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42 IGUANA  •  VOLUME 13, NUMBER 1  •  MARCH 2006 RYAN

Stan Rand recording Túngara Frogs (Physalaemus pustulosus) on Barro Colorado Island, Panamá, in the 1960s.

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IGUANA  •  VOLUME 13, NUMBER 1  •  MARCH 2006 43

There is a hole in my chest where my heart used to be, and
a chasm in tropical biology the size of the Panama Canal.

Stan Rand died on 14 November 2005, and we are all the worse
off for it, personally and professionally.

Dr. Austin Stanley Rand, Stan to all who knew him, was
born on 29 September 1932. Perhaps destiny dealt him no
choice but greatness in biology, as his father Austin was already
a famous ornithologist at Stan’s birth. Stan was never far from
field biology or museums when he grew up — he published his
first paper when he was 12!

Stan began his graduate studies with Dr. Ernest Williams
at Harvard, where he became an early member of that awesome
lineage of students of Anolis biology under Williams’s tutelage.
He received his Ph.D. in 1961, and remained at Harvard on a
postdoc for a year. That year included an intensive period of
study of Jamaican anoles at the University of the West Indies.
He and his wife Pat then relocated in São Paulo, Brazil, for two
years while Stan was a postdoc with Dr. Paulo Vanzolini.
Initially, Stan was paid from royalties from a song that “Vanzo”
wrote. That tune must have been quite a hit in Brazil — or
maybe Stan just lived modestly (money was never a big issue to
him). Regardless, the Rand’s first child, Hugh, was born there
and it appears he didn’t starve.

In 1964, Stan received an invitation from Dr. Martin
Moynihan to join a new cadre of impressive young scientists that
Moynihan was assembling at the Smithsonian Tropical Research
Institute in Panama. Stan joined the staff. He, Pat, and Hugh
moved to Panama, where their two daughters, Margaret and
Katherine, were born. They lived on STRI’s Barro Colorado
Island (BCI), where the forest was inhabited by anoles and igua-
nas during the day and the Rands were serenaded by choruses
of Túngara Frogs at night.

Stan published early and often — over 100 scientific papers
in 60 years of publishing. Note that he is not yet done; Túngara
Frog papers with his data will continue to bear his name for
some time, and I predict that one will appear in 2014 to mark
Stan’s 70th year in the scientific literature. Many of Stan’s scien-
tific contributions can be partitioned into studies of Anolis
lizards, iguanas, and crocodiles, and, as a grand finale, his study
of Túngara Frogs. Stan explored many fascinating tidbits of
nature in addition to those projects, but I will briefly review only
these contributions.

Stan published a series of studies on Anolis biology in the
1960s that had an immediate and lasting impact on tropical
biology, ecology, and behavioral ecology. Prominent among
those contributions was his notion of the “ecomorph.” The gen-
esis of this concept originated when his data showed that on
each island of the Greater Antilles, different Anolis species had
diversified and adapted to nearly identical niches on separate
islands. During that time, Stan also investigated dominance
interactions among lizards, showing that, if the size difference
between males was sufficient, the larger male won, but, if the
size difference was minimal, the resident won. He demonstrated
the “residency effect” in 1967, well before this became an impor-
tant issue in behavioral ecology. Stan also integrated physiolog-
ical ecology into his studies. Critical to the ecomorph concept
was the climatic, not just the structural, habitat the lizards occu-
pied. At the behavioral level, he showed that lizards’ responses
to predators were strongly influenced by their body tempera-
tures. All of this foreshadowed Stan’s continuing emphasis on
the organism’s entire biology and the necessity of viewing it in
its natural context.

P R O F I L E

A. Stanley Rand (1932–2005)1
Michael J. Ryan

University of Texas at Austin

PROFILE

Stan and Pat Rand in front of their house in Gamboa, Panama, in 2003.

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1 Iguana has not and does not plan to use obituaries as “profiles” again
in the future. However, Stan Rand’s contributions to herpetology
were so monumental that we felt that we could not, in good con-
science, let his death preclude the opportunity to feature him and
his impact on our discipline. — The Editors

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44 IGUANA  •  VOLUME 13, NUMBER 1  •  MARCH 2006 RYAN

Stan made at least two other major contributions during
his work with anoles. One was a paper with Williams in 1970
on signal redundancy in communication systems. They used
information theory to estimate the quantity of information
about species identity that potentially could be communicated
in a lizard community. They calculated that the total amount far
exceeded what was necessary. This study was a wonderful
demonstration of how animals use multiple aspects of their dis-
plays to reinforce the same message, and it was one of the first
applications of information theory to animal communication in
the wild. Stan also considered the relationship between ecolog-
ical space and predator-prey interactions in the context of “aspect
diversity,” arguing that variation among species could result from
predator-driven selection that causes species to diverge in the
“escape space” available to them. This concept greatly aug-
mented studies of apostatic selection and anticipated much of
the work we see today in sensory ecology.

Another stage in Stan’s work dealt with social behavior in
and between Green Iguanas and crocodiles. Much of this work
was with Gordon Burghardt and his students, centered on the
small island of Slothia, a mere stone’s throw from Barro
Colorado Island. Female iguanas swim to Slothia from BCI and
nest communally. Stan and his brother Will wrote an insightful
paper on conflict resolution. It analyzed the competition
between female iguanas over the burrows they dug for nesting.
The analysis combined stochastic processes and energetic con-
straints and showed that females took into consideration the
amount of energy they had expended in building the burrow.

This study later led to some consternation among theoreticians
interested in honest signals who were convinced that the
“Concorde Effect” (adding more investment only because past
investments have been made) should be maladaptive. I remem-
ber one night on BCI when, over some rather mediocre
Panamanian rum, John Maynard Smith (who to me was always
the “Stan of theoretical biology”) asked Stan just how this could
be. Although Stan knew the theory about games, the data
reigned supreme — this is what they do, he replied.

Some interesting interactions between crocs and female
iguanas had been witnessed by few besides Stan. A female croc
had earlier nested at the same site where the fecund iguanas
sought shelter for their eggs. The female croc, Natasha, as the
several-meter-long crocodile was affectionately known, rushed a
nesting iguana, and grabbed the expectant mother in her mouth.
Instead of devouring her, or at least dismembering her, as any
protective maternal archosaur should do, Natasha delicately car-
ried the iguana back to the water and released her. Crocodiles
carry their newly hatched young to the water, so Stan thought
that having a small squirming baby reptile in her mouth released
Natasha’s maternal instincts. Some of this crocodile work is pub-
lished, and much more is oral history on BCI. When great sci-
entists such as Stan pass, we marvel at the accumulation of
knowledge they left us, and we lament the untapped knowledge
that went with them.

Stan began to study acoustic communication in frogs in the
forest of Boracéia in Brazil while conducting his postdoctoral
research with Vanzolini. That interest continued when he moved
to Panama. He immediately set out to document the vocal
diversity of these gnomes of the Panamanian nights, but he also
turned his considerable nocturnal skills towards one species, the
Túngara Frog (Physalaemus pustulosus, which at the time had the
more melodious generic moniker of Engystomops).

I went to BCI to begin my studies of sexual selection and
communication in Red-eyed Treefrogs (Agalychnis callidryas) in
1978. These frogs proved intractable for the study I had
planned, and I quickly switched my attention to Túngara Frogs.
At this point, Stan had published one paper on their foam-nest-
ing behavior, although nothing on their communication.

Embarking for a 1990 field trip to the mountains of western Panamá
(from left: Debbie Greene, Walt Wilczynski, Stan Rand, Mike Ryan,
Kyra Mills, Ulysses [uncertain of last name], and Frederico Bolanos).

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Stan Rand wearing his “Túngara team” hat during the field season in
2004.

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However, he gave me a manuscript that had been written in the
late 1960s or early 1970s that described the complex calling of
these frogs: a simple call, or “whine”, that could be produced
alone or could be followed by one-to-many secondary compo-
nents, or “chucks.” The manuscript was filled with incredibly
interesting and detailed natural history as well as experimental
studies of female phonotaxis. Among other things, Stan was
interested in whether female frogs were more attracted to the
complex call over the simple call. These experiments were con-
ducted when the emphasis of mate recognition was focused at
the species level and was concerned with how it contributed to
speciation through behavioral isolation. In frogs, this work was
being carried out in exemplary fashion by such luminaries as
Murray Littlejohn and Carl Gerhardt, both of whom were
inspired by the earlier studies of Frank Blair. So, at that time,
Stan was working in the intellectual shadow of the Modern
Synthesis and its emphasis on speciation, but instead he was
addressing questions about female choice and sexual selection.
This was some time before Robert Trivers (who also worked on
Anolis and was advised by Williams during his Ph.D. studies)
wrote his paper on parental investment and sexual selection in
1972. I added some to that early manuscript of Stan’s and we
published it in 1981. I finished my thesis in 1982 while Stan was
still concentrating on reptiles.

Stan and I remained in touch during the next few years,
although neither of us worked on Túngara Frogs. In 1985, we
met in the halls of the Smithsonian’s Natural History Museum
in Washington, D.C. and I suggested we start up a joint project
with those little beasts. In 1986, we began a now 20-year col-
laboration on what became known to some as the “Túngara
Frog Project.” Our initial interests were modest as we began test-
ing female phonotaxis in a carport using a plywood and burlap
testing chamber in Gamboa, Panama. Stan and family had just
moved from Panama City to Gamboa, and STRI was planning
on having some laboratory facilities there soon. The first ques-
tion we addressed was what aspects of the mating call made it
attractive to females, and the first studies we published in 1990
argued that, in this system, sexual selection was generated by
sensory exploitation. That idea, convergent with and inspired by
others such as MJ West Eberhard at STRI, has generated some
interest. Our interests in sexual communication in these frogs
continued to expand in concert with the lab facilities in
Gamboa, which in turn accommodated a more sophisticated
approach to our own research questions.

Stan was always interested in the entire biology of an ani-
mal, and our studies soon grew to embrace additional aspects of
communication, comparative studies of populations and related
species, and, through our collaboration with Dr. Walt
Wilczysnki, the neural mechanisms that controlled mate choice.
Memorable during those years was a 5000-km transect in which
Stan and I sampled and collected Túngara Frog calls and tissues
throughout the entire range of the species. Most of the data were
collected when we departed from Austin, Texas after the meet-
ings of the American Society of Ichthyologist and Herpetologist
(ASIH) that was hosted there in 1993. We drove from Texas to
Panama, collecting all the way. Those samples were supple-
mented by numerous trips to South America, where we studied

other populations of Túngara Frogs and their relatives. The data
from that transect have already provided the grist for at least four
separate studies, and their uses are still not exhausted.

In the year 2000, Stan, Walt Wilczynski, David Cannatella,
and I were PI’s on a multidisciplinary grant. This grant, which
involved collaborations among more than a half dozen labs,
addressed issues from phylogenetics to molecular neurobiology,
all emanating from the basic biology of the Túngara Frog that
Stan first glimpsed in 1964. As the grant ended in 2004, we were
asked to organize a two-day symposium on “Sexual
Communication in Túngara Frogs” at the Animal Behavior
Society meetings in Oaxaca, México. To kick off that symposium,
numerous attendees, many with little or no interest in frogs, sex-
ual selection, or communication per se, packed the room to hear
Stan Rand present what ended up being his last scientific pres-
entation: “Natural History of the Túngara Frog.” When Stan fin-
ished that talk, I detected in the sustained applause a tone of rev-
erence and appreciation for someone special.

The work on Túngara Frogs will continue unabated, serv-
ing as a scientific legacy to Stan, but it also will be a continuing
collaboration from the grave. Although no longer with us, we
will never outlive Stan’s inspiration or exhaust his insights.

Having now provided a most cursory summary of Stan’s
scientific career, I would like to end by saying more about Stan
the person, which is the real reason why so many of us mourn
his passing. Above all, Stan was a naturalist. His eyes, ears, and
mind were focused on the organism in its environment. He was
well schooled in theory, but not terribly impressed by it. He was

IGUANA  •  VOLUME 13, NUMBER 1  •  MARCH 2006 45PROFILE

Holding a microphone and a Marine Toad (Bufo marinus) in place of
the crucifix and the dove, Stan Rand impersonates Saint Francis of
Assisi. Stan was passionate about fieldwork and deeply appreciated the
beauty of toads. The picture was digitally modified at Stan’s request to
create a monk’s hairstyle.

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a great experimenter, but was always a bit cynical about how
such results might apply in the wild. Once we were waiting out
a drought in a small dusty town in the bush in Brazil. We went
to see a movie, “Edward Scissorhands.” An old woman narrates
the story in which, as a youth, she befriends a boy who has scis-
sors for hands. Afterwards, I asked Stan how he liked the movie.
He said it was totally unrealistic, the chronology didn’t match,
and the old woman could not have been a youngster when she
first met Edward. I said, “Stan, for chris-sakes, the boy had scis-
sors for hands! How realistic is that? So what if the chronology
was off.” He wouldn’t budge. The opening of the movie was
built on a house of cards and he would contemplate no farther.

For most, Stan is intricately associated with STRI. The
Smithsonian Tropical Research Institute is a great institution,
and Barro Colorado Island is its crown jewel. We celebrated
Stan’s retirement from STRI with a symposium in his honor at
the 1998 meetings of the ASIH in Guelph, Canada, the pro-
ceedings of which are published in Anuran Communication
(Smithsonian Institution Press, Washington DC, 2001). All of
the more than 20 authors I first invited to participate said yes;
so much for a list of back-ups! When I introduced the sympo-
sium, I said that, but perhaps for BCI, Stan was STRI’s most
valuable resource. An institution’s greatness is defined, of course,
not just by its physical facilities but also by its humanity — and
that is where Stan made by far his greatest contribution. Because
of his immense knowledge of tropical biology, Stan was often
called upon for advice, especially to initiate novices to this land
of plenty. His generosity knew no bounds, and his humor,
warmth, and enthusiasm were contagious. He readily extended
this generosity into the personal realm. He and Pat formed the
social hub of Gamboa for the last 20 years, and their house was
a scientific salon. Pat’s famous “frog dinners” for many of the vis-
iting researchers in Gamboa (regardless of whether or not the

scientists worked on frogs) were the social highlights of our sum-
mers. These dinners were also an incubator of scientific ideas.
Stan was not a “science nerd,” he was broadly informed and
could entertainingly engage guests with a broad array of topics,
but his insights and wit were always sharpened and ready to be
applied to the next scientific question.

I remember once bemoaning that, because Stan was not at
a university, numerous students missed out on all that he had
to offer. Wrong, wrong, wrong! STRI offers a wide array of fel-
lowships for researchers at all stages of their careers. They all
need STRI sponsors. I have counted more than 50 students
that Stan sponsored before 1990; surely I have missed many. In
addition, since 1986, Stan acted as sponsor to more that 70
interns and associates who have worked with us on the Túngara
Frog project. He visited my lab in Austin twice a year for a long
time; those visits were so heavily booked that I had to sequester
Stan at a local pub to have time with him — but even that hid-
ing place was discovered all too quickly. Finally, literally hordes
of students owe Stan deeply. Two now rather famous biologists,
one studies monkeys and the other ants, told me long before
they were famous that Stan was crucial to the early develop-
ment of their research forays on BCI. When I informed by
mass e-mail numerous colleagues of Stan’s death, I received a
plethora of responses in which the word “love” was used much
more than one might associate with “macho” (and “macha”)
field biologists.

So, now we say good-bye, Stan. Thanks for all that you
shared, the family, the friends, the tropics, and the frogs. You will
not be forgotten. Next rum’s on me.

Acknowledgements
I thank both Ray Huey and Jonathon Losos for their thoughts
on Stan’s Anolis work.

46 IGUANA  •  VOLUME 13, NUMBER 1  •  MARCH 2006 RYAN

When a student asked Stan about chemical defenses of Túngara Frogs
(Physalaemus pustulosus), he volunteered to lick some to find out if he
could detect any chemicals. Stan had a humorous disposition and
enjoyed making people laugh in the lab.

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Stan getting ready to fly in an ultralight plane in 2003 over the area of
Chilibre, Panamá. Stan was an adventurous man always looking for-
ward to new experiences.

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