February 2005

I'm sad about Ernst's death, and worried about Carolyn's mom, who is in the hospital.

And I'm thinking of the day that Carolyn's mom met Ernst.

Ernst and Gretel had invited Carolyn and me to visit them at their "farm" one Saturday during the fall of 1982. (This was the "summer place in New Hampshire" from which the warbler migrated south in Ernst's famous 1961 paper, "Cause and Effect in Biology.")

Carolyn's mom was visiting from Texas, so they welcomed her as well.

Things didn't get off to a perfect start. Upon entering the house, Carolyn's mom admired a houseplant that Ernst and Gretel had received as a gift. Ernst said, "Yes, it is a beautiful [I can't remember what]." Carolyn's mom said, "No that's not what it is, it's a [again, I can't remember what]." Of course, Ernst the systematist (if not botanist) would not let it go, and neither would Carolyn's mom. Finally, Ernst suggested putting the matter to rest by consulting a field guide, which he pulled from the shelf. Carolyn's mom was right.

Ernst then suggested that, before we got settled in, we should take a walk. It was as usual a wonderful walk, with as good a natural history guide as we could have had (I mean Ernst!).

Along the way, Ernst collected chestnuts. It was a chilly day, and when we got back to the house, Gretel had a nice fire going. Carolyn's mom promptly asked if we could roast the chestnuts that Ernst had gathered, since she had never eaten roasted chestnuts. Ernst started to blurt something out. But Gretel interrupted him and said that that was a wonderful idea and got a pan for the purpose. An hour or so later, the chestnuts were gone. At which point Ernst could no longer contain himself. He explained that we had just eaten seeds from very, very rare American chestnut trees, a species which had gone almost completely extinct. He explained that he gathered the chestnuts every year and sent them to a place where they were carefully cultivated, with the hope that they would someday be transplanted in large numbers back to their original habitat. "I hope we have not eaten them into extinction!" he complained. But by then he was himself amused by the whole affair.

Carolyn's mom was better behaved the rest of the day (!). Just before leaving, we picked some Concord grapes to take home. The next day, Carolyn and her mom made grape jelly and I took some to Ernst in his office. One or two days after that, Carolyn and her mom and I ate some of the jelly and discovered that it was really spicy, and then realized that it had been packed in jars that contained jalapeño peppers that Carolyn's mom had brought from Texas! Carolyn and her mom had packed the "gift" jelly in a jalapeño jar as well.

I saw Ernst a few days later and the first thing he said was that if the gift had been intended as a practical joke, it had backfired, because he thought the hot pepper jelly was delicious!

I miss you Ernst and Gretel.

Get well Joan!

Posted by: John Beatty | February 12, 2005 10:37 AM

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