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Terry L. Erwin: She Had a Black Eye and in Her Arm She Held a Skunk



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Terry L. Erwin: She Had a Black Eye and in Her Arm She Held a Skunk 9
Terry L. Erwin:
She Had a Black Eye and
in Her Arm She Held a Skunk
Re-published from "American Entomologist" Vol. 61 (No. I)
on the occasion of ZooKeys 500
Marlin E. Rice
1 DuPont Pioneer, Johnston, Iowa, USA
Corresponding author: Marlin E. Rice (
Received 9 April 2015|Accepted 9 April 2015|Published 27 April 2015
Citation: Rice ME (2015) Terry L. Erwin: She Had a Black Eye and in Her Arm She Held a Skunk. ZooKeys 500: 9–24.
doi: 10.3897/zookeys.500.9772
Terry L. Erwin is Curator of Coleoptera at the Smithsonian Institution, National
Museum of Natural History, in Washington, D.C. and editor-in-chief of ZooKeys. He
generated signicant controversy in 1982—which continues to this day—when he
published an estimate of 30 million species on Earth, which was substantially more than
the nearly one million described species. He was born 1 December 1940 in St. Helena,
California and spent his youth trout shing with his maternal grandfather in the High
Sierra near Lake Tahoe. As a teenager, with prodding from his father, he built hot rod
cars and was a founding member and later President of the California Conquistadores,
a hot rod club in the San Francisco Bay area. Erwin earned his B.S. (1964, Biology) and
M.A. (1966, Biology) degrees from San Jose State College. With a desire to learn from
the three greatest living carabidologists, he rst obtained a Ph.D. (1969, Entomology)
from University of Alberta under the direction of George Ball. is was followed by
a post-doctoral fellowship at Harvard University’s Museum of Comparative Zoology
ZooKeys 500: 9–24 (2015)
doi: 10.3897/zookeys.500.9772
Copyright Marlin E. Rice. This article was originally published in American Entomologist, volume 61, number 1, and is republished here with permission..
Launched to accelerate biodiversity research
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Marlin E. Rice / ZooKeys 500: 9–24 (2015)
with Philip J. Darlington, Jr. During that
year, a position opened at the (then) United
States National Museum in the Department
of Entomology, which he accepted, but two
months after taking the job, he departed for
a year-long sabbatical at Lund University in
Sweden, where he completed the carabidology
“trifecta” under the mentorship of Carl H.
Lindroth. While in Sweden, the Chairman of
Entomology at the USNM changed from Karl
Krombien to Paul D. Hurd, who saw on his
desk a proposal left by Terry to study California
carabid beetles. Learning that grant money
was available for research in Central America,
Hurd crossed out “California” and wrote in
“Panama.” Terry returned to Washington in
1971, as the second coleopterist within the
USNM, and was greatly surprised to nd
that his proposal had been changed, funding
had been secured, and he was scheduled for
the next ight to the Canal Zone. us began a lifetime career on studies of insect
biodiversity in neotropical forests.
is interview began in Austin, Texas on 13 November 2013 with e Macallan
18 (a single malt scotch) and a toast “to all things on six legs.” It concluded in Portland,
Oregon on 17 November 2014; Erwin was two weeks short of 74 years old.
Rice: What I want to do, Terry, is interview you for a new column in American Ento-
mologist called Legends. And I’ll do this column for ve years, or until I run out of energy.
Erwin: [Laughs.] is column is a good idea; an excellent idea for ESA.
You are still active in biodiversity and conservation, but I really wanted to narrow this
down and look at the entomological aspects and to communicate to entomologists broadly,
so some of these questions will be elementary, but some will be philosophical and you just
run with it any way you want.
Who is the person, or what was the event, that motivated you to study entomology?
ose are always great questions, and I know there are just all kinds of diverse an-
swers you get from everybody, but probably mine is kind of like a common one, and
that is J. Gordon Edwards. He was a professor at San Jose State College; it was college
then and not university, ’cause this was back in the 60s. My father was a race driver—a
tin knocker—and he didnt nish high school, and when he retired from Mare Island
Naval Shipyard, he was a nuclear engineer building atomic submarines, and that was
my path. My grandfather worked at Mare Island, my father worked at Mare Island,
Terry Erwin, senior, Vallejo Senior High
School, 1958.
Terry L. Erwin: She Had a Black Eye and in Her Arm She Held a Skunk 11
my mother, my uncle. Vallejo was a very small town, so that was it. You grew up and
worked at Mare Island. I actually did four summers there to help pay for my college.
You built atomic submarines to pay for college?
I was working in the atomic reactor room of the Polaris missile [USS] George Wash-
ington submarine. My job was to carry buckets of asbestos mud, so they put it on the
preformed stu, then they would wrap it with a berglass cloth. Anyhow, I was waiting
for my call. I was just sitting there in the lower level and I leaned up and it was all wet; on
my arm was chewing tobacco where somebody above had gone “pitooey.” I looked at that
and wiped it o and said this is not for me. [Laughs.] So, that was it. I was in junior col-
lege taking electives and because I had read a book by James Michener about Hawaii and
the Polynesians and their teeth problems, I decided to become a dentist. So I took zool-
ogy and thought it was pretty cool. en I was in a discussion with somebody and they
said, “Do you want to spend your life like this?” [Mimics a dentist staring into a mouth.]
Leaning over, putting a drill into somebodys mouth!
Exactly. I just realized I really didnt want to do that, so I went to San Jose State.
I had a favorite English professor in junior college, so I minored in English and ma-
jored in life science teaching. During that time, I had to take two life science classes,
one of which was marine biology with Polly McMasters and the other was ento-
mology from Gordon Edwards. Polly would get her class up at four o’clock in the
morning and go over to Moss Landing and dig up polychaete worms. Frozen ngers
and just…gawd! en we would go back to San Jose and in the afternoon, Gordon
Edwards would get out the buttery nets, and we would go out to Allen Rock Park
and collect insects in the warm sunshine. Didnt take me long to gure out what I
wanted to do. [Laughs.]
Denitely not a marine biologist?
Denitely not. And also Gordon was just a really dynamic personality, just fantas-
tic. He recruited maybe seven or eight students per year. I just switched to entomol-
ogy and the interesting thing during that phase was my English classes were dragging
me down. I was on probation with a D average and I aced Entomology 51. at was
back in the “Pleistocene” when it was 51, not 101. en I got As the entire rest of my
student career. It was because of Gordon and his professionalism as a professor and the
fact that [when] you brought your insect collection and if there was a Musca domestica
there, he would just salivate, “how great that’s pinned; thats really a great specimen!” A
student just jumps on all that kind of feedback. And that was it.
It is usually one individual and it was Gordon Edwards for you.
He was the one.
Being an “A” student, did you have any challenges during graduate school?
No, actually everybody wanted me to go to Berkeley. ere were some great co-
leopterists there like E. Gorton Linsley. But I said as soon as I walk on campus, I’m go-
Marlin E. Rice / ZooKeys 500: 9–24 (2015)
Terry Erwin, Curator of Coleoptera, Smithsonian Institution, 2004.
Terry L. Erwin: She Had a Black Eye and in Her Arm She Held a Skunk 13
ing to be the carabid expert, so I’ve got to go somewhere else. I wrote to Carl Lindroth
in Sweden, who had just published his volume three of the carabids of Canada and
Alaska. So I wrote to him and he said, “Well, you are already working on bombardier
beetles, and if you want to do that for your Ph.D., I’m not the right person. You really
should go to George Ball.” George who? [Laughs.] I wrote George and got back a let-
ter; he was on an 18-month sabbatical in Mexico collecting carabids. He said, “Okay,
I’m going to be down here for a little while, but why dont you just drive on up [to
Edmonton, Alberta] and nd a place to live and I can support you the rst year with
pinning my Mexican carabids.” He said just check in with Brian Hocking, the Chair
of the department. So my [ex-]wife and I arrived at the Hocking house, and Jocelyn,
the wife, opened the door, and she had a black eye and in her arm she held a pet skunk.
[Laughs.] She was this little British woman with a very nice accent who, unfortunately,
connected with a badminton birdie in her eye! “Welcome. ey said you were com-
ing.” And so they helped us through the rst week and we got a place to live. George
supported me for the rst year; then I got a Queen Elizabeth Scholarship for the next
two years. I nished it in three years.
What was the Queen Elizabeth Scholarship? Was that a full ride?
Yeah, a full-ride scholarship of $2,600 a year. [Laughs.] It did ne and that’s actu-
ally a Canadian grant. e idea was to nish o as soon as possible. en Phil Darling-
ton gave me a post-doc and I went from Edmonton to MCZ [Museum of Compara-
tive Zoology]. en Oscar Cartwright, the old coleopterist at the Smithsonian, retired
and they asked George Ball, who was visiting there, “Can you recommend anybody?”
He said, “Well, yeah, I just had a student graduate. He’s at Harvard right now, and
I’d recommend him.” So they called me and I said, “No, I dont want to come. I want
to do a post-doc with Carl Lindroth in Sweden.” ey said, “You can do that too, so
come on down.” at’s when I had my rst sabbatical. I was in Washington for two
months; then I went to Sweden for a year. After that, I had worked with three of the
top carabidologists in the world, and that really was my objective.
Lets jump forward and look at your career. What do you consider your most signicant
contribution to the eld of entomology?
I think this one-page Coleopterists Bulletin paper, for one thing, started a cottage in-
dustry in fogging, so that became a real technique to look at the forest canopy, and the
second thing was a cottage industry in shooting me down [laughs] from my naïve hy-
pothesis built on some naïve assumptions, and naïve arithmetic, and coming up with
the 30 million [species estimate]. But the point is that most people never realized, that
wasnt the point of the paper. at was a throwaway last paragraph. e point of the
paper was that Peter Raven [then Director, Missouri Botanical Garden] called me and
he was doing something with the National Research Council, where they needed to
know how many species were in an acre of Panama. at was the question. And I said,
“Peter, nobody knows that stu about insects. It’s just impossible.” I had done Panama
fogging in the tree Luhea semannii, and I said, “Well, give me some time and let me see
Marlin E. Rice / ZooKeys 500: 9–24 (2015)
what I can do.” And so I went through and analyzed all that stu with those numbers
and I came up with 46,000 species per hectare in Panama. He took that and that was
great; so let me put this in a little paper for Coleopterists Bulletin. Well, if we know this
for one tree, how many trees are there in the world? Fifty thousand? Okay, how many
insects are host specic? Who knows, but try 13 percent, and so that came to the 30
million. Several people came and said, “Well, what if its ve percent? What if it’s 20
percent?” And so forth. ose numbers have been batted around and theyre still bat-
ted around. e really interesting thing was that Yves Basset, from STRI [Smithsonian
Tropical Research Institute], just published a paper last January in Science, where he
had 110 taxonomists and 10 years of collecting with several dierent kinds of methods.
He came up with a minimum of 28,000 and a maximum of 44,000, based on all of
that. And I did it on one tree and some simple math and came up with 46,000! Actu-
ally, thats probably pretty close. We now know that there are probably over 100,000
[species] per hectare in the western Amazon Basin.
I checked on the paper in e Coleopterists Bulletin; it has been cited, according to
Google Scholar, 835 times.
Yeah, I think it just hit 848.
920 citations as of 1 January 2015.
Terry Erwin fogging for insects in the Amazon Basin, 2014.
Terry L. Erwin: She Had a Black Eye and in Her Arm She Held a Skunk 15
As long as we are on this number of species, you had estimated 100,000 species per
hectare based upon your work in western Ecuador. You have also mentioned 17 billion
hectares in the Amazon Basin. Did you provide a number for species?
No. My usual throw-away line is 100,000 species per hectare and 3.2
per hectare in the western Amazon Basin. ere are 17 billion hectares and 450 dier-
ent kinds of forest. Do the math. So that’s my line—do the math.
You are not going to lay a number out there and be quoted?
Right. No. [Laughs.] My point this morning [during the symposium] was that the
way we collected those things using the garden hose to wash them o the [fogging]
sheets, that’s the sample I used to get there. So what if I missed 50 percent of the speci-
mens because they got washed away or…then the 30 million would have been higher.
You have had a tremendous career studying carabids, but why study beetles, and espe-
cially beetles that inhabit the rain forest canopy? What got you into the rain forest high up
in a tree?
Why dont we just step back to beetles? Gordon Edwards was the coleopterist and
he had a very nice collection and he encouraged us. I started with cerambycids with
three of my buddies, who are all cerambycidologists and they were very competitive; I
mean, really competitive. So I asked Gordon if I could have another family. He said,
“Well, Carl Lindroth had just published volume three on Bembidion and I have been
collecting at Glacier National Park and the Tetons and I have lots; you could key those
out.” So, that’s how I got into carabids. Before I left for Sweden, Karl Krombein was
the Chair [of the department] and he said, “Leave me some proposal about what you
are going to do when you come back.” So I wrote a little proposal to do the carabids
of California, because I had a hundred thousand [specimens] that I had collected as a
student. When I came back, Paul Hurd had taken over as chair and he found out that
there was some money to work in Panama, and so he got my proposal—he crossed out
California and wrote in Panama. So I ended up, for seven years, working and going
back and forth to STRI on Barro Colorado [Island] and that is how I got involved in
the fogging. As I said, when those things came down and I saw these rare carabid bee-
tles on the sheet, I said thats how I have got to collect the carabids of the canopy. But
it sort of went out of the box from carabids into biodiversity because of the 30-million
paper and then that went into conservation and so forth, and the box just kept getting
bigger because of that rst fogging event.
It was like opening a Christmas package.
Oh man, its just unbelievable. And the genus that I’ve been working on for a
number of years, which is the genus Agra [and] which is strictly canopy—it has turned
out to be a lot of fun naming things in the genus Agra, but there are just over 500 spe-
cies described, many of which by me, and in the museum from all my borrows and
all my foggings, I have over 2,000 species. So that means I’ve got like 1,600 species
that need names. And you know, it’s the last biotic frontier. Until we started fogging,
Marlin E. Rice / ZooKeys 500: 9–24 (2015)
nobody knew what was up there. Now we
know the average size of a canopy beetle is
3 millimeters. So you think of architecture:
well, the ner you get, the smaller the twigs,
the smaller the insects.
Two thousand species of Agra! at has
been the focus of your research. I pulled three
names o the web: Agra cadabra, Agra vation,
and Agra katewinsletae. Give me some context
to those names.
Agra vation you could guess right away.
Agra cadabra is a play on words. Agra da-
ble—my [current] wife is Peruvian—and
this was a very pleasing, nice species. So
we speak Spanish and “agradable” means
very pleasing. [Laughs.] Agra katewinsle-
tae for Kate Winslet, Agra liv for Liv Tyler,
Agra catbellae, which is Catherine Bell, so
all of my heart-beating [pats his heart and
sighs] female movie stars can get a name if
they star in a movie where there is a disas-
ter. Okay, so the Titanic goes down; in my
etymology, the analogy is the destruction of
the rainforest and the Titanic going down.
Liv Tyler was in Armageddon, so they
are the same thing; the destruction of Earth,
the destruction of the rainforest. So all of
those celebrity names have to have some-
thing to do with disasters. Catherine Bell is
a star of JAG, a lawyer—she is just luscious.
But her nickname was Cat. Did you know
Frank Hovore?
Yes, I knew Frank.
Well, Frank used to hit golf balls at
the driving range with Cat Bell. He always
promised that he would introduce me to
her; unfortunately, he died rst. But anyhow, he told me her nickname is Cat. So cat-
bellae is the species name and I turned that into “the belly of the jaguar” and that was
in relationship to the demise of the jaguars’ rainforest.
Agra dax.
Agra sasquatch.
Terry L. Erwin: She Had a Black Eye and in Her Arm She Held a Skunk 17
ats clever. You have given the names some thought.
Yeah, all of those. We did a Smithsonian Channel hour, two years ago. My post-
doc, C. J. Gerasi, and I went into the studio, sort of like a Jay Leno kind of set up,
and Susan Spencer is the interviewer. So I come in rst and we are chatting a little bit,
then C. J. came in and we sat there for a little while and Susan says, “I understand
youve named some species after movie stars. Did you ever name one after a man?” I
said, “Well, of course, after my professor and people who have collected.” “Well, any
movie stars?” I said, “Yeah, I did and I had this one species of Agra that had this middle
femur; big, big femora, so I named it after Arnold Schwarzenegger—Agra schwarzeneg-
geri. at was the one man, and then he became the governor of California.” [Laughs.]
en she said, “Oh, yes.” She pulled out this beautiful blue folder with gold lettering
from the Oce of the Governor of California. My students had done an image of
schwareneggeri and sent it out to him and he signed it, “anks for thinking of me—
Arnold.” [Laughs.] Anyhow, that is the only one I’ve named after a male star and its
physical attributes had nothing to do with movies.
What is your passion in entomology—the thing that most motivates you or brings you
the greatest joy?
Curating the national collection. I’m the only coleopterist on the Smithsonian
side. I have four USDA colleagues in Coleoptera and each one of them is a contact
person for their family. Sasha Konstantinov has chrysomelids, Steve Lingafelter has cer-
ambycids, Lourdes Chamorro is our new curator of weevils, and then Nat Vandenberg
is the identier person. So I have all 165 other families in my responsibility and thanks
to David Furth, many of those have now been deactivated. So they and my research as-
sistant, Charyn Micheli, and the collection manager for Coleoptera are in charge of 12
million specimens, and of course, nobody can handle that, but then when you bump
it down to my responsibility with carabids, we have a little bit over one million carabid
beetles. So my goal in my career is to leave that collection just immaculate; as many
identied as possible, new species identied as new species, but maybe not described,
but everything sorted, everything in perfect order and it is great therapy—just to go
in and curate drawers of Coleoptera. Of course, I actually start with the groups I am
actively revising and get those done, but then I’m doing a series of books now—the
Carabidae of the Western Hemisphere. It’s going to be 10 volumes, three are published,
the fourth is almost done, I’m starting on ve, and theres 40,000 species of carabids
described; just over 10,000 from the Western Hemisphere. So the idea is that’s one
legacy project I’m working on, is the ten volumes. To support that is the other legacy,
which is to get the collection in perfect shape. at is what I enjoy most.
Describe the experience; when people hear the word Smithsonian, something majestic
comes to mind, and for somebody to work there, it’s probably like working in a royal palace.
It is, except the clothes of the royalty are tattered, hand-me-down pants and shoes,
[laughs] and it’s absolutely awful. My departmental budget, annually for each curator
throughout the seven departments, is $2,000. at’s all we get: $2,000. at $2,000
Marlin E. Rice / ZooKeys 500: 9–24 (2015)
brings me to the ESA meeting every year, and if I want forceps, I have to buy them out
of my own pocket. ey give us a phone—no charge—and every three years we get
an updated computer system—Dell—and updated is not quite correct. What we have
to do is take our old ones, turn them back to Dell, then they give us last years model.
Anyhow, royalty is all a façade, but just the fact is that we have the greatest, accessible
collections in the world. Paris [Musée National d’Histoire Naturelle] probably has
more specimens, but really not very accessible. You have to go into the attic and look
in old boxes and stu like that. So in that sense, it [the Smithsonian] is a great place
to work. e downside is that we cant go to NSF [National Science Foundation] for
funding. I’ve never been able to actually get nice big bunches of money where I could
do a ve-year project and expect to do it each year, and I just have to beg and borrow
year after year after year to do anything. So that part of the Smithsonian really sucks.
Everyone has a story to tell. What is a favorite memory of your career?
In 1976, I think, was the International Congress [of Entomology] in Washington.
I decided to do the rst international symposium on carabidology and set it up for
three days; a symposium with lots of talks. All the carabidologists came; Darlington
and Lindroth came over, and I used the State Department to bring some of our folks
from behind the Berlin Wall; Fritz Hieke from the Humboldt [University]. In those
days, it was really dicult to get those people out [of East Germany]. We had more
than a hundred people interested in carabidology and David Maddison was our young-
est at 17. Phil Darlingtons talk was about standing on the shoulders of giants. It was
just really a dynamic time. I was living in an apartment at that time and Dave Ka-
vanaugh and a couple of my colleagues from Europe were sleeping on the living room
oor. You know, it was just really an excellent time. I mean, we’ve had a lot of good
times after that, but I think that was a special time.
Back to the Amazon. What do you hope will be the outcome or the long-term impact
of your research?
at’s a good question. I’m hoping that as we get the rest of the 2005-2006 and the
current samples from this year, get that all in so that we have a 20-year image of what’s
going on, then I can tie down these numbers, like the 100,000 species per hectare in
Science or Nature or something like that. at will wake people up to the fact that,
yeah, we might have 30 or 50 million species or a lot more on the planet, but we are
knocking them o a million at a time. So I think just awareness.
Give me a perspective. When you fog, how much diversity or numbers of things do you
nd in a year, or how much have you collected in total over all of your eorts over all the years?
Okay. In fogging, the important thing to do is to ask the question and then design
the experiment using the fogging system to answer that question. So that may mean you
climb a tree and fog just the canopy of that particular tree, or in this recent thing where
we are doing bio-monitoring of the oil company road, we wanted actually a picture of
the entire forest and see what the impact of the road building and the use of the road by
Terry L. Erwin: She Had a Black Eye and in Her Arm She Held a Skunk 19
oil trucks is, and so that stretched over twenty years. We just nished up last year; three
intense years to start when they were building the road, then the 10-year follow up and the
20-year follow up. We have about nine million specimens from twenty-four hundred sam-
ples. Each sample, when you fog standing on the ground up into the canopy, each sample
has an average of about 2,800 specimens on a sheet that is three meters by three meters.
Wow! Nine million specimens.
We have 100 sheets for each seasonal visit and we have no idea how many species
on that particular sheet, but now as a result of 20 years of looking at everything, doing
some extrapolation looking at some taxa, we know how many species there are and the
relationship of that taxon with all the published ones. We suspect now that theres over a
hundred thousand species in one hectare of equatorial rainforest in the Western Hemi-
sphere; a hundred thousand species of insects and their relatives and the real Carl Sagan
number, the individuals in that hectare, [is] 3.2 times 10 to the 10th individuals. So that
gure has no name; you just have to say 10 to the 10th and thats what were getting in a
long-term study. If you just go out and fog one tree (one tropical tree, for example), you
get about seventeen hundred species; depends on the tree, of course—the next tree might
have three thousand species. It just depends on the toxicity of the tree and all kinds of
questions like that, but the main thing is, for this oil company road, the rule was that the
road could only be 27 meters wide because of various problems Ecuador had with previ-
ous oil companies. ey put rules and the virgin rainforest had to be intact on both sides
of the road, so for insects, after 20 years, there was little or no impact on the entomofauna
from that road. However, all the bushmeat was gone in three years. I started with ve
species of monkeys in the plot; at the end of three years there were no monkeys, no tapirs,
no cats, no crassid birds—currasows—anything that was edible was gone.
How far away from the road was this megafauna depleted?
is is the territory of the Huaorani indigenous folks, and before the road, there
were 70 dispersed families across two million hectares. [Here’s] a picture of the Huao-
rani: they have big wooden disks in their earlobes, some of them le their teeth, they
dont have very many clothes, they have blowguns, and they go o for days trying to
hit a monkey with a dart. Once the road came in, they dressed in western clothes,
they had ries and they knew how to hitchhike on oil company trucks and this road
is 121 kilometers long, so driving back and forth every day hunting, they wiped out
the megafauna, or the bushmeat, as we call it, for one to two kilometers back from the
road on both sides. e good thing was, 10 years later, most of those old hunters were
a little too decrepit to go hunting, and the teenagers—I actually had two teenagers,
Huaorani, helping me on the project—they didnt remember or were never trained on
how to follow an old machete trail. So ten years in, my plot had grown in and I asked
them to go and clear out this thousand-meter trail to the back of my transect. I was
teaching my students how to tie knots and hang up sheets and stu like that. After half
an hour I followed the two [Huaorani] guys and the trail was curving. What’s going on
here? I nally caught up with them and they had no idea how to follow scars on the
Marlin E. Rice / ZooKeys 500: 9–24 (2015)
little bushes that were cut ten years before [by machete]. And so, I had two monkey
species back in my plot; so theres hope.
So the monkeys are moving back into the plots.
I want to take you back to the nine million specimens you collected. What is one of the
most unusual things, dramatic things, exciting things that you caught—insect-wise—in
your nearly 40 years of fogging in the Amazon?
One of the very interesting things about these 2,800 specimens that come down, on
average per sheet, is once you start parsing out the individuals and looking at the same sheet
through the dry season, rainy season, and transition season, which is what we did for each
time that we went down, 51 percent of the catch across all 2,400 samples—51 percent
were ants. So the majority of abundance is ants no matter where you go. ats amazing,
absolutely amazing. But the next thing that is really, really interesting is you get walking
sticks and praying mantids of such camouage that you just cant image how these things
evolved to blend in with their tree trunks and the leaves and lichens; its just amazing. But
for me, the most very interesting thing, and I tell this to the hymenopterist at [University
of California] Riverside—Heraty, John Heraty. I’ve admitted this to John Heraty and I hate
to put it in print, but if the micro-hymenopterists would get o their lazy asses and start
describing species, there would be more micro-Hymenoptera than there are Coleoptera.
Really! You think so?
Absolutely, because every beetle, every weevil, has a parasite and those little tiny
micro-hym parasites have hyper-parasites of littler micro-hyms. I mean, its a no-brain-
er. But what I wanted to say about that is, when you look under the scope at this
tremendous biodiversity thats in the canopy, the neatest thing architecturally are the
micro-hyms—theyre just unbelievably fantastic. And dont tell John, but if I had to do
it over again, I might have been a hymenopterist. [Laughs.]
Well, it’s unfortunate that the entomological community can’t see this diversity that you
are talking about to learn to appreciate whats out there.
at’s the real thing, when you actually get one of these canopy samples and get
little spoonfuls in a little plate under the microscope to see the incredible diversity of
forms and species and all that kind of stu that’s in the canopy, that’s actually when
you appreciate how much biodiversity’s out there and this hundred thousand species
per hectare. Now thats the Western Hemisphere; the Amazon Basin has 17 billion
hectares, and in those 17 billion hectares there’s 450 dierent types of forest, and each
of those forests have subtypes of forest within them, so my 30 million estimate is so
conservative that it’s just hard to imagine what’s really out there.
So, what’s your new estimate?
It’s impossible to say, absolutely impossible.
Terry L. Erwin: She Had a Black Eye and in Her Arm She Held a Skunk 21
I can’t get you to give me a number, can I?
No, no! [Laughs.] Its just impossible to say, but the thing is, I’m getting them on
the hoof and were looking at morphospecies, but then the gel jocks are going into a
species—quote unquote—and nding out that actually that maybe theres ve or six
molecular species within that taxonomic species. And so that makes even my samples
more diverse than just what you can see with your eyes, and so then, that gives me
pause to make another estimate, because they are just getting started with how many
siblings are in a morphospecies. So, no—impossible.
You spent time in the Amazon Basin over a period of several decades. Did you ever
encounter a dangerous or threatening situation?
e rst time I was in the Amazon was 1977, so thats probably 30-plus something
years. [Laughs.] It seems like longer than that. You know, I’ve seen snakes and all that
kind of stu and been stung by Paraponera.
Really? Let’s stop there. Describe being stung by the bullet ant.
It’s a real shock when you get stung and you know immediately what it was. e
rst time was on the back of my arm.
Youve been stung more than once?
Yeah. So I grabbed that thing and pulled it out, and I forget who was with me, and
they looked all over and there was another one on my leg and they ipped that o, so I
didnt have any problems. I was just squeezing and squeezing, and then it dropped out,
but they are so hard I didnt kill it and it was crawling away. at lasted for about half
an hour, and by day two, oh, then after the re, it goes to a feeling like a dull toothache
and the toothache kind of goes for a couple of days and then it’s gone.
You mentioned a re. Do you mean that the sting felt like you had been burned?
Right. It’s a severe burning sensation. e second time was in this oil company
transect on the road and the oil company lm team had come out and they were
doing interviews in the forest and they wanted me to stand over there. I was just
standing and not paying attention, but [I] was next to a Paraponera nest at the base
of a tree and one crawled up, went out on my arm while I am giving the interview,
and it stung me in the thumb. Being stung on any of the ngers is the worst thing
Because why?
I think we have more nerve endings in our ngertips. It’s a nerve agent, what they
are actually putting in there. at lm has more four-letter words than I [laughs] prob-
ably even I know in my conscious. I jumped up and ran in circles and they were lming
me and wondering what the hell I’m doing, and I’m cussing and swearing and shaking
my hand. Of course I knew exactly what it was because I had a previous experience. So
those are two of my Paraponera experiences.
Marlin E. Rice / ZooKeys 500: 9–24 (2015)
What is the most dangerous thing in the rain forest?
e most dangerous thing is actually a tree fall, or a branch fall. A good-sized
branch comes down pretty fast and if youre under it and get hit on the head—youre
dead. at’s it. A tree fall, it takes a while, and you hear the crack, you look and if its
coming toward you, you just step one meter [aside] and it misses you. If you run, you
dont know where it is coming down and it could just clobber you. It’s the branches
that come down that are more dangerous. One time in Tambopata [Peru] I was tak-
ing down my pulleys for pulling up the fogger, and I was pulling it out and a branch
broke about that big [makes a circle with his hands the size of a baseball], and when they
break, they kind of have a pointy thing on them and it came down and went through
the hair here [points to his forehead], didnt hit my nose, but the branch went down,
ripped my chest clothes a little bit and then ripped the material in my crotch and stuck
in the ground between my legs. It was a long branch, so I’m sort of looking through
the foliage and all my colleagues are standing around kind of laughing a little bit until
they actually realize what happened. If it had hit me in the head, I’d be dead. When I
got back [to camp], I noticed my underwear also was ripped right out, but nothing on
my body. No scratches. It was so close it just took out my clothes. So that is the only
time in 40 years that I have been doing eldwork in the rain forest that anything close
to being a disaster happened.
Do you have a favorite insect species? It has to be a carabid.
Oh, absolutely. My license plate says AGRA DAX, and Dax is from Star Trek: Deep
Space Nine, and the actress, Terry Farrell—beautiful woman, absolutely beautiful. She
played Dax, and actually it was Terry Farrell’s body, but Dax is actually an alien parasite
that lives in her, but the alien was so ugly that it had to have a dierent body, and what
a body! Anyhow, Agra dax is my favorite. It’s actually a very large Agra with a heavy
body from Panama and metallic green with a rufous head with black antennal seg-
ments, so it’s quite colorful, and this particular group has attened tibiae and femorae,
which means it probably lives with ants, but we dont know too much about it. Its sister
species are Agra sasquatch and Agra yeti. Why? Because they have these really expanded
tarsal segments, so then it’s like Bigfoot.
What do you consider to be your legacy, or how do you want others to remember you?
I guess maybe by what my students do. If I’ve inuenced my students in a good
way and they go on to do stu, then the unbroken chain just keeps going. So thats
George Ball; he had 40 Ph.D. students, not just in carabids, but in other taxa, as well,
and many of us have gone on like Dave Kavanaugh—my best friend and [previous]
Chair and Science Director at the California Academy of Sciences. So you go back to
George and to Cornell, you have Forbes, and you go back from him to Cuvier and Buf-
fon, so theres this chain all the way from the great old-timers down through George
and his students and what I’d like to do is to keep that chain going with my current
student, Laura Zamorano from Colombia, and others.
Terry L. Erwin: She Had a Black Eye and in Her Arm She Held a Skunk 23
I hope I have that much energy when I reach your age.
I’ve now lived in the Amazon for 16 years of my life with the various expeditions
all put together, so for 16 years I breathed absolutely pure oxygen. [Laughs.] So thats a
plus. And beetles are my hobby, as well as profession. I never have any stress. If there’s
something not quite going right, I go curate a drawer of beetles, you know. My current
wife, Grace Servat, is Peruvian and is quite a bit younger than me and she kicks my
ass if I’m lying around, or something. [Laughs.] Shes an avian ecologist that specializes
in the high Andes. So shes up at 4,500 to 5,000 meters in her cushion-plant zone at
the very top breathing more than pure oxygen. I’ve been with her a couple of times
when there is no oxygen for my lowland Amazon lungs, so now I just have her show
me pictures and tell me about it. She does the same for my lowlands; she hates it down
where theres biting bugs and [it’s] hot and sweaty. So we do our own research, then
come back to talk about it, which is exciting to hear.
What is the compromise?
e compromise is our house in Washington; we come back to the home base and
our garden.
Terry, I greatly appreciate your candidness in answering my questions.
Well, it was fun, and e Macallan 18 single malt scotch helped, too!
Whenever I see you at an ESA meeting, you always have a cloud of people hovering
around you.
Most of them are students; younger people. e students keep you young. Like I said,
all my students want me for another 30 years. “You cant go!” [they say]. And I’m not!
anks to George L. Venable ( for providing the Agra beetle illustrations
and Terry Erwin for providing photographs of himself.
is interview is republished with the kind permission of Marlin Rice, and Lisa
Junker, CAE, Director of Publications & Communications, Entomological Society
of America.
Basset Y et al. (2012) Arthropod diversity in a tropical forest. Science 338: 1481–1484. doi:
Erwin TL (1982) Tropical forests: their richness in Coleoptera and other arthropod species.
e Coleopterists Bulletin 36: 74–75.
Marlin E. Rice / ZooKeys 500: 9–24 (2015)
Marlin E Rice is a senior re search manager with DuPont Pioneer in Johnston, Iowa.
He is a past President and Fellow of the Entomological Society of America.
... First pioneered by Southwood (1961) in temperate forests and then later adapted for tropical research by numerous others (Lowman and Wittman 1996), canopy fogging methods use insecticide to collect arthropods from the upper architecture of the forest habitat. These methods have been used in surveying arthropod diversity, particularly in lowland rain forests and at several neotropical localities (Erwin 1982;Basset 2001;Rice 2015). These data in part were used to make Erwin's (1982) bold estimate of 30-50 million arthropod species which was a stark contrast to previous estimates of 1.5-10 million (Erwin 1982). ...
... Sampling occurred during dry, wet, and transitional seasons to measure temporal turnover in species composition. Over nine million specimens were collected from these fogging events (Rice 2015) and many new taxa have been described from the specimens (e.g., Erwin 2010). ...
Full-text available
Canopy fogging was used to sample the diversity of bark and ambrosia beetles (Coleoptera, Curculionidae, Scolytinae) at two western Amazonian rainforest sites in Ecuador. Sampling was conducted by Dr Terry Erwin and assistants from 1994–2006 and yielded 1158 samples containing 2500 scolytine specimens representing more than 400 morphospecies. Here, we analyze a subset of these data representing two ecological groups: true bark beetles (52 morphospecies) and ambrosia beetles (69 morphospecies). A high percentage of these taxa occurred as singletons and doubletons and their species accumulation curves did not reach an asymptote. Diversity estimates placed the total scolytine species richness for this taxon subset present at the two sites between 260 and 323 species. The α-diversity was remarkably high at each site, while the apparently high β-diversity was an artifact of undersampling, as shown by a Monte Carlo resampling analysis. This study demonstrates the utility of canopy fogging for the discovery of new scolytine taxa and for approximate diversity assessment, but a substantially greater sampling effort would be needed for conclusive alpha as well as beta diversity estimates.
... Erwin, come raccontato da lui stesso in un'intervista con Rice (2015), pubblicò nel 1982 un articolo sulla rivista "The Coleopterists Bulletin", a seguito della richiesta del Direttore del Missouri Botanical Garden, il quale stava lavorando per un progetto per il National Research ...
Full-text available
Terry Erwin's influence on amateur entomologists is described by means of a personal experience form a field trip to Ecuador in 1988.
Full-text available
Background: We challenge the oft-repeated claim that the beetles (Coleoptera) are the most species-rich order of animals. Instead, we assert that another order of insects, the Hymenoptera, is more speciose, due in large part to the massively diverse but relatively poorly known parasitoid wasps. The idea that the beetles have more species than other orders is primarily based on their respective collection histories and the relative availability of taxonomic resources, which both disfavor parasitoid wasps. Though it is unreasonable to directly compare numbers of described species in each order, the ecology of parasitic wasps-specifically, their intimate interactions with their hosts-allows for estimation of relative richness. Results: We present a simple logical model that shows how the specialization of many parasitic wasps on their hosts suggests few scenarios in which there would be more beetle species than parasitic wasp species. We couple this model with an accounting of what we call the "genus-specific parasitoid-host ratio" from four well-studied genera of insect hosts, a metric by which to generate extremely conservative estimates of the average number of parasitic wasp species attacking a given beetle or other insect host species. Conclusions: Synthesis of our model with data from real host systems suggests that the Hymenoptera may have 2.5-3.2× more species than the Coleoptera. While there are more described species of beetles than all other animals, the Hymenoptera are almost certainly the larger order.
Full-text available
Most eukaryotic organisms are arthropods. Yet, their diversity in rich terrestrial ecosystems is still unknown. Here we produce tangible estimates of the total species richness of arthropods in a tropical rainforest. Using a comprehensive range of structured protocols, we sampled the phylogenetic breadth of arthropod taxa from the soil to the forest canopy in the San Lorenzo forest, Panama. We collected 6144 arthropod species from 0.48 hectare and extrapolated total species richness to larger areas on the basis of competing models. The whole 6000-hectare forest reserve most likely sustains 25,000 arthropod species. Notably, just 1 hectare of rainforest yields >60% of the arthropod biodiversity held in the wider landscape. Models based on plant diversity fitted the accumulated species richness of both herbivore and nonherbivore taxa exceptionally well. This lends credence to global estimates of arthropod biodiversity developed from plant models.
In the genus Agra, the virgata group is a circum-Amazonian-Middle American lineage, consists of 10 highly characteristic species that have a composite range extending from the tropics of México to Bolivia, east as far as the Brazilian State of Goiás. The majority of species are found on the periphery of the Amazon Basin, the rest in Middle America. Members of the virgata group and some brentid beetles (e.g., Arrhenodes gnatho (Licht.)) have similar bright and contrasting colors and elytral pattern and are sometimes found together. No connection or interaction between these two is known at present that could explain this similarity. Agra mime, new species, is described and attributed to the virgata group. The type locality is: Ecuador: Napo Province, 20 km east of Puerto Napo, Alinahui, 450 m, 01°00′S, 077°25′W. A revised key and checklist of the virgata group species are provided. A map of all known localities for the 10 species in the group is provided. A general discussion is included of the new species, as well as biogeographical notes for 6 additional species for which new locality records were found (Agra vate, A. nola, A. virgata, A. chocha, A. phainops, A. imaginis). The ohausi group, a southeastern Brazil lineage, consists of 4 species. Here I report a new synonymy for A. ohausi Liebke (=Agra rugipunctata Straneo, n. syn.). full page charges borne by the author
Tropical forests: their richness in Coleoptera and other arthropod species
  • Erwin
Erwin TL (1982) Tropical forests: their richness in Coleoptera and other arthropod species. The Coleopterists Bulletin 36: 74-75.