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OCR for page 159
ALFRED EDWARDS EMERSON
December31, 1896-October3, 1976
BY EDWARD O. WILSON
AND
CHARLES D. MICHENER
SOMETHING ABOUT ants, termites, and other social insects
attracts generalists, scholars who begin with a deep inter-
est in basic entomology, or else acquire it, and who restlessly
probe far beyond into such fields as evolutionary theory,
biogeography, the history of science, and philosophy while
conducting otherwise ordinary research. In the eighteenth
century it was Rene Antoine Ferchault de Reaumur; in the
nineteenth century, John Lubbock, Auguste Forel, and that
famous amateur myrmecologist, Charles Darwin. In our own
time William Morton Wheeler has been followed by Karl von
Frisch, Cary! P. Haskins, and Theodore D. SchneirIa.
Into the last group must be placer! Alfred E. Emerson.
Until his cleath he was the leacling authority on termites, a
restless technical expert who contributed massively to their
classification, anatomy, and biogeography. He was also an
important contributor to modern ecology, one of the synthe-
sizers of the 1940's and 1950's who brought the large quan-
tities of new data on adaptation, physiology, behavior, and
distribution into line with the emerging principles of the
"New Synthesis" of evolutionary theory. He was a biogeog-
rapher of importance; his cietailecI knowlecige of the worIc!
distribution of genera and species of termites helpec! bring
insects into the mainstream of general theory in biogeog-
159
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160
.
BIOGRAPHICAL MEMOIRS
raphy. Ant! not least, Emerson cleveloped the concept of the
superorganism to its extreme degree on the basis of his
knowledge of the workings of termite colonies; in the course
of this effort he helped to establish the importance of be-
havioral traits in classification ant! phylogenetic reconstruc-
tions.
Alfrec! Emerson was born in Ithaca, New York on De-
cember 3 I, IS96, the youngest of four children of a Cornell
professor of classical archeology. He moved with his family to
Chicago in 1905 when his father became curator of antiqui-
ties at the Art Institute of the University of Chicago. His
mother was a professional concert pianist and instructor in
the history of music at the University of Chicago; his brother
ant! two sisters all enjoyed successful academic careers. One
sister, Gertrude, became editor of Asia Magazine, settIec3 in
India, and was responsible for drawing Emerson into a
friendship with Indira Gandhi later in his life.
In the midst of this rich early cultural environment, with
its emphasis on the humanities, Emerson flirted briefly with
the idea of a career in music. Then, while a student at the
InterIaken School in Rolling Prairie, Incliana ~~9~O-l914),
he built and ran the school poultry farm the first odd
circumstance In a train of events that led to his career as an
entomologist. He was to become the family's scientific "mu-
tant," as he later (lescribecl himself. Upon reaching college
age in 1914, he went to Cornell University with the intention
of specializing in poultry science. But the courses were too
elementary and clull, causing him to try out the beginning
course in each science department of the university in turn.
When the time came to choose a major subject in his junior
year, Emerson picked entomology, principally as he once
said because the Department of Entomology was at that
time the best of its king] in the world. He became the personal
OCR for page 161
ALFRED EDWARDS EMERSON
161
frienc! of John H. Comstock and his wife, Anna Botsford
Comstock, as well as of James G. Needham, all large figures
in the history of the field. The Cornell entomologists stressed
depth of training and (letaile(1 expertise in individual groups
of insects, and Emerson clearly benefited from this experi-
ence through all of his subsequent research. At the same time
he formed a close friendship with another student, the her-
petologist Karl P. Schmidt, who later became curator-in-chief
of zoology at the Chicago Natural History Museum ant! a
fellow member of the National Academy of Sciences.
While at Cornell Emerson met his first wife, Winifred
Jelliffe, the (laughter of Smith Ely Jelliffe, a leading psy-
chiatrist. The couple became engaged in 1918, and soon af-
terwarct Emerson left for nine months service in the army
(clischarged in December, he did not see combat). Next
Emerson made a trip to the New York Zoological Station at
Kartabo, British Guiana, where, at the suggestion of William
Beebe, he began stuclying termites, anc! thus began his life's
work.
In 1920, as Emerson completect his M.A. at Cornell, he
married Winifred and took her on his second! trip to Kartabo.
A third expedition to British Guiana followed in 1924 and
then a six-month sojourn on Barro Coloracto Island, Panama,
in 1935. The termite collections that Emerson assembled and
the experience he obtained during these early visits to the
American tropics were a rich source of data and ideas on
which he drew during the rest of his life.
In 1921 Emerson acceptec! an instructorship at the Uni-
versity of Pittsburgh. After completing the requirements for
a Ph.D. at Cornell in 1925, he held a Guggenheim Fellowship
in 1925 and 1926 and then an associate professorship at the
University of Chicago. There he stayed for the remainder of
his professional career. The new associations that he formed
OCR for page 162
162
BIOGRAPHICAL MEMOIRS
at Chicago were decisive in the broadening of his interests
and the achievement of theoretical contributions in ecology
ant! behavior.
During the 1920's the Emersons had two chilciren:
Helena, who became the wife of Eugene Wilkening, a pro-
fessor of sociology at the University of Wisconsin; and
William ~elliffe Emerson, employed in the Department of
Anatomy of the University of Chicago. In 1949, following a
summer in which Alfred served as a visiting professor at the
University of California, Berkeley, Winifred diec! suddenly
from the effects of a heart defect acquired during childhood.
In 1950 Emerson married Eleanor Fish, whom he hac! known
for years and with whom he had collaborates! on a children's
book, Termite City (19371. Those of us who knew this couple
in later years were impresser! by the closeness en c! warmth of
their marriage.
By his own testimony, Alfred Emerson's principal contri-
bution to science was the more than one hundrec! articles that
aclded vastly to our knowledge of the systematics, phylogeny,
distribution, and natural history of termites around the
worm. In fact, he may well have been the most productive
researcher on this subject who ever livecI. By 1969, 1,914
species of termites had been clescribed by termitologists.
Emerson's collection, which was (lonated to the American
Museum of Natural History, container! about one million
specimens representing 1,745 species, or 91 percent of the
known worIc! fauna. No less than 80 percent of the species are
.
represented by primary type specimens. His personal library
on termites is virtually complete to the late 1960's, constitut-
ing an important bequest to future investigators. Emerson
remained active right through the later years of his life, as
evidenced by his excellent review of the Mastotermiticiae
(1965), description of the first Mesozoic termite (1967), re-
views of the fossil Kalotermiticiae (1965) and l~hinotermiti-
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ALFRED EDWARDS EMERSON
163
dae (1971), and analysis (with Kumar Krishna) of the rare
and little known Serritermitidae. Emerson's carefully re-
searched and cautious studies are the most authoritative
sources of information'on the classification and evolutionary
biology of the termites. His monographs on the termites of
Kartabo and the BeIgian Congo (Zaire) and Cameroon re-
main after many years the most valuable field guides for
entire tropical faunas. They are so well written and illustrated
as to be useful to anyone with an elementary knowledge of
entomology.
In ~ 949 Emerson coauthored the major synthetic work on
ecology to that time, Principles of Animal Ecology, an influential
textbook known lightly among students and other biologists
as the "The Great AEPPS" after the initials of the authors'
last names (W. C. Allee, A. E. Emerson, Orlando Park,
Thomas Park, and Karl P. Schmidt). This massive work col-
lected much of what was known about animal ecology at that
time, making full use of current evolutionary theory and the
still fragmentary principles of population biology. Emerson's
main contribution was to summarize knowledge of the social
insects, demonstrating with numerous examples the diverse
and often bizarre ways that features of social behavior adapt
species to particular challenges in the environment. In gen-
eral, Principles of Animal Ecology stimulated a great deal of
rigorous research in ecology and helped set the stage for the
surge in population and community ecology that occurred
during the 1950's and 1960's. (A commentary on Emerson's
eminence as an ecologist was published by T. Park in 1967
[Bulletin of the Ecological Society of America, 48: ~04-7].) Emer-
son's scholarly treatment of the social insects was the best
since the monographs by W. M. Wheeler twenty years pre-
viously, and they helped to keep these creatures in the midst
of developments in the major topics of ecology and the
remainder of evolutionary theory.
.
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164
BIOGRAPHICAL MEMOIRS
On top of Emerson's cumulative work concerning ter-
mites, his most striking single contribution, in the opinion of
many, was his use of behavioral traits as taxonomic char-
acters. Emerson referred to the structure of termite nests as
"frozen behavior" that could be weighed anct sketched with
the same reliability and quantity of information as many ana-
tomical traits. He showed that certain species of Apicotermes
can be distinguishes! more reactily by the architecture of their
nests than by the anatomy of the termites themselves. His case
seems exceptionally strong today, because termites have rela-
tively complex, stereotyped behavior, anct as subsequent in-
vestigators have shown, these insects use nest structure to
regulate the microclimate of the colony. It is fair to say that
what Konrad Lorenz and other vertebrate ethologists clid for
the use of behavior in bird systematics, Emerson helped to
accomplish for the use of behavior in the systematics of ter-
mites and other social insects.
Alfrec! Emerson is also well known for his espousal of the
superorganism concept, in which the castes anc! functions of
the insect colony are compared with the anatomical and phys-
iological features of single organisms. This method! of anal-
ogy, first put in concrete form by Wheeler and highly popu-
lar in the first half of the century, was perhaps carried to its
extreme by Emerson. He saw in the social insects the exempli-
fication of"clynamic homeostasis," which he believed to be a
new unifying principle of evolutionary theory. This part of
Emerson's thought has had relatively little impact, principally
because during the period of his most assertive articles
(1952-1958) the pendulum had begun to swing away from
holistic conceptualization and toward piecemeal, experi-
mental analysis of indiviclual physiological mechanisms and
patterns of behavior. But at the very least, however much out
of focus, and even cluring this period of its waning, the super-
organism concept remained} a stimulating distant goal toward
OCR for page 165
ALFRED EDWARDS EMERSON
165
which many younger entomologists felt themselves to be
working.
In spite of being a harci-working scholar in an exacting
specialty, Emerson was a gregarious man, exceptionally gen-
erous with his energies and time. He was friendly not only
with his intellectually gifted associates, but also with less
gifted persons with whom he willingly discussed everyday
topics. He wrote long letters of advice and encouragement to
younger entomologists, never displaying the protectiveness
or hardening of opinion that afflicts some established scien-
tists. in different years he served as president of the Ecolog-
ical Society of America, the Society for the Stucly of Evolu-
tion, and the Society for Systematic Entomology and was a
vice-president of the Entomological Society of America.
Among his honors were an honorary D.Sc. from Michigan
State University in 1961, received after his service as a dis-
tinguished visiting professor in 1960, and the Eminent Ecol-
ogist Award for 1967 from the Ecological Society of America.
He was elected to the National Academy of Sciences in 1962.
On Sunday, October 3, 1976, Alfrec] Emerson diec] of a
heart attack near his summer home at Huletts Landing, on
Lake George, New York. He will be remembered for the
magnitude and rigor of his scholarship, his uncompromising
and lifelong devotion to science, his interest in the relevance
of science to humane learning, and, especially by those who
knew him best, the largeness and generosity of his spirit.
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166
BIOGRAPHICAL MEMOIRS
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1938
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1958
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