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CHARLES GALD SIBLEY
August 7, 1917-April 12, 1998
BY ALAN H. BRUSH
CHARLES GALD SIBLEY WAS born in Fresno, California, on
August 7, 1917, en cl cliecl at age 80 in Santa Rosa, Cali-
fornia. He was not a small-town boy who simply movecl upstate.
Between his early years in Fresno en c! his ultimate move to
Santa Rosa, Charles travelecl worIc~w~cle to conduct en cl report
on his research. He was one of the leacling ornithologists
cluring the latter half of the twentieth century, one of the
founders and a major player in the emerging fielcl of
molecular systematics, en cl contributed significantly to our
knowledge of the evolutionary relationships among the higher
avian taxa.
CharIes's intellectual intensity en cl excitement touched
the lives of many of his contemporaries in ways both good
en cl bacI, en cl he influenced several generations of students.
Few ornithologists have so polarizecl their students en cl
colleagues. Ultimately his greatest impact may be the trans-
mission of his icleas en cl intellectual fervor to students, which
he clicl with an evangelical intensity, sometimes threatening
his wrath but usually with the grace of a master communicator.
Charles was an exceptionally well-organized person,
blessed with a fine intellect and an unyielding belief in
himself. Those at the receiving ens! of one of his famous
217
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B I O G RA P H I C A L
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verbal debates or attacks may not have Tookoc! beyonc! their
own bruised egos to appreciate his finer qualities. He was a
generous person, giving freely and frequently of his time to
students en c! colleagues, particularly if it involves! cliscus-
sions of science. He took pricle in his broacl unclerstancling
of biology en cl its processes, but he stuck to his own beliefs
en c! unclerstancling of biological "facts" until presenter! with
unequivocal information that he was wrong. Then, immecli-
ately, he wouIcl champion the new information, never looking
back to c~well on the fact that he may have been wrong.
This contrary nature of being dogmatic on the one hancI,
while always welcoming new information on the other, macle
it clifficult for some people to clear with Charles en c! his
science, but for his students he was an encIlessly variable,
fascinating, en cl challenging role moclel.
Charles was associates! with six universities over the course
of his academic career. His first appointment was a one-
year assistant professorship in 1948 at the University of Kansas.
A year later he returnee! to his native state to join the faculty
of San lose State College (now California State University
at San loser as an assistant professor of zoology. In 1953 he
went to Cornell University as curator of bircis en c! associate
professor of zoology in the Department of Conservation.
During his 12 years there Charles acivancecl to professor,
taught ornithology to overflowing classes of both graduate
en cl unclergracluate students, clevelopecl Cornell's scientific
collection of bircl specimens, en cl mentorecl nine graduate
students and one postdoctoral fellow. In 1959-60 he took a
sabbatical year at Oxford University as a Guggenheim fellow.
Back on the Cornell campus cluring the summer of 1962,
he oversaw the activities of the 13th International Ornitho-
logical Congress. Broadly speaking, his research during the
Cornell years clealt with hybridization between species-pairs
en c! the molecular systematics of avian orders en c! families.
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CHARLE S GALD SIBLEY
219
Charles, who priclec! himself as an ornithologist, joiner!
the American Ornithologists' Union (AOU) in 1939, became
an electecl member in ~ 949, en cl a fellow in ~ 955. He served
as treasurer for I! years, from 1953 to 1963, en c! as presi-
clent cluring the 1986-~S term. Before becoming president
Charles served twice as vice-presiclent en cl was electecl to
several terms on the Council. In 1971 he was awarclec! the
Brewster Memorial Mecial by the AOU, en cl in 1986 both he
en cl his wife, Frances, became patrons of the organization
he hac! server! so often en c! well.
In aciclition to his AOU activities Charles was a secretary
of the Cooper Ornithological Society, a fellow or corre-
sponcling fellow of six foreign societies, en c! an officer or
council member of five societies. From 1958 to 1962 he
served as the secretary-general of the ~ 3th International
Ornithological Congress, en c! from 1986 to 1990 he was
president of the 20th International Ornithological Congress.
Altogether he was a member of about 15 scientific societies,
inclucling all major ornithological societies of the Uniter!
States, as well as Deutsche Ornithologen-GeselIschaft, Societe
Ornithologique cle France, Asociacion Ornitolog~ca clef Plata,
en c! Suomen Lintutieteellinen ~clistys. He server! on the
eclitorial boards of Evolution, Journal of Molecular Evolu-
tion, en cl Molecular Biology en c! Evolution
In 1965 Charles mover! to Yale University as a professor
of biology, the William Robertson Coe Professor of Orni-
thology, en cl curator of bircis of the Peabody Museum of
Natural History. In 1970 he was appointee! director of the
Peabody Museum of Natural History. During his years at
Yale Charles advised another seven graduate students and
three postdoctoral fellows. In 1986 he was elected to the
National Academy of Sciences. That same year Charles retired
and was named a professor emeritus of Yale University. Later
that year he ant! Fran again mover! back to California. There
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B I O G RA P H I C A L
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he became affiliates! with San Francisco State University as
a Dean's Professor of Science en cl Professor of Biology. In
1988 Charles en cl colleague {on E. Ah~quist received the
Daniel Girauc! Elliot Mecial from the National Academy of
Sciences in recognition of their contributions to our knowI-
ecige of avian systematics, en cl in 1991 Charles was awarclecl
the ATessanciro Ghigi Mecial by the National Institute of
Wildlife Biology (Italy). His final appointment occurred in
March 1993 after moving to Santa Rosa. There he was namecl
adjunct professor of biology at Sonoma State University, in
part so that he couIcl have continual access to his extensive
personal library that he hacl given to the university.
In his conversations with students en c! colleagues Charles
couIcl generate great excitement about the potential of his
research. He clelightecl in invitations as plenary or keynote
speaker ant! he occasionally organizer! mini-symposia at
scientific meetings, where he en cl his students wouIcl give
papers upciating their current research. Throughout his
career he attracted! inclivicluals upon whose lives he macle
an inclelible mark. Among those who stucliecl with him are
four AOU elective members, eight AOU fellows, an AOU
secretary, an editor of The Auk, en c! an AOU treasurer.
Every project that Charles undertook clemonstratecl his
talent for enlisting the help of an extraordinary diversity of
people en c! expertise. For example, in 1961 when he first
conceived of a DNA hybridization facility at Cornell, he
sent K. W. Corbin to Bethesda to learn the techniques from
the three investigators who hac! only months earlier clevel-
opecl the methoclology. In 1966 when Sibley wan tell avian
bloocl samples from European species, he contacted a num-
ber of friencis who wouic! be at that year's International
Ornithological Congress in Oxford, asking for their aicl in
that early work on hemoglobin, Charles was never hesitant
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CHARLE S GALD SIBLEY
221
to enlist knowlecigeable inclivicluals well outsicle academia
in order to achieve his goals in fieldwork.
No fieldwork of his illustrates this better than the immense
effort he put into planning for the 1969 National Science
Foundation expedition to Papua-New Guinea aboard the
research vessel Alpha Helix. A year prior to that expedition
Sibley en c! Prof. George A. Bartholomew (of the University
of California, Los Angeles) macle a comprehensive assess-
ment of the potential fielcl facilities, logistics, en cl personal
contacts in that vast region. There they enlister! the coop-
eration en cl help of an amazing group of inclivicluals, some
of whom were local officials, administrators, ministers of
either the Lutheran or Catholic churches, an archbishop,
ranchers, pilots, local scientists en cl educators associated
with the Australian National University facilities, members
of the Australian Bush Patrol, telegraph operators, directors
of sanctuaries, en cl native Papua-New Guineans.
As a youngster Charles was an avicl bircler en cl kept precise
records of his observations very early on. He was introclucec!
to natural history by reacting John Burroughs en cl Ernest
Thompson Seton. Close friend Robert Failing encouraged
his interest in bircis, en c! high-school teacher Jean M. Nelson
was particularly supportive of his interests in natural history.
Together they founded the natural sciences club at Oakland
High School. In the mic! 1930s as an unclergracluate at the
University of California, Berkeley, he gravitated to the
Museum of Vertebrate Zoology (MVZ). MVZ hacl become a
major center for the stucly of natural history uncler the
direction of Joseph Grinnell, whose fielcl notebook methods
Charles wouIcl later use to fill 15 volumes that cletailecl years
of fieldwork in his precise, uneclitec! script. The MVZ main-
tainecl an emphasis on the fauna of the region, as well as an
association with the museum of paleontology. Accordingly,
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B I O G RA P H I C A L
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his first publications were on fossil bircis obtainer! from the
tar pits at Rancho La Brea in Los Angeles.
After graduation from Berkeley in 1940 (A.B. in zoology),
Charles worker! one year for the U.S. Public Health Service
on plague suppressive measures. Military service intervened,
en cl he was commissioned as an ensign in the U.S. Navy
reserves. During the later stages of Woric! War II he was
callecl for active duty en cl rose to lieutenant as a communi-
cations officer in the Pacific theater cluring the last 19 months
of the war. His primary station was on Emiru IsTanc! in the
St. Matthias group, 75 miles off the northern tip of the
Bismarck Archipelago. During his off-duty time he collectecl
locally en c! sent scientific specimens back to the MVZ. That
effort on Emiru was supplementecl while on rest-ancI-
relaxation expeditions to the Solomon Islancis and the
Philippines.
This combination of travel en cl the collection of scien-
tific specimens was pure pleasure for Charles en cl wouIcl
typify family travel experiences over his lifetime. As the years
passed, his collection of museum specimens was replacecl
by the collection of egg-white en cl bloocl samples for serum,
hemoglobin, en c! ultimately the extraction of DNA. For
example, following the ~ 4th International Ornithological
Congress in Oxford, England, Charles organized a month-
Tong European vacation arounc! visits to zoological gardens,
aviaries, and the homes of European colleagues in an ongoing
effort to obtain critical species for his research.
After the war en c! now married, Charles returnee! to
Berkeley in 1946 to pursue a cloctoral degree uncler the
direction of Alclen H. Miller, who was himself a protege of
Joseph Grinnell. By the micI-1 940s Miller hac! follower!
Grinnell into the directorship of the MVZ en cl was particu-
larly interested in species-level taxonomic problems. At that
time Charles met John Davis, another incoming Miller cloc-
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CHARLE S GALD SIBLEY
223
toral student, whom he joiner! on a series of collecting trips
to Mexico. As a result Charles became fluent in Spanish,
Earned the ropes of carrying out fieldwork in Mexico, en cl
was introclucec! to some peculiar Mexican birc! specimens
collectecl by Helmuth Wagner.
Those specimens turned out to be hybrids between two
species of towhee in the genus Pipi~o. Subsequently, for his
cloctoral research Charles cleciclecl to examine the complex
patterns of plumage variation causecl by hybridization en cl
the breakdown of species-specific reproductive isolating
mechanisms between the recI-eyocl towhee, P. erythrophthalmus,
en cl the colIarecl towhee, P. ocai, along the transvoIcanic
plateau of Mexico. This was a zone of hybridization that
stretched nearly 500 miles from southeastern {alisco to the
states of Veracruz en cl Puebla. His thesis "Species Formation
in the RecI-eyoc! Towhees of Mexico" was publisher! as
volume 50 of the University of California Publications in
Zoology en cl was the first of 17 of his publications that clealt
with avian hybridization.
A major contribution of his doctoral work was the appli-
cation of a method for summarizing the plumage variation
among hybridizing inclivicluals as a single number, a hybric!
inclex value. The establishment of a species-specific hybrid
inclex scale was an extraorclinarily powerful en cl ingenious
method for analyzing complex, multigenic traits whose mor-
phological patterns shifted geographically due to hybridiza-
tion between incipient species. The method was later usecl
by his first group of graduate students to study the complex
patterns of hybridization between species-pairs in the Great
Plains of North America. In retrospect, CharIes's cloctoral
research can best be clescribec! as an early descriptive stage
in the clevelopment of his unclerstancling of the role playocl
by hybridization, both during the process of speciation and
as a result of the breakdown of reproductive isolating mecha-
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B I O G RA P H I C A L
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nisms. These were significant conceptual en c! methoclological
contributions to our unclerstancling of hybridization as a
mechanism of evolution.
After Sibley mover! to Cornell University the hybricliza-
tion studies were extenclecl to inclucle other species-pairs
that hybriclizecl throughout the Great Plains of North America.
They incluclec! Bullock's en c! Baltimore orioles, yellow-shaftec!
en cl recI-shaftecl flickers, indigo en cl lazuli buntings, en cl
rose-breastecl en cl black-heaclecl grosbeaks. Those years were
heacly, exciting times for him, involving his first graduate
students, Davicl A. West, Lester L. Short, Frecl C. Sibley
(unrelatecI), en cl Paul A. {ohnsgarcl in many fielcl trips to
collect hybrids along the Platte River en c! elsewhere in Colo-
raclo, Kansas, Nebraska, en cl the Dakotas. In aciclition he
revisited the Mexican highIancis to extend his earlier work
there.
Although the hybrid inclex method hacl proven to be a
powerful tool for studying the complexities of hybricliza-
tion for the breakdown of reproductive isolation, by 1958
Charles was looking for better ways to quantify the degree
of introgression between species-pairs. Simultaneously Paul
Johnsgarc! was in neec! of financial support to complete his
own cloctoral thesis. In an attempt to resolve both issues
Charles wrote a small proposal to the National Science Foun-
ciation to examine the possibility of using the new technique
of paper electrophoresis to study species-specific variation
in the serum proteins of game bircis. If successful, it might
be applied to the analysis of genetic variation in hybrid
populations.
As the research assistant in this small study {ohnsgarcl
followed Charies's instructions to the nth degree almost.
It was the "almost" that wouIcl prove to be serendipitous.
Like most of CharIes's students both then en cl subsequently,
Paul stood in mortal fear of invoking his wrath. Departure
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CHARLE S GALD SIBLEY
225
from the laboratory protocols was a carclinal sin. Paul, how-
ever, hacl react McCabe en cl Deutsch's earlier paper on the
electrophoresis of egg-white proteins. Out of curiosity en cl
a broacler interest, but without Charies's consent, Paul
incluclecl a few egg-white samples along with the serum
samples cluring his electrophoretic analyses.
At it turner! out, even with the crucle technique of paper
electrophoresis, the serum protein electrophoretic patterns
seemed much too variable among inclivicluals to be appliecl
to the hybridization studies. (Recall that at that time nothing
was known about protein variation, either within or between
species.) Lamenting this en cl greatly cliscouragecI, Charles
began to write up the results as a report to the National
Science Foundation. It was then that Paul mustered the
courage to reveal his covert analyses. The egg-white electro-
phoretic patterns were consistent among inclivicluals of a
species en cl clifferecl among the few species that hacl been
examined. Charles instantly recognized the implications of
those observations. A powerful new too! en c! a new set of
characters were awaiting application by systematists. Almost
overnight he put aside his plans for using serum proteins
to study the variation among hybrids and began to lay plans
for an electrophoretic study of egg-white protein variation
in bircis. Over the subsequent clecacle en cl a half that research
wouic! become a massive comparative taxonomic stucly of
the higher avian taxa. IncleecI, the relationships among avian
orders en cl families wouIcl be at the forefront of his research
interests for the remainder of his life. Thus began the next
phase of CharIes's research, which wouIcl overshadow the
earlier work throughout the 1960s en cl into the early 1970s.
The move to electrophoretic analyses of egg-white pro-
teins involvecl a major shift in CharIes's career. Along with
Herb Dessauer of Louisiana State University, who stucliecl
reptiles en c! amphibians, en c! Morris Goodman of Wayne
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B I O G RA P H I C A L
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State University, who stucliec! primates, Charles became one
of the founders of molecular systematics. For each of these
men this shift required a great clear of retooling both men-
tally en c! in the laboratory. The transition involves! a move
from activities that primarily usecl classic fieldwork couplecl
with comparative morphology to one of ciaily laboratory
analyses using the methods of comparative biochemistry. As
one might expect, the new approach was also encumbered
with some of the oIcl thinking.
A peculiar bias that Charles carrier! concerned! the genetic
variation of structural proteins versus enzymes en cl the ways
that natural selection wouIcl constrain the latter. He, along
with one of his colleagues at Cornell, believer! that enzymes
wouIcl be invariant in their amino acid sequences clue to
evolutionary constraints on their activity. Enzymes, in their
view, functioned only at specific temperatures and pH values,
en cl natural selection wouIcl weecl out all but the most ef-
fective structure for each enzyme en cl species. IncleecI, clur-
ing the early 1960s Charles en c! his colleague believer! that
an enzyme's primary structure might prove to be iclentical
both within and among species. Any variation in an enzyme's
structure wouic! rencler it inactive according to their logic,
en cl they knew little about the newly cliscoverecl phenomenon
of allozymes being stucliecl by AlIan C. Wilson at the University
of California, Berkeley, en c! Clement C. Markert at Johns
Hopkins University. Thus, in their view enzymes wouIcl be
unlikely to carry phylogenetic information en cl wouIcl be
useless for both systematic en c! population genetic studies.
Throughout much of the 1960s, informal debates on this
issue occurred between Charles en cl Wilson.
WiTson's careful studies of allozyme variation, coupled
with Markert's research on picine lactate clehycirogenases,
eventually convinced Charles that enzymes clicl in fact vary
within species. This conversion proviclec! the basis for another
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CHARLE S GALD SIBLEY
229
the taxonomic comparison of the egg-white proteins. The
early electrophoretic methods for the separations of proteins
on paper strips soon became obsolete. Paper electrophoresis
gave way to starch gel electrophoresis, whose relatively crucle
resolution potential was supplantecl by polyacrylamicle gel
electrophoresis en cl eventually by isoelectric focusing in either
polyacrylamicle gels or agarose plates. In an ongoing attempt
to refine en cl improve his comparative ciata, Charles acloptecl
each new improvement almost as soon as it became com-
mercially available.
Early on he was convinced that the comparative stucly of
protein variation couIcl aicl significantly in determining avian
phylogenetic relationships at the higher levels of classification.
He was equally certain that the methods wouIcl not be much
help at the levels of species en cl genera. Although protein
differences were basically phenotypic characters, they clifferec!
in one significant way from the traclitional morphological
characters usecl by most systematists at that time. Namely,
protein structure, cleterminec! by amino acic! sequences, was
only one step removed from the genetic code itself. Conse-
quently, differences among proteins were a more direct
reflection of the unclerlying genetic similarities en c! cliffer-
ences among species than was gross morphology. It was this
relationship between genes en cl the traits they encoclecI, in
this case the primary structure of proteins, that convincer!
Charles he was on the right track.
The first results of the early electrophoretic studies sug-
gested that the relationships among the higher taxa might
be cleterminecl with relative ease. The protocols were simple:
obtain egg white from the species of interest, separate the
proteins of each sample on either starch or polyacrylamicle
gels uncler appropriate controls en cl stanciarcl electrophoretic
conditions of wattage en cl time, stain the gels with amigo
blue black, photograph the gels, and then compare the
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B I O G RA P H I C A L
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resulting patterns. Voila! Evolutionary relationships were
revealecl like never before. It was a heacly time, en cl the
worIcl was watching en cl waiting for the results. Some were
envious that Charles was making such headway in solving
age-oIcl taxonomic problems, others were bitter that their
own expertise was being eclipsecI, but most ornithologists
were enthusiastic about the progress being macle.
By as early as 1959 the Cornell laboratory was deeply
involvecl in a comparative study of the egg-white proteins
by means of acrylamicle gel electrophoresis in small glass
tubes. Soon thereafter, en cl with his usual skill, energy, en cl
enthusiasm, Charles was extolling the virtues of those ciata
in resolving Tongstancling systematic problems. At annual
scientific meetings en cl through invited lectures in North
America and Europe he spreacl the message about the
wonclers of the new comparative methods. In 1960 he eagerly
presented ciata that clemonstratecl the affinities of the OIcl
WorIcl sylviicis en cl muscicapicis in contrast to their more
distant New Woric! cousins, the paruTicis. By the time of the
13th International Ornithological Congress, which was helcl
in 1962 in Ithaca with Sibley as secretary-general, there were
electrophoretic ciata bearing upon the relationships of many
· · ~
more Ivan fame. yes.
The methods of electrophoretic analysis may have been
relatively uncomplicatecI, but the effort to examine the evo-
lutionary relationships of all the higher avian taxa by means
of electrophoresis was daunting. There were the nests of
thousands of species to find. Each egg-white specimen had
to be compared electrophoretically over en cl over again.
Thousands of analyses were carried out over almost two
decades. Nothing but unequivocal data would satisfy Charies's
objectives. How else couIcl one compare all of the higher
avian taxa by means of this new technology? The museums
of the woric! houses! the scientific specimens neeclec! for
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CHARLE S GALD SIBLEY
23
comparative morphological studies, but there were no
depositories of egg-white specimens. Every species usecl in
CharIes's research program hacl to be collectecl by him en cl
his collaborators.
Charles set out to clo that, encouraging volunteers from
throughout the worIcl to collect samples en cl ship them to
Cornell University. The effort was massive en c! profouncITy
successful. For over a clecacle the samples came in from
every continent. Willing students acquired collecting per-
mits, risker! their necks climbing trees en c! cliff faces, comber!
forests, prairies, en cl tundra, all in search of samples from
both common en cl rare species. Hosts of both professional
ornithologists en c! amateur birclers collaborates! in the effort.
Along the way more than a clozen technicians carried out
the lab work that was completecl at Cornell en cl Yale. The
effort was monumental en c! culminates! in two monographs
publishecl by the Peabody Museum of Natural History at
Yale University: the first authored by Charles alone (1970)
en c! the seconc! coauthored! with l. E. Ah~quist (1972~. Charles
was proucl of these publications, as well he shouIcl have
been. Many taxonomic problems were resolvecI, although
others remained.
In aciclition to the egg-white protein studies there were
sicle excursions to utilize other protein systems either by
way of confirmation or for specific taxonomic problems.
One of these, coauthored with A. H. Brush, involvecl an
extensive stucly basecl on the electrophoretic variation of eye
lens proteins. Another, coauthored with H. T. Hendrickson,
involved the plasma proteins. Two particularly intractable
taxonomic problems, one involving the relationships of the
flamingoes en c! the other the relationships of the seer! snipe,
were tackled by using ion-exchange column chromatographic
techniques to examine variation in the tryptic peptizes of
hemoglobins. Other studies were never publishecI. The most
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B I O G RA P H I C A L
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important of these was a massive database clevelopec! at Yale
clearing with the electrophoretic variation of avian hemo-
gIobins Sanples were obtained from over half of the then
recognizes! birc! species. Another stucly involves! the use of
serology to examine the bloocl serum proteins of muscicapicis
en cl sylviicis. Ultimately it was the stucly of the egg-white
proteins that pair! the highest cliviclencis.
The egg-white studies of the bircis of the worIcI, follow-
ing those of avian hybridization on the Great Plains, wouIcl
have been a life's work for most inclivicluals in academia
but not for Charles. As the successes of the electrophoretic
analyses of the egg-white proteins began to accumulate, a
new technique was being tested in his laboratories at Cornell
en cl later at Yale. The methocl's early clevelopment by others
was an attempt to examine differences in DNA molecules
by means of annealing, or hybridizing, short fragments of
DNA to one another. The technique soon became known
as DNA-DNA hybridization. Although CharIes's laboratory
at Cornell began to explore the potential of the methoc! as
early as 1963, another clecacle wouIcl pass before Charles
hacl perfected the "DNA machine" in his laboratories at Yale.
The DNA-DNA hybridization studies involves! the clevel-
opment of another tissue collection. Initially, while at Cornell,
an attempt was macle to use tissue culture methods to grow
avian fibroblasts obtainer! from embryos. This methoc! was
soon abanclonecl clue to technical problems en cl the avail-
ability of a more direct method. Because bircis have nucleatecl
rec! blooc! cells, blooc! samples were the obvious en c! expedient
source of DNA. By the mid-1970s studies of the proteins of
egg white, bloocI, en cl eye lenses were all but complete, it
was time for the DNA studies to begin in earnest.
The years at Yale were some of the best for Charles en cl
some of his worst. The best saw the publication of his egg-
white monographs by the Peabody Museum of Natural His-
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CHARLE S GALD SIBLEY
233
tory en c! the clevelopment of the DNA-DNA hybridization
database. By 1986 the latter was being usecl to piece together
a comprehensive phylogeny of the orders en cl families of
the bircis of the woricI. In printer! form the clencirogram
spanned more than 20 feet along the walls of poster sessions
helcl in conjunction with annual scientific meetings cluring
the 1980s. It thus became known as the tapestry en c! was a
phenomenon in itself, as groups of people simultaneously
examined its cletails.
The worst moments at Yale involves! allegations against
Charles for two kinds of scientific impropriety. The first
was a fecleral indictment alleging that he hacl illegally
importer! the egg white of six European species, inclucling
one that was wholly fictitious en cl contrived by unknown
inclivicluals, either within or outside the U.S. Fish en cl WilcI-
life Service. After a goof! clear of meclia attention en c! the
paying of a substantial fine, this episode eventually lecl to
CharIes's resignation of the directorship of the Peabody
Museum of Natural History. It was a sac! moment, incleecI,
for a man who hacl priclecl himself for following the fecleral
guidelines regarding the necessary scientific collecting per-
mits here en c! abroad. It was simultaneously a black mark
against the scientific community that clicl so little to protest
this injustice. Sibley never explainecl why he chose to pay
the fine uncontested.
From a scientific point of view the second allegation was
much more serious. It involvecl the informal charge that
the analyses of DNA-DNA hybridization ciata hac! been
manipulatecl to yielcl results that conformed with precon-
ceivecl notions of phylogenetic relationships. One couIcl argue
that the methods of ciata analysis were not as rigorous as
they might have been. There were certainly differences of
opinion among the members of Sibley's own research group
on how best to quantify and summarize the data, however,
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234
B I O G RA P H I C A L
EMOIRS
this was an aspect of natural growth en c! clic! not constitute
fraud. In fact, the issue probably wouIcl never have arisen if
Charles en cl his group hacl not ventured into the treacher-
ous waters involving human evolution. The debates in that
arena are legenciary, beginning with Raymond Dart en cl
leacling up to tociay's antagonists. In Sibley's case the issue
revolver! arounc! rates of genetic change along different
phylogenetic lineages: specifically, the one that led to the
genus Homo, the other leacling to the remaining higher
primates. It was this debate that focuses! the attention of
the scientific community on CharIes's preferred methods
of analysis of the DNA hybridization ciata. At its heart the
issue was whether the entire genome of an organism evolves!
at a constant average rate, as Charles maintained. Although
there is solicl evidence to suggest that rates of change clo
cliffer among different lineages, the issue is still unresolvecI.
As in all other matters of his life Charles believecl in
himself. He believed unequivocally that his analyses of the
relationships of the bircis of the woric! were correct. In 1990
Yale University Press publishecl two massive scientific con-
tributions. One, in collaboration with his close friend en cl
colleague Burt Monroe, Jr., was Distribution en c! Taxonomy
of the Bircis of the Worm, a comprehensive treatment of all
avian species recognized as of ~ 990. The other, with his
longtime associate Jon E. Ah~quist, was Phylogeny en c! C]assi-
fication of the Bircis of the Worm: A Stucly in Molecular
Evolution This was the tapestry, along with all of the support-
ing data.
Charles knew the history of systematics well. He knew
better than most that classifications were always uncler review
ant! mollification, ant! he clic! not cleTucle himself into
believing that his classification wouIcl be the final worcl on
avian taxonomy. One of his cireams, however, cluring the
early phase of the DNA research was to be able to react off
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CHARLE S GALD SIBLEY
235
nucleoticle sequences from a DNA molecule. That was the
kind of precision he sought, knowing full well that the tech-
nology of the 1970s en cl 1980s was not up to that task.
Today automatic DNA-sequencing methods produce Tong
sequences of nucleoticles, en cl several genome projects are
at or nearing completion. AIreacly his students en cl their
students have built upon the contributions macle by Sibley
en cl his group. The possibility of eventually reaching a con-
sensus with regard to the phylogenetic relationships of bircis
is certainly obtainable, something that wouic! give Sibley
immense satisfaction.
Charles passed away at his home in Santa Rosa on Easter
Sunday, April 12, 199S, from myelogenous leukemia. He is
survival by Frances, his wife of 56 years, whom he met as
Frances Louise Kelly, en cl their daughters, Barbara Susanne,
Dorothy Ellen, en c! Carol Nacline.
THE TEXT OF THIS biographical memoir was modified from one pub-
lished in The Auk (116~19993:806-14), coauthored by Kendell C.
Corbin and Alan H. Brush. I thank both Corbin and Jon Ahlquist
for their contributions.
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SELECTED BIBLIOGRAPHY
1939
Fossil fringillids from Rancho La Brea. Condor 41:126-27.
1950
Species formation in the red-eyed towhees of Mexico. Univ. Calif
Publ. Zool. 50:109-94.
1954
Subspecies and clines: The contribution of avian taxonomy. Syst.
Zool. 3:105-10.
1955
Hybridization in the red-eyed towhees of Mexico. Evolution 8:252-90.
Ornithology. In A Century of Progress in the Natural Sciences 1853-
1953, pp. 620-59. San Francisco: California Academy of Sciences.
1957
The evolutionary and taxonomic significance of sexual dimorphism
and hybridization in birds. Condor59:166-91.
1958
With D. A. West. Hybridization in the red-eyed towhees of Mexico:
The eastern plateau populations. Condor 60:85-104.
1959
With P. A. Johnsgard. Variability in the electrophoretic patterns of
avian serum proteins. Condor 61 :85-95.
1960
The electrophoretic patterns of avian egg-white proteins as taxo-
nomic characters. Ibis 102:215-84.
1962
The comparative morphology of protein molecules as data for classif~ca-
tion. Syst. Zool. 11:108-18.
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CHARLE S GALD SIBLEY
1968
237
With K. W. Corbin and J. E. Ahlquist. The relationships of the seed-
snipe (Thincoridae) as indicated by their egg-white proteins and
hemoglobin. Bonn. Zool. Beitage 19:235-48.
1970
A comparative study of the egg-white proteins of passerine birds.
Bull. Peabody Mus. Nat. Hist. 32:1-131.
1972
With J. E. Ahlquist. A comparative study of the egg-white proteins
of non-passerine birds. Bull. Peabody Mus. Nat. Hist. 39:1-276.
1973
With J. E. Ahlquist. The relationships of the hoatzin (Opisthocomus).
Auk 90:1-13.
1974
With K. W. Corbin, A. Ferguson, A. C. Wilson, A. H. Brush, and
J. E. Ahlquist. Genetic polymorphism in New Guinea starlings of
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1980
With J. E. Ahlquist. The relationships of the "primitive insect eaters"
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1983
With J. E. Ahlquist. The phylogeny and classification of birds based
on the data of DNA-DNA hybridization. Curr. Ornithol. 1:245-92.
1984
With J. E. Ahlquist. The phylogeny of the hominoid primates, as
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1985
EMOIRS
With J. E. Ahlquist. The phylogeny and classification of the Australo-
Papuan passerine birds. Emu 85:1-14.
1986
With J. E. Ahlquist. Reconstructing bird phylogeny by comparing
DNA's. Sci. Am. 254:82-92.
1987
With J. E. Ahlquist, A. H. Bledsoe, and F. H. Shelton. DNA hybrid-
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1988
With J. E. Ahlquist and B. L. Monroe, Jr. A classification of the
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1990
With J. A. Comstock and J. E. Ahlquist. DNA hybridization evidence
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1994
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Representative terms from entire chapter:
natural history
