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WARREN H. WAGNER, JR.
August 29, 1 920-January 8, 2000
BY D O NALD R. FARRAR
IN HIS PH.D. RESEARCH Warren ("Herb") Wagner was intro-
duced to classical methods of systematic botany, and found
them wanting. He was disturbed by the frequent absence of
quantitative data and the generally untestable hypotheses
of traditional reconstructions of species' evolutionary rela-
tionships. At the time, the latter was based largely on the
expert's weighing of the evidence and authoritative state-
ment of an opinion that could be argued but not easily
tested. Herb was determined that in his own research
monographing the endemic Hawaiian fern genus Die]]ia,
he would use evidence from all sources and explicitly state
the relative influence of each in an objectively constructed
illustration of phylogenetic relationships. The result was the
birth of his groundplan divergence index, for which he
soon became widely known. Herb's insight and instigation,
coupled in ensuing years with computer-assisted analysis of
comparative data, revolutionized the fundamental methods
and concepts of phylogenetic reconstruction, leading directly
to the burgeoning field of cIadistic analysis of evolutionary
relationships among plants. For his seminal contributions
Warren H. Wagner, {r., is generally considered a founding
father of modern plant systematics.
301
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B I O G RA P H I C A L
EMOIRS
Warren Herbert Wagner, Jr., was born on August 29,
1920, and was raised in Washington D.C., the son of Warren
Herbert Wagner and Harriet Claflin Wagner. His early
interests in natural history took him frequently to the
Smithsonian Institution, where he became acquainted with
the experts, including the eminent pteridologists William
R. Maxon and Conrad V. Morton and lepidopterist Austin
Clark. In college at the University of Pennsylvania he became
the enthusiastic field companion of Edgar T. Wherry, author
of the The Fern Guicle (paperback, Dover Publications, 1995~.
Wherry was a mineralogist who became an expert on fern
habitats and the first to point out the important associa-
tions of epipetric ferns with particular rock types. This
undoubtedly nurtured Herb's enthusiasm for mineralogy,
later his extended field trips with students often included a
day of mineral collecting. When as a student I brought
back an unusual form of cliff-brake fern from Missouri,
Herb was anxious to visit the site, not so much for the fern
as for the barite crystals I had found there. His fascination
with butterflies (he authored or coauthored 20 papers on
Lepidoptera) he called them "flying flowers) dictated
that he carry a butterfly net on field excursions, thus pre-
senting the archetypical layman's image of a biology professor.
I vividly recall stopping at a fast-food restaurant in the Missouri
Ozarks, where after ordering, Herb headed for a nearby
field filled with flowers and butterflies. The curiosity of the
restaurant staff was definitely aroused by the spectacle of
this man running through the field swinging a net at prey
invisible to them. After we explained, our waiter walked
. .
into the field to shout, "Hey perfesser, your lunch is ready!"
Graduating from the University of Pennsylvania in 1942,
Herb entered the U.S. Navy Air Corps, serving first in the
Atlantic, then in the Pacific Fleet, where he was a naval air
navigator. In the Pacific islands he spent his off-duty hours
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WARREN H. WAGNER, JR.
303
collecting ferns en c! butterflies, later publishing (with Davic!
Grether) "Ptericlophytes of Guam" as well as articles on the
ptericlophytes en cl butterflies of the Acimiralty Islancis. During
this time he also flew into California, taking his specimens
to E. B. CopelancI, renowned expert on Philippine ferns, at
the University of California, Berkeley. This was the beginning
of an association that wouic! bring him back to Berkeley for
graduate stucly. While in the Navy, he also began what was
to become a lifelong study of the ferns of the Hawaiian
IsTancis.
At Berkeley in 1945 Herb joined an exceptional group
of graduate students returning from WorIcl War II that formecl
fertile grounds for growth of new concepts in botany, evo-
lution, en cl systematics. His student colleagues from 1945
to ~ 950 incluclecl Charles Heiser, Ernest GifforcI, lack
Rattenbury, Isabella Abbot, Frank Ranzoni, Verne Grant,
Art Krukeberg, en cl Ecl Cantino. Their teachers incluclecl
Melvin Calvin, Richard GoIcischmicit, Curt Stern, G. Leclyarcl
Stebbins, en c! Herb's major professor, Lincoin Constance.
Copeland, though retired, was still active and served on
Herb's Ph.D. committee.
Also among Herb's student colleagues was Florence
Sign aigo, who was studying the systematics of reel algae with
George Pappenfuss. Herb en cl Florence were introclucecl by
fellow student Charles Heiser in the elevator of the herbarium.
Florence recalls,
Herb and I used to go over to San Francisco, to various bars, where we
would order a beer, and after a while Herb would ask the bartender if it
was all right if he played the piano. Sometimes the bartender would show
up later at the piano with two free beers. Once one had to ask Herb to stop
playing a piece because it was making a woman at the bar cry. And once he
was offered a job as a piano player.
Herb en c! Florence were marries! in 1948. They hac! two
chilciren, Warren Charles Wagner (b. 1953) en cl Margaret
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B I O G RA P H I C A L
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Frances Wagner (b. 1957~. Florence switcher! her allegiance
from algae to ferns en cl together she en cl Herb comprised a
formiciable research team both in the lab en cl in the fielcI.
Their home in Ann Arbor was a busy en c! warm environ-
ment, frequently hosting receptions for visiting botanists
en cl on holiciays wonclerful clinners for any of his graduate
students who were in town. Herb continues! to clelight aucli-
ences in informal gatherings en cl sometimes at bars with
the flamboyant piano playing that reflected his personality.
It was always fun to watch the bar manager's expression
change from skepticism to astonishment as Herb began to
play. Once, after several evenings of this in a hotel bar,
Herb was refuses! permission to play because the house
pianist, embarrassed by the contrast with his own lackluster
style, was threatening to quit.
Herb actively pursues! his research en c! teaching until
just weeks before his cleath on January 8, 2000, at the age
of 79 from suciclen cardiac arrest. He hacl experienced symp-
toms of heart failure for a few years before his cleath, but
not enough to incapacitate him. Although officially retired,
he hacl continual teaching his courses on woolly plants
en c! plant systematics en c! maintainer! a rigorous schecluTe
of invited lectures to institutions around the worIcl as well
as national and international meetings and symposia. In
the summer prececling his cleath Herb en c! Florence con-
cluctecl fielcl work in Alaska en cl in southwestern Canada,
from both places returning with, of course, new species of
Botrychium.
After receiving his Ph. D. in 1950 Herb spent a year as a
Gray Herbarium fellow at Harvard, then moved to the Uni-
versity of Michigan in 1951, where he remainec! throughout
his career. From 1966 to 1971 Herb served as director of
the University of Michigan's Matthaci Botanical Garden.
He chairec! the Department of Botany in the Division of
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WARREN H. WAGNER, JR.
305
Biological Sciences from 1974 to 1977, en c! chairec! many
aciclitional department en cl college committees, inclucling
the University of Michigan's Tropical Studies Committee
from 1983 through 1997. He was chairman or president of
nine professional societies, inclucling the American Fern
Society, American Society of Plant Taxonomists, en cl the
Botanical Society of America, en c! council member, trustee,
or Divisor to clozens of organizations. He was in clemancl as
an external reviewer of departments of biology en cl botany
across the country. He server! as an editor for the Univer-
sity of Michigan Press, The Indian To urn e] of Ptericlology,
en cl The Flora of North America (coecliting "Ptericlophytes"
in volume two tI9931~. He reviewer! countless journal manu-
scripts en cl grant proposals. To these causes en cl many more
he gave freely of his time while continuing to teach en cl
while maintaining a research program that generates! over
250 publications. He was electecl to the National Academy
of Sciences in 1985. His official retirement in 1991 proved
to be only a formality, as his research en c! teaching continued
unabated.
One of Herb's first endeavors as a young professor at
the University of Michigan was probing the origin and
relationships of the Appalachian Aspleniums, a confusing
group of ferns to which he hacl been introclucecl years earlier
by E. T. Wherry. The keys to solving this puzzle of starkly
different species with a seemingly complete array of inter-
mecliates lay in (~) examination of chromosome numbers
en c! their pairing behavior in meiosis, (2) relating this chro-
mosome ciata to spore abortion en cl intermediate morpholo-
gies, en cl (3) appreciation of the fact that fertility couIcl be
restored to "sterile" species hybrids through allopolyploidy,
a simple doubling of the basic number of chromosomes
(1954~. Thus the now well-known Appalachian Asplenium
triangle was resolver! into three cliploic! species (the corners
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B I O G RA P H I C A L
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of the triangle), three fertile allotetraploic! species (origi-
nating as hybrids between the three cliploicis), en cl numerous
backcross hybrids that occurred wherever a tetraploicl en cl
diploid species grew together. Subsequently verified through
artificial crosses, flavonoicl chemistry, en cl allozymes, this
moclel of reticulate evolution quickly became the basis for
making sense of similar species complexes in other fern
groups en cl in seecl plants.
Revolutionary in its time, attributing an important role
to species hybrids in plant evolution (196S, 1969) contra-
dicted the long-held notion of species hybrids being evolu-
tionary cleacI-encis. Herb's studies clemonstratecl that plant
species hybrids conic! in fact be the initial step in the
formation, through allopolyploicly, of new species that con-
tinuccl to participate in subsequent evolution of the genus
(19801.
Sterile Fit hybrids also proved to be much more common
in plants than in animals, en cl Herb was on a mission to
spreac! the news. His seminar presentations always worker!
in a series of hybrids demonstrating wicler and wicler crosses,
encling with a wittily misshapen fern that he procIaimecl to
be a cross between a woos! fern en c! a rec! oak! Such exaggera-
tions drove home the point that hybrids were to be expected
in nature en cl recognized as a component of the flora at
any given time en c! place. Although most of these hybrids
might be sterile cleacI-encis, their presence constituted part
of the "evolutionary noise" (his term) through which the
systematist must trace "signal" lines en c! processes leacling
to long-term persistence en cl divergence (1970~.
A part of Herb's diatribe on hybrids was that they were
easy to detect, because they were invariably intermediate
between their parents. Because clevelopment of most mor-
phological traits wouIcl be uncler the control of a set of
genes representing a combination of the two parents, not
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WARREN H. WAGNER, JR.
307
only could one predict the morphology of hybrids but given
a hybrid and one parent, one could also predict the other
parent. In the case of allotetraploid species that may have
formed in the ancient past, it was possible that one or both
of the diploid "parents" might now be extinct. This was
Herb's conclusion relative to the wood-fern genus Dryopteris,
which seemed to lack an extant diploid needed to form two
of the allotetraploid species (1970~. His naming of this extinct
species was hard to accept by many and led to a decade of
alternative proposals designed to avoid postulation of a
missing species. As with the Asplenium triangle, subsequently
derived molecular evidence supported Herb's conclusion.
Herb's persistent proclamation of hybrid intermediacy
set up a straw man easily knocked down by later studies
showing transgressive hybrid morphologies in traits controlled
by one of a few genes. This didn't phase Herb. His goal was
always to understand and promote the "big picture," the
principles that explained most of nature and natural pro-
cesses. His procedure though was to study the knowable
details. Through accumulation of details the big picture
would emerge. Thus he produced exhaustive studies of foliar
dichotomy ~ ~ 952), heteroblastic leaf morphologies ~ ~ 957),
paraphyses ~ ~ 964), spore structure ~ ~ 974), and vein reticu-
lation ~ ~ 979) . He compiled detailed floristic analysis of the
areas in which he worked the southern Appalachians (1963,
~ 970), Hawaii ~ ~ 999) and distributional analyses of species
and genera he studied. From the latter he became convinced
that pteridophytes, despite their ease of dispersal by spores,
for the most part showed the same distribution limitations
as seed plants (1972~. Subsequent research demonstrating
the general out-breeding nature of ferns provided the
explanation two or more spores germinating in interactive
proximity being required for sporophyte production and
thus for migration of most diploid species.
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B I O G RA P H I C A L
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Although I hac! taken Herb's course in plant systematics
and had been exposed to his philosophy for several years, I
clicin't come to appreciate his truly comprehensive knowI-
edge of pteridophytes until 1967, when Herb participated
in the offering of a fern course for graduate students in
Costa Rica. For two weeks he lecturecl ciaily, not only on the
morphology en c! systematics of tropical ferns but also on
the ecology, distribution, and occasionally the physiology
of the thousand or more species we were likely to encounter,
all seemingly without resort to notes. My feeling then en c!
now was that one couIcl hope to contend with Herb's analysis
of the big picture only with a similar comprehensive knowI-
edge of the parts.
Though a comprehensive Wagnerian treatment of the
ptericlophytes was not proclucecl cluring his lifetime, Herb's
influence on ptericlology in the last half of the twentieth
century was enormous, through his own studies en cl those
of his students en cl their students. He was coeditor of the
"Ptericlophyte " volume of The Flora of North America ~ ~ 993)
en cl author or coauthor of treatments on OphiogIossaceae,
Lycopocliaceae, Schizeaceae, Aspleniaceae, en cl Dryopteris.
At the time of his cleath Herb en c! Florence Wagner hac!
largely finished "The Ptericlophyte Flora of Hawaii" (it is
now being completed by Florence Wagner). That flora, in
its treatment of the remarkable evolutionary patterns of
Hawaiian pteridophytes, will reflect their lifetime accumu-
lation of knowledge of pteridophyte biology.
Herb hac! a passion for studying the small. In 1963 with
Aaron I. Sharp he publishecl a paper in Science describing
"a remarkably reclucecl vascular plant" the fingernail-size
gametophyte of the fern genus Vittaria. The reduction to
which the paper referred was not the size of the gametophyte
plant itself but its failure to ever produce a sporophyte, the
larger en c! more familiar phase of the fern life cycle. Though
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WARREN H. WAGNER, JR.
309
well clocumentec! in bryophytes, indefinite persistence of
the supposedly ephemeral gametophyte phase through
vegetative reproduction was an unheard of phenomenon in
ferns. Furthermore, these inclepenclent gametophytes were
very common in the southeastern Unitecl States, covering
square meters of moist cliff surfaces much the same as bryo-
phytes. The paper in its initial submission was titles! "The
Most Reclucecl Vascular Plant," but the reviewers cautioned
that still greater recluction might be found. True to this
precliction Wagner ant! Robert Eve rs shortly thereafter
clescribecl from the canyons of southern Illinois the gameto-
phyte of Trichomane~another inclepenclent gametophyte,
this one reclucec! to a mere branching filament of cells.
I arrivecl in Ann Arbor just at the time of these discoveries
en cl was fascinated to fincI, on my first trip with the Wagners
to southern Ohio en c! Kentucky, both genera of inclepenclent
gametophytes growing in luxuriant abundance. With Herb's
enthusiastic encouragement en cl my own love of exploring
cliffs en c! rockhouses, I was powerless to resist a lifelong
enchantment with the evolution en cl ecology of these plants.
The existence of inclepenclent fern gametophytes is now
well clocumentec! in North America, Hawaii, en c! Europe
en cl probably occurs worIc~wicle as a natural result of the
preaciaptation of certain tropical species for vegetative
reproduction en c! clispersal in the gametophyte stage, a habit
evolvecl to promote cross-fertilization in epiphytic habitats.
The other small plants to attract a disproportionate
amount of Herb's attention were the moonworts, diminutive
plants of the genus en cl subgenus Botrychium. Generally
less than 10 cm tall, these plants produce but one leaf per
year of very simple (reduced) morphology, usually well hidden
among associated vegetation. When Herb first turned his
attention to this group, six species were recognized worIcI-
wicle, five in North America. With Florence's expertise in
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B I O G RA P H I C A L
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cytology en c! their combiner! ability to coax hoards of students
en cl amateurs to crawl through meadow vegetation on hands
en cl knees they began detection of a much larger complex
of species than ever imaginecI.2 Their uncanny ability to
discern sub tie morphological clifferen ces reve ale cl a diversity
of cliploicI, tetraploicI, en cl hexaploicl species now totaling
303 en c! illustrating as well as any organisms the concepts of
cryptic speciation ~ ~ 983) .
Of Herb Wagner's many contributions to plant systematics,
he is most wiclely known for his early conceptual contribu-
tions to cIaclistic methods of analysis en cl representation of
phylogenetic relationships, now the method of choice for
research into evolutionary relationships among organisms.
After first conceiving en cl applying his grounciplan cliver-
gence inclex method in his dissertation work,4 Herb spent
the next two clecacles analyzing, perfecting, en c! promoting
it, while applying it to more and more complex systematic
problems (1964, 1969~. Ultimately he convinced most of
his colleagues that his objective en c! testable methods yielclec!
results more satisfying than the subjective judgments of
experts, and with their adoption by the new breed of
computer-sawy systematists, "ciaclistics" was off en c! running.
Later reflecting on the struggles of this period, he com-
mented that "most active taxonomists are so busy that they
have little time to contemplate the philosophical founcia-
tions of their calling. They are too preoccupied with the act
of classification to be burclenecl with the icleas behind it or
to devote themselves to developing a consistent theory"
(1969~.
Askocl to review his clevelopment of the grounciplan
divergence inclex ~ ~ 969, ~ 980), Herb acknowlecigec! that
no one part was new and that his thinking was initially
influenced by the writings of Benedictus Danser5 regarding
detection of the ancestral form or "grounciplan" of phylo-
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WARREN H. WAGNER, JR.
311
genetic groups. Herb's contribution was putting philosophy
and method together to yield a diagrammatic depiction of
phylogenetic relationships based on explicit data and assump-
tions. Herb's method consisted of five steps: (1) identifying
the taxa to be considered, (2) selecting characters that showed
evolutionary trends, (3) determining the ancestral state for
each character, (4) finding the degree of advancement of
each taxon, and (5) connecting taxa by their degree of
shared derived characters. Each of these steps required careful
objective analysis with no a priori assumptions. "Homology
in ~ r^=rillei^m LEA Hat ~ A~til~
`~ ~ ~~` ~~ `~ ~ .... Only trends and
patterns shown by the data themselves can be applied" (1969~.
Most basic was the use of in-group and out-group compari-
sons to objectively determine ancestral character states and
the Occam's razor principle of assuming an overall dia-
gram (tree) requiring the fewest character changes (steps)
as being the most likely. Herb's method was soon comput-
erized to produce "Wagner trees" as they became known.6 7
With many subsequent modifications and increasing sophis-
tication, Wagner trees continue to appear in systematic lit-
erature. Along with the awards for Herb's many contribu-
tions to systematic botany (Will) Hennig fellow, National
Academy of Sciences, American Academy of Arts and Sci-
ences fellow, Asa Gray Award of the American Society of Plant
Taxonomists), the Wagner tree inscription appropriately rec-
ognizes his profound influence on modern phylogenetic
reconstruction.
Herb emphasized that a major goal of his groundplan
divergence index was to teach concepts in systematic botany.
It "forces us to investigate the nature of character states
and to evaluate all of the available characters." He admon-
ished that "the systematist should not simply 'plug in' his
data set and allow the computer to come up with the
cladogram. He should think it out himself, and this, scien-
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B I O G RA P H I C A L
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tifically, may be one of the most useful rewards of following
each of the procedures of the Grounciplan-clivergence
MethocI" (1980~. He clicl more. He creates' an entirely new
family of plants, the Dencirogramaceae (also known as the
Wagneraceae), to teach the principles involvecI. The species
(illustratecl on 5 x 7 carcis) clemonstratecl evolution from
normal to fleshy stems, simple to compounc! leaves, free to
fusecl petals (or possibly the reverse of all of these) as well
as other variations. In the classroom these "plants" stimulatecl
hours of discussion (sometimes fierce arguments) over the
direction en cl pattern of their evolution en cl which was the
most parsimonious solution.
The exercise proved so effec-
tive that through the 1960s new species of Dencirogramaceae
continual to be cliscoverecl (as well as fossil ancestors). They
also reproduced vigorously and dispersed, ultimately achieving
much the same distribution as Herb's students en c! grancI-
stuclents. Publications appeared analyzing their systematic
relationships using an array of computerized methods. They
became as well recognizes! en c! as important in systematic
lore as real plant families.
Such was Herb Wagner's talent for getting students
immerses! in systematics en c! plant science in general. He
hacl little sympathy for those who complainecl of the cliffi-
culties of academia or who clicl not pursue their studies
with a strong, honest effort. For students clisplaying genuine
interest in their research discoveries he quickly multipliecl
that interest through his own. His clear excitement over
discoveries large and small was the genius of his inspira-
tional leaclership. He couIcl make hard work not only palatable
but also fun. My recollection of lunchtime discussions among
Herb en c! us students is that always there was the sense of
examining breaking news at the forefront of scientific
discovery. Importantly, it was the science behind those clis-
coveries, rather than the people, that was the focus. He
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WARREN H. WAGNER, JR.
313
cultivates! the attitude that all research was worthwhile en c!
that the goal was advancement of knowlecige, not personal
glory. He applied this philosophy to encourage reluctant
students to publish their work, saying that they "owes! it to
science" to communicate their finclings. This particular ploy
workocl to keep me in school when I was contemplating
taking time out for a stint in the Peace Corps.
Herb received well-cleservecl awards for en cl acknowlecige-
ments of his gift for teaching both insicle en cl outside the
classroom, but his influence certainly was not limiter! to the
classroom. In aciclition to numerous research fielcl trips,
Herb selclom macle a seminar visit to a new or botanically
interesting area without insisting on an accompanying field!
trip. These trips invariably incluclecl a retinue of local amateur
botanists as well as students en cl academic professionals.
From their "Wagner experience" huncirecis of students, pro-
fessionals, en cl amateurs became hookocl on science, not
because they wan tell to please Herb, although that was always
fun, but because they became genuinely infuser! with the
excitement of scientific discovery. Herb's ability to inspire
others through his interest in their studies en cl their knowI-
ecige not only fosterec! inclepenclent research but also creates!
a legion of professionals en cl amateurs eager to contribute
ciata to Herb's projects as well. The total productivity of
this synergism, though unquantifiable, remains hugely visible.
Herb's clistinguishecl career at the University of Michigan
incluclecl chairmanship or cochairmanship of over 45 cloctoral
committees en c! membership on more than 235. He taught
a variety of courses, inclucling systematic botany en cl biology
of woolly plants, both of which he continual to co-teach
after "retirement" in 1991 through the fall of 1999. Teach-
ing was as much a joy to Herb as it was to the students who
continued to pack his courses. His outrageous performances
en c! exaggerations clelightec! his audiences. It was always of
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B I O G RA P H I C A L
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great interest to his teaching assistants to see who among
the students clicl en cl who clicin't believe that Raffiesia was
pollinatecl by elephants, Wolffia by mosquitoes, en cl Poclo-
phy]]um by turtles. Herb's public lectures en c! seminars were
equally popular. Few biologists have been in such demand
as a visiting speaker. His curriculum vitae list of invited
lectures totaTec! ~ 69 after retirement!
Warren H. Wagner, fir., wall be remembered as a wonclerful
teacher en cl inspirational leacler whose legacy lives on in
huncirecis of inclivicluals whose lives he touched. His com-
mancl of the principal subjects of his research, his belovecl
ferns, was excellecl by none. He usecl intimate knowlecige of
cletaiT to synthesize big-picture principles that withstood! the
scrutiny his flamboyant style invited. His contribution to
plant systematics en cl evolution en cl to the biology of ferns
profoundly influencer! the direction of these fielcis into the
twenty-first century.
Aciclitional biographic information on W. H. Wagner,
Jr., with more complete bibliographies has appeared! in
obituary publications in Taxon (49 E2000] :585-592) and
American Fern Journal 92~20001:39-49~. The photograph
en c! information on early years were graciously proviclec! by
Florence Wagner. Factual information is taken from Herb
Wagner's 1999 curriculum vitae. Other anecdotes en cl observa-
tions extent! from my Tong association with the Wagners, as
a graduate student in Ann Arbor en cl in many subsequent
fielcl trips en cl discussions of plants, people, en cl philosophy.
NOTES
1. W. H. Wagner. 1996. Flying flowers! Butterflies and their
foodplants. LSA Bulletin (published by the College of Literature,
Science, and the Arts, The University of Michigan) 9:4-9, cover.
2. W. H. Wagner and F. S. Wagner. 1998. Moonwort madness: A
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WARREN H. WAGNER, JR.
315
reply. Am. Fern Soc. Bull. (Fiddlehead ForumJ 25:30-31. The Wagners
recount how they became interested in Botrychium and some of their
adventures in searching for these reclusive plants.
3. This number includes several species not yet officially published.
4. W. H. Wagner, Tr. The fern genus Diellia: Structure, affinities,
and taxonomy. Univ. Calif Publ. Bot. 26 ~ 1 ~ ~ 1952~: 1-212.
5. B. H. Danser. A theory of systematics. Bibl. Biotheoret. 4~1950) :1-20.
6. A. G. Kluge and T. S. Farris. Quantitative phyletics and the
evolution of anurans. Syst. Zool. 18 ~ 1969~: 1-32.
7. T. S. Farris. Methods for computing Wagner trees. Syst. Zool.
19 (1970) :83-92.
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B I O G RA P H I C A L
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B I B L I O G RAP H Y
1952
Types of foliar dichotomy in living ferns. Am. f. Bot. 39:578-92.
1954
Reticulate evolution in the Appalachian Aspleniums. Evolution 8~2~:103-18.
1955
Cytotaxonomic observations on North American ferns. Rhodora 57:219-40.
1956
The morphological and cytological distinctness of Botrychium minganense
and B. Iunaria in Michigan. Torrey Bot. Club Bull. 83:261-80.
1957
Heteroblastic leaf morphology in juvenile plants of Dicranopteris linearis
Gleicheniaceae ~ . Phytomorphology 7 :1-6.
1963
With A. J. Sharp. A remarkably reduced vascular plant in the United
States. Science 142:1483-84.
Pteridophytes of the Mountain Lake Area, Giles Co., Virginia, including
notes from Whitetop Mountain. Castanea 28:1 13-50.
1964
Paraphyses: Filicineae. Taxon 13 (2) :56-64.
The evolutionary patterns of living ferns. Torrey Bot. Club Bull. 21:86-95.
1968
Hybridization, taxonomy, and evolution. In Modern Methods in Plant
Taxonomy, ed. V. H. Heywood, pp. 113-38. New York: Academic
Press.
1969
The construction of a classification. In Systematic Biology, pp. 67-90.
Publication number 1692. Washington, D.C.: National Academy
of Sciences.
OCR for page 317
WARREN H. WAGNER, JR.
317
The role and taxonomic treatment of hybrids. BioScience 19:785-89,
795.
1970
Biosystematics and evolutionary noise. Taxon 19:146-51
Evolution of Dryopteris in relation to the Appalachians. In The Distri-
butional History of the Biota of the Southern Appalachians, ed. P. C.
Holt, pp. 147-92. Part II: Flora, vol. 2. Blacksburg, Va.: Virginia
Polytechnic Institute and Southern University Research Division.
With D. R. Farrar and B. W. McAlpin. Pteridology of the highlands
biological station area, southern Appalachians. 7. Elisha Mitchell
Sci. Soc. 86:1-27.
1972
Disjunctions in homosporous vascular plants. Ann. Mo. Bot. Gard.
59~2) :203-17.
1974
Structure of spores in relation to fern phylogeny. Ann. Mo. Bot.
Gard. 61 (2) :332-53.
25 years of botany. Ann. Mo. Bot. Gard. 61 ~ 1 ): 1-2.
1979
Reticulate veins in the systematics of modern ferns. (In Rogers McVaugh
Festschrift) . Taxon 28 ~ 1,2,3) :87-95.
1980
With F. S. Wagner. Polyploidy in pteridophytes. In Polyploidy. Biological
Relevance, ed. W. H. Lewis, pp. 199-214. New York: Plenum Press.
Origin and philosophy of the groundplan-divergence method of
cladistics. Syst. Bot. 5 (2) :173-93.
1983
Reticulistics: The recognition of hybrids and their role in cladistics
and classification. In Advances in Cladistics, vol. 2, eds. N. I. Platnick
and V. A. Funk, pp. 63-79. New York: Columbia University Press.
With F. S. Wagner. Genus communities as a systematic tool in the
study of New World Botrychium (Ophioglossaceae) . Taxon 32:51-63.
OCR for page 318
318
BIOGRAPHICAL MEMOIRS
1984
Applications of the concepts of groundplan-divergence. In Cladis-
tics: Perspectives on the Reconstruction of Evolutionary History, eds. T.
Duncan and T. F. Steussy, pp 95-118. New York: Columbia University
Press.
1993
With A. R. Smith. Pteridophytes of North America. In Flora of North
America North of Mexico, vol. 1, eds. Flora of North America Edito-
rial Committee, pp. 247-56. New York: Oxford University Press.
1 995
Evolution of Hawaiian ferns and fern allies in relation to their con-
servation status. Pac. Sci. 49:31-41.
1 999
With D. D. Palmer and R. W. Hobdy. Taxonomic notes on the
pteridophytes of Hawaii. II. Contrib. Univ. Mich. Herb. 22:135-87.
OCR for page 319
Representative terms from entire chapter:
species hybrids
