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SAMUEL PROCTOR ORAL HISTORY PROGRAM at
the University of Florida
K: My name is Christopher Koehler. This is March 14, 1994. I am interviewing
Professor Lincoln Brower. I was wondering if you would please state your full name
and provide spellings.
B: My name is Lincoln Pierson Brower. I live at 9615 SW 43rd Terrace in Gainesville,
Florida 32608, on the edge of Paynes Prairie preserve.
K: I was wondering if you might start by telling me where you were born.
B: I was born in Summit, New Jersey, September 10, 1931.
K: What were your parents names?
B: My mother's name was Helen Romer Pierson and my father's name was Bailey
Brower. My father's family came from New York on one of the Dutch immigrant
vessels and my mother's family traces back to the French Huguenots who were
religiously persecuted, left France, and came to England. On my mother's side, I
come through the D.A.R. (the Daughters of the American Revolution), and on my
father's side through the Holland Society. [My father's] family left Holland following
the damming of the Zieder Zee, which destroyed the seaport of a small town of
Holland called Enkhuizen, a fishery village. After the dam was built up to reclaim
more land, they left. That is the story of my family. My father's side stayed in the
New York area. My mother's side was near New York on the Hudson River, then
moved out to Morris County, New Jersey on the edge of the Great Swamp, which is
a magnificent wildlife preserve now.
K: That sounds like a very distinguished lineage. Was your family much involved with
the D.A.R. and the Holland Society?
B: I was a Depression baby and my family went through financial hell in 1929 with the
collapse. So all my involvement with those august societies was pretty much
secondhand. I remember in Morris County, New Jersey a fantastic Fourth of July
parade and my grandmother, who was very much involved with the D.A.R., was
riding on one of the floats. That was a long time ago.
K: Did you also grow up in that same city in New Jersey?
B: The homestead that I lived on came down through my great grandfather's family and
it is in western New Jersey in an area called Chatham Township. It is near Madison
which was on the Lackawana Railroad [that] opened up western New Jersey early
on for commuting to New York City. When I was a young lad, my family had this
enormous farm and they grew roses and they also had milk cattle--a dairy. The
houses of the different family members were around this farm, and I think it was
originally about a thousand acres, and now it has been totally decimated and wiped
out and there is a very well-to-do residential suburb of New Jersey or New York
[there], as the case may be.
K: This is really a case then of it being a family seat, as it were?
B: It was the family seat for four generations.
K: Can you recall any experiences that strike you as characteristic of that place or that
B: I was born in 1931, so the 1930s were taken up with the whole problem of the
Depression and surviving economically. Then World War II kind of dominated my
early youth and following World War II there was economic recovery. We are talking
about 1939 to 1945 and that was a period of really dramatic change in New Jersey
from what had been a very rural area to an area that was perceived by developers
as ripe for economic development of fairly well-to-do upper middle class
My grandfather had been in the greenhouse business. He was quite an inventive
man and a very fine grower of roses. If you go back far enough prior to World War
II, he developed the beautiful roses that [had] four-foot stems. We had this
wonderful old pond that we used to go swimming in. It had been the ice pond for the
rose company prior to the advent of refrigeration and the old ice houses were still
there. It was really amazing. What they did was cut blocks of ice with horses that
walked out on this pond with great mechanical saws and they then ground the ice,
chopped it up, and packed it in the big boxes. And they shipped these four-foot long
roses to Buckingham Palace in London by ship. During the height of the winter
when things were at a low time for flowers in England, the royalty would buy these
roses. That was very exciting.
At the end of World War II, there was a boom. The family got back on its feet
economically toward the end of the war because roses could be grown and sold and
there were not a lot of material goods after World War II. Well, that boom suddenly
burst because of the development of the airplane industry. It made it feasible to fly
flowers in from Hawaii, and from Central and South America. So shortly after World
War II the rose company went down the drain.
By about this time now, 1949, I went to Princeton University, receiving my degree in
1953. I was off to another life. But in 1936, [when] I was five years old, we used to
have this wonderful tennis court on our land and my family spent a lot of time
playing tennis and we all got really good at it. But at the age of five, it was a little
more than I could take, so I would lie down on the lawn. Those were the days when
the lawns were full of weeds. They were very well manicured, but they were
biologically diverse and so there were all kinds of insects crawling around and flying
around that lawn. We had a lot of clover in that grass, and I remember seeing this
beautiful little Lvcaena hypophleas, which is a little copper butterfly about as big as
your thumbnail. I was just intrigued as I was watching this thing as I was lying in the
grass. That sparked my interest in butterflies.
There had been an old German immigrant who settled in a nearby village called
Green Village. Green Village was on the edges of the Great Swamp of New Jersey.
This man's name was Charles Rummel and he was a lepidopterist, a student of
butterflies and moths. He spent his entire life collecting butterflies and moths during
the Depression and afterwards. He had retired and he had all these wonderful trail
systems throughout the Great Swamp. I used to walk out with him. He had these
traps that he baited with a mixture of beer and molasses and rum and what have
you. All these wonderful moths and insects would come into these things. So I
became very interested in moths and butterflies at the age of seven. With the
exception of about a two-year hiatus in high school when I became interested in
young ladies, [Laughter] I have stuck with this interest in butterflies and moths all my
life. And I decided before I went to Princeton when I was a junior in high school--I
found a lady who was also interested in butterflies so that made it even better--that I
would major in biology and go into the biology profession. This was at the age of
about sixteen years old, seventeen years old. And that is what I did.
K: So your choice of career was sparked when you were five and set by time you were
sixteen years old?
B: Yes. It was also sparked by my family's interest in growing things because, while I
never had anything to do with growing roses, it was always fascinating to watch
them do it. And the family used to talk about that and I learned about variability in
plants. My grandfather did not have a college education, but he was a brilliant
person and he dimly understood heredity. He used to talk about "sports" of flowers--
suddenly there would be a stalk that had a different color rose on it. And they would
develop those and sell them. He actually was responsible for developing what came
to be called the American Beauty Rose by selection. So I became interested in that.
Plus this area of New Jersey that we lived in was very interesting geologically
because it was on the edge of the last advance of the Wisconsin glaciation. So
about ten thousand years ago, Madison, New Jersey was right at the border of the
ice and in that area you have a very dynamic terrain. Madison, New Jersey itself is
right on the edge of a huge glacial moraine and within that glacial moraine are these
kettleholes. (A moraine is formed by the outflow of water carrying large amounts of
sediment. The sediment then builds up on the land, but there are some big chunks
of ice that get left there--tons and tons and tons--and these get buried by the sand.
Because they are buried, they are insulated. Then the rest of the glacier retreated
and eventually the big chunks of ice melt and they left huge depressions. These
depressions are called kettleholes.)
There is a whole series of kettleholes in that part of New Jersey and one of the
things that is really interesting about [kettleholes] is they are full of really beautiful
wild flowers. As a young lad, my mother used to take me down into these
kettleholes. There is one in particular that we used to visit. So she got me
interested in looking at biological diversity of that sort at a very early age. I used to
work and help [my grandfather] a lot as a kid. He loved to garden as an old man
and I learned a lot about gardening. I also learned probably more about how to do
science working under him than I ever learned in college. [Laughter] He was very
fussy about, "Now Linc, you have to line up all these tools in a row before you do
anything. No, no, no. You line them up in the order that you are going to use them."
And if you ever put a rake upside down that was a virtual death sentence. Then we
would start spading the garden and you had to throw the dirt ahead and turn it and if
you did not do it right you would get a long lecture on how to do it right. And my
grandfather did everything right. Eventually, we would have to rake the whole thing
and then get on top of it with our hands and get out all the stones. The soil was
incredibly receiving when it was finally planted. And anything you grew, grew
beautifully. And so I learned orderliness and scientific predictability and so forth
from him. I think that that later on played a very important role in my own training of
students and insisting that they do things right.
It is really interesting to think back on that because my grandfather would say we
had to do things. [And] he was not the sort of man that you would ask "Why?" That
was what college was for. The teachers were paid to answer your questions when
you asked "Why?" [Laughter] But with my grandfather, you just did it that way, but
clearly there was a good reason for it. He proved his point by the results that one
got. And just about a year ago I was thinking about that sort of philosophical
approach of orderliness toward life. We were trying to raise a difficult butterfly here.
It is a really interesting one genetically and in the evolution of mimicry. Anyway, we
were trying to raise [this butterfly] in the lab. I know how to raise butterflies; I have
done it all my life. But all of a sudden, these butterflies stopped eating. The
caterpillars stopped eating their leaves. And I had done everything according to
Hoyle. My students were there. I had a couple of really good students who were
disappointed by the complete collapse of our project because all the caterpillars
stopped eating and then they just gradually starved to death. As a scientist, I did not
have a clue what was killing them. We had done it the way we ought to have done
it. We followed the precepts that I learned at the age of five [that were] reinforced in
college and graduate school and still [it] did not work.
It was really interesting. We took these larvae over to the entomology department
here. One of the things that is nice about the University of Florida is that you have
got all these different colleagues in different fields who are specialists and really
know their fields. So we took the caterpillars over to Professor [Drion G.] Boucias
[Associate Professor, Entomology and Nematology], who is an insect pathologist. I
thought we had changed the food plant on these butterflies. By that I mean rather
than taking the leaves off tree A from my back garden, which I was decimating
because we had so many caterpillars, [Laughter] we would switch to tree B which
was along Williston Road coming into Gainesville. I thought, "Well, maybe that tree
had been sprayed by insecticide." That was the most likely explanation of why
everything suddenly started dying. Well, Boucias said that he did not think it was
insecticide. We still do not know why they died, but probably it is because we
changed the leaves from a tree that had grown in the shade of the forest, to one
growing along the side of the road that was in the sun. They are tougher and here is
what I learned: If you do anything to caterpillars, and I have constantly told my
students my whole lifetime, "If you are going to raise caterpillars, you have to keep
them really clean and you have to make sure that they do not run out of food. Do it!"
[Just like] my grandfather, you see.
Well, it turns out that if caterpillars run out of food for just a few hours, the leaves
that they have eaten have bacteria on the surface. The bacteria go in with the
leaves that they chew up and if they do not constantly pass food through their guts
what happens is the bacteria take them over and actually kill them. Getting back to
the whole point of the story, sometimes you do things and get results that,
scientifically, you do not know exactly why. But the point here was that if you do not
take care of them carefully and they get deprived even for a few hours, like coming
in at 11:00 on Sunday morning instead of 8:00, they run out of food, [and] then
bacteria will take them over, and in fact, will kill them. The point I am trying to make
here is that there are people who are really experienced from a pragmatic point of
view of doing things [and] there is usually a solid, scientific explanation behind it.
And so I kind of "learned that instinct," if you can learn an instinct. My colleague [H.]
Jane Brockmann [Professor of Zoology] would give me holy hell if she heard me say
that. But anyway, [Laughter] those precepts of the way you go about doing things
are scientific and ultimately they really pay off.
K: What was there left to learn in college? You said that you went to Princeton and
majored in biology?
B: Yes. I had gone to Chatham High School during my formative years and I took a
biology course there which was just an ordinary, not very interesting, biology course.
And that was it. Then I went on and took chemistry and physics and math and what
have you to prepare for college. So I went to college. Meanwhile, I had been
raising butterflies all over the place. In fact, I got kicked out of school for it once.
Did I say any of that on my curriculum vitae?
K: No. [Laughter]
B: It was a wonderful spring day and there is this wonderful green moth called Feralia
iocosa which is a really interesting moth because its larvae feed on a conifer. They
feed on hemlock trees, and they are among the very first moths to hatch in the
spring. They pupate in the ground and then they crawl up the tree trunks [during]
the first week of spring. They do not occur in Florida, they are northeastern.
Anyway, they are really beautiful moths because the front wings are green and
resemble lichens on the bark. When I first saw one of these things pinned out in
rows of a butterfly collection I thought I just had to have one of those. So, spring
called, and Jane [my wife-to-be] and I took off that afternoon. I was seventeen
years old and I had just gotten my driver's license and everything was just right in
order. Anyway, at Chatham High School, there was this principal [who] was an old
fart as I look back at him, [who] decided the only way to handle this kind of
revolutionary behavior was to expel us for a day. Expulsion for a day was that we
had to sit in one classroom all day long and be humiliated by the students passing
through that classroom. So I never forgot that. We did get the moths and they
proved far more interesting than what I would have learned in that old high school.
K: Talk about encouraging intellectual curiosity.
B: [Laughter] Well, I suppose I could see their point. We were renegades at that point.
But anyway, so I went on to college. If I take a step back, since I am supposed to
be giving some insight into what drives me, my family was Methodist. My mother's
side is [Methodist]. It was a matriarchy that dominated the male side of the family.
Most of my development came through that strong, female side. We had a very
strong family in New Jersey. My grandfather was a very strong person in his own
right. Religious proselytizers used to come out all the time and try to convince these
people to join this, that, and the other Protestant sect, the latest one that broke off.
There is this old family story where the preacher came to my grandfather's house in
New Jersey, and my grandfather gave him dinner and something came up. I never
did get the story quite straight, but the upshot of it was the preacher said that my
grandfather was going to hell. My grandfather came back with some retort that
decided him against religion for all time into the future ad infinitum and none of his
lineage would have anything to do with it. My grandmother, [who was] a good
Methodist, sort of listened quietly and did her own thing and had three daughters,
one of whom was my mother, another one of whom was my aunt, Ruth Churchill,
who was a wonderful lady, an amazing influence on everybody's life around her.
She continued the matriarchy and still does at the age of ninety-five. I will say this
now just so it is in the record. As a life-long Republican, Ruth Churchill, when
Clinton first ran (hopefully it will be the first of at least two times), switched her vote
from having voted Republican her entire life until she was ninety-six years old and
she decided she had enough of them. I guess you can see where I stand by
listening to it.
But anyway, this daughter, Aunt Ruth, had enough church training that she felt
obligated that church was going to be an important part of her family. She took the
route out and joined the Unitarian Church and so we were shepherded off to Sunday
school at the Unitarian Church. My mother could not care less about religion, but
she went along with Auntie Ruth.
All I remember about this whole experience was the very first two weeks when my
Aunt Ruth--this must have been when I was about seven years old--started telling
about the origin of the universe. She had all these pictures about how planets were
formed and all this. That was all I heard and it is all I remember about my entire
Sunday school experience was they got really interested in astronomy at that point.
[Laughter] When I went to Princeton I took two courses in astronomy which were an
absolute lark. I am not a mathematician. I probably would have gone much more in
that direction had I been better at math. Biology really appealed to me, obviously
because of my experience.
When I went to Princeton, there was a man there who had been born in Wales, in
England, and his name was Colin S. Pittendrigh. Pittendrigh, in turn, had been a
student at Columbia University, a post-doc under Theodosius Dobzhansky
[American geneticist]. Dobzhansky was one of the great twentieth century
evolutionary biologists. Pittendrigh was a very articulate man and he really loved
talking about evolution and natural selection. My mother and sisters, and her sister
Auntie Ruth, went to Smith College, so they were pretty well educated, but as far I
can gather, I do not think Smith College really taught that much about evolution and
natural selection. It would be kind of interesting to go back and look at why that is.
Anyway, I was obviously very interested in adaptation, the design of nature. The
design of nature had been used in the nineteenth century by the parson naturalists
of England who would put up all these wonderful examples of natural history to
prove how wonderful God is, and everybody listened to them in England and the
result of that was they all got interested in natural history. Then came along Darwin
in 1859 with his theory of natural selection which provided an alternative explanation
for how these adaptations came into being. I went to Princeton with a good sense of
biology and variability and I never had really asked the question about how did this
design come about. In other words, even though I had it in high school, this pathetic
high school, I really did not have a clue what natural selection was. When I went
into Pittendrigh's class and heard him start lecturing about evolution and natural
selection, everything kind of fell into place for me. It was a really interesting
experience. And from that point on, I really have been a student of evolutionary
biology, trying to understand how things evolve and more of the ecological context of
how evolution occurred, trying to measure the forces that are involved in bringing
about adaptive changes or maintaining certain adaptive states through time rather
than looking at it paleontologically. I was also very interested in [paleontology] when
I took a course on [it in] graduate school at Yale four years later. Anyway, Princeton
was very interesting and Pittendrigh was one of the teaching giants at Princeton
University and was awarded year after year [as] best teacher.
K: You have mentioned Pittendrigh, and now you have gone on to Yale University for
the Ph.D. Was that immediately after taking the bachelor's degree?
B: Yes. Because of the Depression and so forth, we were pretty low on funds and our
family, my brother and myself and my cousins, really did not have the opportunity to
interact with the art museums to any degree. We were not really interested in it. At
Princeton, I became very interested in the history of fine arts. I found it interesting in
terms of the historical overlap of the development of ideas, techniques and art
history with the development of science, and the evolution of evolutionary thought
and natural selection. So I became very interested in that. Evolutionary biology
dominated my interest, and my interest in the lepidoptera. My senior year I spent
raising, [Laughter] much to my roommate's chagrin (back in those days we had one
person to a room, so it is not quite so bad as it is now where they shove three or
four people into the same room), caterpillars. I was really primed.
A new society had been founded in 1947, the Lepidopterist Society. A young, very
talented man by the name of Charles Remington, who had gone through Harvard in
record time as a Ph.D. student, ended up as professor of entomology at Yale
University. Charles and a friend of his formed what was called the Lepidopterist
Society. They developed a journal. Charles was very active in the Lepidopterist
Society at that time. I went to a meeting in New York City, and I think that is where I
first met Charles Remington. Charles was very eager to get new graduate students.
And Jane, my wife-to-be, was interested in lepidoptera as well. We went to this
meeting, met Charles, and eventually went to Yale and became his students.
At that time, there were three areas of interest that fascinated me, partly left over
from Princeton and partly based on my own interests. One was the evolution of
mimicry in butterflies, which really is a paradigm of how evolution works. It still is a
really fine paradigm for asking questions right on the forefront of current evolutionary
thought today. I thought I would like to do that as a thesis, but Jane decided she
wanted to, so she did that for her thesis. I, meanwhile, got very interested in the
question of the origin of species. How do new species originate? Charles had this
group of butterflies that he had been looking at, the Papilio glaucus group (Tiger
swallowtail butterflies). It has been a long time since I studied these things. There
is a group of closely related species that occur out in the Rocky Mountains. In the
East, there is only one species, so you have this interesting ecological contrast of
the single species in the East, and in the West you have got these three species that
are living together and intensely interacting with each other. It was a really
interesting evolutionary thesis.
Jane and I would head off to Colorado and then Utah during the summers and then
in the spring we would go down to the Archbold Biological Station and work on her
thesis. She was my assistant and I was hers. We got very much involved with each
other's research. I was very interested in the mimicry experiments and helped
design them and helped execute them. At that time, the mimicry system that Jane
became famous for studying was the Monarch butterfly and the Viceroy. The
Viceroy is a classical mimic of the Monarch, and it was thought to be a palatable so-
called Batesian mimic, named after Walter Henry [Henry Walter] Bates [English
naturalist, 1825-1892], who had gone up the Amazon in 1850. [His trip] overlapped
with Darwin's developing ideas of natural selection.
Anyway, so we did the first experimental tests of so-called Batesian mimicry. The
idea there is a palatable species of butterfly which is persecuted by birds and other
predators has its own colorations but through time changes that coloration to mimic
another butterfly which for some reason is toxic or poisonous and that is called the
model. So the mimic comes to look like the model butterfly in evolutionary time.
And the advantage is that a palatable butterfly masquerades under the color pattern
of the unpalatable one and so gains a mimetic advantage. That had never been
proven, never been tested experimentally. There are all kinds of correlative
arguments and biogeographic arguments and indirect, strongly reasoned arguments
that mimicry has to be. Just looking at totally distinct groups of butterflies with the
occasional one that looks like some other group, there is a very interesting
intellectual pattern if you look at it.
So we tested this and it turns out that Monarchs were just completely rejected by
these birds, whereas other swallow-tail butterflies--in fact related to the ones that I
had been studying for my thesis--were palatable. We were lucky because there
were lots of palatable butterflies at the Archbold Station and nearby, and here is
where I really got interested in the Monarch butterfly. I went out and collected and
raised the larvae which were then used in the feeding experiments, and so I did a lot
of the field work of collecting the butterflies and getting them ready for Jane's
In the course of this, I was interested in species interaction. There is a related
butterfly in the same family which is called the Queen butterfly, so you have the
Monarch and the Queen. One is Danaus plexippus, and the other is Danaus
gilippus. These are really interesting butterflies, from a lot of points of view. But
now, both of these butterflies feed on milkweed plants, genus Aclepias, which is a
derivative from the Greek god Asklepius, who was the Greek god of medicine. That
implies that milkweeds are medicinal plants. Milkweeds are called milkweeds
because they have latex in their leaves and the latex spews out if you damage a
leaf, and if you taste it it has an incredibly bitter taste. In fact, the plants are avoided
by cattle and they are inedible. The only way you can eat milkweed plants is to boil
them and throw the water away two or three times, because what you are doing is
extracting the poisons in the water and getting rid of them. This is why cooking is a
big selective advantage for humans. There are not that many vegetables that you
can eat. There are not that many plants that you can eat. Most of them, in fact,
contain horrendous numbers of different chemicals that have been designed in the
course of evolution to avert caterpillars, to avert browsing mammals and so forth
from eating them.
Anyway, the hypothesis goes back to the nineteenth century where Bates' original
observations in the Amazon led him to the hypothesis that brightly colored butterflies
that are flying around and very conspicuous are behaviorally different from other
groups of butterflies which are much less conspicuous in their color patterns and
their behavior. But every once in a while within the inconspicuous group, you find a
butterfly that looks like one of these wildly colored ones over here. In fact, they
originally could not tell them apart [because] they were so similar until you started
looking at the details of their morphology. Well, then it turned out that this kind of
relationship of mimicry, the model butterfly and the mimic models are common,
brightly colored, often gregarious, [and] they expose themselves. They are very
easy to catch and so forth. They must be unpalatable, Bates reasoned. Then as
more knowledge was gained about what these butterflies lay their eggs on and what
their caterpillars eat, it turns out that the different taxonomic groups of model
butterflies are tightly associated with specific groups of poisonous plants. Monarchs
and the related species in that family, the Danaidae, feed on milkweed. A group of
swallowtail butterflies, which are unpalatable, are called the Troidine swallowtails,
and they feed on a group of plants in the Aristolochiacae. Aristolochiacae are called
birthworts because they were used by the nuns in the Middle Ages to induce
abortion. There is a nunnery in England that has a lot of these plants. That is
another story. The hypocrisy goes back through the history of humanity. So much
for the religious right. [Laughter]
So it turns out that the different groups of butterflies that are thought to be
unpalatable are associated with toxic plants. So you have a taxonomic group of
butterflies associated with a taxonomic group of plants. It is a correlation. And so a
German by the name of Hess proposed that the butterflies that are feeding on these
plants are deriving toxic properties from the plants.
K: They are manufacturing it themselves?
B: Right. The plants are manufacturing the poisons, and rather than the butterflies in
some way generating their own toxins, they are sequestering them from the plants,
and that would explain why these different groups of models were associated with
these poisonous plants. They were getting the poisons out of the plants and hence
were protected against being attacked by birds.
K: That means that butterflies would have to have evolved a defense against those
B: It would, exactly. So then the palatable groups of butterflies which are feeding on
plants that are not toxic--or if they are toxic, they are in some way destroying the
toxins and not storing them in their bodies--come to look like the poisonous ones
and take advantage of the fact that the birds have learned to leave alone the
So that is how mimicry is thought to have evolved. This was a great theory that had
been spun in the nineteenth century and had not really been tested--whether or not
mimicry is effective in the laboratory. It is very hard to test it in the wild. We have
done that, but it was a very difficult thing to do. Jane's experiment showed that the
Viceroy was an effective mimic of the Monarch.
Now in the process of my raising Monarchs, I tasted one of these milkweed plants.
It just about knocked me over. It was so foul tasting. I drooled and I almost
vomited. I realized that pharmacologists, when they talk about poisonous plants,
know what they are talking about. So that kind of sparked my interest in the
relationship of toxic butterflies to their food plants.
When I took my first job, I went for a year to the great seat of evolutionary biology--
Oxford University--to study under an incredibly eccentric man whose name was
Edmund B. [Briscoe] Ford. Edmund B. Ford and I and Jane became very good
friends. E. B. Ford was not a misogynist, but women did not exist for him.
Somehow my wife, Jane, and Henry, as everybody called him, struck it off really
well. He really liked her. We all got along very well.
E. B. Ford had written a book on mimicry with Carpenter, George [Geoffrey] D.
[Douglas] Hale Carpenter [Mimicry, London, 1933], who had been at Oxford. Oxford
was a great center of mimicry. Before Carpenter, [there] had been E. B. Poulton,
who was one of the great promoters of Darwinism at the end of the nineteenth
century and on into the twentieth century. Ford was the first one to break from the
tradition of just anecdotalism and of correspondence with the naturalists in the field
that did this, that and the other thing. There was some experimental evidence, but
really weakly designed, sort of off-the-cuff experiments. Mimicry was an enormously
complex and beautiful house of cards which could have been destroyed if it had
been shown that the models were not unpalatable. And Jane's birds, the scrub jays,
would not eat the butterflies, would not eat Monarchs.
It is interesting. These were wild-caught birds from south central Florida [at] the
Archbold Biological Station. [They were] brought into the laboratory, [and] they were
pretty tame because they had been feeding them there, so they were easy to work
with. Until that point, I had never held a bird in my hand. I learned not only how to
deal with birds, but also, to come to respect them tremendously. They are very,
very interesting animals. I think they are far more intelligent than anybody even
knows. I do not know how you measure intelligence in birds, any more than in
students, but there is something there. There is a lot more there that is going to
come out in the next fifty years--if the planet is not totally decimated in the process
and there are no birds left in the next fifty years, which is very likely. (My prediction.)
I got interested in the pharmacology (i.e. investigating the effect of the plant toxins
on the butterflies and the birds). I went to Amherst College (Amherst,
Massachusetts) in 1958 after a year at Oxford. I stayed there for twenty-two years.
Amherst College is a really fantastic institution. It is not a bureaucratic morass, the
way the University of Florida is. The bureaucracy of this place is absolutely
destructive. I am thoroughly fed up with it. I am sure that anybody who is faculty at
this place is just utterly frustrated by this system that is designed on the predication
of "everybody's a thief and a cheat." But anyway, that is another story. Amherst
College was a small, wealthy, liberal arts educational institution and they had the
highest rejection rate of any college in the United States, or any university in the
United States. The biology department at Amherst College had a really long
tradition which was greatly fostered shortly after World War II by grants from the
Rockefeller Foundation. Research in the biology department became a tradition.
Good quality, mostly experimental research. It is very interesting. I did not realize
this until I was doing the history of Monarch butterfly migration a while back. One of
the very early famous butterfly biologists, William J. Holland, who wrote The Butterfly
Book [New York, 1898] and The Moth Book[: A Popular Guide to the Knowledge of
the Moths of North America, New York, 1903], went to Amherst College, and
graduated from Amherst College. Two of the other very famous lepidopterists went
to Williams College [Williamstown, Mass.] which of course is over the hill on the
other side of the Berkshires and is a great rival of Amherst College. The biology
department was so away from its own history that that had been long forgotten when
I got there.
It was an interesting experience for me, because I came in as the field biologist and
natural historian and I developed it at Amherst College in the context of an
experimental biology department. And I liked experimental biology because I think
that it allows you efficiently to answer questions--more so than by just muddling
around in the field and making all these historical correlations that have been done
in much of field biology. The whole thing that mimicry did was lead to an enormous
house of cards which was beautiful, but had to be put together with solid,
experimental research. Well, that is basically what we did when I went to Amherst
starting in 1958. For those twenty-two years, the biology department had what is
called an honors program. Anybody who went to Amherst College and majored in
biology, unless they were really deficient mentally, was able to get into a good
medical school. It was a passport to Harvard Medical School. So we had really
good students at Amherst College. When I first started teaching there, naturally I
was on the outside as the field biologist. But gradually--and it was not even
gradually, it was about two years--I began attracting students who were not
necessarily totally oriented toward medical school. And it turns out that there were a
lot of kids who came to Amherst College who had been brought up in families that
were really into birding. One of my most famous students at Amherst College was
John Alcock, who is now at Arizona State University where he is really quite a
famous biologist in his own right. I recognized John in my natural history class when
he wrote his essays on his exams. Each one of them was a gem. John was a
disciplined, bright kid and he was going to major in English and I seduced him over
into majoring in biology and he went on to great things.
Well, anyway, over the years I had a whole bunch of really good students
serendipitously falling into what could have been a mire of molecular biology,
biochemistry mindsets. I managed to keep my head above water, but at the same
time the students in the honors program--and we would have anywhere from eight to
twelve honor students a year--fed on each other and learned from each other year
after year. Some of them were doing high-level biochemistry stuff, some were right
on the forefront of biology. It was a really good department. My students could
bring into my lab things that I myself would not have been able to do. For example, I
got interested in spectrophotometric measurements of the chemicals in milkweed
plants. One of these students actually happened to be a Mount Holyoke College
[South Hadley, Mass] student, and we had lots of interactions with Mount Holyoke, a
women's college just over the Mount Holyoke range to the south. There are five
colleges in this region now, and when I was there there were four. But the four
represented a very strong, intellectual powerhouse of interaction possibilities.
So this one woman, whose name was Maureen Flannery, was a student of an
organic chemist at Mount Holyoke College by the name of Kenneth Williamson.
One way or another, Ken and I got together with Maureen. We had tried with other
students previous to that, but it did not work out because it was too tough. But
Maureen Flannery was one of these kids who was just incredible. She was almost
totally deaf. She majored and honored in history, chemistry, and biology and
received summa cum laude in all three areas. Anyway, she was also a wonderful
person. Maureen and Ken and I worked out what we called the TNDP assay--
Tetranitrodiphenyl--which, to make a long story short, was a reaction which was
discovered by a nobel prize chemist in Switzerland by the name of Tadeus
Reichstein [Swiss chemist, Nobel Prize in Physiology and Medicine, 1950].
Reichstein was interested in synthesizing cardiac glycosides. These are the
poisonous substances that are in milkweeds that the Monarch butterfly takes up. In
fact, I am getting ahead of myself here because I want to go back and tell you how
we led up to that. I will pause here for a minute and come back to the TNDP
This was about half way through my career at Amherst that we got into the
chemistry of it. So we collected blue jays in the wild and brought them into the lab
and made our own cages up and had a really elaborate and fantastic laboratory for
doing bird feeding experiments. I did very well in chemistry in college, in fact, I loved
it, but I really did not know the chemistry of these toxins that were in the milkweed
plants yet. Anyway, I reasoned that if the hypothesis is correct that monarchs are
getting their poisons out of their milkweed plants, then one way to test this would be
to try to raise Monarch butterflies on a plant which is innocuous, that does not
contain poisons. This is all a bit naive, but we settled on lettuce, thinking that that is
fairly innocuous. [laughter] Although, if one had kept up on Beatrix Potter [English
author, 1866-1943] and Peter Rabbit, one would realize that lettuce itself is toxic
because if you eat too much of it, it will put you to sleep, which is what put Peter
Rabbit to sleep in Mr. Macgregor's Garden. It has substances in there that are sort
of sleep-inducing drugs.
B: They are in low concentration. Well, it turns out that the monarch caterpillars would
not touch lettuce. So what we did was--and this is a typical selection experiment--
we took about three thousand eggs of Monarch butterflies and hatched them out
and put them on lettuce seedlings and they all died. So we did the same thing and
put them on cabbage seedlings, Burpy's Fine Garden Seeds, and we figured they
must have been selected for low bitterness and what have you. I mean there was
all sorts of qualitative thinking. Well, after putting several thousand eggs on these
plants, two or three caterpillars, lo and behold, started eating the cabbage, which
goes on to show you how specific these butterflies are for what foods they will eat,
milkweeds, and you can hardly get them to eat anything else. We did this basically
by very strong selection. We raised these butterflies through to the third instar and
then they stopped eating. And having my grandfather's training, I put them back on
milkweed, took them through, finished them off on milkweed, bred them and then
took the eggs and put the offspring of the next generation on the cabbage, and now
they would eat cabbage. We had selected a strain which could eat cabbage. So we
reared through enough butterflies, we set up our bird experiments and the strategy
was to feed the birds cabbage-reared Monarchs, which we did. Lo and behold, the
blue jays ate them, one after the other.
Now when Jane had done her experiments at the Archbold Station, we had used
these scrub jays, which are jays, but they are a different species. These are wild
birds which had probably had experience with these butterflies. They pecked them,
but not one of those scrub jays ever ate a Monarch. They pecked them and rejected
them after they pecked them. I think one bird, maybe in the total of 400 trials with
four or five birds, ever ate a Monarch, if that.
K: An optimist?
B: An optimist, or a stupid bird, or something. So anyway, now we were in a really
interesting position because we had trained our birds that Monarchs are okay, and
they were eating them. Now the test. Now we offered the birds that we had induced
to eat cabbage-reared Monarchs, Monarchs that have been fed on their natural
milkweed plant. And the birds ate the butterflies, and within twelve minutes began to
get violently sick and started vomiting all over the place and threw up everything that
they had eaten. We repeated that experiment several times with larvae, with pupae,
and with adults and lo and behold, all stages of the Monarch butterfly induced
emesis (vomiting), if they had been reared on milkweed, but not on cabbage.
This very strongly supported the old hypothesis of Haas and Bates and the rest of
them that the butterflies are somehow, as caterpillars, deriving the toxins from the
food plants. Well, we elaborated on those experiments. I went back to Oxford in
1963. I was introduced to a man by the name of John Parsons. John Parsons was
a pharmacologist who had some horrendous disease and he was very brilliant man.
He was fast upon trying to find a cure to his own disease before it killed him. John
is a wonderful person and he is still alive, but I have lost track of him, which is
But anyway, John and I really hit it off, and John was interested in digitalis-derived
poisons, which are the poisons that are very similar to those that are in milkweed
plants. And John Parsons did an experiment in which he fed Monarch butterflies
that I had raised in Trinidad. We did a lot of field research in Trinidad and the West
Indies, which is really South America, an island off the mainland. John Parsons
made extracts of Monarchs and so forth and fed them to starlings and showed that
the starlings vomited. Then he tested these compounds that he had extracted from
Monarchs with commercially available, pure digitoxin-like compounds, and results
were similar. His pharmacological experiments then meshed in with ours.
It was quite clear the next step was to develop a chemical assay, and I am going to
tell you something here now that I am going to write up. I have not written it up yet.
There developed a competition between John Parsons and Miriam Rothschild.
Miriam Rothschild is very famous, and one of the richest women in the world. Her
family was more responsible for the development of nuclear power than any other
family in the world because the Rothschild family had been the big bankers of
England. One way in which Israel saw itself as energy independent was by the
promotion of nuclear power. Well, anyway, it is a long, long story and I do not have
to go into it. Miriam Rothschild and I became very competitive, and I invited John
Parsons to come to my laboratory at Amherst College and got an NSF [National
Science Foundation] grant to bring him over. I probably cannot prove this, but John
Parsons was told by his boss in England--he worked at Mill Hill in London--that if he
came over for a year to my laboratory it would be the end of his job. So John
Parsons did not come. And we would have worked on the chemistry together and
would have done great things. [I was] frustrated by that.
Now I get back to Maureen Flannery and my friend Kenneth Williamson of Mount
Holyoke College. I decided it was time to get into the chemistry of these things. So
we had these resources at Amherst, the students and colleagues, who were
interested and we went to Tadeus Reichstein. I got myself in real trouble with
Miriam Rothschild. Miriam Rothschild is what I consider a master puppeteer of
scientists. She is very interested in evolutionary biology and ecology, and
particularly insect natural history and chemical defense, which she has done as a
profession. She also is a professional taxonomist of fleas. She is involved in all
kinds of activities. She is an amazing woman, a great heroine. What I am talking
about now is really stuff that will eventually be published. It probably should have
been a long time ago.
I had sent these butterflies that I raised on different milkweed plants to Oxford, to
Miriam Rothschild and either directly to her or via her to Tadeus Reichstein, who
was the Nobel prize chemist on analyzing cardiac glycosides. I was to be coauthor
of that paper. Miriam Rothschild and I disagreed on certain aspects of the
ecological interpretations of that paper. When I got the proof of the paper, where I
was a coauthor with Reichstein, Rothschild, Brower and a couple of other people
[who] were involved in this study, I withdrew my authorship, and in the process
massively insulted Miriam. She has been on my case ever since [the] late 1970s.
Was that a mistake? In my opinion, it was not because I kept my soul intact. No
matter what it cost me, I will go down to the grave with my soul intact. It was
probably a petty disagreement and should not have been worried about, but there is
an enormous amount of politics which in many respects is analogous to the double
helix bullshit that went on, if you have ever read that.
K: Oh, yes.
B: It was a great story.
K: Not necessarily a high point for science, but a great story.
B: Well, it is the way things seem to happen in this world because humans are very
political animals. Basically, having been screwed in terms of dealing with John
Parsons, what I did was set out to develop a new technique by involving three or
four sets of honor students who tried to work this out, [but] we did not get very far
with it. We did not exploit the students because we did it over the summer and if it
worked, then they would have that as a thesis. Clearly, two sets of students simply
were not up to it. Maureen Flannery, this wonderful, brilliant young woman, came
along and she and I and Ken worked together. Chemists and pharmacologists are
interested in identifying compounds that are in plants that are useful for the
development of drugs for the treatment of human and animal disease. Basically, the
process is that you go out and you get fifty pounds of leaves and you dry them and
you bring them back to the laboratory and you extract them. I was interested in--and
this is one of the places where Miriam and I parted ways intellectually-- population
biology. I am a population biologist. Populations are made up of individual
butterflies. Yes, you can identify what is in Monarch butterflies by grinding up ten
thousand of them and analyzing it. What I was interested in was if Monarchs really
do contain these poisons, how much poison is there in one butterfly? And it was
known at that point, and we now know that there are 108 species of milkweed plants
in the United States, Canada and Mexico. It is a very interesting evolutionary story.
They are chemically diverse. And it turns out that Monarchs that have fed on some
milkweed plants are palatable. We know this now. We have determined that
experimentally by feeding the butterflies raised on various plants to birds. It turned
out that some of the ones that we thought were going to be most poisonous were
not toxic at all. Well, it turned out they were not toxic because the plants they fed
on, although they were milkweeds, lacked these poisons.
We did develop this tetranitrodiphenyl assay--TNDP assay--and the object was to
have an assay which was sensitive enough so that we could take a butterfly, dry it
and grind it, and divide it into two parts. One part would then be extracted to get the
chemicals out to measure. The other part would be saved to feed the bird to see
what the biological response to the dry butterfly powder was. So on the one hand,
we could get a bioassay, and on the other, we could use the other half of the
butterfly to get a chemical assay. We did it. Maureen and Williamson and I did it,
which then established, basically, the population ecology of naturally occurring plant
chemicals as sequestered by butterflies. In 1969, I wrote an article which I called
"Ecological Chemistry" which was published in the Scientific American. Actually,
that article was published before we developed this chemistry. And that set the
stage for Maureen's and our collaboration. Once we developed this assay, we
started doing the analysis of populations. This could go on forever, so I am going to
abbreviate things here. We can now do population biology, how many Monarchs
are poisonous, how poisonous are they and are they poisonous to different degrees
if they fed on different milkweeds, not only poisonous versus non-poisonous. But
what we showed was that, in fact, there is a spectrum of palatability. There is
enough poison in one butterfly to make a dozen blue jays vomit. Or they are
completely palatable. So you have an emetic spectrum or a toxic spectrum and we
call this the palatability spectrum. One of my first awards in biology was from
Esquire magazine for defining the blue jay emetic unit. It is interesting that we have
defined it in a way that caught the public eye because most of my scientific
colleagues still do not have a clue as to what the hell an emetic unit is. [Laughter] It
But anyway, so one thing led to another and now we have come to another great
conflict. Having gone into chemistry, it is pretty obvious to me that I am not a
chemist. And I am not going to go zooming into that field, getting very far. And
anyway, my inclinations are to do field biology and then bring it into the lab and do
tests on them. I started getting really interested in variability in different populations
of Monarchs. What we predict here from our laboratory experiments is that if the
butterfly populations are heterogeneous, some poisonous and some not, then there
is a possibility that birds could figure that out and break through and eat Monarchs
which are not toxic. If you go back through the literature, going back into the last
century, you have got one school that maintains birds do eat butterflies, another
school that maintains birds never eat butterflies, because ornithologists do not often
see them eating butterflies for one reason or another. Actually, I have not seen very
many birds eat butterflies in the wild. I think there is a perfectly simple explanation
for that. When they are flying around, they are just too hard to catch. And a bird is
probably going to end up spending more energy trying to get a butterfly morsel than
it is going to get out of that morsel.
We had predicted that there was a palatability spectrum in the wild. Where to test
this? I had gone in 1973-1974 on a sabbatical to the University of California at
Davis to work with another set of biochemist colleagues. Not only did we develop
the method to measure how much chemical was in the butterfly, but I wanted to see
if we could relate the poisons that are in the butterflies to the particular milkweed
plants that they had eaten. California is an interesting place to do this, because it
turns out that California has some milkweeds that are very toxic and others that are
not. Working with my colleague, James Seiber, who was at the University of
California at Davis, and a woman who was working with him by the name of Carolyn
Nelson, togetherwe developed the ideas of visualizing the poisons. This is basically
thin layer chromotography, which is a straightforward chemical procedure where you
take an alcoholic extract of the butterfly and you actually measure, using the TNDP
test on a small aliquot of that sample. You measure how much poison there is in the
butterfly, and knowing that, then you take the residue, which is in the alcoholic
extract, and you clean it up using different solvents. It is an elaborate and very
scientific procedure which basically gets rid of a lot of gunk. Then you take a fancy,
ten thousand dollar device and you spot the extract on a plate, which is basically an
elaborate, fancy piece of blotting paper. So you take that extract of that butterfly
and you spot it on the bottom of this thin layer chromotography plate. It is slow [and]
laborious, and you have to be very careful not to mix things up. Again, orderliness in
science is so, so important. I just cannot emphasize how important it is to have an
orderly approach doing it. If people do not want to be orderly, they should not even
think of going into science, because it will be sheer luck if they get anywhere.
Because otherwise there is going to be so much noise, they are never going to get
anywhere. Seiber and Nelson were really precise.
Anyway, so you spot this material, and you can spot fourteen extracts of about
fourteen butterflies on one plate. Or butterflies and extracts of the plants they ate.
Well, to make a long story short, it turns out that different species of milkweeds have
very different arrays of cardiac glycosides in them. It is not just one chemical.
[There are] probably four or five hundred separate chemicals represented by all
these different milkweeds. What happens is you take this plate that you have
spotted and you put it in a solvent, the solvent gradually creeps up the plate, and it
carries the poisons with it. Depending upon the chemistry of the poisons, how many
hydroxyl side groups each has, they will go up and so the different chemical
mixtures separate out so that you have a series of spots in the channel for each of
these butterflies or plants. Well, to make a long story short, each butterfly,
depending upon which milkweed it has, has a fingerprint, a cardenolide or cardiac
glycoside fingerprint. And you can match that fingerprint to the plant that it ate.
Some of the butterflies will not have anything, and all you know is that they ate
plants that lacked these poisons. Others that have eaten Asclepios humistrata will
have a pattern that is totally different. If they have eaten Asclepios syriaca or
Asclepios viridis, or whatever plant it is that you are interested in.
Well, the Monarch has a whole other aspect to its biology, which I also got
interested in when I was helping Jane with her thesis work in Florida. And that is
that it is a migratory butterfly. The butterfly, in the fall, migrates by the millions
southwestward. And a biologist by the name of Fred Urquhart at the University of
Toronto in Canada had developed a tagging program where they actually glued little
blue labels on butterfly wings, released the butterflies and then other people caught
them further south and southwest. Urquhart involved over three thousand people--
mostly school teachers and so forth--in this tagging program, starting in about 1942.
This led Urquhart to produce a series of publications on where Monarchs are
migrating. This was a real mystery originally developed by C. B. [Carrington Bonsor]
Williams, who was another one of these wonderful, old English entomologists.
Williams had written a book in 1930 on the migration of butterflies [British Immigrant
Butterflies and Moths, London, 1930]. In that, he spent a great deal of time going
back into the nineteenth-century literature on Monarch butterfly migration. The
Monarch butterfly, in my opinion, is the most interesting insect in the world. The
different aspects of its biology are incredible, and I refer to the butterfly as the
magical bottle of wine. You have a beautiful glass on the table and you fill it up with
chateau, bordeaux and shall I say .. Rothschild? [Laughter] Then you drink it and
you pass some over to your friend, and you put the bottle down, and you pick it up
and it is full again. And studying the monarch is like that. I mean, it is a never-
ending, fascinating creature that has so many dimensions to its biology. But I
suspect that any species that is really carefully studied also will have those
dimensions. Which is why I am really concerned about man and woman's
decimation of the landscape of the earth. Maybe we will talk about that later, some
other time. It is going to hell in a handbasket, without any doubt because of human
So it became clear that Urquhart would tag thousands of butterflies every year, and
he would get back two or three tags. Milkweed species A grows in the Northeastern
United States, milkweed species B grows in the Midwest. Butterflies which eat
milkweed species A are going to have a different cardenolide, fingerprint pattern,
than B that is grown in the Midwest. And so if there is a common point where they
are assembling, and if you collect the butterflies and look at them, then you should
be able to use the fingerprint, collect the butterflies, analyze them and tell where
they have come from, because the plants have discrete biogeographic distribution.
Basically, Urquhart's and my interests converged.
I had hypothesized when I was doing my research in Florida that Monarchs were
wintering and breeding in Florida and in fact, they were not really spending the
winter--like they do in California--in clusters, but they were spending the winter
active and breeding on milkweed. So where do they go? Nobody knew where they
went. It became clear that the tagging recoveries that Urquhart was getting were
concentrating in places like Texas and that the pattern indicated they were flying
southwesterly toward Texas and then disappearing. Two butterflies were recaptured
in Mexico, one in a city north of Mexico City called San Luis Potosi, which is actually
a city out in the middle of the desert, and another was captured near Mexico City.
Urquhart, in 1974, published an article simultaneously in several Mexican
newspapers and magazines in English, and some in Spanish. I think I had
convinced him that Monarchs were not overwintering. He would never admit that.
And I am not sure I did [convince him], but there were other indications that
Monarchs were breeding in Arizona. Somebody else had discovered that. And so
in the Southern range, they were breeding. He had gone to look for Monarchs
overwintering in Florida in the late 1950s. He found only one overwintering site and
no others were ever found. So the question is, where are all these millions of
butterflies that are migrating south spending the winter?
Well, he ends this advertisement, which was a call for people to join his tagging
team, to help try to figure out if the Monarchs are breeding and which generation is
coming back to North America the following spring. A man by the name of Kenneth
Brugger, who was an American salesman living in Mexico, read this advertisement
and wrote to Urquhart, and they struck up a correspondence. Ken Brugger fell in
love with a Mexican woman, whose name was Kathy Aguada. Ken, in November of
1973, was driving through the mountains west of Mexico City, about eighty miles
west, and suddenly, there was a hail storm up in the mountains. And this must have
been in the fall; it could have been in the spring, it is not definite when it was. I was
unable to pin down the exact time. But anyway, out of the sky in the hail storm were
thousands of Monarch butterflies being pelted down. So Ken wrote to Urquhart, and
Urquhart encouraged him to look on.
To make a long story short, [on] January 2, 1975, Ken and Kathy (Kathy was
Mexican so she spoke Spanish) started asking all over the place [about Monarch
butterflies] and started exploring this area. Ken and Kathy discovered an
overwintering colony of Monarch butterflies on a mountain called Sierra Pelon.
Urquhart has never published exactly where. A lot of detective work had to go into
figuring this out. It was on the Sierra Pelon, which is west of Mexico City. And then
we reasoned, by reading various things that had been written in recapture records,
and so forth, that Brugger found a second site--which was called the Sierra Chinqua,
not too far, maybe thirty miles away from the first site--another big colony.
This discovery was made in 1975. Urquhart's research had been supported by
National Geographic magazine. Rumor had it that the discovery had been made. I
wrote to Urquhart and asked him if he would share the location of the site with us.
And he wrote me back that they were keeping it a secret until it was published. I
wrote to National Geographic magazine and asked them, and they were not going to
divulge anything until they published it. To make a long story short, and this is a
really intense story, in August of 1976, National Geographic published Urquhart's
discovery of Monarchs overwintering in Mexico. Then in October of that year, the
journal of the Lepidopterist's Society published a more detailed article by Urquhart.
Meanwhile, I had been talking to Urquhart by phone and writing him and asking him
if he would let us go to the site [and] tell us where it was, because I realized now--
having found that there is a palatability spectrum in butterflies--[that] here was
obviously an incredible natural laboratory. The article that was published in National
Geographic showed millions and millions of Monarch butterflies in a very small area.
And so here [was] this aggregation behavior, going back into the nineteenth-century
observations of Bates that aggregating butterflies are chemically protected and so
forth. Here was the most fantastic natural laboratory experiment in the wild that one
could dream of. [It was an opportunity] to go down there and see if the birds were
eating the butterflies and finally test this whole thing and also to fingerprint the
butterflies and see if the birds were eating the ones that were non-toxic. I got this
very supercilious letter from Urquhart saying that Monarchs migrate through Florida.
I had moved to Florida now, and this was right between my move from Amherst to
the University of Florida, which took place in 1980. I had opened the
correspondence with him at Amherst, then I moved to Florida. He had said that I
could collect all these butterflies migrating along the Gulf coast of Florida, which of
course peaked my imagination even further. I was still at Amherst at this time. It
was the fall of 1976 [when] this intense correspondence was going on.
There was a post-doctoral student at the University of Massachusetts in entomology
and his name was William H. Calvert. And Calvert was really interested in butterfly
biology, and he was also a real easy-going sort of hippie. He really did not give a
damn whether he had a job or not, as long as he was managing to survive. But he
really liked doing biology. He came from Austin, Texas, and he was a native Texan,
and he had spent quite a bit of time with friends exploring North[ern] Mexico. He
had a friend who was fluent in Spanish. So Calvert and I talked and thought it would
really be fun if we could find these butterflies. I very carefully read the two articles
that Erkhardt had published.
[National Geographic's] policy--as having been stated to me in a letter in response to
my writing them--said that they were not going to make the sites available to the
public until after the article was published. After it was published in August 1976
Urquhart Erkhart still would not. They had a policy meeting. And I just found out
last January 1994 that National Geographic had had a policy meeting where they
decided not to divulge the location of the sites to the public, for fear that they would
be overrun and destroyed. That was their policy, and so in the National Geographic
article and in the Lepidopterist's Society article there were two little descriptions that
Urquhart gave. One was that they were huffing and puffing at above ten thousand
feet altitude, and he was older and developing a heart problem, if he [did not already
have one]. This was pretty serious business. That comes out in the article very
clearly. Secondly, in one of the articles, it said it was in northern Michoacan. Now
the National Geographic published the location as the Sierra Madre. Now that
sounds nice and it sounds like a Humphrey Bogart movie or some damn thing. But
the Sierra Madre is literally scores, if not hundreds of miles away from where the
butterflies were actually discovered.
It is not completely clear to anyone at this point in time until somebody directs a
question--which I intend to do in the not-too-distant future because I have just
written the history of this thing--whether that deception was purposeful or whether it
is because Urquhart, who is definitely ignorant of [the] geography of Mexico (you
can read into that whatever you want to, and I will not elaborate on that) settled on
that name serendipitously.
Well, if you look at a map of Mexico, [or] if you look at the National Geographic map
of Mexico, which shows the topography, there are three major mountain ranges: the
Sierra Madre Occidental and the Sierra Madre Oriental, the eastern and western
Sierras. They go down the eastern and western sides. One is an extension of the
Rockies, the eastern one. The western one is more or less an extension of the
western mountains of the United States. They peter out approximately at the
latitude of Mexico City. There is another range that goes across Mexico, and it
actually goes all the way across Mexico and intersects the Sierra Madre Oriental
and Occidental. That is where the big volcanoes of Mexico are. It is called the
Sierra Volcanica or the Sierra Transvolcanica. It is a transvolcanic mountain range
and it is very recent. It only goes back probably to the Miocene, which has
interesting implications for the evolution of Monarch migration. But that is another
Having completely deceived the world as to where the butterflies were, but having
also given these two clues of the altitudes in the northern part of Michoacan, Calvert
and I got another old topographic map of Mexico and we simply laid out all the
altitudes that were above ten thousand feet in Michoacan. It turns out that there
were only a few mountains that were above that altitude, and they were all in this
part of northern Michoacan, or seemed to be corresponding to what had been stated
in these articles. I funded Calvert's expedition, and he left Austin, Texas. He went
home for Christmas, and he and his buddies got together on the 26th of December.
We were having a rousing New Year's Eve party at Amherst that night, December
31, 1976. The telephone rings at around 11:00 at night, and there was a terrible
connection. He [Calvert] found the butterflies. [He found them in] four days.
Actually, they found them on the 30th of December. It took them literally four days,
three to four days, to find the butterfly site. Bill had lost his notebook on the trail,
which had a lot of notes about his first observations on the butterflies. Kenneth
Brugger, meanwhile, had discovered this site, which turned out to be the Sierra
Chinqua site and picked up Bill's notebook and gave it to Urquhart. And we never
saw that notebook again.
B: Really. I will not dwell on it.
B: So Bill came back to the United States. He had to come back for school. I
organized a second expedition with Bill, and I think it was on the 26th of January,
1977. Bill and I and my assistant, who was working for me in the lab at that time,
Lee Hedrick, drove in to the butterfly site. We parked our cars. It was very difficult
to get into this place because it was back through logging roads and the most
amazing, beautiful forest you can imagine. [It was] very high; you have to go over
11,000 feet and then down again to get into this place. It was one hell of a trip. We
almost did not make it. We had two Volkswagen vans. I hired two in case one of
them died. Actually, I had enormous respect for Volkswagen vans after that trip.
Anyway, we made it. We walked down. It was about 3:00 in the afternoon, and who
should be sitting in the forest tagging butterflies, but Professor Urquhart. [Laughter]
That generated one hell of a storm, which hit all the newspapers in the world and
painted me to be black-souled. Urquhart accused us of following him in and all that
stuff, which was untrue. It is a long story.
Over the years after that, Bill and I mounted a whole series of expeditions to Mexico.
Bill did all the heavy field work because I was teaching. He--being footloose and
fancy free--could go down there for months on end. He was really good at camping
in Mexico, which is not easy, to say the least. He discovered about twelve different
overwintering sites in Mexico. The thing is, tragically, they are all within a very
small--thirty by seventy mile, thirty by one hundred and twenty kilometer, I forget--
area where there are half a dozen mountain ranges where these Monarch colonies
occur. Here in these colonies, we have estimated that there are ten million
butterflies per hectare and some of these colonies are up to ten, fifteen acres in
extent. Five hectares. I think that would be 12.5 acres of butterflies. That is ten
million times twelve acres: [whistles] 125 million butterflies. They densely cluster on
All my predictions about this being a fantastic area to do field experiments turned out
to be absolutely true. Over the years one of my Amherst College students, Linda
Fink, who became a senior honor student under me, [went with me] to Mexico. She
did her thesis there in 1979-1980. We combined the field observations and
binocular observation of birds, which we watched catching the butterflies in the
colony. It turns out the birds are eating up to a million butterflies per year, per
colony. It turns out [that] Linda and I were able to show that the butterflies that the
birds are eating--at least one of the two species are eating--they are doing it by
picking out those Monarchs which had fed on non-toxic milkweeds. The whole
thing, all these predictions that came through over the years, we were able to
confirm in spite of old uncle Fred.
So the story goes around and around and around, and the Mexico studies which I
began in earnest in 1977 have occupied probably 85 percent of my research time
since. Now--as is true of any almost any field biologist working in fairly pristine
environments--the big problem is these environments are being destroyed. It turns
out the butterflies go into what is a very remote, relict forest ecosystem. Basically, it
is a Canadian fir forest ecosystem where warming [occurred] following the retreat of
the glaciers. These forests migrated up to a band on the mountains which
corresponds to where, in the summertime, there are clouds all around there. So it is
a fog forest, a fog belt. That is where these fir trees live. During the winter, it is
drier, actually, and [there are] a lot of clear days. The butterflies go up. These
millions of butterflies coming out of all of North America, east of the Rocky
Mountains, millions if not billions of them migrate into these few sites.
These sites are threatened by lumbering, which is accelerating. [They are also
threatened] just by overpopulation of Mexicans, breeding with an enormously high
birthrate in that area. I perceive the demise of this entire oyamel fir forest
ecosystem of the butterflies in the next ten to twenty years. As I talk to you now, the
phone is probably ringing as we are trying to get some action at high levels to try to
save the forest ecosystem. We have been working on it for fifteen years, but it is
futile. What we have done has been futile. It has been very small efforts. One of
the things we did was to try to adapt the United States system of national parks and
national forests with a core area which would be comparable to totally protected
national park land, surrounded by national forests where there would be controlled
cutting. Well, the President of Mexico made an edict, and these areas were
"protected." It turns out that this was, in retrospect, an enormous mistake because it
focused interest on these areas that are now controlled by the government. So,
typical of Mexico bureaucracy, all the permits for logging are now being issued in the
peripheral zone, which is the national forest zone. Once they are in there, they do
not give a damn where they cut. So it is just chaos now.
We are trying to battle this. We are faced with this perennial, false argument that
people are more important than butterflies, and we cannot deprive the poor Mexican
of the resources of the lumber. Well, what about the future Mexicans? The forest
will be completely gone in twenty years, and not only will they have absolutely no
wood to use or to enjoy, [but] what they will lose is a very large proportion of their
water supply because these mountains are in an area which is a very, very dry
season for six months of the year. In the summer it rains, and in the winter it is dry.
I had been in those forests when, if you are standing in the forests, it is raining. If
you stand in the clearing, fifteen feet away, it is not raining. The trees actually
capture the rain out of the clouds as the clouds move through the forests.
K: If there is nothing there to filter the rain, it is lost.
B: Right. It is called horizontal rainfall. It occurs in high, mountainous areas. It
accounts for a very substantial proportion of the total of rainfall which all the desert
towns around the base of these mountains--they are not desert--are dependent
upon. Just yesterday, I was talking on the telephone to a man who has lived in
Mexico for thirty years who told me [about] an absolutely incredible further
encroachment that is happening in these forests. And that is that the lumber people
go in and they cut down the trees and after they cut down the trees, truckers go in
and dig up the topsoil and they take out the topsoil and they sell it to people who live
lower down for developing their farms. So not only is the place being forested, the
soil is being mined. It is just an impending disaster.
I wrote a paper with another post-doc, Steve Malcolm, who was here for several
years working in my lab and who is now at Western Michigan University. Monarch
butterflies are one of the most common butterflies in the world. But this migration to
Mexico and overwintering there, and then flying back into the United States and then
breeding for several generations and then the great-grandchildren flying back to
Mexico and how do they get there? They have not learned it, so this is a genetically
inherited pattern of how they do it. So this whole complex migration and chemical
ecology, the relationship of the milkweeds and the oyamel fir forests and the history
of the evolution of the forests of North America and Central America and so forth, is
all tied up in the migration biology and in my judgement, the eastern migration of the
Monarch butterfly is an endangered biological phenomenon. People have to start
looking beyond just this sort of naive approach of endangered species. Yes, there
are lots of endangered species and that is another big problem. But you can also
have the biological manifestation of that species, which is even bigger than the
species itself, such as the Monarch migration or the blue whale migration or the
white whale migration. And almost all those animals have been obliterated and we
stand by as the Japanese are still out there killing whales. It is intolerable. Modern
society is destroying the earth and the biological diversity of this earth, and biological
phenomena are going left and right.
K: Much of biology has left a certain reductionism, going toward a holistic sense. It is
time to do the same with conservation.
B: I think so.
K: An ecological perspective in conservation.
B: I think so. But you know, you start talking about the problems of conservation at
academic meetings, with the exception of the Conservation Biology Society, and a
lot of people are really uneasy about raising these thorny social issues when you are
supposed to be doing science, as a scientist. In my opinion, it is all over with. We
have to do that, scientists, biologists--nobody is in a better position to understand
the destruction that is going on in the natural ecosystems of the world. I had a
horrendous argument with a very prominent head of the genetic engineering unit at
a cocktail party here about a year ago. This man was for genetically engineering all
the way. And I said to him, "Have you considered the possibility that if you can
design crops that can grow in marginal habitats like corn growing in saline soil, for
example, then what you are going to do is spread the amount of food into yet more
marginal habitats, reducing the complexity, and eliminating the animal life from those
ecosystems, and most of the plant life, and what have you got for it? What you are
doing is trying to solve the problem of feeding the world now with the inevitable
result that you will encroach on marginal habitats, provide the food for yet another
generation of humans, in other words sustaining what already is an unsustainable
overpopulation crisis." I just cannot differ more strongly with these people. And his
reaction was, "Well, you must be against people." And I said to him, "I am only
against certain people."
K: [Laughter] There is no shortage of people on the planet with five billion of them. I
have a few more questions, if I may.
B: Okay, I will stop now and just answer your questions.
K: No, this has been wonderful. I really appreciate you.
B: Well, it has been fun.
K: It is almost as if you have been able to read my list of questions. It has been great.
What brought you from Amherst to the University of Florida?
B: Basically politics and money and the opportunity to be closer to Mexico. During the
1970s, ecology was really on the rise at Amherst College. I had a lot to do with that
and I became perceived as a real threat. [The biology department was a] very small
department of eight people. When I first started teaching at Amherst College, I had
four students in my natural history course. The natural history course, at that time,
was a course which was a biology major elective. Over the years, my colleagues
managed to manipulate that course, which is what they did with other professor's
popular courses. They manipulated them so that they became non-credit biology
courses, simply electives. Therefore, the better students would not take these
courses. Well, that was one little manipulation. The other was that my course
became so popular, particularly the ecology [course]. I taught natural history in the
spring, and I taught ecology in the fall. Ecology went from an enrollment initially of
four students to, when I left there, over seventy students. Two of the more politically
prominent members of our department--one of whom was a very, very good teacher,
and I will never knock him on that basis--felt really threatened because ecology was
attracting enormous numbers of students. The most popular science course had
been their genetics course. The enrollments in genetics got up to about sixty to
seventy students. The year before I left Amherst College, the enrollments in
ecology surpassed them.
We had developed team teaching at Amherst College--two people teaching, which
has enormous advantages because if you want to go off and lecture, you can. The
other person can take it up for you. Well, we went through a series of appointments
as a second ecologist at Amherst College who did not get tenure. And they did not
get tenure for completely spurious reasons which basically was political undermining
of these young people's careers. That happened to a man called Doug Schemsky.
He actually threw in the towel and went to the University of Chicago. It happened to
Stuart Fisher, who was a really fine limnologist who had been educated at
Dartmouth. Stuart went to Arizona State University, where he is now a very
K: So what is a limnologist?
B: [He] studies fresh water habitats. In the desert, you got all kinds of interesting and
peculiar fresh water [habitats]. Anyway, Stuart Fisher has been very successful.
And it happened to two other people. One woman who was so humiliated, she
finally just threw in the towel and quit. And I just got fed up with it. I felt that I was
spending too much energy. I was chairman of the department, and every move I
made was being countermoved and undermined. And the worst thing of all was that
the students got involved in the politics. Here is the final coup de gras. One of the
women that was filling in between these other people had gone to Connecticut
College. She was really a very bright woman. She had more honor students that
year than anybody else in the department. Even more threatening than the fact that
she had these honor students was that three of the four of them were the best honor
students in the department. They were intellectually strongest. That was too much
for these guys to take. There was a question as to who was going to get this next
position. It was not even a permanent position at the time, as I remember. These
two guys actually manipulated these really bright students to vote against their own
K: That is disgusting.
B: So it became so politically rigged. There is a lot of heavy-duty stuff that you have to
put up with in academia. The president of Amherst College, John William Ward,
was not really in touch with the faculty. There was a faculty clique at Amherst
College that kind of advised the president, and people had time to spend politicking
and maneuvering and perceiving and advancing policies which they felt were
important to the college. If they were in the position to influence the president of the
university, they had enormous power. There was what was called a committee of
six, which was six faculty members who were nominated by the faculty and elected
by the faculty to advise the president. They ultimately oversaw all tenure decisions.
And I will never forget in my lifetime an interaction with the committee of six, one of
whom was one of these two colleagues of mine. One of these people I was telling
you about--not the woman, one of the two men--came up for faculty promotion. One
of my colleagues said, "Well, he is not doing science, he is doing astrology." I was
so dumbfounded at the time that I felt like I was sitting around [thinking], "One of you
tomorrow will be responsible for my death, which of you will it be?" Nobody said
anything. And I analogize historically to a similar kind of human behavior in my own
Anyway, so I just got so damn fed up that when the University of Florida offered this
really nice position down here, a senior professorship, and I would be closer to
Mexico, I would not have quite as much teaching, I would have the opportunity, for
the first time really, to have graduate students. I had one graduate student at
Amherst. I decided to leave. I do not regret leaving that political morass. I never
have been able to regain the incredible strength of students that I had at Amherst
College because they were so highly selected and because we had developed such
a strong program in ecology. I also got very involved in the environmental
movement at that point, which was very threatening to these people too, because it
was becoming a significant political force in the Connecticut valley. I probably
should have sweated it out and stayed there, as I think back on the career, because
the bureaucracy at the University of Florida, in my opinion, is another kind of very
destructive influence on trying to be really creative at this place. It really makes it
difficult for individuals to shine, in my judgement. Some do, in spite of it, but it is an
irritation that should be removed. And I do not know how you do that. I think it is
K: I know a little bit about what the Linnean Society is. Tell me what the Linnean
Society was and then maybe go into a little more [about] why you were so honored.
B: The Linnean Society of London was named after the Swedish naturalist Carl
Linnaeus, who invented the binomial nomenclature system. That was important
because its species were seen as created entities but the minute you start lumping
them together and what Linnaeus called a genus, then you have got several closely-
related species in a single genus. The minute you do that, you are on the slippery
slope of evolution. [Laughter] Anyway, Carl Linnaeus, to the end, denied evolution,
but he probably would have been burned at the stake if he had maintained it at that
point in history, which was in the eighteenth century.
Well, anyway, I am not sure how the Linnean Society came to be formed in England,
and I ought to know. But it is the oldest biological scientific society in the world. And
it has been in London for almost 200 years. Well, it is not that old, 150 years old.
One of the very first people to be awarded the Linnean medal was Alfred Russel
Wallace [English naturalist, 1823-1913]. And Alfred Russel Wallace was the co-
discoverer of the theory of natural selection. Historically, everybody knows about
that--that Darwin fiddled and could not bring himself to publish the theory of natural
selection until the theory was laid out, practically verbatim, sent to him by Wallace
from Malaysia. Following which, the British, in gentlemanly fashion, published the
two papers back to back in the journal of the Linnean Society. Anyway, Wallace
eventually got the medal, and then Theodosius Dobzhansky [American geneticist]
did and Steven Jay Gould [American paleontologist, Harvard] recently got it, and a
couple of years ago, I got it. It was probably because of my putting together natural
history with experimental biology and then going back in the field and testing things.
That plus my interest in conservation biology, which is very nice. I am glad to see
recognized in the statement of the award my interest in conservation biology, being
willing to take strong positions and let the chips fall where they may. So I guess
those are the reasons. I am sure having lived in England for two years did not hurt,
because I made a lot of good English friends. I tremendously enjoy English people.
They love to talk and they are not very inhibited. If they disagree with you it is a
challenge both ways. Instead of getting emotionally wrought about things, you get
really interested in the arguments and sometimes you can change them 180
degrees, and vice versa. So it is really fun and open and free. I enjoy that kind of
really strong, intellectual activity that is true of the British intellectual community.
K: Good. Judging from your curriculum vitae, you have earned what looks to be a fair
amount of mass media attention, everything from Reader's Digest popularizations to
"20/20". To me, that says that non-academics, non-scholars, non-scientists are
picking up a great deal from your work. It is speaking to something. I was
wondering if you might know what that is, or might care to speculate. What is this
B: Well, as I said before, I think biologists are in a position to recognize the degree to
which the planet earth's biota is being destroyed. The only other time anything like
this probably happened is when the asteroid hit the earth and nailed the dinosaurs.
That obviously happened at a faster rate than what we are doing. But almost all of
evolution has been slow and gradual. Animals and plants simply cannot respond to
the rate at which we are decimating environments. Something is going to fall apart.
The Monarch butterfly has certainly captured my own imagination, in terms of how it
is like the canary and the mine syndrome. The biology of this organism is
integrating such an enormous area. Millions of square miles are involved in its
breeding and migratory range. Millions of square miles, just in eastern North
America, let alone the fact that this butterfly has colonized Australia historically and
has developed its migration south of the equator. So the thing is 180 degrees out of
phase, which goes back to Colin Pittendrigh at Princeton University, who was
interested in clocks in biology and if you are in the Northern hemisphere in the
Southern hemisphere you have got different kinds of seasons and so forth.
So my interest in the Monarch has gotten me involved in, on the one hand, following
my grandfather's edicts and taking the microscope down to see what is going down
at this end, but also taking the telescope of the astronomer and turning it around
backwards and looking at it so you see the big picture. On the one hand, I have
intensely gone at it analytically, but on the other hand, I stand back and I see this
enormous picture. And I see the onslaught of herbicides in the United States, which
I consider to be another enormous threat to biological diversity, not because they kill
animals, but because they kill the food of animals. The traditional way to starve out
and kill people is to deprive them of their food supply. And that is exactly what
herbicides are doing to the biota. I was in Minnesota chasing Monarchs last
summer and what did I see? Thousands and thousands of acres of nothing but
corn, nothing but soybean and two or three other crops. There are no weeds
anywhere, there are no wildflowers. There is no biological diversity. What was once
a prairie grassland has turned into a petrie dish, a growing fodder for humans. It
has gotten so bad that there is some deal going on between the farmers and the
county who sprays the roadside, these little dirt roads that go back and forth which
divide up these old farms of the nineteenth century. There are thousands of farms
out there. You drive along these farms and you look on the side of the road and
they are growing one species of grass right up to the side of the road. Then they
harvest that. So they spray herbicides on these roadsides and there is no biological
diversity anywhere out there. Everywhere I look: [In] Mexico they are cutting down
the forests because of human overpopulation or other economic forces that are
really a product of too many people in this world. So the Monarch then becomes for
me a flagship of environmental deterioration. Kids love the Monarch butterfly.
Teachers love to talk about it. There is a rich enough biology that people can
understand the mystery of its migration in the fall. There are people all over the
world who are fascinated by this creature.
By working with the various television companies and so forth, I am able to get the
message out to a wider audience. But I will tell you something. It is done at a price.
For one thing, it takes a tremendous amount of time. And the other is it is going to
be done right. We are talking about information here, biological information. Okay,
there is the reality of what is going on, which we probably see dimly, even though we
know more about this butterfly than any other of the thousands of species in the
world. We literally know more about this butterfly than any butterfly. We do not
know much about them. There are 156 other species in the same family. We only
know the food plants of twenty-three of them. So we know very little, but in spite of
that, with the Monarch we know enough to capture the imagination of the people.
Now, what I have been trying to do as a scientist is to make sure that the media get
their facts straight and at the same time, keep it in the context of (A), a beautiful
story, which it is. It is intrinsically beautiful, intellectually and physically. I mean, the
mountains of Mexico, the places where the Monarchs migrate through, are
wonderful. Just seeing these millions of butterflies is unbelievable. So this is a
matter of conveying information. There isthe biology which is there and is real, part
of which we understand. There is our perception of it. Then there is the media's
perception of it--how can we make a story out of this that is going to sell our
products on "Good Morning America" or whatever--and then getting this back to the
people so that people become more aware of conservation.
So I have done that, and tried to make sure that that information does not get
corrupted and twisted. I mean, it is unbelievable how the media will twist these
stories to make it in their perception more saleable. Anytime you bring up the sex
life of the Monarch butterfly, which is a whole other really interesting story, they can
just lose all focus on the whole damn picture. They go berserk, and then they build
out of it a completely stupid and unrealistic story. I have been trying to keep the
hand on the tiller of the information that goes out through the public channels which
is the main reason I have involved myself with the media. But it takes an awful lot of
time, and perhaps our historical colleagues would be interested in this [story]. A
couple of years ago, we applied for a cooperative, multi-venture grant from NSF to
buy some fancy equipment for analytical chemistry. I signed onto that grant
application and you have my curriculum vitae there. In it, I have listed the number of
different television companies I have been associated with. Coming back as a
critique of my credentials was this blistering statement of sarcasm about Brower's
interest in the medium and, "What the hell does this have to do with science
K: Oh my..
B: So when I say there is a cost, there are these myopic, narrow-minded analytical
idiots out there who are unwilling to step back and say, "Well, not only is this bio-
diversity thing important, but after all, where the hell are the tax dollars coming
from?" They are coming from the public. I have always felt that I have been lucky in
my life to have been supported and educated by tax dollars or by people's gifts to
private institutions and I would like to make sure that the people who are sponsoring
me have an idea of what I am actually doing in science. So I would like to get that
son of a bitch and ring his neck, whoever it was, because he does not know what he
is talking about.
K: Is there anything that I have not given you a chance to say?
B: Probably not. I think you have been very generous.
K: Well, I want to thank you very much, Professor Brower, for agreeing to be
interviewed for this very good interview.
B: Thank you very much, Chris. It has been a pleasure meeting you and it will be
interesting to see what happens with this.