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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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302 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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304 B I O G RA P H I C A L EMOIRS 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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306 B I O G RA P H I C A L EMOIRS 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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308 B I O G RA P H I C A L EMOIRS 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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310 B I O G RA P H I C A L EMOIRS 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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312 B I O G RA P H I C A L EMOIRS 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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314 B I O G RA P H I C A L EMOIRS 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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316 B I O G RA P H I C A L S E L E C T E D EMOIRS 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.

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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.

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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.

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