AMERICAN Volume 92 FERN Number J O U R N A q January—March 2002
QUARTERLY JOURNAL OF THE AMERICAN FERN SOCIETY
Obituary: Rolla Milton Tryon, Jr. (1916-2001) Gerald J. Gastony, David S. Barrington, and David S. Conant 1
a Associated with the Intertracheid Pit seas) of the Woody Fern Botrychium multifidum ela C. Morrow and Roland R. Dute 10
A New Filmy Fern from the Dominican Republic Carlos Sanchez 20
Adiantum argutum, an Unrecognized Species of the A. latifolium Group Jefferson Prado and David B. Lellinger 23
Polypodium cseapage Plants Sporulate viacpecsndaeed in a Non-seasonal Glasshouse Envi- ronmen a E. Simdn and Elizabeth Sheffield 30
The American Fern Society
Council for 2002
CHRISTOPHER H. HAUFLER, Dept. of Botany, University of Kansas, Lawrence, KS 66045-2016. President TOM RANKER, University Museum, Campus Box 265, University of Colorado, Boulder, art eu ‘65. President W. CARL TAYLOR, 800 W. Wells St., Milwaukee Public Museum, Milwaukee, WI eg 1478. Secretary JAMES D. CAPONETTI, Dept. of Botany, University of Tennessee, Knoxville, TN 37916-1110. Treasurer GEORGE YATSKIEVYCH, Missouri Botanical Garden, P. O. Box 200, St. oe MO 63166-0299. rship Secretary JAMES D. MONTGOMERY, Ecology HI, R.D. 1, Box 1795, Berwick, PA 18603-9801. Back — Curator
R. JAMES HICKEY, Botany Dept., Miami University, Oxford, OH 45056. rnal Editor DAVID B. LELLINGER, U.S. National Herbarium MRC-166
Smithsonian Institution, Washington, DC 20560-0166. Memoir Editor CINDY JOHNSON-GROH, Dept. ‘of pom — Adolphus College,
800 W. College Ave., St. Peter, MN 56082 Bulletin Editor
American Fern Journal EDITOR
R. JAMES HICKEY Botany De ami University, Oxford, OH 45056 ph. (513) £5 Sone e-mail: hickeyrj @ muohio.edu
ASSOCIATE EDITORS GERALD J. GASTONY .............. Dept. of Biology, Indiana University, ages aig IN 47405-6801 CHRISTOPHER H. HAUFLER ...... Dept. of Botan get University of Kansa 66045-2106 ROBBIN C. MORAN ew York Botanical Gecten: Bike NY 10458-5126 JAMES H. PECK ser of — University of Arkansas—Little Rock, 2801 S. University Ave., Litthe Rock, AR 72204
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Tnonin lA hy - > thn < me 4) te OCCretary
American Fern Journal 92(1):1—9 (2002)
MISSoy
Ay Obituary: Rolla Milton Tryon, Jr. g4p 1 4 2099 (1916-2001) DEN LiBRap
GERALD J. GASTONY Department of Biology, Indiana University, Bloomington, IN 47405-3700
Davip S. BARRINGTON Department of Botany, University of Vermont, Burlington, VT 05404-0086
Davip S. CONANT
Department of Natural Sciences, Lyndon State College, Lyndonville, VT 05851
Rolla Tryon, a member of the American Fern Society since 1932 and one of the twentieth century’s most eminent students of pteridophytes, was born on August 26, 1916 in Chicago, Illinois. His father, a professor of American history and education at the University of Chicago, maintained a summer cottage in Chesterton, Indiana in addition to his home in Chicago. Rolla’s fascination with ferns and fern allies developed during boyhood forays from that Ches-
The photograph was taken by Dr. Walter H. Hodge in Mexico City in December, 1972 and was made available by the Hunt Institute for Botanical Documentation.
2 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
terton cottage into sand dune habitats along Lake Michigan in the northwest of Indiana. At the age of 18, he published his first paper, relating his obser- vations on Osmunda plants in the Indiana Dunes (see complete bibliography below). As a boy, Rolla was greatly influenced by, and in turn influenced, Charles Deam (author of the 1940 Flora of Indiana), advising Deam about fern species he had found in the dunes area. When a doubting Deam appeared at the cottage door one day asking to meet Rolla and to be shown these ferns in situ, he was surprised to learn that Rolla was not the adult of the family but a mere boy of 14. Thus began a productive friendship documented in Rolla’s correspondence with Deam from May, 1935 to January, 1953. All the penny postcards and letters he received from Deam have been carefully maintained in one of Rolla’s files, now archived at Indiana University—fascinating reading. Rolla’s insatiable boyhood appetite for ferns got him into a bit of trouble at home, however, when his father learned that he had charged Bower's three volumes on The Ferns to his account at Brentano’s bookstore in Chicago.
Rolla built a solid academic superstructure on the foundation of these boy- hood experiences. Among his academic accomplishments were an A.A. degree in 1935 and a B.S. degree in 1937, both from the University of Chicago, and a Ph.M. in 1938 from the University of Wisconsin. In 1940 he earned an MS. and in 1941 a Ph.D., both from Harvard University. During his days as a Har- vard student, he contracted malaria in South Carolina while collecting plants for M. L. Fernald, and during the war-torn year following completion of his Ph.D. he served as a lab technician in the U. S. Chemical Warfare Service at Massachusetts Institute of Technology. His father thought he should follow his Ph.D. in botany with another, this time in chemistry, so that he could earn a living, but instead Rolla became an Instructor in Botany first at Dartmouth College, then at the University of Wisconsin before becoming an Assistant Professor in Botany at the University of Minnesota in 1945. While an Assistant Professor at the University of Wisconsin, Rolla met Alice Faber. Their marriage in 1945 initiated not only a happy and enduring domestic partnership but also a research synergism whose productivity has nourished pteridologists through- out the world.
In 1947 Rolla became Associate Professor in Botany at Washington Univer- sity, St. Louis and Assistant Curator of the herbarium of the Missouri Botanical Garden, positions he held to 1957. During this appointment, he and Alice were the original organizers of the Missouri Botanical Garden’s annual Systematics Symposium, whose 48th meeting was held 12-13 October, 2001. This highly successful annual meeting has received continuous support from the National Science Foundation from its second year (1954) to the present (with the lapse of a single year). From 1946 to 1957 Rolla served as curator and librarian of the American Fern Society’s library and herbarium, responding to members’ requests for loans of materials. That herbarium and library was subsequently entrusted to Warren H. Wagner at the University of Michigan. Following a year as Research Associate at the University of California, Berkeley, Rolla went to the Gray Herbarium of Harvard University as Associate Curator and Curator of Ferns in 1958 and became Curator of the Gray Herbarium in 1967. Rolla
ROLLA MILTON TRYON, JR. 3
and Alice traveled the world extensively, attending international meetings, conducting field work, studying specimens at major herbaria in the Americas, Europe, and Africa, and conducting field courses on the ferns. In addition to other services to professional societies, Rolla served for many years as Asso- ciate Editor of Rhodora and the American Fern Journal, as Associate Editor of Brittonia (1961-1964), as Editor-in-Chief of Rhodora (1977-1982), and as Pres- ident of the New England Botanical Club and the American Fern Society.
A framed photograph of his revered mentor, Charles A. Weatherby (see American Fern Journal 40[1] for a remarkable series of papers honoring this unusually respected and beloved botanist), was always prominently displayed on Rolla’s desk at Harvard, undoubtedly inspiring his own welcoming, patient, and supportive response to all who entered his office seeking counsel. In 1970 Rolla initiated an annual New England Fern Conference at Harvard Forest. For 20 years this provided a stimulating intellectual setting in which students of fern biology discussed and developed their ideas. In 1972 he became Professor of Biology at Harvard University, holding both the Curatorship and Professor- ship until his retirement in 1987. He remained at Harvard as Professor Emer- itus from 1987 to 1989 when he moved to the University of South Florida in Tampa as Adjunct Professor, bringing with him his extensive library of fern and biogeographic literature. To mark the retirements of Alice and Rolla Tryon from Harvard, a festschrift of 13 papers plus introduction was published in their honor in the Annals of the Missouri Botanical Garden (vol. 77: 225-339. 1990).
At the University of South Florida, Rolla and Alice helped found the Insti- tute for Systematic Botany and endowed the Tryon Lecture Series that brings several internationally known botanists to the university each year. In their research-active office on the Tampa campus, he and Alice continued their pter- idological work, as his following bibliography indicates.
Rolla Tryon’s publication list exceeds 100 titles and includes a great breadth of topics. Papers ranged from articles on pteridophytes for the 1943 Encyclo- pedia Britannica to a glossary of terms relating to the fern leaf, discussions of the history of pteridology and fern classification, a remembrance of his grad- uate mentor and counselor Charles A. Weatherby, discussions of the formali- ties of fern nomenclature, and many book reviews. His monographs and re- visions focused mostly on ferns but also included angiosperms Convolvulus and Elymus. Signal among these were his revisions of Pteridium, Doryopteris, the Selaginella rupestris group, American Notholaena, and the Cyatheaceae. His papers on fern biogeography began with Doryopteris in 1944, matured in his exposition of geographic speciation in Selaginella in 1971, and continued to his and Alice’s 1999 discussion of the phytogeography of eastern North American ferns (honoring Ching Ren-Chang). Floristic and taxonomic notes on ferns ranged from simple observations of growth forms and hybrids to eluci- dations of complex taxonomic and nomenclatural issues. For his 1955 publi- cation on the taxonomy of cycads (coauthored with students in his Washington University class) he was awarded the 1956 Robert Montgomery award of the Fairchild Tropical Garden for distinguished achievement in the world of palms
4 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
and cycads. At the time of his death, he had a book review in press in Rhodora and a paper in press in Bradea (coauthored with his former student Paulo Windisch).
In addition to his numerous papers on ferns and other topics, Rolla is no- table for his books. Among these are two editions of his Ferns and Fern Allies of Wisconsin (1940, 1953) and Ferns and Fern Allies of Minnesota (1954, 1980). He is renowned for his knowledge of the ferns of Peru, first expressed in his 1964 Ferns of Peru (250 pages in the Contributions from the Gray Her- barium of Harvard University). This treatment was updated and completed in six subsequent parts entitled Pteridophyta of Peru between 1989 and 1994, mostly coauthored with Robert Stolze but with several portions contributed by other pteridological specialists. His monumental 1982 book Ferns and AIl- lied Plants with Special Reference to Tropical America, coauthored with Alice Tryon, is an encyclopedic treatment of this subject that continues to stimulate new research, as does his treatment of Pteridaceae, with Alice Tryon and Karl Kramer, in volume 1 of Families and Genera of Vascular Plants edited by K. Kramer and P. S. Green.
Rolla’s kindly and perceptive mentoring and his outstanding contributions to our knowledge of ferns is signaled by having the following four fern taxa named in his honor.
1) Asplenium tryonii Correll. In describing this species, Donovan Correll (1961) said “It is a pleasure to name this species for Dr. Tryon, who has always been most gracious in helping his fellow-workers with their never- ending problems in the study of ferns.” Known only from Chihuahua, Mex- ico, this species was further discussed and illustrated in Ferns and Fern Allies of Chihuahua by Knobloch and Correll (1962).
Alsophila tryonorum Riba. The eminent Mexican pteridologist Ram6n Riba (see American Fern Journal 90:112—118, 2000) stated that “this species is named after Dr. Rolla M. Tryon and Dr. Alice F. Tryon for their contributions to the taxonomy of the ferns” (Riba, 1967). The plural specific epithet rec- ognizes the close professional relationship between this highly productive research team. This tree fern species is now known as Trichipteris tryono- rum (Riba) R. Tryon following its transfer by Rolla in his 1970 paper on the classification of the Cyatheaceae.
Nephelea tryoniana Gastony. ‘“‘I am pleased to name this species for my mentor, Dr. Rolla M. Tryon, in recognition of his outstanding contribution to the understanding of the systematics and evolution of the family Cy- atheaceae” (Gastony, 1973). Subsequent research by Conant (Conant and Cooper-Driver, 1980; Conant, 1983) revealed that this tree fern species is a reproductively stabilized diploid hybrid species that is now regarded as Alsophila tryoniana (Gastony) Conant.
Tryonella Pichi Sermolli. This new generic name was established by Pichi Sermolli (1974) “in honour of the eminent pteridologist R. M. Tryon, Jr., author of many important papers on ferns, who, inter alia, supported the distinction of the present genus from Doryopteris, though without giving it
i) —
ioe) —
ns —
ROLLA MILTON TRYON, JR. 5
anew name.” This name is currently regarded as a synonym of Doryopteris by Tryon and Tryon (1982) and Tryon, Tryon, and Kramer (1990).
Among the doctoral graduate students he trained, Rolla counted the follow- ing (those with asterisks received their degrees from other institutions): Alice F. Tryon, *Karl Kramer, *Ramon Riba, Gerald Gastony, Lawrence Palkovic, David Barrington, David Conant, Paulo Windisch, *R. James Hickey, *Robbin Moran, Sonia Sultan, and Calvin Sperling. He also mentored Robert Stolze in his taxonomic revision of Cnemidaria at the Field Museum. Always available to his students, he modeled his supportive and insightful mentoring on his experiences with his own graduate mentor, Charles Weatherby. For this he has earned our love as well as our respect. His impact on his students, and their students, and their students is incalculable.
In 1978 Rolla M. Tryon, Jr. was elected to honorary membership in the Amer- ican Fern Society, a special category of membership for persons who have made outstanding contributions to the study of ferns. In 1984 he received a Merit Award from the Botanical Society of America “In recognition of distin- guished achievement in and contributions to the advancement of botanical science. Pre-eminently knowledgeable in matters of taxonomy and nomencla- ture, this foremost pteridologist is a perceptive student of phytogeography and of the evolutionary impact of the selective process during plant migration.”
In addition to Rolla’s botanical activities he was also highly skilled in run- ning a family farm in Knox County, Indiana for many years. He visited the farm, overseeing its management, a few times each year. The extensive records he kept in managing crops and livestock illustrate his practical ability in man- aging business as well as scientific data.
On August 20, 2001, six days before his eighty-fifth birthday, Rolla Milton Tryon, Jr., left us to continue our work with the pteridophytes of the world, and to delight in them, without him. We do this fortified by his writings, the echoes of his encouraging words, and his everlasting example. He was the beloved husband of Alice Faber Tryon, the benefactor of countless students of pteridophytes, including many who never knew him personally, an inspiration and counselor to many collaborators and coauthors, the advisor of doctoral students, the teacher of innumerable undergraduates, and our dear friend and mentor. He will be deeply missed. He already is.
LITERATURE CITED
CONANT, D. S. 1983. sis revision of the genus Alsophila (Cyatheaceae) in the Americas. J. Arnold Arbor. 64: 333-—
Conant, D. S. and : ones DRIVER. 1980. Autogamous allohomoploidy in Alsophila and Ne-
phelea (Cyatheaceae): a new hypothesis for speciation in homoploid homosporous ferns.
mer. J. Bot. 67: 1269-1288.
CorreLL, D. S. 1961. Two Texas-Chihuahuan ferns. Wrightia 2: 200-203.
— G. J. 1973. A revision of the fern genus Nephelea. Contr. Gray Herb. 203: 81—
eure _ I. W. and D. S. CorreLL. 1962. Ferns and fern allies of Chihuahua, pace —
a Foundation, Renner, Tex PICHI pameene R. E. G. 1974. seit a Webbia 29: 1-16.
6 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
Risa, R. 1967. New taxa in the genus Alsophila. Rhodora 69: 65-68.
BIBLIOGRAPHY OF ROLLA M. TRYON, JR. (1916-2001)
TRYON, R. M., JR. 1934. Some observations on Osmundas. Amer. Fern J. 24:
TRYON, R. M., JR. 1936. Ferns of the dune region of Indiana. Amer. Mid. en 17: ite
TRYON, R. M., JR. 1936. Botrychium dissecturn and forma obliquum. Amer. sg n J.
TRYON, R. M., JR. 1938. The phenomenon of forking in ferns. Amer. Fern J. 2
TRYON, R. M., JR. 1938. Recent additions to the flora of Indiana. Proc. “la poe Sci. 47: 76— 17
TRYON, R. M., JR. 1939. Notes on the ferns of Wisconsin. Amer. Fern J. 2 9, TRYON, R. M., JR. 1939. The varieties of Convolvulus spithamaeus and a C. sepium. Rhodora 41: 5—423.
TRYON, R. M., JR. 1940. Notes on some Indiana plants. Proc. Indiana Acad. Sci. 49: 89-90.
TRYON, R. M., JR., N. C. Fassett, D. W. DUNLOP, and M. E. 6 EMER. 1940. The ferns and fern allies of Sip — Dept., Univ. of Wisconsin, Madiso
TRYON, R. M., JR. 0. An Osmunda hybrid. Amer. Fern J. 30: 65-66.
TRYON, R. M., JR. 1 mi a revision of the genus Pteridium. Bhai 43: 1-31, 37-67. Reprinted as Contr. Gray Herb. 1 -7
bia R. M., JR. 1942. ae Roland, A. E., The ferns of Nova Scotia. Amer. Fern J. 32: 73—
Perey R. M., JR. 1942. A new Dryopteris hybrid. Amer. Fern J. 32: 81—
TRYON, R. M., JR. 1942. A revision of the genus Doryopteris. Contr. G ae 80. TRYON, R. M., JR. 1942. Review of Brown, C. A. and D. S. Correll, The ferns ak 7e allies of Louisiana. Rhodora 44: 484—485.
sia TRYON, R. M., JR. 1943. Several articles on Pteridophyta for Encyclopedia Brittanic TRYON, R. M., JR. ma Abstracts of the American Fern Jour nal for Biological Abstracts.
TRYON, R. M., JR. and J. W. Moore. . Notes on aquatic and prairie acca in ane i 6. TRYON, R. M., JR. and J. W. Moore. 1946. A preliminary checklist of the flowering plants, ferns and fern allies of Minnesota. Univ. of Minnesota, Minneapolis. TRYON, R. M., JR. 1946. A new Doryopteris hybrid. Amer. Fern J. 36: 4 OorE, J. W. and R. M. TRYON, JR. 1946. A new record of Isoétes inalsinpod. Amer. Fern J. 36: 89-91 vista F, K and R. M. TRYON, JR. 1948. A fertile mutant of a Woodsia hybrid. Amer. Fern J. 35: 13 TRYON, M., JR. 1948. Some Woodsias from the north shore of Lake Superior. Amer. Fern J. 38: 159-170. Boouer, L. E. and R. M. TRYON, JR. 1948. A study of Elymus in Minnesota. Rhodora 50: 80-91. TRYON, R. M., JR. 1950. Charles Alfred Weatherby—teacher and counselor. Amer. Fern J. 40: 9-10. TRYON, R. M., JR. 1950. A new erect species of the Selaginella rupestris group. Amer. Fern J. 40:
69-74. TRYON, R. M., JR. — Ferns of the Missouri Ozark region. Missouri Bot. Gard. Bull. 39: 136-138.
TRYON, R. M., JR. 1951. Review of Manton, I., Problems of cytology and evolution in the Pterido- phyta. Ecology 32: 769-770. TRYON, R. M., JR. 1952. A sketch of the history of fern classification. Ann. Missouri. Bot. Gard. 39:
6 TRYON, R. M., JR., N. C. Fassett, D. W. DUNLOP and M. E. DiEMER. 1953. The ferns and fern allies of Wisconsin, Ed. 2. Univ. Wisconsin Press, Madison. TRYON, R. M., JR. 1954. The ferns and fern allies of Minnesota. Univ. Minnesota Press, Minneap- olis
TRYON, R. M., JR, E. BARBOUR, E. Davis, H. Kipp, R. Lonc, C. Marvin, B. MIKULA, and R. MOHLEN- BROCK. 1955. Ancient seed plants: the cycads. Missouri. Bot. Gard. Bull. 43: 65-80.
ROLLA MILTON TRYON, JR.
TRYON, iy M., re ney ep ae ager vee its allies. Ann. Missouri. Bot. Gard. 42: 1-99. KRAME N, JR. 1955.
eeu. new species of Doryopteris from Surinam. Ann. Missouri Bot a < _ ner oe Sti . — JR. 1956. A revision of the American species of Notholaena. Contr. Gray Herb. 179.
aa. . a JR. 1957. Adianturn in Peru: new species and combinations. Amer. Fern J. 47: 139— ah
KosuskI, C. E., C. V. MorTON, M. Ownsey, and R. M. TRYON. 1958. Report of the committee for recommendations on desirable procedures in herbarium practice and ethics. Brittonia 10: 93-95.
TRYON, R. M., JR. and A. F. TRYON. 1959. Observations on cultivated ferns: the hardy species of tree ferns (cha and Cyatheaceae). Amer. Fern. J. 49: 129-142. Reprinted in Lasca Leaves 10: 26-33. 1960.
TRYON, R. M., Ir. 1960. The ecology of Peruvian ferns. Amer. Fern J. 50: 4 TRYON, R. 1960. A glossary of some terms relating to the en — Taxon c 108-109, Reprinted
in Russian iio in Bot. Jour. Akad. Nauk, U.S.S 736-739. TRYON, M., JR. 1960. Review of Pteridophyta in Munz, P. a o balou rene Amer. Fern J. 50: seit
TRYON, R M. JR. 1960. A review of the genus Dennstaedtia in America. Contr. Gray Herb. 187:
23—
TRYON, * 4 JR. 1960. New species of ferns from serait mei South America. Rhodora 62: 1-10.
TRYON, R. M., JR. 1961. Taxonomic fern notes. I. Rhodora 63: 7-88
TRYON, R. M., JR. 1962. The fern genus Doryopteris in aa ee and Rio Grande do Sul, Brazil. Sellowia 14: 51-59
TRYON, R. M., JR. 1962, Review af Wherry, E. T., The Fern Guide. Amer. Fern J. 52: 89-91.
TRYON, R. M., JR. 1962. A note on Nephrolepis pa ie cv. Duffii. Amer. Fern J. = 153-155.
TRYON, R. M., JR. 1962. Taxonomic Fern Notes. II. Pityrogramma (including Trismeria) and Ano- gramma. Contr. Gray Herb. 189. 52-76.
TRYON, R. 1962. Taxonomic fern notes. III. Contr. Gray Herb. 191: 91—
TRYON, R. M., JR. 1962. Review of Knobloch, I. W. and D. S. Sessil ow and Fern Allies of Chihuahua. Rhodora 64: 347-348
TRYON, R. 1962. A commentary o axpeetiudus names. Taxon 11: 116—120.
TRYON, R. 1963. Nomenclatural er a Taxon 12: 28-285.
TRYON, R. M. 1964. Evolution in the leaf of living ferns. Mem. Torrey Bot. Club 21: 73-85.
TRYON, R. 1964. erie ferns of Peru. Polypodiaceae (Dennstaedtieae to Oleandreae). Contr. Gray Herb. 1
194: 1-— TRYON, R. 1964. rite ie Fern Notes IV. Rhodora 66: 110—117. PICHI-SERMOLLI, R. E. G., F. BALLARD, R. E. HOLTTuM, H. IT, F. M. JARRETT, A. C. JERMY, E. A. C. L. E. SCHELPE, M.-L. TARDIEU-BLOT, and R. M. TRYON. 1965. Index Filicum Supplementum
Quartum pro Annis 1934-1960. Internat. Bureau for Plant Taxonomy and Nomenclature, Utrecht, Netherlands.
TRYON, R. 1965. rig. te in the ferns. Taxon 14: 213-218.
TRYON, R. and D. M. BRITTON. 1966. A study of variation in the cytotypes of Dryopteris spinul Rhodora 68: 59-92.
TRYON, R. 1967. Taxonomic fern notes V. New combinations in Peruvian species of Thelypteris.
odora 69: 5- TRYON, R. 1967. paviine of Hara, H., The flora of eastern Himalaya. Rhodora 69: 456-457. Boum, B. A. and R. M. TRYON. 1967. Phenolic compounds in ferns. I. A survey of some ferns for
c TRYON, R. M. and A. F. TRYON. 1968. Edith Scamman kage 1967). Amer. Feri 5 58: 1-4. TRYON, R. 1968. Nomenclatural proposals. Taxon 17: 588-590. TRYON, R. 1969. Pteridology, Pp. 97-102 in J. Ewan (ed.) A short history of botany in the United States. sone Publishing Co., New York. TRYON, R. 1969. Taxonomic problems in the geography of North American ferns. BioScience 19: iat
8 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
paige R. 1970. aes of Correll, D. S. and M. C. Johnston., Manual of the vascular plants of xas. Rhodora 72: 533-535. Pines R. 1970. The le of the Cyatheaceae. Contr. Gray Herb. 200: 3-53. TrYON, R. 1970. Development and evolution of fern floras of oceanic islands. Biotropica 2: 76-84. Reprinted 1971 as Pp. 54-62 in W. L. Stern (ed.) Adaptive aspects of insular evolution.
Wash
TRYON, R. 1971. The American at ferns allied to i trata seeps ang 73: 1-19.
TRYON, R. 1971. Ferns of the Andes and Amazon. Morris Arbor. Bull. 2
TrYON, R. 1971. The process of evolutionary migration in species oe sae Brittonia 23: 89— 100.
TRYON, : 1972. Endemic areas and geographic speciation in tropical American ferns. Biotropica 4: 121-131.
TRYON, s 1972. Taxonomic fern notes, VI—New species of American Cyatheaceae. Rhodora 74:
TRYON, R. M., Jr. and A. F. TRYON. 1973. spac 7 - evolutionary relations in the
cheilanthoid ferns. Bot. J. Linn. Soc. 67: Suppl. 1: TRYON, A. and R. TRYON. 1973. Thelypteris in se ae ee America. Amer. Fern J. 63: 65— 76
TRYON, R., B. VOELLER, A. TRYON, and R. RIBA. 1973. Fern biology in Mexico (a cl field program). BioScience 23: 28-33.
TRYON, R., C.-J. WIDEN, A. HUHTIKANGAS, and M. LOUNASMAA. 1973. pony nana derivatives in Dryopteris parallelogram and D. patula. Phytochemistry 12:
TRYON, A. and R. TRY 974. Ge ogaph — in temperate rence fant and some rela- tionships in Galas Amer. Fern. J. 64: 99-104.
TRYON, R. 1975. The Benjamin D. G Gilbert fern nea Amer. Fern J. 65: 60. TRYON, R. 7 and G. J. GAsToNny. 1975. The biogeography of endemism in the Cyatheaceae. Fern
Gaz 573-79. TRYON, R. 4 and D. S. CONANT. 1975. The ferns of Brazilian Amazonia. re paiitias 5: 23-34. TRYON, R. 1976. A oa of the genus Cyathea. Contr. Gray Herb. 206: GASTONY, G. J. a . M. TRYON. 1976. Spore morphology in the Cyatheacoe Il. nae genera Lophosoria, soe Sphaeropteris, Alsophila, and Nephelea. Amer. J. Bot. 63: 738-758.
aug R. M. 1977. Studies on the American Cyatheaceae and on the pie ees of on family. Males. Bull. 30: 2839—
ete R M. and G. VITALE. 1977. dua for antheridogen production and its mediation of a mating system in natural populations of fern gametophytes. Bot. J. Linn. Soc. 74: 243-249.
TRYON, R. 1979. Biogeography of the Antillean fern flora. Pp. 55-68 in D. Bramwell (ed.) Plants and islands. Academic Press, New York.
TRYON, A. F., R. TRYON, and F. BADRE. 1980. Classification, spores, and nomenclature of the marsh fern. Rhodora 82: 461-474
TRYON, R. 1980. Review of Sepensnck, J. A. and O. Huber, Flora of Avila. Amer. Fern J. 70: 79.
Tryon, R. M., JR. 1980. The ferns and fern allies of Minnesota, Ed. 2. Univ. Minnesota Press, Minneapolis.
wer - ft A. TRYON. 1981. Taxonomic and nomenclatural notes on ferns. Rhodora 83: 133-—
wee - 1982. — “e a A. R., Pteridophytes in D. E. Breedlove (ed.) Flora of Chiapas Part 2. Madrofio TRYON, R. pane ALF. aly a Additional taxonomic and nomenclatural notes on ferns. Rho-
TRYON, R. M. and A. F. TRYON. 1982. Ferns and allied plants with special reference to tropical America. Springer-Verlag, New
TRYON, R. 1984. An unusual new Bapheeyplesenctin from Peru. Amer. Fern J. 74: 108-110.
TRYON, R. 1985. Fern speciation and biogeography. Proc. Roy. Soc. Edinburgh, B. 86: 353-360.
TRYON, R. 1986. Dicksoniaceae. Lophosoriaceae. Metaxyaceae. Cyatheaceae. Pp. 1-59 in G. Harling and L. Andersson (eds.) Flora of Ecuador, No. 27. Swedish Research Councils, Stockholm.
TRYON, R. 1986. Some new names and combinations in Pteridaceae. Amer. Fern J. 76: 184—186.
ROLLA MILTON TRYON, JR. 9
TRYON, R. nip — biogeography of species, with special reference to ferns. Bot. Rev. (Lancaster) S2otks—
TRYON, R. i pee Pp. 327-338 in H. Lieth and M. J. A. Werger (eds.) Tropical rain forest ecosystems. let Amster
TRYON, R. M. and R. G. STOLZE. 1989. Pied eae of Peru. Part I. 1. Ophioglossaceae—12. Cy- eee. Fieldiana, Bot ‘New eries 20: 1-145.
Tryon, R. M. and R. G. STozE. 1989. Pteridophyta of Peru. Part II. 13. Pteridaceae—15. Dennstaed-
tiaceae. Fieldiana, Bot. New Series 22: 1-128. TRYON, R. M., A. F. TRYON, and K. U. KRAMER. 1990. nbpigh Sues Pp: 230-256, in K. Kubitzki (ed.) ee eee and genera a a plants. Vol. 1. oo and gymnosperms. Vol.
K. U. Kramer and P. S. Green. Springer-Verlag, a6 as RM and R. G. SToLzE. 1991. Pteridophyta of Peru. a IV. 17. Dryopteridaceae (with collaboration of] T. Mickel and R. CG. Moran). Fieldiana, Bot. New Series 27: 1-176. TrYON, R. M. and R. G. STOLZE. 1992. Pteridophyta of oF has Ill. 16. Thelypteridaceae (con- tributed ‘ne A. R. Sensi Fieldiana, Bot. New Series 2 TRYON, R. M. and R. G. STOLZE. 1993. Pteridophyta of a < V1 ie ariaernepRi Poly- podiaceae (with gepacieiers of B. Leén). Fieldiana, Bot. New Series —190.
TRYON, R. M. and R. G. STOLZ 1994. Pteridophyta of Peru. Part VI. 22. spe akan sir (with collaboration of = \.H ckey and B. Ollgaard). ne apr Bot. New Series 34: 1— TRYON, R. 1997. Systematic ee on Oleandra. Rhodora 9 -3
TRYON, R. 1997. Proposal to reject the name Acrostichurn a a (Pardaceas), Taxon 46: 339— 340
CHURCHILL, H., R. TrYON, and D. S. BARRINGTON. 1998. Development of the sorus in tree ferns: Dicksoniaceae. Canad. J. Bot. ne 1245-1252.
TRYON, R. M. and A. F. TRYON. 1999. Observations on the phytogeography of eastern North Amer- ican ferns. Pp 250-273 in X-C Zang and K-H Shing (eds.). Ching Memorial Volume. Institute of Botany, Chinese Acad. Sci., Beijin
TRYON, R. 2000. Systematic notes on the Old World fern genus Oleandra. Rhodora 102: 428-438.
TRYON, R. M. 2001. Review of Hoshizaki, B. J. and R. C. Moran, Fern grower's manual, revised and expanded edition. ego 103: 34
Winoiscu, P. G. and R. TRYON. 2001. The pane Ricardo Franco (State i “is Grosso, Brazil) as probable migration ean and its present fern flora. Bradea 8: 267—
American Fern Journal 92(1):10—-19 (2002)
Crystals Associated with the Intertracheid Pit Membrane of the Woody Fern Botrychium multifidum
ANGELA C. MORROW AND ROLAND R. DUTE Department of Biological Sciences and Alabama Agricultural Experiment Station, Auburn University, Auburn, Al 36849, USA
ABSTRACT.—CALCIUM-containing crystals have been found in the lumens of secondary tracheids in the rhizome of the woody fern Botrychium multifidum. These crystals are styloids with rough, pyramid- shaped ends. The crystals are usually single; however, conjoined or grouped crystals were also found. Crystal formation apparently has no constant relation to the pit membrane, but crystals of mature tracheids are often associated with the pit membrane or are located in the pit areas. Crystals were also located between the helical thickenings of the lumen walls. No crystal chamber or crystal sheath was found in association with the crystal body.
Crystals are a common feature in many plant tissues (Scurfield and Mitchell, 1973), and more than 1000 crystal producing woody plants, spanning 160 fam- ilies, were described at the light microscopic level by Chattaway (1955, 1956). Scanning electron microscopy has allowed for more rapid identification of crystals in plant tissues and a clearer picture of their morphology (Scurfield and Mitchell, 1973). Although crystals in xylem tissue have been reported in the vessels of Intsia Thouars (Fabaceae; Hillis, 1996), Torreya yunnanesis C.Y, Cheng & L.K. Fu (Taxaceae; Kondo et al., 1996), and Polyalthia Blume (An- nonaceae; Scurfield and Mitchell, 1973), they are most commonly found in the xylem parenchyma, septate fibers, or vessel tyloses (Scurfield and Mitchell, 1973).
The formation of crystals by Botrychium, the only extant fern that produces wood (Gifford and Foster, 1989), has not been previously reported. During our studies of the torus-bearing pit membrane in the tracheid of Botrychium mul- tifidum (S.G. Gmelin) Rupr., we discovered occasional instances of crystals associated with the pit membrane. This paper describes the morphology of these crystals as observed with SEM. The discoverey of these very small crys- tals was unexpected, and our exploration of them to this point has been strictly descriptive. However, in our discussion we explore several possible reasons for crystal formation in this wood.
MATERIALS AND METHODS
Rhizome samples of upright or orthotropous rhizomes of Botrychium mul- tifidum. were collected by Dr. D. W. Stevenson (New York Botanical Garden, New York City, U.S.A.) from Plumas County, California and fixed in FPA. The collection site (elevation 2000 m) is rocky mountain soil at the edge of a mead- ow and the ground is frozen for much of the year. The samples were typical rhizomes selected as random samples and representative of the population. In
MORROW AND DUTE: CRYSTALS IN BOTRYCHIUM MULTIFIDUM 11
our lab, samples were cut transversely into 1-2 mm pieces that were placed into 50% ethanol and then dehydrated through a graded alcohol series. Sam- ples then were cut into small wedges, placed into hexamethyldisilazane (HMDS) for 2 hours (Nation, 1983), and subsequently placed under a chemical hood overnight to dry. Dry samples were attached to aluminum stubs with double-sided sticky tape and coated with gold-palladium. For comparison pur- poses, samples of Botrychium dissectum Sprengel and B. virginianum (L.) Swartz from Lee County, Alabama were prepared in the same manner as B.multifidum. Specimens were viewed with a Zeiss DSM 940 at 5,10, or 15 kV. Qualitative element identification was performed using energy dispersive spectroscopy (Tracor Northern Micro Z II) coupled to the SEM.
RESULTS
Secondary xylem tracheids of Botrychium multifidum contain helical wall thickenings and intertracheid circular bordered pits (Fig. 1). Thickenings, as seen in longitudinal section, are uniform neither in height nor in distance between gyres, and thickenings are sometimes branched (Fig. 1). The pit mem- brane is almost always differentiated into a torus and margo (Fig. 2). Microfi- brils of the pit membrane are loosely woven in the margo region, but tightly woven in the torus. Tearing of the pit membrane was sometimes evident in the margo (Fig. 2). Crystals were found in association with torus-bearing pit membranes of tracheids (Fig. 11), as well as in tracheid lumen (Figs. 1, 3). These crystals were not apparent at the light level. Crystals associated with these tracheids are styloids (Frey-Wyssling, 1981; Carlquist, 1988); they are rectangular columnar with pyramidal ends. Intact crystals have columns that are four-sided and are smooth-surfaced. The pyramidal crystal ends consist of four equilateral triangles, although wedge-shaped ends also were observed (Fig. 4). Crystal ends, when visible, typically appeared to be rough (Fig. 3), although some crystals with smooth ends were observed (Fig. 4). Crystals ranged in size from 4.3 to 12 wm in length, and 1.14 to 2.4 pm in width (N = 12). The mean crystal length is 7.27 »m, mean width is 1.55 pm, and mean ration of width-to-length is 1: 4.7. The crystals were not always regular in shape and sometimes appeared to have their growth modified by the presence of a helical wall thickening (Fig. 3). By energy dispersive spectroscopy (EDS), these crystals were found to be composed of a calcium compound, most likely calcium oxalate (Fig. 13).
Although single isolated crystals were most commonly found, joined double crystals with U-shaped conjoined end were also observed (Fig. 4). Crystals in groups of two or more were also encountered and had either parallel or per- pendicular orientation to each other (Figs. 1,5). Crystals were found in various positions within the tracheary lumen. They were located between either the helical thickenings of the wall material, laying flat on the inner cell wall, or projecting out from the pit membrane (Fig. 1).
It was clear from some specimens that the crystals were composite structures (Figs. 4, 6-8). In some instances the subunits resembled raphides (Figs. 6,7),
12 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
Fics. 14. SEM micrographs of intertracheary pit membranes and crystals. 1) Tracheary lumen with helical wall thickenings (W), crystals (arrow), pit aperture (A), and circular bordered pit membrane (P); scale bar = 5 um. 2) Intertracheary pit membrane with torus (T) a margo (M).
e pit border was removed when the wood was split during preparation; scale = 2 pm. 3) Crystal entering a pit aperture (A). Note how = crystal appears to have grown tsa the helical thickening to the right (double arrow). R = rough end of crystal; scale bar = 2 um. 4) Double crystal joined at one end (arrow); scale bar = m.
MORROW AND DUTE: CRYSTALS IN BOTRYCHIUM MULTIFIDUM
FIGs
SEM micrographs of crystals in which crystals revea
Multiple crystals with parallel orientation and perpendicular orientation. Note subunits crystal (arrow); scale bar = 5 pm. 6) E
| their subunit composition. 5)
s in broken view of composite crystal formed by smaller raphide shaped crystals (arrow); scale bar = 500 nm. 7) Composite crystal w ith styloid (arrow) and raphide crystal (double arrow) shaped subunite: scale bar = 2 pm. 8) Composite crystal with styloid crystal subunits (arrow); scale bar = 2 pm.
14 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
whereas in others they resembled small styloid crystals that were fused to form one large crystal (Figs. 7, 8). Both types of subunits appear to integrate into one another (Fig. 7). One shattered example had a hollow center (Fig. 9).
Crystals were observed to traverse the pit aperture (Figs. 3, 10) and contact the pit membrane (Figs. 11, 12). These did not appear to penetrate the pit membrane, but we are uncertain of this point due to the poor preservation of the pit membranes in our samples. Fig. 12 demonstrates a unique occurrence in which a pit membrane is approached by a crystal from either side. Due to the position of the crystal relative to the pit membrane, we were unable to confirm the presence of a torus in each crystal-associated pit membrane; how- ever a torus was present in the samples that exhibited a crystal behind the pit membrane (fig. 11).
No noticeable chamber or crystal sheath was ever observed in association with a crystal. No evidence of a surrounding membrane was discovered, al- though the ends of the crystals often were rough (Fig. 3). Efforts to examine crystals with TEM to determine the presence or absence of a chamber or crystal sheath were not successful. Crystals were not observed in either Botrychium dissectum or B. virginianum.
DISCUSSION
Three major systems of mineralization occur in plants. These include silic- ification, calcium carbonate crystallization, and calcium oxalate crystallization (Grimson et al., 1982). Calcium oxalate crystals, either in the monohydrate or polyhydrate state, are the most common mineral deposits (Webb and Arnott, 1982). EDS evidence indicates that our crystals contain calcium. The bipyra- midal shape of the crystals’ ends, and the rectangular columns, suggest that they are crystals of calcium oxalate in the polyhydrate form (Frey-Wyssling, 1981). Usually, acid solubility tests are used to confirm crystal composition in plants (Webb and Arnott, 1982). In addition, the oxalate nature of a calcium crystal can be tested with cupric acetate and ferric sulphate (Deshpande and Vishwakarma, 1992). However, due to the small size and sparse number of crystals found in Botrychium multifidum, these tests were not performed.
The location of crystals in tracheid lumens in unusual. Crystals in wood are most frequently found in ray or axial parenchyma cells (Chattaway, 1955, 1956), although they may also be found in septate fibers, vessel tyloses, and even in vascular cambia (Deshpande and Vishwakarma, 1992). In Polyalthia, vessels contained a crystalline mass (Scurfield and Mitchell, 1973). In the cur- rent study, crystals were isolated within the tracheary lumen, and there was no evidence suggesting attachment to cell walls. Some crystals appeared to be touching, but were not attached to, the pit membrane. Crystals also were found that had no apparent association with a pit membrane. Therefore, it appeared that crystal formation was not directly related to pit membranes.
Crystals in plants often are formed in membrane-bound compartments with- in the vacuole (Arnott and Pautard, 1970; Franceschi, 1984: Webb et al., 1995). As proposed by Arnott and Pautard (1970), the cell membrane may control
MORROW AND DUTE: CRYSTALS IN BOTRYCHIUM MULTIFIDUM
cv
Fics. 9-12. SEM micrographs of crystals in association with pit area and pit membranes. 9 Fractured crystal with hollow center (arrow). Note the aperture behind the crystal (double arrow); scale bar = 5 jm. 10) Crystal entering a pit aperture. Note shaping of the crystal around helical thickening (arrow); scale bar = 2 1) Crystal behind pit membrane; scale bar = 2 pm. 12) Pit membrane associated with crystals from contiguous tracheids; A = aperture; C = crystal; P = pit
membrane; scale bar = 2 pr
16
A. Experimental
AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
PEAK LISTING RGY AREA
cp wh
EL. AND LINE - KA
B. Control I
PEAK LISTING RGY AREA
OU bh oN
EL. AND LINE a I KA
7
Pic. 13.
EDS of tracheid. A. Spectral tracing of crystal within a tracheid. The calcium compone
nt of the spe ectrum is conspicuous and is indicated by the peak labeled CA. B. EDS of tracheid without
represents calcium in the middle lamella. AU
= palladium; SI =
MORROW AND DUTE: CRYSTALS IN BOTRYCHIUM MULTIFIDUM 17
both shape and growth of crystals. There was no direct evidence that crystals of B. multifidum were once enclosed in a membrane, but Scurfield and Mitch- ell (1973) suggest that a rough area on a crystal is indicative of the adhering remnants of membrane. In crystals of B. multifidum only the membrane’s im- pression on a crystal would be evident because living portions of the tracheid has undergone autolysis and no membrane remains. If the vacuole with its membrane-covered crystal pressed against either the cell wall or a cell wall thickening as a crystal formed, this contact could explain the shape of these crystals.
Water flow though the xylem could also deposit crystals (no longer enclosed by cytoplasm) randomly throughout a tracheid, including on top of a pit mem- brane or between wall thickenings. Due to erosion, water flow might also change crystal shape.
Another aspect of crystal development in plant cells in isolation of a crystal by wall material or a suberized sheath after the crystal has formed within a vacuole. This process would, in essence, externalize the crystal (Frank and Jensen, 1970). In Agave (Agavaceae), crystals are produced in such extraplasm- ic compartments (Wattendorff, 1976a, b). Wattendorff (1976b) found that all styloid idioblast of Agave, where they did not touch the wall, were surrounded by a suberized sheath. Although crystals of B. multifidum are styloids, they appear neither to be associated with a sheath of any sort nor to be isolated by cell wall material.
The reason a cell forms a crystal is not well understood. Crystal formation may represent a crystallization of waste material or storage of minerals (Desh- pande and Vishwakarma, 1992). Crystal formation also may be associated with ionic balance, and therefore, the formation of a crystal could be a form of osmoregulation (Franceschi and Horner, 1980). Franceschi and Horner (1979) correlated the amount of calcium in the growth medium and the number of crystals formed in Psychotria L. (Rubiaceae) callus. Lane (1994) has suggested that calcium oxalate crystals may promote the polymerization of lignin which of course, would be occurring in the developing tracheids of B. multifidum.
It is evident at times that crystal formation in plants is under genetic control (Frey-Wyssling, 1981; Webb, 1999); however, genetic control of the formation of all crystals has not been proven. The cell in which a crystal is produced undergoes many changes at macro, micro, and ultrastructural levels, as well as, changes in cell chemistry. These changes, documented in other plant taxa during crystal formation, make it unlikely that crystal formation could be sim- ply the result of precipitation or crystallization (Franceschi and Horner, 1980), although crystal formation may represent crystallization of waste material or storage of minerals (Deshpande and Vishwakarma, 1992). Deshpande and Vish- wakarma (1992) also identified seasonal fluctuation in crystal formation after the cessation of cambial activity. Gourley and Grime (1994) described crystals that were more commonly found in the late wood of Acacia Mill.(Fabaceae). The availability of water was also determined to be a factor in crystal formation (Gourley and Grime, 1994).
It was impossible to determine for certain whether crystals in the tracheids
18 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
of B. multifidum formed before or after cell death. Perhaps due to greater water flow resistance occurring at the pit membrane, there would have been a greater chance for calcium precipitation in the pit area rather than in the tracheary lumen. If this were the case, crystals could at the pit membrane form after the death of a tracheid. However, the rough ends observed on some crystals sug- gest they may have been enclosed at one time by a membrane. Additionally, the crystals appear to conform to the shape of the pit aperture or cell wall thickenings and do not appear to have been randomly distributed by water flow. Perhaps the best explanation of where these crystals develop is in mem- brane compartment within vacuoles of living tracheids. The enlargement of a crystal in a plane perpendicular to the cell’s axis would result in its abutting a wall or pit membrane, thus influencing crystal shape. Crystals that elongated parallel to a cell’s axis would not encounter these boundaries and would not be shaped by them. Crystals that were not pressed into a cell wall or pit mem- brane also would not develop this shaping and might settle between wall thickenings, or after cell death, move with the xylem water stream. However, the positions of crystals in Figures 3, 10, and 12 with respect to the pit mem- brane suggest that crystal position is not the result of water flow.
Crystal formation has also been associated with the products of fungal me- tabolism within the plant cells (Scurfield and Mitchell, 1973). In our study, no fungal hyphae were found near any of the crystals. Therefore, this possibility in B. multifidum seems unlikely.
If crystal manufacture is under genetic control, what advantage does the cell gain from its production? This is an especially intriguing question with regards to Botrychium as crystal production would be occurring in cells about to die. Lane (1994) has suggested that calcium oxalate crystals play a role in lignin polymerization and perhaps this may be true in these lignified tracheids. How- ever, the lack of crystals in other Botrychium species indicates that this would be true only under certain environmental conditions. As previously men- tioned, some authors believe that crystals represent the storage of calcium that could be either reserve calcium or waste calcium (Deshpande and Vishwak- arma, 1992; Webb, 1999). Storage of needed calcium in a near-death cell would be unlikely. However, these crystals may have been produced by a cell for ionic balance or osmoregulation. Ionic balance and osmoregulation are critical for immature cells (Franceschi and Horner, 1980). If a plant were growing on soil with high nitrate levels, assimilation of this compound would increase cell pH, and oxalic acid might be produced to counter this effect. The oxalate anion could then react with calcium to form a crystal that would remove the excess anion from cell sap (Franceschi and Horner, 1980).
Another explanation could be protection from herbivores, although crystal production in a leaf cell would be more plausible for defense purposes. The crystals in B. multifidum are too small and too few in number for this type of protection. Fire protection was listed as an explanation by Gourley and Grime (1994) for crystals in Acacia, but again this is unlikely for a rhizome.
Based on our data the best explanation for crystal formation in the xylem of B. multifidum is that crystals are the result of excess calcium precipitation,
MORROW AND DUTE: CRYSTALS IN BOTRYCHIUM MULTIFIDUM 19
which could represent either waste, storage, or osmoregulation in the plant. Because the crystals are located in dead cells, active resolubilization by a cell would be unlikely; however, if crystals were dissolved by water flow in the xylem, their minerals could be carried in the transpiration stream. Deshpande and Vishwakarma (1992) have suggested that formation of calcium crystals
may be a reversible process in some tissues. Therefore, these crystals do not necessarily represent a calcium loss for the plant.
LITERATURE CITED
ARNOTT, H. J., and F. G. E. PAUTARD. 1970. Calcification in plants. Pp 375-446, in maida saledicatian: Cellular and molecular aspects. H. Schraer, ed. North-Holland, Amsterdam
CHaTTaway, M. M. 1955. Crystals in woody tissues. Part I. Trop. Woods 102:55—74, ———. 1956. Crystals in woody tissues. Part II. Trop. Woods 104:100—24. DESHPANDE, B. P., and A. K. VISHWAKARMA. 1992. Calcium beam a in the fusiform cells of the pli es of Gmelina arborea. [AWA Bull. N.S.. isa V. R. 1984. Developmental — : calcium ee an sand depositions in vulgaris L. leaves. Protoplasma 12 H. T. HORNER. 1979. Use of aera eis callus in study of calcium oxalate crystal idioblast formation. Z. Pflanzenphysiol. 92:1-75. 1980. Calcium oxalate crystals in plants. Bot. Rev. (Lancaster) 46:361—42 FRANK, E. and W. A. ae 1970. On the formation of the pattern of crystal idioblast in ade ensiformis D.C. IV. The fine structure of the crystal cells. Planta 95:202—217. FREY-WYSSLING, A. 1981. Crystallography of the two hydrates of crystalline calcium oxalate in plants. Amer. J. Bot. 68:130-141 GIFFORD, E. M., and A. S. FOSTER. 1989. oe and evolution of vascular plants. 3rd ed., eeu and Co., Salt Lake C Gour.ay, L. D., and G. W. GRIME. an. Calc cium oxalate . in african Acacia isa and their says by scanning proton a (SPM IAWA Bull. N.S. 15:137-1 ee M. J., H. J. ARNOTT, and M. A. WEBB. 1982. A scanning electron eo ins of 71133-1140.
winged point — in the bean legume. Scanning Electron Microscopy III Hits, W. E. . Formation of robinetin crystals in vessels of Intsia species. ore ng vie N.S. x reaione Konpo, Y., T. Fuyl, ¥. HAYASHI, and A. Kato. 1996. Organic crystals in the tracheids of Torreya
yunnanenss IAWA Bull. N.S. 17:393—403.
Lang, B. G. 1994. Oxalate, germin, and the extracellular matrix of higher plants. F.A.S.E.B.J. 8: 294-301.
NATION, J. ca 1983. A new method using sre ea ca for preparation of soft insect tissues for scanning electron microscopy. Stain Technology
ScurFIELD, G., and A. J. MITCHELL. 1973. Crystals in woody ely Bot. J. Linn. Soc. 66:277—289.
WarTTENDoREF, J. 1976a. A third type of raphide crystal in the plant kingdom: six-sided raphides
i 4a
ioe animate of the suberized svioid crystal cells in pam leaves. Planta (Berl.) ee pier
Wess, M. A. see Cell-mediated crystalliztion of calcium oxalate in plants. Pl.Cell 11:751—761.
, and H. J. ARNOTT. 1982. A survey of calcium oxalate crystals and other mineral inclusions
in seeds. eres Electron Microscopy III:1109-1131
ALETTO, N. C. Carpira, L. E. Lopez, and H. J. ARNOTT. 1995. The intravaculolar
oat fae associated with calcium oxalate crystals in leaves of Vitis. Plant J. 7:633-648
American Fern Journal 92(1):20—22 (2002)
A New Filmy Fern from the Dominican Republic
CARLOS SANCHEZ Jardin Botaénico Nacional, Carretera del Rocio, km 3.5, Calabazar, Boyeros, C. P. 19230 Ciudad de la Habana, Cuba
ABSTRACT.—A new species of Hymenophyllum subg. Hymenophyllum with entire involucral valves is described from the Dominican Republic on the island of Hispaniola.
During the prepartion of a revision of the filmy ferns (Hymenophyllaceae) for the Flora of the Greater Antilles project, a peculiar species was discovered among the undetermined specimens in the Gray Herbarium.
The taxon belongs to Hymenophyllum subg. Hymenophyllum, following Morton (1968, pp. 162-164), a subgenus represented by only two species in the Antilles. Hymenophyllum tunbrigense (L.) Smith is known only from Ja- maica and Hispaniola (Proctor 1985, p. 90), whereas H. fucoides (Sw.) Sw., is more widely distributed (Proctor, 1985, p. 92; 1989, p. 58). The members of this subgenus are characterized by having toothed segment margins, and most have sinuous to toothed involucral valves as well. The new species is de- scribed as
Hymenophyllum integrivalvatum C. Sanchez sp. nov. Fig. 1
Ab speciebus aliis antillanis subgeneris Hymenophylli valvis integris, stip- itibus brevissimis, segmentis pinnarum paucis (1 vel 2), necnon laminis gla- berrimis diversa.
TypE—Dominican Republic: Pcia. La Vega: Near the pyramid ca. 13 km from Valle Nuevo on the road to San José de Ocoa, ca. 2500 m elev., 22 August 1957, Gastony, Jones & Norris 740 (GH; isotype US).
Rhizomes creeping, filiform, 0.1-0.3 mm in diam., clothed with deciduous, brownish, pluricellular trichomes, with a few conspicuous, straight roots ca. 5 mm distant. Fronds small, erect, determinate, approximate, 1.15—2.1 cm long.; stipes 0.1-0.3 mm long., 0.2 mm in diam., very narrowly alate thorough- out, dark brown, glabrous or with a few brownish, often 2-celled trichomes; laminae narrowly ovate, lanceolate or oblong 1.4-1.8 cm long, X 0.8-1 cm wide, pinnate-pinnatifid; rachises notably flexuous, narrowly and evenly alate, the alae less than 0.1 mm wide, dark brown, glabrous; pinnae 5-8 pairs, spreading to ascending, mostly with 2 acroscopic segments; segments narrowly elliptic, oblong, or linear-oblong, 1.2-1.8 mm wide, glabrous, the margins dis- tantly toothed, the teeth usually more distant than their length and ascending, the midvein dark brown, the lamina tissue olivaceous-green when dry; sori conspicuous in size in comparison with the length of the lamina, borne at the lamina apex or in the distal half, subaxillary on the acroscopic side of the
SANCHEZ: A NEW FILMY FERN FROM THE DOMINICAN REPUBLIC 21
B22,
4. Cale ry Se Fic. 1. Holotype of Hymenophyllum integrivalvatum (GH). A. Habit of plant. B. Detail of an ultimate segment. C. Detail of a sorus.
pinnae; involucres 1.6—2.2 mm long, 1.4 mm wide, broadly elliptic or broadly ovate, bivalvate, the valves wider than the sterile segments, the margin entire, the filiform receptacle included.
DISTRIBUTION.—Endemic to Hispaniola (Dominican Republic), known only from the type collection.
Hasirat.—Epipetric in very moist burned and timbered pinelands, forming thick mats on rocks along streams in very moist ravines, according to the in- formation on the label.
The new species is most closely related to H. tunbrigense (L.) J. E. Smith, which is also known from a few collections from Hispaniola. Hymenophyllum tunbrigense differs in having larger fronds, straight rachises with wider alae, more divided pinnae, narrower segments, and sinuous involucres. The entire involucral valves, very small fronds, and the absence of trichomes on the re- mainder of the lamina separate the new species from all other Antillean spe- cies of subg. Hymenophyllum.
ACKNOWLEDGMENTS
I am indebted to the following individuals for their help and hospitality during my visit to different North American herbaria to study Greater Antillean pteridophytes: John T. Mickel and Thomas Zanoni (New York Botanical Garden), David B. Lellinger (United States National Herbar- ium), and Emily Wood and David E. Boufford (Gray Herbarium, Harvard University). I am grateful to the New York Botanical Garden and The John D. & Catherine T. MacArthur Foundation for financial support. Thanks also to David B. Lellinger for revising the English version of the man- uscript. I also wish to thank Manuel G. Caluff for the illustrations.
22 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
LITERATURE CITED
Morton, C. V. 1968. The Genera, Subgenera, and Sections of the Hymenophyllaceae. Contr. U. S. Natl. Herb. 38:153-214.
Proctor, G. R. 1985. Ferns of Jamaica: A guide to the pteridophytes. British Museum (Natural History), London,
. 1989. Ferns of Puerto Rico and the Virgin Islands. Mem. New. York Bot. Gard. 53: 1-389.
American Fern Journal 92(1):23—29 (2002)
Adiantum argutum, an Unrecognized Species of the A. latifolium Group
JEFFERSON PRADO Secdo de Briologia e Pteridologia, Instituto de Botanica, Caixa Postal 4005, 01061-970 Sao Paulo, SP, Brasil Davip B. LELLINGER Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0166
ABSTRACT.—The present paper distinguishes A. argutum, an unrecognized but widespread species from South America, from the related A. Jatifolium, and designates a lectotype for A. argutum.
Several pinnate or bipinnate Adiantum species have an indument like that of the A. serratodentatum group, but differ in having fewer, larger, less di- midiate pinnules and thin, very long-creeping rhizomes. Among the species of this group are A. argutum Splitg., A. incertum Lindm., which is based on Lindman Regnell Exped. I. A2083 (S not seen; isotypes B, GH) from Paraguay, the widespread A. Jatifolium Lam., and A. viviesii Proctor, which is based on Proctor 41389 (US; isotypes IJ, SJ) from Puerto Rico. Adiantum glaziovii Baker, which is based on Glaziou 13345 (K, isotype US) from Rio de Janeiro, Brazil, is a synonym of A. Jatifolium. The early, unplaced name A. elatum Desv., which is based on a Brazilian specimen from the Herb. Dombey (P-Herb. Juss. Cat. 1421 not seen Morton photo 3153) will likely displace one of the later named species found in Brazil. The specimen was said by Morton to have almost glabrous segments; it needs to be examined critically.
Adiantum argutum Splitgerber, Tijdschr. voor Natuurl. Gesch. en Physiol. [Leiden] 7: 427. 1840. Figs. 1, 2.
Lectotype (chosen here): “in sylvis montosis Surin. prope Bleauwe Berg,” Surinam, May 1838, Splitgerber 891 (L not seen, photo US). Other syntype: idem, id., Splitgerber 290 (L not seen, photo US).
Adiantum fovearum Raddi var. reductum Jenm. Ferns Br. W. Ind. & Guiana 87. 1899. TYPE: Not clearly stated, but presumably Guyana, Jenman (NY? not seen).
Splitgerber (1840) described A. argutum based on material he collected in Surinam. The author did not mention any specimens in the protologue, only the locality. Original materials were found by Morton at Leiden (Morton photos 193 and 194, both US), and they may be considered the syntypes of Splitger- ber’s taxon. The lectotype selected here (Splitberger 891) has an original label, handwritten by the author with the same information he published with the
24 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
Ss ° J gv - is ° \ 10 o 2 ¢ 3 o “Sa ‘4 e e o —— a : ° e t eveinaph Ottis Pl ° en Me ind <— aa yy. ‘, ane = Be = é bad e \ a \ ° r bane 0 , ° a ° f eS @
Fic. 1. Range of Adiantum argutum Splitg.
original description. The photograph of the other syntype (Splitgerber 290) resembles A. Jatifolium Lam., although we can not place it there with certainty.
According to Splitgerber (1840), A. argutum has a long-creeping rhizome; the laminae are lustrous adaxially and have 3 or 4 pinna pairs, acuminate pinnules, a subrhombic terminal pinnule, reduced and flabellate basal pin- nules, sparse, minute setiform scales abaxially, and oblong sori. In fact, these characters distinguish this species from all others closely related to it. Other important features to recognize A. argutum are the distant fronds and the id- ioblasts on the abaxial surface of the pinnules.
Unfortunately, over the years, the Splitgerber species was included within the concept of A. Jatifolium by Vareschi (1969, p. 734), Kramer (1978, p. 91), Tryon and Stolze (1989, p. 66), and Smith (1995, p. 259). In other cases, the
PRADO AND LELLINGER: ADIANTUM ARGUTUM 25
name was synonymized under this species, by Posthumus (1928, p. 105), Lel- linger (1989, p. 148), and Cremers and Hoff (1990, p. 19).
Adiantum argutum demonstrates its close affinity to A. Jatifolium mainly in its slender, long-creeping rhizome, 2-pinnate fronds, and stipe and rachis cov- ered by deltate to lanceolate scales with a pectinate base. However, A. latifol- ium differs by its smaller, obtuse to subacute pinnules with a roundish apex that are abaxially glaucous, glabrous, and without idioblasts.
Adiantum incertum Lindm. differs from A. argutum and A. Jatifolium in having the scales on the abaxial surface of the pinnules hairlike and with a few basal processes, rather than having such scales with a pectinate base or totally lacking scales. In addition, it is a species restricted to Paraguay and extra-Amazonian Brazil: Goids (Maurilandia, Rio dos Bois, Hatschbach 34271, MBM, MO, NY, UC), Mato Grosso (Santa Terezinha, 21 km SW of Portal da Amazonia, Thomas et al. 4334, NY, US), SAo Paulo (Sao Carlos, 9 km NNE of the BR Station at Santa Eud6xia, Eiten & Eiten 3488, US), and Parana (Foz do Iguacu, Parque Nacional das Cataratas, Hatschbach 23171, HB, MBM, MO, UC, UPCB).
Adiantum obliquum Willd., although its fronds look much like those of A. argutum, can be distinguished by its short-creeping rhizomes, approximate and usually 1-pinnate fronds, and pinnules with conspicuous idioblasts on both surfaces. It may be more related to A. Jucidum Cav. and perhaps to A. petiolatum Desv., with which it hybridizes. These three species may form a separate group.
Adiantum argutum has a more restricted area of distribution than its closest relative A. Jatifolium. It occurs in northern South America (Colombia to French Guiana) and in the Amazonian regions of Peru, Bolivia, and Brazil. It grows in primary and secondary forests, on dark red lateritic clay soils, from 50 to 1000 m elevation.
Representative specimens of A. argutum studied:
COLOMBIA: Meta: Sierra de La Macarena, Cajfio Entrada, Philipson & Idrobo 1748 (US); Villavicencio, Pennell 1607 (GH). Boyaca: Los Llanos, Haught 2833 and 2844 (both GH). Vichada: San José de Ocumé: near Rio Vichada at Botomi, ca. 14 km NW, Hermann 11107 (US); NE de Pto. Infrida, 3°58’N, 67°50’W, Churchill et al. 17748 (NY).
VENEZUELA: Bolivar: La Tomasa, Williams 1295 (US); Rio Paragua, Isla El Casabe, Killip 37301 (US); Salto Alta, Alto Orinoco, Croizat 486 (NY); Dtto. Sifontes, Concesién Minera Oro Uno, 7 km NW of la Clarita, 6°13'N, 61°27'W, Aymard et al. 3976 (NY); Sierra Imataca betw Rio La Reforma and Puerto Rico, N of El Palmar, Steyermark 88012 (US). Amazonas: Around the margin of the Rio Orinoco above Tamatama, Williams 15199 (GH); Cuenca del Rio Manapi- are, 5°5’N, 66°03’W, Huber 435 (NY). Delta Amacuro: Rio Cuyubini, Cerro de la Paloma, Steyermark 87649 (NY). Mérida: Near border Rio Grande de Toro, 61°44’W, 80°4'N, Breteler 3781 (US).
TRINIDAD: Fendler 2 (NY).
GUYANA: Cuyuni-Mazaruni: 8 km N of Bartica on W bank of Essequibo River, 06°29’N, 58°38’W, Henkel & Chin 297 (US). U. Takutu-U. Essequibo:
i io” ” a INS
PRADO AND LELLINGER: ADIANTUM ARGUTUM 27
Rupununi area, Surama Village, 04°08’N, 59°04’W, Acevedo et al. P3297 (US); Marudi River, 02°11’N, 59°11'W, Henkel et al. 2902 (NY), 3032 (US); Kuyuwini River, 02°11'N, 59°11’W, Henkel et al. 3022 (US); NW Kanuku Mts., 3°21'N, 59°30'W, Hoffman & Foster 3510 (US); Rupununi River, Jansen-Jacobs et al. 4207 (US). Potaro-Siparuni: On 0.5 km island in Essequibo River, 1 km S of Fairview, 4°40'N, 58°40'W, McDowell 3371 (US); Iwokrama Mts., Annai-Ka- rupukari Rd., 04°19’N, 58°51’W, Hoffman et al. 1409 (US); River Isherton, 2°20'N, Smith 2432 (GH, NY). Barima-Waini: Head of Barima River, Ayamba Falls, 4.5 mi W of Eclipse Falls, ca. 10 km W of Arakaka, 7°39’N, 60°09’, Pipoly & Lall 8200 (NY); Head of Barima River, NW of Kariako River, 7°30'N, 60°35’W, McDowell 4393 (NY); Labbakaka Creek, Tiger Creek, Sandwith 1209 (K, NY).
SURINAM: Haut Litany: Basin du Litany, 2°31'N, 54°45’W, Granville et al. 12040 (US). Nickerie: Area of Kabalebo Dam project, Lindeman & Roon 884 (US); Area of Kabalebo Dam project, 4°—5°N, 57°30’—-58°W, Lindeman et al. 165 and 343 (both NY); Sectie O, along railroad, vic. Km. 70, Maguire & Stahel 23605 (GH, NY). Brokopondo: 2.4 km S of village Gansee, Donselaar 1189 and 1276 (both GH); Zuid River, 3°10’—3°20’N, 56°29’-56°49’W, Kayser Airstrip, 45 km above the confluence with Lucie River, Irwin et al. 57697 (NY).
FRENCH GUIANA: Cayenne, Inini River, 3°28’N, 52°36'30’"W, Cremers et al. 8781 (US); Camp Eugene, Basin du Sinnamary, 4°51’S, 53°4'W, Cremers & Granville 13727 (NY); Gobaya Soula, Basin du Maroni, 53°58’W, 3°37’S, Cre- mers et al. 10125 (US); Saoul, 3°37’N, 53°12’W and vicinity, Route de Bélizon, N of Eaux Claires, Heald & Yahr 56 and 65 (both NY); Comté., degrad auprés de Crique Martineau, Oldeman 1426 (NY); Mt. Balbao, Secteur Sud, 3°35’N, 53°20'W, Granville et al. 8958 (NY).
PERU: Madre de Dios: Near the confluence of Rio Tambopata and Rio La Torre, 39 km SW of Puerto Maldonado, 12°50’S, 69°20’W, Smith & Condor 1114 (US) and 1363 (NY); Tambopata, Vargas 18577 (GH); Tambopata, vic of Moho towards Piedra Redonda, at the Bolivian frontier, 12°30'S, 69°40’W, Nu- fiez et al. 9695 (GH, NY); Tambopata, SSW of Pto. Maldonado at the confluence of the R. La Torre and the R. Tambopata (SE bank), Tambopata Nature Reserve, 12°49’S, 69°17'W, Barbour 4763 (NY), Loépez 4585 (GH).
BOLIVIA: Beni: Pcia. Ballivian: Rio Colorado, Collegio Técnico Agropecu- ario de Rfo Colorado, 15°00’S, 67°10'W, Fay & Fay 2105, 2640, 2652, 2654, 2681 (all US); 18.4 Km E of Riberalta, then 1 km NE on old road to Cachuela Esperanza, 11°05’S, 65°50’W, Solomon 7804 (NY); Isla Capanario, 50 m from the San Borja—San Ignacio de Moxos road, 212 km from Campoamento Totai- zal, Rolleri 140 (NY); Pcia. Moxos: Chimanes Forest, 15°10’S, 66°37’W, Fay & Fay 2794 (US). Sta. Cruz: Bella Vista, Rio Blanco, Scolnik & Luti 681 (US); Pcia. Ichilo: Old meander loop of the Rio Ichilo, 1-1.5 km SW of the Buena Vista—-Villa Tunari Hwy., 17°18'S, 64°12’W, Nee & Moran 45225 (NY). Pando: Nicolas Suarez, SW of Cobija on the Rio Naraueda, 11°08’S, 69°08’W, Sperling
oo
Fic. 2. Adiantum argutum Splitg. Fig. 2A. Habit. Fig. 2B. Abaxial surface of some pinnules. Fig. 2C. Abaxial surface of a pinnule showing the scales.
28 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
& King 6475 (GH, NY, UEC); Ca. 20 km from Cobija towards Castro Erifia, Casas & Sussana 8123 (NY); W bank of the R. Madeira betw Cachoeiras Madeira and Misericordia, Prance et al. 6612 (NY). La Paz: Pcia. Iturralde, Siete Cielos, R. Manupare, 12°27'S, 67°37’W, Solomon 16947 (NY).
BRAZIL: Amapa: Serra do Navio, bank of the Rio Amapari, Emmerich & Andrade 745 (HB, R). Roraima: Posto Mucajai, Rio Mucajaf, vic of Mucajai airstrip, Prance et al. 10991 (GH, R, UC). Para: Lageira, airstrip on Rio Mai- curt, 0°55'S, 54°26’W, Strudwick et al. 3088 (NY), 3129 (MG, NY, US), 3580 (US); Curud S.A., near Alenquer, Santarém, Piggott 2547 (K, NY); Breu Branco, ca. 40 km S of Tucurui, 4°03’S, 49°40’W, Daly et al. 1376 (GH, MO, NY, US); Serra dos Carajas, Serra Norte, ca. 15 km W of AMZA Exploration Camp., 6°S, 50°15’W, Berg & Henderson BG472 (GH, NY, UC, UEC, US); Serra dos Carajas, 6°4'S, 50°8’W, Secco 286 (GH, K, MG, NY, SPF); BR-163, Cuiabé—Santarém Highway, km 885.5, Prance et al. P25171 (MG, NY, UC, US); Parque Indigena do Tumucumaque, Rio Parti de Oeste, Miss&o Tiriyo, 2°20’N, 55°45’W, Caval- cante 2401 (K, MG, NY, US); Conceigaéo do Araguaia range of low hills ca. 20 km W of Redengao, near Sado Jodo and Troncamento Santa Teresa, 8°03’S, 50°10’W, Plowman et al. 8635 (GH, NY), 8757 (GH, NY, US); Rio Xingu, Balée 2398 (NY); Confluéncia com Rio Pardo, Vasconcelos et al. 260a (NY); Rio Ita- caiunas, affluent of the Rio Tocantins, Serra Buritirama, 50°15’W, 5°30’S, Pires 12427 (NY); Rio Cumina, Ducke (Hb. Mus. Goeldi 8885, 15163) (both HB, MG). Amazonas: 1—5 km road Boca do Acre to Rio Branco, Prance et al. 2533 (GH, MG, NY, R, US); Vic. of Tototobi, Basin of the Rio Demeni, Prance et al. 10208 (NY, UC, US); Rio Curuquete, Providencia, Prance et al. 14632 (NY, UC); Sao Paulo de Olivenga, 30 km above the mouth of the Rio Coti, Prance et al. 14444 (B, NY); Vic. of Macujai airstrip, Prance et al. 10991 (MG, NY, UC); Borba, 4°02'S, 59°06’W, W side of the Rio Cunama, Hill et al. 12868 (MO, NY). Ron- dénia: Mineragao Campo Novo, BR-421, a 2 km a Oeste da Mineragéo Campo Novo, 10°35'5"S, 63°37’W, Vieira et al. 517 (NY, US); Basin of Rio Madeira, Trail north of Rio Madeira from 2 km, below confluence of Rio Abuna, Prance et al. 8345 (K, NY, UC, US). Acre: Palacio de Castro, fazenda Mococa, ramal no km 120 da rod. Rio Branco—Pérto Velho, Santos et al. 122 (MG, NY, US); 9 km from Rio Branco on Rio Branco—Pérto Acre road at cut-off for Colénia Cinco Mil, Lowrie et al. 650 (MG, NY, R, US); Sena Madureira, Bacia do Rio Purus, varagao para o Seringal Fonte Boa, 10°07’'S, 69°13’W, Silveira et al. 668 (MO, NY); Xapuri, Seringal Cachoeira, 35 km SE of Xapuri, Pinard 809 (NY), Kainer 126 (NY). Mato Grosso: Colider, Comunidade Sao Francisco, Salino 284 (GH); Serra Ricardo Franco, 15°S, 60°W, Windisch 1503 (HRCB); Rio Pei- xoto de Azevedo, Faz. Sao José (Cachimbo), Bokermann 6747 (UEC); Santa Terezinha, BR-158, Vila Confresa, pr. ao aeroporto da Fazenda Confesa, 10°35'S, 5°35’W, Windisch 5987 (UC).
ACKNOWLEDGMENTS The first author appreciates the financial support of the Brazilian Research Council CNPq (Proc. n. 300843/93-3 and 450658/99-6) and of the Smithsonian Institution, Washington DC (Short-Term Visitor Grant). We thank also Sra. Emiko Naruto for preparing the illustration.
PRADO AND LELLINGER: ADIANTUM ARGUTUM 29
LITERATURE CITED
Cremers, G. and M. Horr. 1990. Inventaire taxonomique des plantes de La Guyane Francaise. I- Les ace ta Museum National d’Histoire Naturelle, Inventaires de Faune et Flore. Fasc
MER, K. U. 1978. ors pteridophytes of Suriname. Uitgaven Natuurw. Stud. Suriname Nederl. Antillen 93:1-19
LELLINGER, D. B. 1989 ais ferns and fern-allies of ssi a Panama and Chocé (Part 1: Psilo- taceae through Dicksoniaceae) haope 2A
PosTHUMUS, O. 1928. The ferns of Surinam. pre ie Java. 1
SMITH, A. R. 1995. poe L. Pp. eae in P. E. Berry, B. K. Holst, ae Vatachievsdhi (eds.), Flora of the Venezuelan Guayana, vol. 2: Pteridophytes, Srempatophytan: Acanthaceae—Ar- aceae. Timber Press, Portland.
SPLITGERBER, F, L. 1840, Enumeratio Filicum et Lycopodiacearum quas in Surinamo legit F. L. iteadies Tijdschr. Natuurl. Gesch. Physiol. 7:391-444.
TrYON, R. M. and R. G. STOLZE. ee ah eae of Peru, Part II. 13.Pteridaceae—15.Dennstaed- tiaceae. Fieldiana Bot., n.s. 2
VARESCHI, V. 1969. Helechos. = oat sod in T. Lasser (ed.), Flora de Venezuela. vol. 1, Tomo 2. Instituto Boténico, Caracas
American Fern Journal 92(1):30—38 (2002)
Polypodium vulgare Plants Sporulate Continuously in a Non-Seasonal Glasshouse Environment
SANNA E. SIMAN AND ELIZABETH SHEFFIELD School of Biological Sciences, 3.614 Stopford Building, The ee of Manchester, Oxford Road, Manchester M13 9PT, U
ABSTRACT.—In their natural environments pteridophytes usually have regular sporing periods, the onset of which is triggered by the interaction of climatic and nutritional factors. Little, however, is known about what changes there may be in the sporing behaviour of a fern when it is transferred from its natural habitat to an artificial environment, such as a glasshouse. We recorded sporing behaviour in relation to vegetative growth in two genetically matched populations of Polypodium vulgare. One population was placed in a controlled-climate glasshouse, the other was left outside. The recruitment of new fronds was significantly higher in the indoor population than in the outdoor population. The indoor population also maintained a high proportion of actively sporing fronds throughout the winter. There was no net recruitment of new fronds in the outdoor ce lation during the winter and early spring. Some elements of the glasshouse envi t, probably the enhanced light and temperature, induced continuous sporing in this fern. Considering the ever-increasing interest in ferns as ornamental plants, and the growing body of evidence of toxic and allergenic effects caused by fern spores, this kind of sporing behaviour may have implications for human health.
Ferns in their natural environments usually have regular and predictable periods of spore production and release. In the temperate zones and the sea- sonal tropics they tend to release their spores towards the end of the growing season. In the wet tropics, where the growing season is much longer or even continuous, initiation of new fronds and maturation of older fronds take place throughout the year (Page, 1979). Most ferns have been said to show very little fluctuation in annual spore output with variations in climate, in contrast with good and bad seed years in angiosperms and conifers (Page, 1979). This does not apply to all fern species. Page (1976) pointed out that for Pteridium aquil- inum (bracken) the spore yield can vary widely between different years. Sim- ilarly, Steeves (1959) noted that, for Osmunda cinnamomea, a hot dry summer is usually followed by a high degree of fertility in the following spring, where- as a cooler moister summer leads to reduced fertility. Furthermore, the onset of the reproductive phase in fern sporophytes can be demonstrated to be reg- ulated by the interaction of several factors, such as light exposure, temperature and the nutritional status of the plant.
Field observations have suggested that the onset of the reproductive phase in ferns (as in many flowering plants) is induced by particular photoperiods, but the evidence so far published is scanty (Wardlaw and Sharma, 1963). periments carried out by Wardlaw and Sharma (1963) indicated that there is a more or less direct relationship between active photosynthesis and/or pho- toperiodic perception in the expanded leaves and the induction and devel- opment of sori in the next inner leaves of Dryopteris austriaca. Harvey and
SIMAN & SHEFFIELD: POLYPODIUM VULGARE SPORULATION 31
Caponetti (1972), however, demonstrated that increasing light intensities in- hibited sporophyll differentiation in Osmunda cinnamomea. Maximal initia- tion of sporangia occurred in total darkness in this species, so it may be that green and non-green spored ferns respond to light in different ways. There may be doubts about the importance of photoperiodic induction, but in deter- mining the extent of the fertility in ferns, photosynthetically available radia- tion and the duration of exposure to light are probably of major importance. Steeves (1959) compared the incidence of fertility between O. cinnamomea plants in heavy wood and open areas and found a greater incidence of fertility in the latter plants. Conway (1957), Dring (1965) and Page (1976) suggested the same property in Pteridium (bracken), in which they found a gradual de- crease in fertility with increasing degree of shade, although vegetative growth in the latter may be little impaired. The enhancement of sporogenesis by high light has recently been confirmed in experiments conducted with clones of bracken grown in high and low levels of photosynthetically available radiation (Wynn et al., 2000).
Temperature also plays a role in the onset of the reproductive phase as shown by Labouriau (1958). He found that initiation of sporangia was stimu- lated by exposure of the developing outermost set of Osmunda claytoniana fronds to a temperature of 26°C; plants kept at a lower temperature remained sterile. Similar trends were reported in Pteridium by Sheffield (1996) and Wynn et al. (2000).
Allsopp (1964, 1965) suggested that nutritional conditions, particularly car- bohydrate supply, appear to be of greater importance for the induction of spo- rangia in pteridophytes than photoperiodic or similar stimuli. Several studies indicate that the nutritional status of the plant is indeed of great importance for the initiation of the spore-productive stage (Wardlaw and Sharma, 1963). Goebel (1887, 1905, 1908) and Atkinson (1896) concluded, from experiments with Onocleoid ferns, that if carbohydrate supplies are inadequate, developing leaves tend to remain in the vegetative state. Goebel (1928) found that imma- ture sporophylls of certain ferns developed as vegetative fronds when the ster- ile fronds of the plant were removed. This has been repeated in numerous fern species (e.g. by Labouriau (1958), Wardlaw and Sharma (1963), and Steeves and Wetmore (1953)), who thus linked induction of sporogenous tissue to car- bohydrate supply. Sussex and Steeves (1958) cultured excised leaf primordia of Leptopteris hymenophylloides, Todea barbara and Osmunda cinnamomea, and found that high sucrose concentrations in the medium was essential for the inception and early development of sori and sporangia in those species. They also showed that an increased supply of inorganic nitrogen promotes the onset and extent of fertility in T’ barbara. According to Wardlaw and Sharma (1963), there is a positive relationship between the amount of photosynthesis- ing leaf surface and the induction of fertility.
It is apparent from the above that we have some limited knowledge of what triggers sporing in ferns in natural conditions. The sporing behaviour of ferns transferred from their natural habitat to an artificial environment, e.g. a glass- house, however, remains largely unexplored. Considering the ever-increasing
32 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
interest in ferns as household and garden ornamentals (Gress, 1996) and the fact that fern spores may cause adverse health effects in humans (Siman et al., 1999), more knowledge in this field is urgently required.
The aim of the current experiment was to compare the reproductive perfor- mance of Polypodium vulgare plants placed in either a seasonal outdoor en- vironment or a constant high-temperature and high-light glasshouse environ- ment.
MATERIAL AND METHODS
Polypodium vulgare plants were collected in March 1998, from stone walls along the south-east side of road B4403, running along the south-east shore of Llyn Tegid (Bala Lake) (N 52°53’, W 3°38’), Wales, U.K. Polypodium was cho- sen as it represents a widespread genus, including a broad range of horticul- tural favorites, such as P. amorphum, P. cambricum and P. interjectum, of sim- ilar morphology and life history (Mickel, 1994).
he plants were potted in commercial potting compost within a day of col- lection. Lengths of rhizome were split into two equal parts (i.e. bearing the same number of fronds in each of the two pots in a pair), and each was placed in one pot. In this way 64 pots were prepared, i.e. 32 pairs of pots containing clones. The length of the potted rhizomes varied from 1 to 5 cm, but was much the same within each pair. In order to minimise the impact on the results of the growth of any apical meristems, only median pieces of rhizome were used. All plants were allowed a six-month settling-in period (mid-March to mid- September) outdoors, after which one pot of each pair was left outdoors, to the north-east of a glasshouse in the Manchester University Experimental Grounds, Manchester, U.K. These were the ‘‘outdoor population”. The other group of the plants was put inside a glasshouse (mean day temperature: 28°C, range 20-38; mean night temperature: 15°C, range 11—27; photosynthetically available radiation: c. 110 pmol m~? s~'), at the aforementioned Experimental Grounds. These were the “indoor population”. At the start, the total number of fronds in each population was very similar. No plants were given any ad- ditional nutrients during the course of the experiment. Those outside were subject to ambient rainfall, those inside were watered regularly. During the settling-in period all fronds in 14 pots, seven in each population, died. Eight of the 14 pots belonged to matched pairs, so the aim of ensuring a genetic similarity between the two populations was still met to a high degree.
Weekly records were taken of the number of fronds in each pot with a) no sori, b) immature (green) sori, c) sporing (yellow-orange) sori and d) empty (brown) sori, from the beginning of October 1998 until June 1999 for the indoor population. The outdoor population was recorded until the end of its growing season in mid-September 1999.
The proportions of recruited fronds during the experimental period by the two populations were compared with a x? test. The differences in numbers and proportions of fronds of each developmental stage in the two populations were compared numerically.
SIMAN & SHEFFIELD: POLYPODIUM VULGARE SPORULATION 33
|= outdoors
mean number of fronds per pot
28 8 oe Cc = ee = = o
18/11/98
= 8 N = ~ Nn i=)
& —~ oO =
30/12/98
8 3
10/02/99 24/02/99 10/03/99
Fic. 1. Changes in the mean number of fronds per pot in two genetically matched populations of Polypodium vulgare. The indoor aaa was placed in a controlled-climate greenhouse (mean day ek anarae 28°C, mean night temperature: 15°C, photosynthetically available radiation: ca 110 pmol m~? s~'); the outdoor seatiitten was left outside (U.K. natural weather conditions). The re- cruitment of new fronds in the indoor population occurred in three waves, one from October 1998 to mid-January 1999, the second from mid-January 1999 to late March 1999, and the third from early April 1999 to the end of the experiment. Error bars show standard error of the mean
RESULTS
The recruitment of new fronds from October 1998 to early June 1999 was significantly higher in the indoor population than in the outdoor population (x2-test, x = 127, df = 1, p<0.01) (Fig. 1). During this time the indoor popu- lation increased its number of fronds more than fourfold. The recruitment of new fronds took place in three waves (Figs. 1 and 2a), each of which increased the number of fronds by a factor between 1.5 and 1.7. The outdoor population increased its number of fronds by a factor 1.2 from October 1998 to June 1999 (Fig. 1). All recruitment of new fronds in the outdoor population took place from mid-April 1999 onwards. The increase in the number of fronds in the outdoor population continued during the summer months until mid Septem- ber 1999 (Fig. 3a).
The majority of the new fronds recruited in the indoor population during the course of the experiment was fertile (Fig. 2a), so the indoor population maintained a high proportion of actively sporing fronds throughout the winter.
ach wave of recruitment in the indoor population began with a sudden in- crease in the number of initially sterile fronds; a number which decreased as sori began to appear. The proportion of fronds that remained sterile throughout the wave decreased with each wave. Thus, at the end of January 1999 the proportion of sterile fronds was 37%, in early April 1999 28% of the fronds were sterile and in early June 1999 the proportion of sterile fronds was 17%
34 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
number of fronds
100
S86&8S88a88
percentages of fronds
co eieresetege 33a 3 g BSses ede Sbaz2 23 5 date non at&éF4C aw FSS Ee
Fic. 2. Sporing behaviour in a population of Polypodium vulgare kept in a controlled-climate
bars represents i) sterile fronds and fronds with ii) green sori, iii) sporing sori and ix) empty sori, as indicated by the key in the figure. (a) Total number of fronds in the population (full bars) and number of fronds in each of the four groups (i.e. sterile, green, sporing, empty) for each week of the experiment. (b) Proportions of sterile, green, sporing and empty fronds, respectively, for each week of the experiment.
(Fig. 2b). The frond mortality represented 7.8% of the total number of fronds gained over the experimental period and was entirely due to old fronds wither- ing and falling off.
In the outdoor population there was no net recruitment of fronds during the winter and early spring. When the new fronds started to emerge, in late April,
SIMAN & SHEFFIELD: POLYPODIUM VULGARE SPORULATION 35
Number of fronds
3 g co. raed 2S
Percentages of fronds
os 8 88 8
$ 3 co
Fic. 3. Sporing behaviour in a population sa painir to vulgare kept outside in the Manchester University Experimental Grounds, U.K. (natural weather conditions) from October 1998 to Sep- tember 1999. The subdivision of the bars nanan i) sterile fronds and fronds with ii) green sori, iii) sporing sori and iv) empty sori, as indicated by the colour code key in the figure. (a) Total number of fronds in the population (full bars) and number of fronds in each of the four groups (i.e. sterile, green, sporing, empty) for each week of the experiment. (b) Proportions of sterile, green, sporing and empty fronds, respectively, for each week of the experiment.
most of them soon turned into fertile fronds, so there was a steady increase in the number and proportion of fertile fronds over the summer (Figs. 3a and 3b). The proportion of sterile fronds decreased simultaneously so towards the end of the growing season in mid-September 1999, the proportion of sterile fronds was 17% (Fig. 3b), i.e. the same as for the indoor population at the end of its
36 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
third wave in early June 1999. The increase in number of fronds in the outdoor population was slightly impaired at two points (24/June/99 and 22/July/99) by herbivory from snails, but the fronds thus lost represented no more than 5% of the population.
DISCUSSION
It is clear that the conditions of a warm and illuminated glasshouse stimu- lated the vegetative growth and spore output of the P. vulgare plants.
In the indoor population, the initial response to the glasshouse conditions was increased vegetative growth. At a time when the outdoor population stopped producing new fronds, the indoor population continued recruiting. Similar continuous growth has been observed in Pteridium grown in glass- houses (Wynn et al, 2000), but it is interesting that Thomson (2000) reports that Pteridium plants from four places (Honshu, Japan; Kiev, Ukraine; Bridg- ton, Maine, U.S.A. and Waterloo, Michigan, U.S.A.) require cold treatment (4°C) for four to eight weeks to ensure successful spring emergence of croziers when cultivated in the relative warmth of Sydney Royal Botanic Gardens (summer temp: max. 25.5°C, min. 18.2°C; winter temp: max. 16.8°C, min. 8.7°C). It seems that fluctuating temperatures warmer than those of the natural environment of a fern can have adverse effects on frond recruitment.
The majority of the new fronds emerging in the indoor population of the present study became fertile, so a high proportion of fertile fronds was main- tained throughout the winter. Steeves and Wetmore (1953) concluded, after experiments with Osmunda cinnamomea, that the factors which determine fertility exercise their effects during the year before the leaves expanded. As- suming, in the present experiment, that each wave of recruitment created in the indoor population mimicked one growing season, we could suggest that the warm and bright indoor climate had an effect on the fertility of the second and third waves of new fronds. The proportion of fertile fronds in each of those two waves was higher than in its preceding wave. This may well be an effect of the enhanced nutritional status of the population, caused by a high production of photosynthate, which, transported as sugars to the bud primor- dia, might induce fertility, as suggested by Harvey and Caponetti (1972).
In its natural environment P. vulgare produces ripe spores from July/August. The ripening of the spores is a gradual process, occupying a period of several months. Within a single sorus some sporangia shed early and others will take longer to ripen and shed later (Wright and Wright, 1999). The persistent pro- portion of 10-20% sporing fronds in the outdoor population from October 1998 to March 1999 is evidence of this behaviour.
Each wave of recruitment indoors, as well as the single period of recruitment outdoors, (i.e. the growing season) increased the number of fronds by a factor of c. 1.5. This suggests that there were, at the beginning of the experiment, an equal number of dormant buds in the rhizomes of the populations and that the number of dormant buds an existing number of fronds can initiate for the next generation is restricted by something other than purely environmental
SIMAN & SHEFFIELD: POLYPODIUM VULGARE SPORULATION 37
factors. This could explain the occurrence of the proportionally similar waves of recruitment.
The present study shows that by enhancing the light and temperature it is possible to interrupt the strict reproductive cycle and induce continuous spor- ing ina pteridophyte population. Evidence of similar behaviour has recently been obtained in another study, in which dormant Pteridium rhizomes pro- duced fertile fronds within 13 weeks of being put into warm, well lit condi- tions (Wynn et al., 2000). Air samples taken in glasshouse and fernery envi- ronments in the UK do include ferns spores at all times of the year (e.g. Win- ston, 1998; Siman, 2000).
This study suggests that transfer of plants to glasshouses could benefit fern spore collectors by inducing continuous sporing in plants. There are less welcome implications of fern spore production in indoor environments, how- ever, especially for species that do generate vastly more spores in glasshouse settings than those in natural environments. A glasshouse is an enclosed en- vironment with little chance of biological particles being blown away by winds. This means that there is a higher risk of inhalation of fern spores in a fern-rich glasshouse than in most places outdoors. Based on the growing body of evidence of toxic and allergenic effects caused by fern spores (as reviewed by Simén et al., 1999, see also Siman et al., 2000), we suggest that some pro- tective measures (e.g. face masks) should be taken by people who regularly work in or visit indoor fern-rich environments.
ACKNOWLEDGMENTS
authors wish to thank David Newton for looking after the plants in the Experimental Grounds, Gareth Ballance for standing in to take records in those weeks when we were away an the University of Manchester Research Support Fund and the F. C. Moore Studentship Fund for financing the study.
LITERATURE CITED
ALLsopp, A. 1964. The metabolic status and morphogenesis. saab aplaiatinigs 14:1-27. ALLSOPP, A. 1965. The significance for development of water supply, osmotic relations and nutri- Handbuch der Pflanzenphysiologie 15:504—552. Srewene G. F. 1896. The probable influence of disturbed nutrition on the evolution of the veg- etative phase of the sporophyte. Amer. Naturalist 30:353-357. Conway, E. 1957. Spore production in bracken (Pteridium aquilinum (L.) Kuhn). J. Ecol. 45:273-
284 DRING, M. Si ba The influence of shaded conditions on the fertility of bracken. Brit. Fern Gaz. 9:2
EVANS, a ie The nature of flower induction. Pp. 457-480, in L. T. Evans, ed. The induction of flov gear Macmillan of Australia, Melbourne.
GOEBEL, K. 18 Uber kunstliche Vergrunung der Sporophylle von Onoclea struthiopteris. Ber. Deutsch. ae Ges. 5:69.
GOEBEL, K. 1905. Onecnouraglhiy of Plants. Part II. (English ed.). Oxford University Press.
GoEBEL, K. 1908. Einleitung in die experimentelle Morphologie der Pflanzen. Druck und Verlag von B. Me ype Leipzig
"sheng K. 1928. Organographie der Pflanzen. Teil I. png Jena.
S, A. ae oo, Hort. Week 28-Mar 1996:2
38 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 1 (2002)
Harvey, W. H., AND J. D. CAPONETTI. 1972. In vitro studies on the induction of sporogenous tissue on leaves of cinnamon fern. I. Environmental factors. Can. J. Bot. 50:2673—2682. LABOURIAU, L. G. 1958. Studies on the initiation of sporangia in ferns. Arq. Mus. Nac. 46:119-202.
PAGE, Cc. = 1976. The taxonomy and phytogeography of bracken—a review. Bot. J. Linn. ae 75s)
Boy Ay e 1979. Experimental aspects of fern ecology. Pp. 105-140, in A. F. Dyer, ed. The ex-
perimental papas of ferns. Academic Press, London bance E. 1996. From pteridophyte spore to lena it in the natural environment. Pp. 541— ar aa and R,J. Johns, eds. Pteridology in perspective. Royal Botanic Gardens,
SIMAN, . E. 2000. Fern spores and human health. PhD dissertation. University of conan eee SIMAN, S. E., A. C. POVEY, AND E. SHEFFIELD. 1999. Human health risks from fern spores?—a revie 87.
SIMAN, S. E, A.C. POVEY, ra Warp, G.P. MARGISON AND E. SHEFFIELD. 2000. Fern spore extracts can damage DNA. Brit. J. Cinice 83:69-73. STEEVES, T. A. 1959. An eriieeanias al two forms of Osmunda cinnamomea. Rhodora 61:223—
230.
STEEVES, T. A., AND R. H. WETMORE. 1953. Morphogenetic studies on eae cinnamomea L.: some aspects . “ general morphology. Phytomorphology 3:339
Sussex, I. M., AND T. A. STEEVES. 19 ie + aust on the control of ya of fern leaves in sterile culture. ot Gaz. 119:203-—
spas J. A. Morphometric and genomic diversity in the genus Pteridium (Dennstaedtiaceae).
n. Bot. 85:77-99. Waou E W., AND D. N. SHARMA. 1963. Experimental and analytical studies of pteridophytes. Bot. ne 101-121,
eho D. 1998. Risk assessment of environmental exposure to fern spores. M.Sc. dissertation, hota! of Mancheste
WRIGHT, B., AND A. WRIGHT. om The “Wright” way to collect and clean fern spores. Pteridologist 3:62 ote
Wynn, J. M., J. L. SMALL, R. J. PAKEMAN, AND E. SHEFFIELD, 2000, An assessment of genetic and envixonmental effects on sporangial development in img (Pteridium aquilinum (L.) Kuhn) using a novel quantitative method. Ann. Bot. 85:113-115.
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ee AMERICAN eee FERN aici
J O UJ R N A 1 April-June 2002
IN MEMORIAM: WARREN HERBERT WAGNER, JR.
Obituary: Warren H. Wagner, Jr. (1920-2000) Donald R. Farrar 39 Bibliography of Warren Herbert Wagner, Jr. David B. Lellinger 50
The Mating Systems of Some stele Polypodiaceae n-Liang Chiou, Donald R. Farrar and Tom A. Ranker 65
Belowground enseicras and Abundance of Botrychium Gametophytes and Juvenile Sporoph
ay Cindy Johnson-Groh, Chandra Riedel, Laura Schoessler and Krissa Skogen 80 Additions to the Fern Flora of Saba, Netherlands Antilles David B. Lellinger 93 Taxonomic Notes on Hawaiian Pteridophytes Daniel D. Palmer 97 Novelties in Pteridaceae from South America Jefferson Prado and Alan R. Smith 105
Is Gametophyte Sexuality in the Laboratory a — Predictor of Sexuality in N. m A. Ranker and Heather A. pnd 112
Additional Support for Two Subgenera of Anemia (Schizaeaceae) from Data for the Chloroplast Intergenic Spacer Region eo and Morphology Skog, E. A. Zimmer and J. T. Mickel 119
Intrafamilial Relationships of the Thelypteroid cy (Thelypteridaceae) n R. Smith and Raymond B. Cranfill 131
Two New Species of Moonworts (Botrychium subg. Botrychium) from Alaska Mary Clay Stensvold, Donald R. Farrar and Cindy Johnson-Groh 150
Isoétes X herb-wagne ri, an [ pecific Hybrid of I. bolanderi x I. echinospora
o
(Isoétaceae) W. Carl Taylor 161
Botrychium alaskense, a New Moonwort ea the Interior of Alask Warren Herb Wagner, Ir and Jason R. Grant 164
A New Name for an Old Fern from North Alabama James E. Watkins, Jr. and Donald R. Farrar 171
Continued Pteridophyte Invasion of Hawaii Kenneth A. Wilson 179
The American Fern Society
Council for 2002
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American Fern Journal 92(2
AUG 2. 9 2002 GARDEN LIBRARY Obituary: Warren H. Wagner, Jr. (1920-2000)
DONALD R. FARRAR
Department of Botany, Iowa State University, Ames, IA 50011
Warren Herbert Wagner, Jr. was born on 29 August 1920 and 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 Smith- sonian Institution, where he became acquainted with the experts, including
40 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
the eminent pteridologists William R. Maxon and Conrad V. Morton. In college at the University of Pennsylvania, he became the enthusiastic field compan- ion of Edgar T. Wherry, author of the “The Fern Guide.”” Wherry was a mineralogist who became an expert on fern habitats and the first to point out the important associations of epipetric ferns with particular rock types.
Graduating from the University of Pennsylvania in 1942, Herb entered the U.S. Naval 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 collecting ferns and butterflies, later publishing (with David Grether) ‘“Pteridophytes of Guam”’ (1948) as well as articles on the pterido- phytes and butterflies of the Admiralty Islands. During this time he also flew into California, taking his specimens to E. B. Copeland at Berkeley, re- nowned expert on Philippine ferns. This was the beginning of an association that would bring him back to Berkeley for graduate study. While in the Navy, he also began what was to become a life-long study of the ferns of the Hawaiian Islands.
At the University of California—Berkeley, in 1945, Herb joined student colleagues Charles Heiser, Ernest Gifford, Frank Ranzoni, Vern Grant, Art Krukeberg, and others in courses instructed by G. Ledyard Stebbins, Adriance Foster, and Herb’s major professor, Lincoln Constance. Copeland, although retired, was still active and advised informally on Herb’s research. Also among Herb’s student colleagues in systematic botany was Florence Signaigo. Herb and Florence married in 1948 and began a lifelong productive partnership in research and publication.
After receiving his Ph.D. in 1950, Herb spent a year as a Gray Herbarium Fellow at Harvard, then moved to the University of Michigan in 1951, where he remained throughout his illustrious career in teaching and research. He chaired or co-chaired the graduate programs of more than 45 Ph.D. students. He taught a variety of graduate and undergraduate courses, including his highly popular “Systematic Botany” and “Biology of Woody Plants,’ both of which he continued to co-teach after ‘‘retirement” (1991) through the fall of 1999. Herb’s public lectures and seminars were equally popular. Few biolo- gists have been in such demand as a visiting speaker. His curriculum vitae list of “invited lectures” totaled 169—since 1992!
From 1966 to 1971 Herb served as Director of the University of Michigan’s Matthaei Botanical Garden. His popularity with garden clubs, amateur bota- nists, and conservation groups made the Gardens a center of outreach activities in this arena, as well as a center for research and display of con- servatory plants. He chaired the Department of Botany in the Division of Bio- logical Sciences from 1974 to 1977 and chaired many additional department and college committees, including the University of Michigan’s Tropical Studies Committee from 1983 through 1997. He was chairman or president of nine professional societies, including the American Fern Society, American Society of Plant Taxonomists, and Botanical Society of America, and council member/trustee/advisor to dozens of organizations. He served as an editor for the Flora of North America, co-editing the Pteridophytes in volume two.
FARRAR: WARREN H. WAGNER, JR. 41
As reviewer of countless journal manuscripts, grant proposals, and botany/ biology programs, he gave freely of his time while continuing to teach and maintain a research program generating over 240 research publications.
“Probably the best-known botanist ever to work at the University (of Michigan),’”’ Herb Wagner’s influence on his science was immense. He has been referred to as “the founder of modern day systematics” in reference to his seminal contributions to cladistic analysis of phylogenetic relationships. His studies in the recognition of species hybrids, their value in understand- ing species relationships and their role in speciation and evolution became classics. His knowledge of ferns worldwide was phenomenal and allowed unique insights into fern ecology and life history, as well as systematics. Along with awards recognizing his contributions to systematic biology (including Willi Hennig Fellow, National Academy of Science, and the Asa Gray Award of the American Society of Plant Taxonomists), his name is indelibly inscribed in cladistic literature via the universally recognized ‘‘Wagner Tree’’ representation of phylogenetic relationships.
In writing about the career of Edgar Wherry, Herb stated that Wherry “was one of those rare individuals—a real naturalist.’’ Extended as the highest of compliments, the same could be said of Herb Wagner. His had an exception- ally keen ability to observe the small and intricate details of plants interact- ing with their environment. Study the knowable. Accumulate information on the parts. With time, dedication, and an open mind, the big picture will emerge. These gems of Herb’s philosophy attributed value to all research, no matter how small the project or whom the researcher. All good data were worth getting excited about—and he did. Herb’s ability to inspire others through his interest in their studies and their knowledge not only fostered independent research, but also created a legion of professionals and amateurs eager to contribute data to Herb’s projects. The total productivity of this syn- ergism, though unquantifiable, remains hugely visible.
Herb maintained a rigorous schedule of teaching, research, invited lectures and symposia presentations until just weeks before his death on January 8, 2000 at the age of 79. He was very much looking forward to participating in the 2000 Botanical Society of America symposium on the ‘Biology and Conservation of the Ophioglossaceae’’ that he helped to organize. In the summer of 1999, Herb and Florence conducted field work in Alaska and in southwestern Canada. From both places they returned with, of course, new species of Botrychium.
The foregoing highlights of the career and achievements of Warren ‘‘Herb”’ Wagner fall short in communicating the extraordinary nature of his per- sonality, his gift for teaching, and his full influence on pteridology and pterido- logists of the last half-century. The more personal reminiscences that follow portray a rare individual whose life we are all so fortunate to have shared.
ACKNOWLEDGEMENTS
Information on the early years of Herb’s career were graciously provided by Florence Wagner. Factual information is taken from Herb Wagner’s 1999 curriculum vitae. Other observations
42 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
extend from my long association with the Wagners, as a graduate student in Ann Arbor and in many subsequent field trips and discussions of plants, people, and philosophy.
Portions of this article are excerpted with permission from Taxon 49:585-592. August 2000, “Warren H Wagner, Jr. (1920-2000),” by Donald R. Farrar. Quotations are by Julian P. Adams and William R. Anderson of the University of Michigan in The University Record 55:2-4. January 17, 2000, ‘Warren H. (‘Herb’) Wagner,” by William R. Anderson.
REMINISCENCES
Meeting Herb Wagner.—I first met Warren Herb Wagner, Jr., on the eve- ning of 13 June 1980, at Flathead Lake Biological Field Station in far north- western Montana. This meeting has stuck in my mind, perhaps because in many respects it typified my relationship with him. I was a new student in pteridology and had come to Flathead Lake to take his (and Florence’s) four- week-long fern course. Earlier that year I had phoned Herb to tell him that I had enrolled in his course and to let him know about a new, unnamed fern hybrid I had found in the Shawnee Hills of southern Illinois, a cross be- tween the walking fern (Asplenium rhizophyllum) and the maidenhair spleenwort (A. trichomanes). As it turned out, Herb was a few days late for the course because of a conflict with the final days of the International Bota- nical Congress in Vancouver. He finally arrived at the biological station about 10:30 p.m., when I was alone in the lab and identifying plants. The lab door suddenly opened and Herb burst into the room. He walked about half its length before acknowledging my presence. We introduced ourselves. He explained that he had been collecting moonworts all day en route to the field station, but it didn’t seem like it to me. Instead of being exhausted after a long day of fieldwork, he was animated and lecturing me about the biology of moonworts and ferns in general. I listened spellbound and thrilled that he would take the time to explain it all to me. “Let’s see your hybrid fern,” he demanded. “It’s in my cabin,” I replied, ‘and my cabin is a long way away, and it’s pitch-black outside, and I’ve lost my flashlight. Can I show it to you tomorrow?” Then—and I'll never forget this—he gave me the most odd sup- plicating look, and with hands clasped together as if praying, he said, ‘““Won’t you please go get it—now!”’ How could I refuse? I stumbled back to my cabin in darkness, found the specimens, and retraced my steps to the lab. Herb examined the specimens. “Yes, that’s it! Asplenium rhizophyllum xX A. trichomanes. Congratulations!’” He shook my hand; I had received his imprimatur. Then, without further ado, he described how he wanted the lab rearranged—by me, that is. Tables, benches, plant presses, cardboard divid- ers, microscopes—all these were to be moved according to a plan that he had devised while I was retrieving the specimens. After explaining where every- thing should be relocated, he said, ‘‘Ok, gotta blow!” and he left the lab as abruptly as he had entered. I was once again alone in the lab, excited by what I had just learned about moonworts and spleenworts, and too motivated to mind having been pressed into service.—Robbin C. Moran, The New York Botanical Garden, Bronx, NY 10458-5126.
FARRAR: WARREN H. WAGNER, JR. 43
Herb at ‘“‘Bug Camp’’.—My first memory of Herb Wagner is from Spring, 1952. During my senior year at the University of Michigan, about to graduate with a major in Biological Science, I worked as an undergraduate lab assis- tant in botany. That job involved lugging five-gallon bottles of distilled water, spraying the greenhouse for insects, and other tasks. One day I was called in to meet a new faculty member, Dr. Warren H. Wagner, Jr. He asked me to be his graduate assistant that summer at the bug camp, the University of Michigan Biological Station at Douglas Lake. With no other plans, I agreed
That marked my beginning as a botanist. I assisted Wagner in courses in Phycology and Pteridology. To be a graduate student, I had to take a course, so I signed up for an independent study with him. Since the genus Equisetum was well represented around the bug camp, Wagner assigned me to work with its taxonomy. Herb showed himself to be an enthusiastic, knowledgeable field botanist. I well remember, after a hard day bent in a half-crouch searching out Botrychium, stopping with him for ice cream on the way back to Douglas Lake. And, I remember evenings all of us gathered around a piano to hear him play. Most evenings were spent in the lab until 11 p.m. or so, working with Herb, on the materials gathered during the long days in the field.
That summer, Herb advised me to go to some other university for my mas- ter’s degree, to widen my exposure to the whole field of botany, with the understanding that I would return to Michigan to work for my doctoral de- gree under him. I went off to Florida State University as a graduate research assistant on a tidal marsh study, then to the University of California, to study the anatomy of Equisetum stomata with Dr. Adriance Foster and get an M. A. in Botany. After a stint in the U. S. Army, I returned to Ann Arbor, to my mentor, Dr. Warren H. Wagner, Jr., and to the taxonomy of Equisetum.—Richard L. Hauke, 900 N. Stafford St., Apt. 1103, Arlington, VA 22203-1844.
Unflappable Herb.—If you know something of Herb’s activities during World War II, it is no surprise that he was unflappable. He had been an offi- cer in the U. S. Navy, a navigator on air flights, and for a time was based on Guam. Like the pilots, he was not called upon to fly every day; on his days off he collected butterflies and ferns. After the end of the war, he and David F. Grether published an account of the pteridophytes of Guam based princi- pally on their collections and those of other U. S. military personnel. (Occ. Pap. Bishop Mus. 19(2):25-99. 1948). He was a native of Washington, DC, and his butterfly collection was given to the Smithsonian Institution.
The best collecting for butterflies, Herb found, was beyond the American defense lines that were maintained by Marine guards and patrols. Some well armed Japanese troops were still at large beyond those lines, along with numbers of their fallen colleagues who were very attractive to butterflies. Because of the danger, the Marine guards didn’t like Herb’s excursions, but as an officer, he was able to go where he wanted. A butterfly net was of the utmost importance to those excursions. It was the equivalent of a white
44 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
flag that signaled the bearer’s non-combatant status. Even so, there must have been a considerable element of danger, although Herb said he was never shot at by the Japanese while carrying a net. (A plant press, even loaded with ferns, did not confer the same protection, presumably because its function was unknown to the Japanese soldiers.
In Ann Arbor, Herb’s spring wildflower course was always popular. His humor, knowledge, and enthusiasm were delightful. Who could forget his earnest description of a hand lens and how it was to be used and worn around the neck? He then pulled from within the lectern a large hand lens attached to a red ribbon so wide that it was practically a sash! Although Herb was generally unflappable, I do remember one exception that was, de- pending upon your point of view, hilarious.
The year I assisted him, his lectures were presented in three classrooms that had been converted into a long, narrow lecture hall. The slide projector, which I manned, was placed well back in the hall, perhaps 40 feet from the desk, lectern and screen at the front. In order to signal for a change of slides, Herb would hold a wooden pointer vertically and drop it dramatically on the concrete floor next to his feet. Although none of us knew it at the time, the repeated thunk was not popular with the Dept. of Geology professors who had offices on the floor below.
One afternoon, Herb called for darkness. The shades were drawn while Herb turned out the room lights. I turned on the projector and put the first slide on the screen. Naturally, I was looking at the projector when Herb drop- ped the pointer to request the second slide. There was a tremendous det- onation in the front of that quiet room. My head snapped up in time to see Herb still coming down behind the desk after what must have been a terrific leap, nearly a pole vault. He strode over to the door, clicked on the lights, took out and lit with trembling hands one of the small cigars he smoked at that time, inhaled deeply, and said ‘‘Take a five-minute break.” Of course, the students could not contain themselves. By evening, most of the graduate students knew what had happened: one of them had taped a few caps from a cap pistol to the blunt end of the pointer. After the recess and cigar, Herb’s lecture and slides continued as before, punctuated only by an ordina thunk. The Geology professors were not at all amused.—David B. Lellinger, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0166.
Graciousness and Support.—Like most graduate students back in the 1970's, it wasn’t long before beginning my study of ferns that I was introduced to Herb Wagner. My first impressions were that he was brilliant, dynamic, mesmerizing, and exactly the type of scientist that I wanted to be. Of course, that wasn’t possible because he was certainly one of a kind. He reviewed one of the first papers that I wrote concerning meiotic studies in Ceratopteris. I remember that the review was less than favorable about some aspects of the study and of the way that it was presented. He was right, of course, but for some reason, perhaps extensive re-writing, it eventually was published.
FARRAR: WARREN H. WAGNER, JR. 45
A couple of years later, | wrote a paper that made an attempt to propose an alternative explanation for the pattern of reticulate evolution that Herb had described for Appalachian Dryopteris. This was the last piece of my doctoral dissertation. I soon graduated and while I was at work as a young assistant professor, I actually received a reprint request for this paper from Hugh Iltis with a big “‘congratulations”’ scrawled on it! But any glory was to be short-lived. A couple of years later, during a meeting of the Southeastern fern group at the Duke Marine lab, I remember sitting quietly in the front row as Herb methodically destroyed my brash interpretation of Appalachian Dryopteris. | was humbled, but not humiliated. He was far too gracious for
at.
Although I was not one of his students in any type of academic lineage, he was a constant and helpful influence in my career. Letters of recommen- dation for jobs and promotions, reviews of my academic department, and oc- casional suggestions spanned the course of over 20 years. My last personal encounter with him was during a visit to Knoxville in the late 1990’s where he gave an inspiring seminar to a group of faculty and students. I was teach- ing a sophomore genetics class that semester and made a point of inviting my students to hear his seminar. Although they weren’t botany majors and generally had very little interest in plants, it was obvious that he had not lost his charm, dynamism, and ability to mesmerize even these supposedly disinterested young people in his story about moonworts! I keep a small pic- ture of Herb behind me in my office and often turn and sneak a peek at him for inspiration when I am trying to figure out how to do a better job of pre- paring a lesson or writing up a laboratory exercise.—Les Hickok, Department of Botany, The University of Tennessee, Knoxville, TN 37996.
Humanity and Professionalism.—He combined intellect with a humanity and friendliness. Herb Wagner, as we called him when he befriended us, traveled widely and infected nearly everyone he met with an enthusiasm for life and for inquiry into the natural world.
His influence upon me came early in my professional career. While a stu- dent at the University of Kentucky between 1961 and 1963, Herb had already visited the campus, and as always, took a field trip into the Kentucky hills. I was told that an old man walking along a road of a cove yelled, ‘“‘Hey, what are you fellers doing down there?’ Herb’s immediate response, ‘‘We’re just botriculating.”” And though suspicious of revenuers, the old man responded, “Oh well, go right ahead.”
It was shortly after I arrived in Cullowhee in 1969 that Herb made a visit to Highlands Biological Station in preparation for the fern conference in 1970. He had come across a paper that Jim Horton and I published listing new county records that included Asplenium heteroresiliens, a hybrid on marl known only from the coast. He visited our herbarium and in his in- structive but gentle way said, ‘I see why you identified Asplenium tricho- manes for heteroresiliens, but it isn’t.” Our somewhat depauperate and poorly dried specimen didn’t exactly match either species. Herb also got me
46 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
to be more careful in my identification of other species, pointing out that what appeared to be Populus grandidentata growing beside the science build- ing was something else, which we later identified as Populus X canescens.
In the early 1970’s Herb was President of the American Fern Society, pop- ularizing the profession. About this time I had a free evening and took in the movie, “A New Leaf,” a spoof of a millionaire and his attraction to a wall- flower type of lady who happened to be a student of ferns. To my amaze- ment, she came to the point of verifying her identity that the tropical fern she discovered in Alaska was confirmed by ‘‘Dr. Wagner at the University of Michigan.” Such was his fame as it expanded beyond the ivory tower into society in general.
Herb visited Cullowhee many times. On his last trip here in April, 1997, not only did he give the visiting scholar lectures but also wanted to return to some nice place before he returned home. His 4 p.m. seminar that Friday afternoon was so well received that a couple of visitors, Bob Dellinger, local naturalist, and Gary Kauffman, USDA Forest Service botanist, asked to tag along with us the next day to the Joyce Kilmer area. Although Herb had in- dulged in country ham the previous days (he was supposed to be on a low salt diet) and was a bit tired from the activities the previous two days, he de- lighted in finding Carex austro-caroliniana with peduncles several centi- meters long. Such was his exuberance over those things he found fascinating.—Dan Pittillo, Department of Biology, Western Carolina Univer- sity, Cullowhee, NC 28723.
In His Element.—When | think of Herb, I tend to think of him in a field trip setting. For instance, when he was invited to the annual meeting of the Field Botanists of Ontario in Midland, Ontario a number of years ago, I re- member the lead car leaving the sandy gravelly country road and entering an open sandy field with a few copses of junipers. In what resembled a “cops and robbers” film on T.V., the lead car had not come to a full stop when Herb rolled out the front door of the car. He took a giant step towards the copse and was almost immediately flat on the ground under the prickly Juni- per. Suddenly there was a victory call for all and sundry within 200 yards— “Botrychiums.” All Herb’s followers were on the spot in a few seconds and each looked in marvel at some very small specimens between the juniper and the sand—a gap of less than a foot. Herb was in his element in the field, and his disciples enjoyed every minute of it!—D. M. Britton, Professor Emer- itus, Department of Molecular Science & Genetics, University of Guelph, Guelph, Ontario, N1G 2W1 Canada.
One of the All-time Giants.—My first contact with Herb Wagner came about in early 1978 when I was at the University of Cambridge, England. I had been intrigued for some time by a note in Herb’s paper on spores in rela- tion to fern phylogeny (Ann. Missouri Bot. Gard. 61: 346. 1974) on the pres- ence of large spores in ferns in the higher altitude rainforests of Hawaii not being linked to polyploidy. I had noted that in the New Zealand species of Grammitis (Grammitidaceae) there is a trend towards the production of
FARRAR: WARREN H. WAGNER, JR. 47
larger spores, and frequently fewer spores per plant, in the species occurring at higher altitudes and latitudes. Although few chromosome counts are avail- able this does not seem to be linked to polyploidy. I had assumed that this wasn’t a parallel of the strategy of higher plant taxa that at higher altitudes produce fewer and larger seeds to contain resources needed during longer periods of dormancy, because dormancy is not an issue with Grammitid spores, which are photosynthetic. Perhaps Herb’s ‘‘selection for precinctive- ness’’ was working here? So I wrote to him concerning my observations; he promptly replied that he thought it was, and that in Hawaii, for example, any spore or any propagule that is especially likely to be carried by wind is going to lose out. His observations on butterflies had some part in his logic, with the most conspicuous Hawaiian butterfly being a very sluggish inhabi- tant in mountain valleys, rather than ridges, where it was likely to be blown away. I’m still very much undecided about selection for precintiveness in Grammitidaceae after further work, as the high altitude New Guinea species do not show the same increased spore size as the New Zealand ones, but Herb’s prompt, encouraging and discursive response to a junior unknown pteridological correspondent made a deep and lasting impression on me that here was one of the all-time giants of pteridology.
Nine years later, when I had moved to the Royal Botanic Gardens, Kew, I finally met both Herb and Florence at the International Botanical Congress in Berlin in 1987, and the impression I had of Herb from his correspondence was strongly reinforced. He enjoyed ferns and people and talk, and I really envied all of his students for having such a stimulating mentor! A talk with Florence at Berlin turned up a surprising coincidence—one of my M. Sc. supervisors at the University of Auckland, New Zealand, Prof. Jack Ratten- bury, had been best man at Florence and Herb’s wedding—it’s a very small botanical world.—Barbara Parris, Fern Research Foundation, 21 James Kemp Place, Kerikeri, Bay of Islands, New Zealand.
Tutor and Friend.—Herb Wagner was my tutor and friend during the last dozen years of his life and he assisted me in gaining the skills needed to study the Hawaiian ferns. Herb worked with the Hawaiian fern flora for more than half a century, during which time he greatly increased the know]l- edge of its diversity and biology. He described more than three dozen new species, varieties, and hybrids. With his contributions, our organized under- standing of the Hawaiian ferns was much advanced.
Herb made several long visits to Hawaii during which he went on exten- sive field trips and spent much time in local herbaria. During these visits he taught courses reviewing the Hawaiian ferns and took his students on field excursions. A remarkable and enthusiastic teacher, the passionate sharing of his deep knowledge of Hawaiian ferns inspired many in Hawaii to study this neglected group of plants. His lectures and seminars were remarkable for his enthusiastic, thorough, and effective presentation of material to students with only a minimal knowledge of pteridophytes. I will remember the ballet like pirouettes he would do at the blackboard when emphasizing a point,
48 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
and I remember the vast collection of anecdotes and jokes he used to enliven a lecture and to help students remember the subject under discussion. He would not let the decreasing physical vigor he experienced in later years interfere with active field work. On his return to Ann Arbor from Hawaii he always left behind a whirlwind of enthusiasm for the study of Hawaiian ferns.
Florence Wagner’s collaboration and organizational abilities allowed the Wagner family to effectively pursue the twin goals of teaching and research. Many of us in Hawaii will fondly remember a day’s end with Herb and Florence, where a good dinner, cocktails, stimulating conversation, and en- joyable company combined to make for a very pleasant evening.
He is missed by all who knew him in Hawaii. His stimulating visits will be missed and the time he spent here will be remembered here with nostalgia.—Daniel D. Palmer, 1975 Judd Hillside, Honolulu, HI 96822.
Missing Herb.—I miss Herb too much to find words to describe how misera- ble I am at knowing he is not at the end of the phone needing a this or a that, or welcoming my weird plant ID, or to confirm one of those peripheral vegeta- tive idioblastic forms that my heart hoped might be something new. I remem- ber the summers he visited Iowa Lakeside Lab, the AFS field trips, particularly the one in northern Michigan. I did not know Herb from Michigan as many did, as he was my academic grandfather—and father in Iowa by remote-control, a fitting way to be for us hybrid studying types. He thought that my following Darwinian fates of individuals and natural history approach to fern reproduction and population structure was significant when most around me spoke only of things homoeologous, bar codes of molecules on gels, boot- strapping clades, and such. In comfort, Herb offered that he often no longer knew what cladistic people meant with the language they used to describe his ground-plan divergence method he created so long ago. I only wanted to be a teacher, and Herb’s way with a gathering of students still is my exemplar. We will all miss him; he will always be with us.—James H. Peck, Department of Biology, University of Arkansas—Little Rock, Little Rock, AR 72204.
A Personal Gift.—I didn’t go to the University of Michigan to study with Herb Wagner. My interests were in plant physiology, but in my first semester a fellow graduate student, Bob Faden, who was then a student of Herb’s, secured an invitation for me to accompany the annual Wagner entourage to the hills and canyons of southern Ohio and Kentucky. Herb had just pub- lished papers on the independent occurrence of gametophytes of Vittaria and Trichomanes in the eastern United States. We had no trouble finding great mats of Vittaria gametophytes and, with some squirming on our backs under overhanging sandstone cliffs, we also found the green, threadlike gametophytic filaments of Trichomanes. Returning with collections of these intriguing plants, I was thrilled and convinced that with my growing physiological expertise I would coax them in culture into producing sporo- phytes and would resolve the mystery of their independent (without sporo- phyte production) occurrence in the wild. Nearly 40 years later I still work on that.
FARRAR: WARREN H. WAGNER, JR. 49
After two years of graduate school I was having doubts about my future in academia. With considerable trepidation I informed Herb of my intention to take some time off and join the Peace Corps. To my surprise, he supported my decision, but then gently reminded me that I was now the ‘world’s expert” on independent gametophytes and I ‘‘owed it to science’ to stay one more year to publish what I had learned. By the time I did that, of course, my thinking had changed and I was forever hooked on fern gametophyte biology. Herb probably expected that outcome, but it wouldn’t have hap- pened without his encouragement and confidence in me.
The excitement of field trips with Herb and Florence remain highlights of my graduate years at Michigan, as do warm memories of holidays at the Wagner home, cutting out snowflakes or whatever project Florence had de- signed for their “extended family” of graduate students. This nurturing of an Ozark farm boy a long way from home made a difference. It was a personal gift. Yet I know it was only one of many such personal gifts, bestowed on many others as well, by Herb and Florence. For those gifts we all say thanks!—Don Farrar, Department of Botany, Iowa State University, Ames, Iowa 50011.
American Fern Journal 92(2):50-64 (2002)
Bibliography of Warren Herbert Wagner, Jr.
Davin B. LELLINGER Department of Botany, Smithsonian Institution, Washington, DC 20560-0166
Asstract.—This bibliography contains 395 entries, all the scientific publications of Warren H.
Exact publication dates were found for most journal articles. However, month or estimated dates were used for other lire eciseem books, which usually are - datable within a = All the entries are contained in a searchable database now maintained by Florence S. Wagne The eral categorizes the ee according to their subject matter and includes a more or less exact publication dates that are the basis for the onan list.
Wacner, W. H., Jk. 1941. New localities for Botrychium matricariaefolium in Maryland. Amer. Fern J. 31:21—22.
———.. 1941. Butterfly hunting: Why and How? The Naturalist 1:6 1941. [Mimeographed publi- cation of the Naturalists Field Club of the University of Pennsylvania]
. 1941. District of Columbia Butterfly Notes (Lepidoptera: weit tale Entomol. News
52: 196-200, 245-249. The parts were published 18 Jul and 8 Oct
. 1942. Bipinnate Christmas ferns. Amer. Fern J. 32:27—29.
9—70.
ia)
1943. New locality for a rare Hairstreak (Lepidoptera: ane Entomol. News 54:11. 1943. scat: sae by remote control. Amer. Fern J. 3 7
944, rms new to Trinidad. Amer. Fern J. 34 oe 1944, pare occurrence of the apparent hybrid Cystopteris. Amer. Fern J. 34:125—127. 1945. Fern hunt in Puerto Rico. Amer. Fern J. 3 1945. Ferns on pacific island coconut trees. Amer. Forn J. 35:74—76. 1946. Fern field notes in the Washington—Baltimore area. Castanea 11:59-60. . 1946. Notes on the protection of rare ferns in the Wochisister Malti asces area. Bull. on dis Washington-Baltimore Area Flora 10:5-6. [Mimeographed]
- 1946. Botrychium multifidum in Virginia. Amer. Fern J. 36:117-1 : 1946. Pteridophyta. Pp. 3-8 in Hermann, F. J. A checklist of olnnts of the Washington— Baltimore area, ed. 2. [Mimeographed . 1947. Tree-climbing Gleichenias. Amer. Fern J. 37:90-95
& D. F. GRETHER. 1948. Pteridophytes of Guam. Occas. Pap. Bernice Pauahi Bishop Mus. 19(2):25-99,
———.. 1948. A new fern from Rota, Mariana Islands. Pacific Sci. 2:214—-21 RETHER. 1948. The Pteridophytes of the Admiralty Islands. Unity, Calif. Publ. Bot. 23(2): 17-110, t. 5-25.
& D. F. GreTHer. 1948 [1949]. The butterflies of the Admiralty Islands. Proc. U. S. Natl. Mus. 98:163-186. ——.. 1949. A reinterpretation of Schizostege lidgatei (Baker) Hillebrand. Bull. Torrey Bot.
Club 76:444—461.
. 1950. The Hart’s-tongue Fern. 18th Annual Calif, Spring Garden Show, April 21-28, pp. i,
=
| . | | | |
Re |
1950. The habitat of Diellia. Amer. Fern J. 40:21—
- 1950. Ferns naturalized in Hawaii. Occas. Pap. ae Pauahi Bishop Mus. 20(8):95— 121.
- 1951. A new species of Diellia from Oahu. Amer. Fern J. 41:9-13.
LELLINGER: BIBLIOGRAPHY OF WARREN HERBERT WAGNER, JR. 51
. 1951; Sie pe a analysis of evolution in Pteridophyta. [Review of: Manton, I. 1950. Problems of Cytology and sig in the Pteridophyta. Cambridge Univ. Press, London
sir sire York.] Ss. oe 5:177—
951. Review of: Manton, I. pi Problems of Cytology and gigerton ie in the Pterido-
ha Cambridge Univ. Press, London and New York. Amer. Fern J. 4 93. . 1952. The flowerless plants. [Review of: Billington, C. 1952. Fems of Michigan. Cran- — Institate oe Science, Bloomfield Hills, MI.] Sci. Monthly 75(5):3 rm Genus Diellia: Its Structure, Affinities and Ao Univ. Calif. Publ. ree a) leah t. 1-21.
. 1952. Review of: McVaugh, R. & J. H. degen 1951. Ferns of Georgia. Univ. of Georgia mae pe Michigan pee 58(21):366— ———. 1952. Review of: Billington, C. 1952. caine of Michigan. Cranbrook Inst. of Science,
Bloomfield Hills, MI. Cranbrook Inst. Sci. News Letter 22(1): ee
. 1952. Juvenile leaves of two Polypodies. Amer. Fern J. 42:8 . 1952. Types of foliar dichotomy in living ferns. Amer. J. i pe 578-59 92. . 1953. The genus Diellia and the value of characters in determining fern affinities. Amer. J. Bot. 40:34—40. 2 1953; An Asplenium prototype ne be genus Diellia. one — Bot. Club 80:76-94. . 1953. Report of the Secretary for 2. A . 1953. A cytological study of the pam Spleonw i re mer. Fern J. 43:109-114 1953. Review of: Hubbard, D. H. 1952. Ferns of Hawaii National Park. Hawaii Nat. Hist.
——— & D. J. HaGenan. 1954. A natural hybrid sy et apaerias lonchitis and P. acrostichoides en the Bruce Peninsula. Rhodora 56:1-6, t. 4. A Bolivian Elaphoglossum of unique “leaf structure. Bull. Torrey Bot. Club 81:
1954. Report of the Secretary for 1953. Amer. Fern J. 44:21-24 1954. Reticulate evolution in the Appalachian Asphiiiana, Evolution 8:103-118 [Reprinted as pp. 136-151 in Orndoff, R. 1967. Papers on Plant Systematics. Little, — Boston. ] oes The evidence used recent classifications of the ferns. Rapports Comm. aux Sect. : , 6, Ville Congr. Int. B WILSON, - ci & W. H. WAGNER he pris a of: Lawalrée, A. 1950. Flore cnn de Belgi- que: Pt spy Jardin Botanique de |’Etat, Bruxelles. Amer. Fern J. 4 Wacner, W. H. Jr. 1955. Cytotaxonomic observations on North American “on phen 57:219- 240. . 1955. Should - American Hart’s-tongue be interpreted as a distinct species? Amer. Fern J. 45:127-1 . 1955. Austin ie Clark. re News 9(4,5):151-157. . 1955. Review of: Manton, I. . A. Sledge. 1954. Observations on the cytology and acne of the sean sh For Ceylon. Phil. Trans. Roy. Soc. London, Biol. Sci. 238:127-185. Amer. Fern J. 4 . 1956. A reoviany rege mii en a C. C. Hall. Madrofio 13:195—205 D. J. Hac . 1956. A diploid variety in the Cystopteris fragilis sienpies Rhodora
58:79-87, t. ee. . 1956. Asplenium ee x ot gla a new triploid hybrid produced under arti- fical conditions. Amer. Fern J. 4 Voss, E. G. & W. H. WAGNER . 1956. in on Pieris virginiensis and Erora laeta—two butterflies hitherto enikciovted from Michigan. Lepidopterists’ News 10:1 & L. P. Lorp. 1956. The morphological and cytological distinctness of Botrychium minga- swe and B. lun in Michigan. Bull. Torrey Bot. Club —280. D. J. emia oe [1957]. a a on some tralia: -producing populations of the Cystopteris fragilis complex. Amer. Fern J. 46:137-146. 1957. Heteroblastic leaf moepholgy in juvenile plants of Dicranopteris linearis (Gleiche- tincead). Phytomorphology 7
52 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
. 1957. Botanical Phase. Pp. 1-14 in J. M. Sheldon & E. W. Hewson. Atmospheric Pollu-
tion by Aeroallergens, Progress Report No. 1. Eng. Res. Inst. 2421-1-P, Ann Arbor, MI
J. DaruING, JR. 1957. Synthetic and wild Asplenium gravesii. Brittonia 9:57-63.
STEINER, E., A. S. SUssMAN & W. H. WacnerR Jr. 1957. Botany Laboratory Manual. Dryden, New York
Wacner, W. H. Jr. & R. S. WHiTMIRE. 1957. Spontaneous production of a morphologically distinct, fertile oo by a sterile diploid of Asplenium ebenoides. Bull. Torrey Bot. Club
seg i is GILBERT, 1957. An unusual new Cheilanthoid fern from California. Amer. J. Bot. : 743. Constance, L., H. Lewis, R. C. Rouuins, R. F. THORNE & W. H. WAGNER JR. 1958. Suggested outline for at aprons botany. Pl. Sci. Bull. 4(1):1-3 esis ni H. Jr. 1958. Notes on the distribution of a kentuckiense. Amer. Fern J. 9-43.
eee Botanical Phase. Pp. 1-13 in J. M. ci & E. W. Hewson. Messiah sone Pollu- tion rc neering a, Progress Report No. 2. Eng. Res. Inst. 2421—-2-P, Ann Arbor, & EA 8. Perennial ragweeds fAnabocetia) in Michigan, with the lem of a new, vito Si ie Rhodora 60:177—204 1958. Review of: Bold, H. C. 1957. Marntolosy of Plants. Harper, New York. Science 128:1079. Dincte, A. N., J. C. Gin, W. H. Wacner Jr. & E. W. Hewson. 1959. The emission, dispersion and deposition of ragweed pollen. Adv. Geophys. 6:367—387 Wacner, W. H. Jr. 1959. Botanical research on atmospheric pollution. Proc. 13th Ann. Meeting Northeastern Weed Control Conf., Rutgers Univ.:262-268. [Mimeographed] K.
wood and its relationship to a peculiar plant from West Virginia. Amer. Fern J. 48:146— 159.
- 1958 [1959]. The hybrid ragweed, Ambrosia artemisiifolia x trifida. Rhodora 60:309-
- 1959. Problems in the classification of ferns. Proc. IXth Intl. Bot. Congr. 2(Abstrs.): 418-419.
CanTLON, J. E., W. H. WAGNER JR. & W. is Riripon 1959. A range extension in Arctic Alaska for cares lunaria, Amer. Fern J. 49:25-29.
Wacner, W. H. Jr. 1959. Hybrid swarms of mittee [Review of: Walker, T. ss 1958. Hybridization in some — of Pteris L. Evolution 12:82—92.] Amer. Fern J. 49:4
, ed. 9. An annotated bibliography of ragweed (Ambrosia). a Ailacay Appl. Immu-
nol. . ae
- 1959. Botanical phase. Pp. 1-22 in J. M. Sheldon & E. W. Hewson. Atmospheric Pollu-
tion ky Aeroallergens, Progress Report No. 3. Univ. Mich. Res. Inst. 2421-3-P, Ann
MI.
. SCHWEMMIN & W. H. Wacner JR. 1959. Pollen release in the common ragweed todanaie cape tagioersd Bot. Gaz. 120:235—243. W. . 1959
can Grapeferns resembling Botrychium ternatum: A preliminary
. 1960. Evergreen Grapeferns and the pres of infraspecific categories as used in North American Pteridophytes. Amer. Fern J. 50:32—45.
. 1960. The proliferations of Asplenium pipet Castanea 25:74-79.
. 1960. Ragweeds. Cranbrook Inst. Sci. po Letter 30(1):2-8.
- 1960. Botanical phase. Pp. 1-25 in J. M. Sheldon & E. W. Hewson. a ao tion by sy gi Progress Report No. es Univ. Mich. Off. Res. Admin. 034 Ann
. 196 ae and pigmentation in aa subg. Sceptridium in the northeastern United ‘Staten, Bull. Torrey Bot. Club 87:3
961. Filicineae, Frond. Pp. 396-398, pooh in P. Gray, ed. The Encyclopedia of the nies Sciences. Reinhold, New York.
ee 9
LELLINGER: BIBLIOGRAPHY OF WARREN HERBERT WAGNER, JR. 53
. 1961. Some new data on the vernation differences of Botrychium dissetum and B. terna- tum. Amer. Fern J. 51:31-— . 1961. Problems in the elaneification of ferns. Pp. 841-844 in Recent Advances in Botany,
———. 1961. On the relative development of the fertile segments in Botrychium dissectum and B. ear Amer. Fern J. 51:75-81.
1961. The Goldback Ferns of California. Review of: Alt, K. S. & V. Grant. 1960. Cytotaxo- nomic observations on the Goldback Fern. Brittonia 12:153-170. Amer. Fern J. 51:105-106. . 1961. Spore studies and speciation in the ferns on Hawaii. 10th Pac. Sci. Congr. Abstrs. Symp. Pap. 135-136 . 1961. Roots and the naa differences between Botrychium oneidense and B. dissec- tum. Rhodora 63:164-1 & K. E. Boypston. io, en new hybrid showing homology between Asplenium ebenoides and A. pinnatifidum. Brittonia 13:286—289.
961. Nomenclature and typification of two Botrychiums of the southeastern United States, Taxon 10:165-169.
1 Review of: Wherry, E. T. 1961. The Fern Guide. Northeastern and Midland Eiesiliie States and Adjacent Canada. Doubleday, Garden City. Amer. Mid]. Naturalist 67:
& D. J. HaGENAH. 1962. Dryopteris in the Huron Mountain Club area of Michigan. Brit- a 14:90—100. 962. Cytological observations on Adiantum xX tracyi C. C. Hall. Madrofio 16:158—161. ae, grein The endemic Botrychiums of the southeastern United States. ASB Bull. 9:40. Morzentl, V. M. & W. H. WacNER Jr. 1962. Southeastern American ‘‘Blackstem spleenworts” of 0-41
Wacner, W. H. Jr. 1962. Plant co —— and leaf production in iene multifidum “‘ssp. icum’ and ‘forma dentatum.’’ Amer. Fern J. 52:1-18. . 1962. The synthesis and expression of nou pennies data. Pp. 273, 276, 277 in L. Benson. 1962. Plant Taxonomy: Methods and Principles. Ronald Press, New York. . 1962. A graphic method for orn inka: based upon group correlations of indexes of divergence. Pp. 415 n L. Benson. Plant Taxonomy, Methods and Princi- ples. Ronald Press, New York. . 1962. Irregular morphological development in hybrid ferns. Phytomorphology 12:87—100. . 1962. Ferns and their allies. [30-minute film] AIBS Secondary School Biological Sciences Film Series, McGraw-Hill 962. oe crosses and spore characteristics in the genus Psilotum (Psilotaceae). ages J.B 962. soiree boschianum in Arkansas. Amer. Fern J. 52:85—86. Geanen, ri I., W. W. Payne & W. H. WacNner JR. 1962. Botanical phase. Pp. 1-16 in J. M. Sheldon & E. W. Hewson. Atmospheric Pollution by Aeroallergens, Progress Report No. 5. Univ sch: Off. Res. Admin. 04202, 04771, 04772, 04773-1-P, Ann Arbor, MI. Wacner, W. H. Jr. & D. E. RAWLINGS. 1962. A sampling of pins agen Subg. Sceptridium in the vicinity of Leonardtown, St. Mary’s Co., Md. Castanea 27:132-1 toa ips and taxonomic categories in lower mated plants. Pp. 63-71 in V. H. Harwue . Léve, eds. Symposium on Biosystematics. Intl. Assn. for Plant Taxono- my, Utrecht, ie ds. . 1963. A biosystematic survey of United States ferns—Preliminary abstract. Amer. Fern J. 53:1—16. . 1963. Gros and herbarium studies, University of Washington, June 23-28, 1963. [Mimeo- — 12p 5 ee raat 1963. gee prothallial clones (Trichomanes?) locally abundant on Illinois Reon Amer. J. B 0:623. 1963. Pteridology in oie Newslett. Hawaiian Bot. Soc. 2(8):117-123. [Mimeo- graphed]
54 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
J. SHARP. 1963. A cnet reduced vascular plant in the United States. Science 142:1483-1484, cover picture . 1963 [1964]. Pteridaphytes of the concn mye area, Giles County, Virginia, including notes from Whitetop Mountain. Castanea 28:1
964. Chemistry and taxonomy. [Review = ie ee ed. 1963. Chemical Plant Taxon- ony Academic Press, New York.] Science 143:1428. . 1964. vig vce Filicinae. Taxon 13:56-64. & K. L. Cuen. 1964. [23 new chromosome records] in Love, A. & O. T. Solbrig. IOPB oe ae eports. I. Taxon 13:99-102 . Fern Geecihen ten in terms of divergence patterns. Abstrs. Xth Intl. Bot. Congr.
a
sie Assessment of species relationships in the Filicineae. Abstrs. Xth Intl. Bot. Congr. 147-14
Evans, A. “ & W. H. Wacner Jr. 1964. me cae i x intermedia—a natural Woodfern cross of noteworthy morphology. Rhodor. 266.
Wacner, W. H. Jr. 1964. vig ps Pteridophyta an cai 1-15.] Pp. 1, 2, 39-48 i
. Rad- ford, H. E. Ahles & C. R. Bell. Guide to the Vascular Flora of the Carolinas. he, a North Carolina, Aran Hill.
Scora, R. W. & W. H. Wacner Jr. 1964. A oo chromatographic study of eastern Ameri- can pce he Hn Fern J. 54:10
Wacner, W. oa JR. 1964. The neha “een of living ferns. Mem. Torrey Bot. Club 21(5):86—95.
. 1964 [1965]. ees creer Copeland (1873-1964) and his contributions to pteridology.
Amer. senha 177-
. 1965. Edwin Bin any Copeland 1873-1964. Taxon 14:33—41.
. 1965. ietitadies maps of Familes 1-15, Pteridophyta] Pp. 1-6 in A. E. Radford, H. E. Abiles & C. R. Bell. Atlas of the Vascular Flora of the Carolinas. Univ. No. Car. Tech. Bull. 165.
& K. L. CHEN. 1965. eee o ie as and sporangia as a tool in the detection of Dryo-
eee hybrids. Amer. Fern J. 5
——— & F. S. Wacner. 1965. Rochester area ee Ferns (Dryopteris celsa) and their hybrids. Proc. yal Sa . Sei. 11(2):59—
STEINER, E., A. S. SUSSMAN & W. H. ia JR. 1965. Botany Laboratory Manual, rev. ed. Holt, Rinehart ; Winston, New York.
Wacner, W. H. Jr. 1965. Pellaea oo in North Carolina and the question of its origin. J. Elisha Mitchell Sci. Soc. 81:9
. 1965. Bibliography. pete BioScience 15:809-810.
. 1965. Fern paraphyses: acipusaaig on recent papers. Taxon 1
» D. R. Farrar & K. L. CHEN. 1 ye A new sexual form of Pellaea sails var. glabella from Missouri. Amer. Fern J. 55:171-178. & J.B. Sm
IN. 1966. ae ie Sections and Societies. Botanical Sciences (G). Science
151:865—866.
MickeL, J. T., W. H. Wacner JR. & K. L. CHEN. 1966. Chromosome observations on the ferns of Mexico. Caryologia 19:95-102.
Wacner, W. H. Jr. 1966. Two new species of ferns from the United States. Amer. Fern 1: 56:
sa
1966. Report of the Librarian and Curator for 1965. Amer. Fern J. 56:45-46,
- 1966. New data on North American Oak Ferns, oo eg 68:121-138.
- 1966. Modern Research on Evolution in the Ferns. Pp. 64-184 i . A. Jensen & L. G. Kavaljian, eds. Plant Biology Today. Advances and os ee ed. : Sn Belmont,
. 1966. gman in the Ferns. Bull. Fairchild Trop. Gard. 21(3):4—8,17-22, cover picture. & F
Wacner. 1966. Pteridophytes of sek Mountain Lake Area, Giles Co., Virginia: Bio- repent beste. 3 1964-65. Castanea 31:1 966. The
e new University of Michigan psoas Gardens. Taxon 15:285—286.
LELLINGER: BIBLIOGRAPHY OF WARREN HERBERT WAGNER, JR. 55
. 1966. Illustrations of transient fern forms. Amer. Fern J. 56:101—107. . 1967. Reports of Sections and Societies. a Sciences (G). Science 155:880. Eva
, J. T. MickeL, D. B. LELLINGER & A. M. 967. Pteridophytes of Costa Rica: Prelimi- nary chockliet ad species known to be or ie occurring in Costa Rica. [Mimeographed, 54 as ]
R. C. Woopsipe. 1967. Ferns for an indoor limestone boulder habitat. Amer. Hort. Mag. oa 219-223 . E967. Rexiow of: R. L. Taylor & R. A. Ludwig, eds. 1966. The evolution of Canada’s flora. Univ. of Toronto, Toronto. Canad. Field-Naturalist 81:206—207. . 1968. Hybridization, taxonomy, and evolution. Pp. 113-138 in V. H. Heywood, ed. Mod- ern Methods in Plant Taxonomy. Academic Press, London
. L. McWituiaMs. 1968. The University of Michigan Botanical Gardens special collec- tions. Arb. & Bot. Gard. Bull. 2:43-46. . 1968. recreate stamens and carpels of a willow from northern Michigan. Michigan Bot. 7:113—120.
196
ge taxonomy and modern systematics. BioScience 18:96—100.
Farrar, D. : & W. H. WAGNER Jr. 1968. The gametophyte of voter holopterum Kunze. Bot. Gaz. 129:210-219.
Wacner, W. H. Jr. 1968. Review of: R. H. Mohlenbrock. 1967. The Illustrated Flora of Illinois: Ferns. So. Ill. Univ. Press, Carbondale. Quart. Rev. Biol. 43(4):461
———. 1968. The role of a botanical garden in a modern marae Scietiee 162:1165-
. 1969. The construction of a classification. Pp. 67-90 in Systematic Biology. or 1692 cit the National Academy of Science, A erscnts DC. (Translated by E. R. de la Sota and published as La Construccién de una Clasificacién. Quid de la Ciencia, la Tecnologia y la
Educacion 2(18):440-448. 1983, haa 532-538. 1984.] oe, es Richard Eric Ho sean sagen kgeray pteridologist. Amer. Fern J. 59:1-3. ——.,F.S. “Wa GNER & D. J. Ha . 1969. The Log Fern (Dryoptreris celsa) and its hybrids in sen ig preliminary a Michigan Bot. 8:137—145. oe . Bio —— important is it to basic evolutionary systematics? Abstrs. xh a Bot. Congr. 230. F. S. WacneR. 1969. A new natural hybrid in the Appalachian Asplenium complex and its taxonomic significance. Brittonia 21:178-1 . 1969. The role and taxonomic treatment of ntti. BioScience 19:785-—789. . 1970. . The Barnes pyres — Populus X barnesii, hybr. nov—A nomenclatural case in eal Michigan Bot. 9:53— ——. 19 Sg-te sane wad evo ia ionary noise. Taxon 19:146—151. . 1970. Natural oe of floating stems of Scouring-rush, Equise- tum hyemale. ‘Miche Bot. 9:166-17 , D. R. Farrar & B. W. McALpPIN. oa epee ae _ eee Biological Station area, ees ee J. Elisha Mitchell Sci. & L. D. Gomez P. . Field research on ferns in co ae Ophioglossum, Dictyoxi- phium, Geetha rae 10-12, 1971. iaiwnsaeiepties 4pp.] 1971. Evolution of naires in ia to the a Virginia Polytech. Inst. State Univ. Res. Div. Mongr. 2:147—19 1971. The Sonthetaions Adder’s- —— Ca epee vulgatum var. pycnostichum, ig for the first time in Michigan. Michigan Bot. 971. Ferns in biology: Some final comments. nose 21:311, 323-324. nt Report of the President for 1970. Amer. Fern Soc. News & Views 2: — . 1971. Lindsaea (Schizoloma) ae Swartz in Hawaii. Amer. Fern J. 61 58. Wuirtier, D. P. & W. H. WaGNER JR. 1971. The variation in spore size and pre he in Dryo- pteris taxa. Amer. Fern J. 61: ofeagee 1972. Botanical research at botanical gardens. Pp. 28-33 in P. F. Rice, ed. agen, of the Symposium—A National Psion Garden System phi Canada. Techn. Bull. N Royal Botanical Gardens, Hamilton, Ontario, Canada.
56 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
snes 2 iy by & W. H. Wacner Jr. 1972. A native Woodfern garden. Amer. Horticulturist
51(1 WAGNER, ae 7 Jr. 1972. Estimation of the evolutionary ane Bibliographic references —— to the en gga Method. [Mimeographed,
Pp.
Cain, S. A. & W. H. WaGNER ash Dale J. Hagenah (1908-1971): An outstanding Michigan botanist. Michigan Bot. 1 ae
Wacner, W. H. Jr. 1972. io pe the President for 1971. Amer. Fern Soc. News & Views 6:1—4.
. 1972. Dale J. Hagenah. Amer. Fern J. 62:2
, F. S. Wacner & L. D. cia P; 1972. The Cantal American fern genus Pleuroderris: its
— yr significance r. J. Bot. 59:677. eo a nonst a little-known fern epiphyte with dimorphic stems. Amer. ae 62:3
oar 1°
. 1972 eal Review of: R. M. Lloyd. 1971. perenne of the Onocleoid ferns. Univ.
Calif. Publ. Bot. 61:1-93. Bull. Torrey Bot. Club 99:
. 1972 [1973]. Disjunctions in homosporous a plants. Ann. Missouri Bot. Gard.
59:203—217.
& D. R. Farrar. 1973. The oo fern genus Hyalotricha and its family relationships.
lage J. Bot. 60(4, suppl.):3
973. An orchid asc for monarch butterflies (Danaidae). J. Lepidopterists’ Soc.
27: apg
. 1973. Reticulation of Holly shea ee in the western United States and ad- janes er ia mer. Fern J. 63
973. Some future Pee of pes systematics and phylogeny. Pp. 245-256 in A. C.
Jermy, J. * Crabbe & B. A. Thomas, eds. 1973. The Phylogeny and Classification of the Ferns. Bot. J. Linn. Soc. 7 Suppl. 1
Tuomas, R. D., W. H. Wacner Jr. & M. R. Masta 1973. Log Fern (Dryopteris celsa) and related species in Louisiana. Castanea 38:269—27
Wacner, W. H. Jr., F. S. WAGNrR, J. A. ata & J. F, MaTTHEws. 1973 [1974]. Asplenium monta- num X platyneuron, a new primary member of the A i te < ltiaiail complex from useing Mountain, N.C. J. Elisha Mitchell Sci. Soc. 89(3):218-223.
——. 1974. es hackberry (Ulmaceae: Celtis tenuifolia) in the a Lakes region. Michigan tg 13:73-9
MEsLeR, M. R. & 4 H. Wacner Jr. 1974. The venation of Ophioglossum dendroneuron and its systematic significance. Amer. J. Bot. ey Suppl.):37
Wacner, W. H. Jr., J. T. Micke. & L. D. z P: 1974. a dubium and its bearing on the penis of generic relationships ey Cyrtomium and Phanerophlebia. Amer. J. Bot. 61(5, Suppl.):39
. 1974. The classification of leaf types of land plants. Amer. J. Bot. 61(5, Suppl.):67. - 1974. Fern. Pp. 237-248 in Encyclopaedia Brittanica, ed. 15, vol. 7. H. H. Benton, Chicas, HL. Hit, R. H. & W. H. hiiggen ag 1974. Seasonality and spore type of the pteridophytes of Michi- gan. Michigan Bot. 13 Wacner, ae H. Jr. 1974. aa years of botany 1947-1972: Introduction. Ann. Missouri Bot. Gard. 61:1-— pe 25 years of botany 1947-1972: Pteridology. Ann. Missouri Bot. Gard. 61:86—111. - 1974. Structure of spores in relation to fern phylogeny. Ann. Missouri Bot. Gard. 61:332-—353. Dancik, B. P., B. V. BARNES & W. H. Wacner Jr. 1974. Aberrant pistillate catkins of Betula alleg- haniensis. Michigan Bot. 13:177-179 Wacner, W. H. Jr. 1975. Review of: Hirabayashi, = sexi Cytogeographic Studies on Dryopteris of Japan. Harashobo, Tokyo. Amer. Fern J. 6 1975. Notes on the floral biology of ci sai ane negundo). Michigan Bot. 14:73-82. 1973 | [1975]. Plant species and ecosystems: a botanist’s viewpoint. Trans. Missouri Acad. 7/8:40.
- 1975. The spoken “x” in hybrid binomials. Taxon 24:296.
LELLINGER: BIBLIOGRAPHY OF WARREN HERBERT WAGNER, JR. 57
& F. S. WacneR. 1975. A hybrid polypody from the New World tropics. Fern Gaz. 11:125— 135.
Wiptn, C.-J., D. M. Brirron, W. H. WaGNER JR. & F. S. WAGNER. 1975. Chemotaxonomic studies on hybrids of neh sea: in eastern North America. Canad. J. Bot. 53:1554—1567.
WAGNER, ke H. Jr. 1975. A bunchberry ‘‘Last Rose of Summer.”’ gen Bot. 14:201-—202.
5. Sex he the angiosperms—another proposition. Sida 6 ; rp Review of: G. L. Stebbins. 1974. Flow st Plants: ae Above the Species Level. Belknap Press, Harvard Univ., Cambridge, MA. Amer. Sci. 702-703
oi J. D., W. H. WaGNeER Jr., D. R. FARRAR & S. W. LEONARD. 1975. Asano sea in the nga Biological Station area, southern Appalachians. reve 40
Wacner, W. H. Jr. 1976. How to find the rare grapeferns and moonworts. thee e oaks 3(2):2.
& F. S. WacNER. 1976. The role of foliar "ial in the systematics of ferns. Bot. Soc. Amer. Abstr., Tulane Univ., New Orleans. L. D. Gomez & W. H. WAGNER JR. 1976. An uk triploid vie ae polypody from the vicinity of Cartago, Costa Rica. Bot. Soc. Amer. Abstrs., Tulane Univ., New Orleans. 41-42 Wacner, W. H. Jr. & F. S. WacNneR. 1976. Asplenium ee een Sim ‘rom gues Gorge, Greene County, Ohio—a second North American record. Ohio J. Sci. 76:9 & D. J. SCHOEN. ape yi Oak (Quercus imbricaria) and its hybride in Michigan. Michigan Bot. 15:14 — & W. C. TAyLor. api yaa x leedsii and its westernmost station. Sida 6:224—234. A. H. SHowa ter. 1976. Ecological notes on Celastrina ebenina (Lycaenidae). J. Lepi- ie ong Soc. 30:310-312. . Systematic implications of the Psilotaceae. Brittonia 29:54—-63. NER. 1977. A diminutive grapefern (Botrychium) of the maple—basswood for- ests in 1 the Wpper Great Lakes region. Bot. Soc. Amer. Misc. Ser. Publ. 1 & D. . 1976 [1977]. a Central American fern genus Hyaloticha aa its family Seeneikieeks Syst. Bot. 1:348-36 Voss, J. H. BEAMAN, E. A. es F. W. Case, J. A. CHURCHILL & P. W. THOMPSON. 1977. Bnduneoced, threatened, and rare vascular plants in Michigan. Michigan Bot. 16:99-110. ——., F. S. Wacner & L. ~ — P. 1977. An enigmatic polypody fern from Cartago, Costa Rica. igegs 12/13:8
——. 1977. A distinctive ae form of be Marbled White Butterfly, Euchloe ae (Lepi- dope bern in the Great Lakes area. Great Lakes Entomologist 10:107-1 . S. Wacner. 1977. Fertile- amet leaf dimorphy in ferns. Gard. Bull. ai Settlem.
sosi-267 , F. S. Wacner, C. N. MILLER Jr. & D. H. WacNeR. 1978. New observations on the Royal Hern iii Osmunda x ruggii. Rhodora 80:92-106. — & T. L. MELLICHAMP. 1978. Foo — habitat, and range of Celastrina ebenina (Lycaeni- — - Lepidopterst Soc. 32:20— eer Great om anita Pieris virginiensis nae Pieridae) in com- prion siti outhern counterpart. Great Lakes Entomologist 11 ——. 1978. Venlo idioblasts in Pteris and their systematic Sag Acta Phytotax. ane 29:3 . 1978. ea of eo veins in modern ferns. Bot. Soc. Amer. Misc. Ser. 156:30 — & K. E. Boypston. 1978. warf coastal variety of maidenhair fern, Adiantum pedatum. Canad. J. Bot. oe . 1978. A probable natural ap tic of ao eurymedon and P. rutulus (Papilionidae) from {dale J. ass eee Soc. 32 . WaGNER & L. D. GO
es 978. The singular origin of a Central American fern, eae bee mic aes ee aes 10:254-264. 1978 [1979]. Hyalotrichopteris, a new generic name for a Central American polypodioid fern. Taxon 27:548. M. BEITEL. 1979. oreo variation in clones of Running Pine, Lycopodium flabelli- forme. Michigan Bot. 18:19—
AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
- 1979. Reticulate veins in the systematics of modern ferns. Taxon 28:87—95. , L. D. Gomez P. & F. S. Wacner. 1979. Coenosori and foliar nen in New World Pleo- peltis and Marginariopsis. Bot. Soc. Amer. Misc. Ser. Publ. 157:48—49.
& F.
. 1979. [Accounts of i silenare or pre ae or extirpated Senet phytes.] $0: 35-41, 108-110 in D. W. L
, LMAN, sa how Ferns (Dryopteris celsa) and their relatives in the Dismal Swany Pp. 127-139 in P. W. Kirk Jr., ed. The Great Dismal Swamp. Univ. Press of Virginia, Oostinc, D. P., D. J. HARVEY & W. H. WAGNER JR. 1979. Euristrymon ontario (Lycaenidae): first re- port in Michigan. J. Lepidopterists’ Soc. 33:151-152. Wacner, W. H. Jr. 1979. New herbarium building in Costa Rica. Taxon 28:358. » F. S. Wacner & J. M. BerreL. 1979. An unusual occurrence of Asplenium heterochroum. Canlele a 174-177. SmiTH, A. R., W. H. WaGNeR Jr. & T. DUNCAN. 1980. Noteworthy Collections. sae ape lusita- nicum L. ee i a Apr, Clausen (Ophioglossaceae). Madrofi Wacner, W. H. & F. S. Wacner. 1980. Polyploidy in Pteridophytes. Pp. mean in an H. aes ed. ae Popo. Biolog Relevance. Plenum Press, New York & London. 1980. Review of: R. L 8. A acne ‘eae of Equisetum subg. Equi- peteati: Nova Hedwigia 30: eae Michigan Bot. 1 - 1980. A probable new hybrid central pera Michigan Bot. 1 , T. F. DANIEL & M. “ HANSEN. vel o Michigan Bot. ie 37-4 Wa ter, K. S., W. H. meet jr. & F
— cesta matricariifolium x simplex, from
ybridizing Verbascum population in Michigan.
- S. Wacner. 1980. Ecological, biosystematic, and nomencla-
tural oo on Scott’s Splocawact, x Asplenosorus ebenoides. Bot. Soc. Amer. Misc. Ser. Publ. 158:123.
Wacner, W. a jn & M. K. Hansen. 1980. Size reduction southward in Michigan’s Mustard White Butterfly, Pieris napi (Lepidoptera: Pieridae). Great Lakes Entomologist 13:77-88
——. 1980 . Dyer, ed. 1979. The Experimental Biology of Pan. Academic Press, — York. Econ. Bot. 34:305-306.
. MAYFIELD. 1980. Foodplants and cocoon construction i
(Lepidoptera: Saturniidae) i in southern Michi
ussion pp. 222, 224, 227, 229. Reprinted as pp.
. F. eds. 1985. Chadinte Theory and Methodology. Van Nostrand eSnips New York.]
Barnes, B. V. & W. H. WAGNER Jr. 1981. Michigan Trees. A ned to vad of Nein and the Great Ladue Region. Univ. of Michigan
THOMPSON, J. W. & W.
WAGNER, w. H. a a F. Dani a "vs EITEL. 1980 [1981]. Studies on Populus heterophylla in southern Michigan. Michigan Bot. 19:269— Duncan, T., R. B. PHitups & W. H. AGNER, JR. ti [1981]. A comparison es —e diagrams derived by various — and cladistic methods. Syst. Bot. 5:264— Wacner, W. H. GNER. 1981. New s chium (phiogonsicon) from —) Ameri D.M SON.
.. W. H. Wacner Jr. & K. S. Wa ter. 1981. Unu
usual frond developmen in the Sensi- tive Fern Onoclea sensibilis L.. Amer. Midl. Naturalist
105:396—400
LELLINGER: BIBLIOGRAPHY OF WARREN HERBERT WAGNER, JR. 59
Wacner, W. H. Jr., M. K. HANSEN & M. R. MayYriELD. 1981. True and false foodplants of Callosamia paced (Lepidoptera: Saturniidae) in southern Michigan. Great Lakes Entomologist
14:159-16 fee Fems in the Hawaiian Islands. i wniengce Forum 8:43—4 ——, F. S. Wacner, S. W. LEONARD & M. R. 981. A ts ai of Ophioglossum
Bsn “ 'p. St. John. Castanea 46: ne on
———. 1982. Ferns, Clubmosses, Spikemosses, Quillworts, and Horsetail. Pp. phe gt in -S.']. yeaa pe — North American Wildlife. Readers’ Digest Assn., Pleasantville, N
—-. F. AGNER. 1982. The taxonomy of Dryopteris x poyseri Wherry. pt Bot.
21; a : Funk, V. A. & W. H. WaGNER Jr. 1982. A bibliography of botanical cladistics: I. 1981. Brittonia
sages ere Wacner, W. H. Jr. . S. PEIGLER. 1981 [1982]. Two notable range extensions for Callosamia securifera lovers in Georgia and ane Carolina. J. Lepidopterists’ Soc. 35:247 F. Wacner & C. HaAuFLeR. 1982. A hybrid population hae the ‘‘all- fertile”
Paths paradoxum and the hemidimerphic B. hesperium (Ophioglossaceae). Bot. Soc.
Wa ter, K. S., W. H. WaGNER JR. & F. S. WAGNER. 1982. erie biosystematic, and nomencla- tural notes on Scott’s Spleenwort, <x Asplenosorus ebenoi mer. F . 72:65-75. Wacner, W. H. M. BeIreL & F. S. WAGNER. 1982. Complex niniees bits in the leaves of puidecine Mogaphs -like leaves in Lycophytes. Science 218:793—794. CARLSON, T. M. & W. H. Wacner Jr. 1982. The North American distribution of the genus Dryopte- s. Contr. “pir Michigan Herb. 15:141—-162. raetens W. H. Jr. & F. S. WAGNER. 1982. Botrychium rugulosum (Ophioglossaceae), a newly rec- ognized species of evergreen Grapefern in the Great Lakes area of North America. Contr Univ. Michigan Herb. 15:315-324 & C. E. NAUMAN. mii setipa tdalhampell, a spontaneous fern hybrid from southern Florida. Amer. Fern J. 7 . 1982 [1983]. Edgar T. eo ian 1885-1982. Bull. Torrey Bot. Club 109:545-548. & F. S. Wacner. 1983. Genus communities as a systematic tool in the study of New World Botrychium (Ophioglossaceae). Taxon 32:51-6 . 1983. Bibliography of Edgar T. Wherry a 49:6-14. . 1983. Plant speciation: the classical approach. [Review of: V. Grant. 1981. Plant Specia- ae! = 2. Columbia Univ. Press, New York] Evolution 37:426-427. 3. Edgar T. Wherry and his contributions to pteridology. Amer. Fern J. 73:1-5. ———, ae SMITH & T. R. PRAY. 1983. A pou ee hybrid, Pellaea bridgesii < mucronata, and its age ic significance. Madrofio 30 . 19 chee stics: The recognition af ae and their role in cladistics and classifica- tion. pale 3-79 in N. I. Platnick & V. A. Funk, eds. 1983. Advances in Cladistics, vol. 2. ee Univ. Press, New York. M. JOHNSON. 1983. Trophopod, a commonly overlooked storage structure of potential systematic value in ferns. Taxon 32:268-269. 983. Homology and the early diversification of vascular plants. Amer. J. Bot. 70(5, part
& F. S. WacNeR. 1983. Two Moonworts of the Rocky Mountains; Botrychium hesperium
and a new species formerly confused with it. Amer. Fern J. 73:53-62.
. 1983. Adiantum concinnum Sana rent cones Common Maidenhair Fern), ir prceene (Helecho Lengua, Paddle Fern, Hart’s Tongue). Pp. 187, 239 in D. H. Janzen, ed. Costa Rican i History. Univ. Chicago Press
—— & L. D. Gomez. 1983. Pteridophytes te Teens Pp. 311-318 in D. H. Janzen, ed. Costa Rican Retinal History. Univ. Chicago Pr
MayFIELD, M. R., M. C. Cote & W. H. WAGNER Jr. Seat Ricciaceae in Michigan. Michigan Bot. 22:145—-150.
MavFIELD, M. R., M. C. Cote & W. H. WaGNeER JR. 1983. Techniques for preparing riccias for herba- rium study. Taxon 32:616-617
60 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
Wynne, M. m W. H. Wacner Jr. & C. B. Beck. 1984. Botany at the University of Michigan. Pl. Sci. Bull. 30:10-11 Wacner, W. Bi Jr. 1984. Review of: E. Hennipman & M. C. Roos. 1982. A monograph of the fern genus Poaaah dei al Konin. Ned. Akad. Watecachn, Afd. Natuurk. II, 80: 1-126. Amer. Fern J . 1984. Review of: a vic umpkin & D. L. Plucknett. 1982. Azolla as a Green Manure. Use and Management in Crop Production. Westview Press, Boulder, Colorado. Econ. Bot. 38:260. - 1984. Applications of the concepts of Groundplan-Divergence. Pp. 95-118 in T. Duncan & T. F. Stuessy, eds. Cladistics: Perspectives on the Reconstruction of Evolutionary History. Columbia Univ. site New York. 4. A comparison of taxonomic methods in Sy aime Pp. 643-654 in W. F. Grant, ed. 1984. Plant a iy Academic sii Canada, Don Mills, Ontar ——, F. S. Wacner, C. HaurLer & J. K. EMERSON. 1984. A new si tates 0 a scaled (Ophioglossaceac, Botrychium). Corned. J. Bot. 62: ee . 1984. Systema ic problems in the fern genus Polystichum. Amer. J. Bot. 71(5, part 2):140. . 1984, Review of J. H. Peck. 1982. Ferns and Fern Allies of the Driftless Area _ — Iowa, Minnesota, eos 1S aig Milwaukee Public Mus. Contr. Biol. Geol. 0. 1982. Michigan Bot. & E. ROuLEAu. 18, a‘ western Holly Fern, Polystichum x scopulinum, in Newfoundland. Amer. Fern J. 74:3 ——,, C. M. ALLEN & Ae i Tatas: 1984. Saks ae a Hook. & Grev. in Louisiana and the taxonomy of O. nudicaule L. f. Cas a 49:9 ——, F. S. Wacener & J. M. BEITEL. 1985. Ev rene for een hybridisation in pterido- phytes i subterranean mycoparasitic gametophytes. Proc c. Roy. Soc. Edinburgh, ser. B, 86:273-2 & V. QuEvEDO. 1985. Polymorphism in ae a — Sw. and the taxonomic sta- tus of A. germanii (Fée) Prantl. Amer. J. Bot. 72(6):9 - 1985. Morphological variation sets hipaa in Pie ele Amer. Fern J. 7 . 1985. Winter Grapefern. P. 13 P. C. H. Pritchard, ser. ed. 1985. Rare and a Biota of Florida, vol. 5 Plants. eae Presses of ie Gainesville. 985. Review of: A. E. Salgado. 1982. Venation pattern in Philippine oe ferns. De La Salle Univ. Monogr. Ser. 3:1-87. Pilipinas (J. Philipp. Stud.) 5:165-16 . 1986. Flying flowers! Butterflies and their foodplants. LSA (Publ. of the feles of Litera- ier Science, and the Arts, Univ, of M Mich.) 9:4—9, cover. 986. Review of: G. R. Proctor. 1985. Ferns of oo A Guide to the Pteridophytes. ice Museum pe ayes mC London. Syst. Bot. 11 J. M. Berret. mparative gai of the i aa ferns. Amer. J. ‘Bot. 65, pea 732-7 Farrar, D. R., W. H. Wacner JRr., N. R. an & C. L. JoHNsoN-Gron. 1986. Subterranean sporo- phytic vised in neg oe subg. Botrychium. Amer. J. Bot. 73(5, on 735-736. —, = 4: JR. 1986. Japanese honeysuckle invasion. Michigan Bot. 2571 6. The New World fern genus Marginariopsis (Polypodiaceae): Pe example “4 leaf di- morphy suse in generic delimitation. Bull. Torr trey Bot. Club 113:15 & Wacner. 1986. Thre ® new species of Moonworts (Botrychium nig nes ao in western North erica. Amer. Fern J. 76:33—47. » F. S. WAGNER & W. C. TAYLor. 106, 7 gti abortive spores in herbarium specimens of sterile one se nergy 76:1 Wac
= co
Paris, C. A \., F. 38 Pain’ <5 eee Cryptic species, species definition, and taxo- nomic Ginaetics es ca homosporous ferns. Amer. J. Bot. 74(5, abstracts):7 WAGNER, a R. The generic concept in pteridophytes. Amer J. Bot. 74(5, Seog oo
menetict Bot. 1987. ae Pp. 137-143 in S. B. Parker, vol. ed. 1987. McGraw-Hill. Encyclopedia = en and Technology, ed. 6, vol. 18. McGraw- Hill, New Yor
- 1987. Some questions about natural hybrids in ferns. Bot. Helvatica 97:195-205.
LELLINGER: BIBLIOGRAPHY OF WARREN HERBERT WAGNER, JR. 61
——. 1988. Review of: F. W. Case, Jr. 1987. Orchids of the Western Great Lakes Region. Cran- brook Inst. Sci. Bull. 48. Jack-Pine Warbler 66:39—40.
. 1988. Status of the Hawaiian fern flora. Fiddlehead Forum 15(2):11— . 1988. ciehere eye vs. divergence in the morphology of Hawaiian a inka Amer. J. Bot. 75(Suppl.):144.
& F. aaa AGNER. 1988. Detecting Botrychium hybrids in the Lake Superior region. Michi- gan Bot. 27:75-80. When can you find them? American ferns and fern allies and their phenologies. Fiddichead Forum 15:26-27.
988. Threatened and endangered (ferns and fern allies of North America). Fiddlehead
sti 15:32—34. , S. TAyLor, G. Grieve, R. O. Kapp & W. K. Stewart. 1988. Simple-leaved Fraxinus (Fraxi- nus: Oleaceae) in Michigan. Michigan Bot. 27:119-134.
. 1988. Herb Wagner, President of IAP, summarizes the programme. IAP News 4(1):1.
& M. K. HANSEN. 1989. An Angulifera dees Kentucky Lepidopterist 15(1):2.
. 1989. Kathryn E. Boydston ponies 8): Michigan’s fern hybridist and two new exam- lis of her work. Michigan Bot. 2 Zou, Xiaoming & W. H. WAGNER Jr. Pring hank A preliminary review of Botrychium in China.
mer. Fern J. 78:122-135. Wacner, W. H. Jr. 1989. Golden anniversary of the first Appalachian Spleenwort triangle. Fiddle-
BEITEL, J. M. & WaGNER, W. H. Jr. 1989. fora American lycopsids and their generic delimita- i rg
A Wacner, W. H. Jr. 1989. The role of new ae of correlating data in pteridology. Pp. 17-20 in K. H. Shing & K. U. Kramer, eds. Proceedings of the International Symposium on Sys- tematic Pteridology. China Science & Technology Press, Beiji mer . 1989. Phylogeny and cladistics in pteridology. Pp. 21-28 in K. H. Shing & K. U. Kramer, eds. Proceedings of the International Symposium on Systematic Pistia: China Science & Technology Press, nag ge & E ENAH. 1989. A synthetic —— spa x Asplenosorus pinnatifidus X Phys. scolopendrium var. americana. Paris, C. A., F. S. WAGNER & W. H. WAGNER - 1989. Cryptic species, —— delimitation, and heen practice in the homosporous ferns. Amer. Fern Wacner, W. H. Jr. & B. G. SCHOLTENS. 1989. Michigan’s ee ae An appeal to birders. poomea Audubon News 37(4):5 . 1989. Botrychium socorrense W. Wagner, sp. nov. Pp. 20-21 in G. A. Levin & R. sige 989. The Vascular Flora of Isla Socorro, Mexico. San Diego Soc. Nat. Hist. Mem. 1 ae M. . Moran. 1989. Lycopodium hickeyi: A new species of North pe Chakenmes. Amer. font 79:119-121 . 1989. Hartford Fern: First tuted plant in America? Fiddlehead Forum 1 & T. Devine. 1989. Moonworts (Botrychium: +e aaa in the =e oo area, Butte and Tehama Counties, California. Madrofio 36:1 . 1990. Ophioglossaceae. Pp. 193-197 in K. U. ak P. S. Green, eds. The Families and Genera of Vascular Plants, vol. 1: Pteridophytes and Gymnosperms. Sco gina Berlin. a i Wetland butterflies in Michigan. Michigan Entomol. Soc. Newslett. 35(3):1 90. Hawaii’s satchel-sorus tree ferns, Cibotium species: What is their aan on Fiddlehead Forum 17:7-8. & F. S. Wacner. 1990. Mocncwrnicts (Botrychium subg. Botrychium) of the upper Great Lakes region, U.S.A. and Canada, with descriptions of two new species. Contr. Univ. Mich- igan Herb. 17:313—325. . More on the Hartford Fern. Fiddlehead Forum 17 & FP. S. bien 1990. Another nothospecies in the fede Asplenium complex. Amer. Fern J. 80:44—49. . 1989 [1990]. spo of: W. J. Cody & D. M. Britton. 1989. Ferns and Fern Allies of Cana- e* a. Canadian Govt. Publ. Centre, Ottawa. Naturaliste Canad. 116:213-214.
62 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
. 1990. cigrieldlig issues from the Pacific standpoint. Abstrs. Papers 156th Ann. Meet-
ing, Amer. Ass . 1990. Biclogical diversity Underlying concepts. Michigan Academician 22(4):311-317. —— &F. S. Wacner. 1990. Notes on the fan-leaflet group of Moonworts in North America
with descriptions i two new members. Amer. Fern J. 80:73-81
Werth, C. R. & W. H. WaGnerR Jr. 1990. (13) Proposal to designate reproductively competent spe- cies of hybrid origin by an x placed in brackets (reword Article H.3, Note 1). Taxon 39:699-—702.
MIcKEL, J. T. & W. H. WAGNER Jr. 1991. Joseph M. Beitel (1952-1991). Brittonia 43:123-125. Wacner, W. H. Jr. 1990 [1991]. A natural hybrid of Gray Dogwood, Cornus racemosa, and Round-leaved Dogwood, C. rugosa, from Michigan. Michigan Bot. 29:131-136. F. S. WaGNER. 1991, Hawaii as an example of the status of Pacific Island pteridophytes. XVIIth Pacific Sci. Congr. Abstrs. 146. MickeL, J. T. & W. H. WacNeER Jr. 1991. Joseph M. Beitel (1952-1991). Fiddlehead Forum 18:
4, 17.
Wacner, W. H. Jr. 1. New examples of the Moonwort Hybrid, Botrychium matricariifolium x simplex Pan Side ae Canad. Field-Naturalist 105:91—94.
MICKEL, . H. WaGNER JR. & E. M. Girrorp. 1991. Ferns and other lower vascular plants. Pp. 163-178. The New Encyclopaedia Ruttaniea, ed. 15, vol. 19. Encyclopaedia Brittanica, Chicago, IL.
Wacner, W. H. Jr. & F. S. WacNER. 1991. The Ophioglossaceae of Mexico. Amer. J. Bot. 78(6, suppl.):149-150.
Morin, N. R., W. H. WaGcner Jr. & A. R. SMITH. 1991. Report on Flora of North America I. Amer. J. Bot. spe suppl.):204
Wacner, W. H. Jr., E. M. Fuss: C. R. WertH & R. L. Bartcis. 1991. First records of — leaved Relioomuruct, Asplenium x alternifolium, in the New World. Castanea 56:128—13
. 1991. i glandular Cinnamon Fern. Fiddlehead Forum 18:2
BRuceg, J. G., H. WAGNER Jr. & J. M. BEITEL. 1991. sh new species a hep eno gah Lycopo- diella Lscopodincne from southwestern Michigan. Michigan Bot. 3
ag 3 3 . H. WAGNER JR. 1991. An eagiiceal leaf of Pees ee. Tete Amer.
Wacner, W. H. Jr., F. S. Wacner, A. A. REZNICEK & C. R. WERTH. 1992. senate singulare Dryoteridacea a new fern nothogenus from Ontario. Canad. J. Bot. 70:24 hp
2. Phytogeography of Hawaiian peridphstes Amer. J. Bot. 79(6, te tae
ese Fy . WAGNER JR. & J. M 1992. Pacific Firmoss (Huperzia si ae (Lycopodiaceas) in eastern North seins at Gros Morne National Park, Newfoundland. Amer. Fern J. 82:63-66.
Wacner, W. H. Jr. & J. M. BEITEL. 1992. Generic classification of modern North American Lyco- podiaceae. Ann. Missouri Bot. Gard. 79:676-686.
1992. Hiemobotrychium, a new section of Botrychium subgenus Sceptridium from the
southeastern United States. Novon 2:267-268.
- 1993. Ophioglossaceae. Adder’s-tongue Family. Pp. 98-100 in J. C. Hickman, ed. The
Jepson Manual. Higher Plants of California. Univ. of California Press, a
- 1993. Problems with biotic invasives: a biologist’s viewpoint. Pp. 1-8 in B. N. McKnight,
ed, ri Biological Pollution: The Control and Impact of Invasive Paes Species. Indiana
Academy of Science, Indianapolis, IN.
- 1993. New species of Hawaiian Pteridophytes. Contr. pent ea Herb. 19:63-82.
. 1993. Fern — and allied plants. Fiddlehead Forum 2 A. ITH. 1993. Pteridophytes of North America. ete ae in Flora of North
America Editorial Committee, eds. 1993. Flora of North America North of Mexico, vol. 1. Oxford Univ. Press, New York.
Taytor, W. C., W. H. Wacner Jr., R. W. Hospy & F. R. WARSHAUER. a Isoétes hawaiiensis: a previously untleecribed Quillwort from Hawaii. Amer. Fern J. 8
=e, LORD,
:67—70. rns and fern allies of Kauai. Bull. Natl. Trop. Bot. mia 23(3):51-54. . 1993. . new combination for a North American lycopod. Novon 3:305.
LELLINGER: BIBLIOGRAPHY OF WARREN HERBERT WAGNER, JR. 63
Montcomery, J. D. & W. H. WaGNer Jr. 1993. Dryopteris Adanson. Pp. 280—290 in Flora of North erica Editorial Committee, eds. 1993. Flora of North America North of Mexico, vol. 2.
Oxford Univ. Press, New York.
Wacner, W. H. Jr. 1993. Schizaeaceae Kaulfuss. Curly-grass Family. Pp. 112-113 in Flora of
North America Editorial etna eds. 1993. Flora of North America North of Mexico, vol. 2. Oxford Univ. Press, New York.
J. M. BEIrEL. 1993. Lyco taihnd Mirbel. Club-moss Family. Pp. 18-37 in Flora of North America Editorial Committee, eds. 1993. Flora of North America North of Mexico, vol. 2. Oxford Univ. Press, ork.
& F. S. WaGNER. 1993. Pek pas C. Agardh. Adder’s-tongue Family. Pp. 85-106 in Flora of North America Editorial Committee, as 1993. Flora of North America North of Mexico, vol. 2. Oxford Univ. Press, eee rk.
, R. C. Moran & C. R. WERTH. 1993. Aspleniaceae Newman. Spleenwort Family. Pp. 228- 245 in Flora of North America Editorial Committee, eds. 1993. Flora of North America North of Mexico, vol. 2. Oxford Univ. Press, yes be
Mepeiros, A. C., R. W. Hoppy & W. H. WAGNER JR. 1 Stee on the rediscovery, status, and
ecology of the very rare Hawaiian fern Gees pene (Thelypteridaceae). Amer. Fern J.
83:86—89 Wacner, W. H. Jr. & F. S. WAGNER. 1994. Another widely disjunct, rare and local oo American coe (Ophioglosacee Botrychium subg. Botrychium). Amer. Fern J. 8
& F. 4. Morphological trends in Ophioglossales. pes + Bot. 81(6,
Aas
. 1995. Evolution : Hawaiian ferns and fern allies in relation to their conservation status.
Pactic Sci. 49:31
5 Bs _ WAGNER cz Fin N. ee Taxonomic notes on the Pteridophytes of Hawaii. Contr. cals. Michigan Herb. 20:241—2
SCHOLTENS, B. G. & W. H. WAGNER JR. om [1995]. Biology of the er Hemileuca (Lepidoptera: Saturniidae) in pb ae Great Lakes ageing 2754
Wacner, W. H. Jr. 1995. ge nore Pp. 44-48 in R. C. Moran: &R. — eds. 1995. Flora ra a vol. 1. Universidad Nal. Aut6noma de México, Méxic . 19 e and sf cua Michigan pteridophytes. Michigan pn 28:207-208.
1996. adie vs. marcescent ferns and fern allies. Fiddlehead Forum 23:32-33.
Mepelros, A. C., W. H. WAGNER JR. & R. W. Hospy. 1996. A new Hawaiian hanging Firmoss (Lyco- podiaceae: Spnctbagt es om the ene Hawaiian Islands. Amer. Fern J. 86:89-97.
Wacner, W. H. Jr. 1997. ona enon 35-36 in J. Boggan, V. Funk, C. Kelloff, M. Hoff, G. Cremers & = Feuillet, eds. 1997. Checklist of the Plants of ai cena ed. 2. Biological Diversity of the Guianas ae erecta Institution, Washington
ScHOLTENS, B. G. & W. H. Wacner Jr. 1997. An example of clinal vari odin in eastern North
merican Buckmoths (Saturniidae: Hemieuc J. Lepidopterists’ Soc. 51:47-56.
Wacner, W. H. Jr. & F. S. WAGNER. 1997. northwestern North American Moonwort metropo- lis and its outliers. Amer. J. Bot. 84(6, Bae ppl.):168.
Noopen, L. D. & W. H. WaGner Jr. 1997 [1998]. Photosynthetic capacity and leaf reorientation in two oa ferns, Polystichum acrostichoides and Dryopteris intermedia. Amer. Fern J. 871
WAGNER, si H. Jr. “1998. Role of disturbed habitats in conservation of pteridophytes. Amer. J. ie eg suppl.):99
. pwd fie hybrids for hardy garden cultivation. Hardy Fern Found. Quart.
0}80-36
F. S. Wacner. 1998. Moonwort madness: a reply. Fiddlehead Forum 25:27, 30, 31. Re-
pie in ge Fern Found. Quart. 9(2):24—26.
Zach, J. C . MANNING & W. H. WAGNER Jr. 1998. : new adder’s-tongue (Ophioglossum: Ophislosacon for North omen Sida 18:307-31 Wacner, W. H. Jr. 1998. Review of: C Page. 1997. Sea of Britain and Ireland, ed. 2. Cam-
bridge Univ. in London and ae York. Syst. Bot. 23:249-250.
64 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
, F. S. Wacner, D. D. PALMER & R. W. Hospy. 1999. oo notes on the Pteridophytes of poe Contr. Univ. Michigan Herb. 22:135-18 A. V. GILMAN. 2001. Dryopteris acti gin nov. (D. carthusiana x goldiana), a rare pip Hil hybrid from Vermont. Amer. Fer 1:9-12. WIDERLECHNER, M. P. & W. H. Wacner JR. 1998 stot Occurrence of European cenie, Rubus caesius (Rosaceae), vein recemi in Iowa and Michigan. Michigan Bot. 37:107-11 Wacner, W. H. Jr. 2002. Leaf forms in southeastern Michigan Sassafras. seo =a 39:70-76. & J. R. GRANT. sens ava alaskense, a new Moonwort hybrid from the interior of Alaska. Amer. Fern J. 9 70. ZikA, P. F., E. R. ALVERSON, ae os i: JR. & F. S. Wacner. in press. Botrychium hesperium in the Wallowa Mountains of Oregon. Amer. Fern J.
American Fern Journal 92(2):65—79 (2002)
The Mating Systems of Some Epiphytic Polypodiaceae
WEN-LIANG CHIOU Division of Forest Biology, Taiwan Forestry Research Institute, 53 Nan-Hai Rd., Taipei 100, Taiwan Donatp R. Farrar’ Department of Botany, Iowa State University, Ames, IA 50011 Tom A. RANKER University Museum and Department of Environmental, pecan & Organismic Biology, University of Colorado, Boulder, CO 803
Asstract.—Genetic loads, estimated from sporophyte production by isolated gametophyte cul-
mating in C. angustifolium (in part), Microgramma heterophylla and Polypodium pellucidum. Polyploidy characterizes the intragametophytic-selfing species, whereas the intergametophytic- mating taxa are diploid. The duplicated loci of polyploid taxa may mitigate the expression of re- cessive lethal alleles caused by intragametophytic selfing, whereas genetic load probably main- tains the mating systems of the intergametophytically mating taxa. Enzyme electrophoretic patterns of fixed heterozygosity support allopolyploid origins of C. phyllitidis and P. aureum and confirm their intragametophytic mating systems. Antheridiogens, present in both groups,
may promote intergametophytic mating in diploids through promotion of the early development
ploids if these gametophytes delay or do not attain insensitivity to their own antheridiogen. In the polyploids, antheridiogens may also alleviate low genetic variability through promotion of occasional outcrossing. The perennial, clone-forming habit of epiphytic Polypodiaceae increases the duration and the physical space occupied by derivatives of a single spore, thus expanding the chance of interaction with a later migrant. Genetic load, duplicated genes, and antheridio- gens, together with a perennial and clone-forming gametophyte growth habit, interact to produce successful breeding strategies of these epiphytic species.
Three mating systems have been documented in ferns: intragametophytic pies 2 cima dla selfing, and intergametophytic crossing (Klekowski, 1979). The more general term intergametophytic mating refers to either or both of the Ge two systems when it is not possible to determine whether cross-mated gametophytes are from the same (intergametophytic selfing) or different (intergametophytic crossing) sporophytes.
Because gametophytes are potentially bisexual, it had been thought that intragametophytic selfing was predominant in homosporous ferns (Klekowski, 1979), and high selfing rates have been reported in some diploid homo- sporous ferns, especially those with subterranean gametophytes (e.g., Soltis and Soltis, 1986a) and in some pioneering species (Crist and Farrar, 1983). Intragametophytic selfing also seems to be the trend in species that are poly-
loid with respect to other species in a genus. Possibly the duplicated loci
’ Author for correspondence.
66 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
of polyploid species mitigate the problem of recessive deleterious allele expression associated with selfing (Klekowski and Baker, 1966: Masuyama and Watano, 1990). Intragametophytic selfing may also reflect a genetic bottle- neck associated with the origin of polyploid species events. However, iso- zyme evidence has shown that intergametophytic mating is the most common breeding system in diploid homosporous ferns with above-ground gametophytes (Haufler & Soltis, 1984; Soltis & Soltis, 1986b; 1990).
Reproductive behaviors are affected by several factors. Inbreeding depres- sion resulting from recessive lethal genes (expressed as genetic loads in this study) seems to be the primary factor preventing intragametophytic selfing in many species (Haufler et al., 1990). Failure to attain bisexuality and asynchro- nous maturation of antheridia and archegonia also promote outcrossing in po- tentially bisexual gametophytes (Klekowski, 1968). Functional unisexuality is augmented by antheridiogen systems that stimulate precocious antheridium formation in some gametophytes in a population while allowing others to re- main unisexually female (Naf, 1979). Antheridiogen can also induce previ- ously buried spores to germinate to form gametophytes that reach the surface and produce sperm capable of fertilizing above-ground plants (Voeller, 1971). Variation in growth habit including indeterminate growth, branching, gemma production, and production of perennial gametophyte clones can also influ- ence reproductive behavior by increasing the probability of interaction be- tween distant gametophytes and between plants established at different times (Chiou & Farrar, 1997b; Chiou et al., 1998; Dassler & Farrar, 2001).
Comprehensive research on fern reproduction is limited, especially in epiphytic ferns (Werth & Cousens, 1990) whose gametophytes exist in very different environments from those of terrestrial species. The habitat of gametophytes of epiphytic species is often within dense bryophyte mats where interaction between gametophytes via antheridiogen or sperm transfer may be significantly hindered compared to gametophytes of terrestrial ferns on less complex surfaces (Dassler, 1995; Dassler & Farrar, 2001). On the other hand, bryophyte mats may maintain free water, the medium for sperm, avail- able for longer periods and thus promote fertilization over longer distances than substrates lacking bryophytes.
Most species of Polypodiaceae are epiphytic. The presence of antheridio- gen systems (Chiou & Farrar, 1997a) and clone-forming, perennial growth
MATERIALS AND METHODS The materials used in this
study follow. G h collected by Chiou and Farr y reenhouse materials and those
ar were used for genetic load studies. Materials
CHIOU ET AL: MATING SYSTEMS 67
for isozyme studies came from the four Florida localities indicated below by two-letter abbreviations. Vouchers are held at Iowa State University.
Campyloneurum angustifolium (Swartz) Fée—ISU greenhouse (origin un- known) (Farrar 02-1-8-1) and Marie Selby Botanical Garden, Sarasota, Florida (origin unknown; Chiou 14337).
Campyloneurum phyllitidis (L.) C. Presl—ISU greenhouse (2 collections, origin unknown) (Farrar 02-1-8-2), Castellow Hammock (CH), Fakahatchee Strand State Preserve (FS), and Jonathan Dickson State Park (JD).
Microgramma heterophylla (L.) Wherry—ISU greenhouse (Florida) (Farrar 02-1-8-3).
Phlebodium aureum (L.) J. Smith—Fakahatchee Strand State Preserve (FS) (Chiou 14342), Jonathan Dickinson State Park (JD) (Chiou 14300), Toso- hatchee State Reserve (TS).
Phymatosorus scolopendria (Burm.) Pic.-Serm.—Hawaii (Farrar 92-8-20-1). Polypodium pellucidum Kaulf.—Hawaii (Farrar 92-8-19-3).
Spores were first sown and gametophytes grown in multi-gametophyte cul- tures on agar-solidified mineral media in plastic petri dishes (Chiou & Farrar, 1997a). Cultures were maintained under continuous, white fluorescent illu- mination of 2000-3000 lux and at a temperature range of 20—24°C.
Genetic load (presence of sporophyte-lethal alleles) was estimated by com- paring isolated-spore and isolated-gametophyte cultures with paired-spore and paired-gametophyte cultures. These cultures were grown in compart- mentalized plastic sheets (‘jelly-molds’’), each with 20 cells containing about 6 ml of agar medium. In each sheet, a single spore was transferred directly into each of 5 cells (isolated-spore cultures), and a one-month-old gametophyte which was still asexual was transferred from multigametophyte cultures into each of another 5 cells (isolated-gametophyte cultures). The other 10 cells of each sheet received two spores (paired-spore cultures) and two gametophytes (paired-gametophyte cultures). Five such culture sheets were separated by clear, flat, plastic sheets and stacked in transparent plastic boxes (vegetable crispers) for a total of 50 replicates of each culture type for each species. Gametophytes grown from spores isolated before germination served to test the possibility that gametophytes transferred from multi- gametophyte cultures to isolate and pair cultures might have experienced gametophyte interactions before transfer. The light intensity of these cultures was maintained between 1500 lux (bottom layer) and 3500 lux (top layer). Plants were watered with distilled water every two weeks after the gameto- phytes were 4 months old.
Whether sporophytes were produced by syngamy was determined by examination with a compound microscope. Genetic load was estimated by counting the percentage of bisexual gametophytes failing to produce
68 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
sporophytes (Peck et al., 1990). The five layers of sheets in each box were designated as blocks, and spore-isolated cultures vs. gametophyte-isolated cultures were designated as split plots. For C. angustifolium, C. phyllitidis, and P. aureum, since two sources of spores were used, a Latin Square was designed. Five treatments were “A” and “‘B” (isolated plants of two different sources), ““AA’’ and “BB” (paired plants from the same source), and “AB” (paired plants from different sources). Isolated gametophytes were removed when sporophytes were formed or 8 months after spore sowing and exam- ined microscopically to determine their sexuality. Development of gameto- phyte sexuality in multi-plant cultures was also monitored (Chiou & Farrar, 1997a).
Isozyme data were analyzed with BIOSYS-1 (Release 1.7, Swofford and Selander, 1989).
RESULTS
aureum and Polypodium pellucidum, when bisexual plants became domi- nant. A relatively small number of male gametophytes (<10%) were present
C. angustifolium, C. phyllitidis, and P. aureum (data not shown). Differences
CHIOU ET AL: MATING SYSTEMS 69
ABLE 1. Sex expression (%) of Polypodiaceae species in isolated spore and isolated gametophyte cultures at 8 months after sowing spores.
Treatment Male Female’ Bisexual!
Campyloneurum angustifolium re
19 0 81 Gametophyte 6 0 94 Campyloneurum phyllitidis Spore 17 41(a) 42(b) Gametophyte 8 11(b) 81(a) Microgramma heterophylla 14 0 86 6 0 94 Phlebodium aureum Spore 0 3 97 Gametophyte 0 3 97 Phymatosorus scolopendria re 0 0 100 Gametophyte 0 0 100 Polypodium pellucidum pore 7 85(a) 8(b) Gametophyte a 35(b) 61(a)
’ Different letters in parentheses for each gender of each species indicate significant differences in Duncan’s multiple test (95% confidence limits).
after sowing spores were present only in C. phyllitidis and in P. pellucidium (Table 1). Whereas few or no female gametophytes remained at 8 months in other species, in C. phyllitidis and in P. pellucidium female gametophytes were still common, especially in cultures from isolated spores.
The sexual expression of gametophytes was not significantly different (95% confidence limits in Duncan’s multiple test) among different layers within cultures boxes except for C. angustifolium, where significantly more male gametophytes developed in the lowest (and darkest) layer.
Genetic Loap.—At 8 months after sowing spores, sexual sporophyte produc- tion was lower in the isolated cultures (of both isolated-spore and isolated- gametophyte cultures) than in paired cultures of M. heterophylla and P. pellucidum (Table 2). There was no significant difference between isolated and paired cultures of P. aureum and P. scolopendria.
In C. angustifolium, gametophytes from source B failed to produce sporo- phytes in both isolated-spore and isolated-gametophyte cultures at 8 months after sowing spores, whereas 83% of isolated plants from source A produced sporophytes. Thus the inferred genetic load of source B was extremely high, whereas the genetic load of source A was quite low.
In C. phyllitidis, no difference was observed in sporophyte production be- tween gametophytes from the two spore sources. There also was no signifi- cant difference between sporophyte production by single gametophytes isolated as young plants from multi-gametophyte cultures and any of the
70 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
TaBLE 2. The percentage of gametophytes producing sporophytes sexually and the estimated genetic load expressed by bisexual gametophytes of 8 month old spore-isolated and gametophyte- isolated cultures of Polypodiaceae species.
% Producing sporophytes Est. genetic load? Source’ Spore® Gametophyte® Average” Spore® Gametophyte® Average® Campyloneurum angustifolium A 77(a) 90 83(ab) 16(b) 8(b) 12(b) AA 92(a) 96(a) 94(a) — - ~ AB 73(ab) 86(a) 79(b) - - - BB 44(c) 48(bc) 46(c) - - o B O(d) O(d) 100(a) 100(a) 100(a) Campyloneurum phyllitidis A 9 40(a) 25(b) 74(a) 38(b) 56(a) AA 48(a) 61(a) 55(a) 2 — - AB 55(a) 64(a) 59(a) - ~ = BB 49(a) 65(a) 57(a) - ~ re ) 61(a) 33(b) 83(a) 31(b) 57(a) Phlebodium aureum 7(b) 82(ab) 79(b) 18(a) 13(a) 16(a) AA 2(ab) 90(a 91(ab) - ~ - AB 98(ab) 80(ab) 88(ab) - - - BB 89(ab) 100(a) 4(a) - - ~ B 2(ab) 97(ab) 95(a) 7(a) 3(a) 5(a) Microgramma heterophylla A O(b) 2(b) 1(b) 100(a) 98(a) 99 AA 32(a) 24(a) 28(a) _ ~ — Phymatosorus scolopendria A 88(a) 90(a) 89(a) 12(a) 10(a) Ei AA 93(a) 93(a) 93(a) - ~ - Polypodium pellucidum A O(c) 2(c) 1(b) 100(a) 95(a) 98 AA 63(a) 26(b) 43(a) - os =
‘ A and B indicate single gametophyte cultures; AA, AB and BB indicate paired cultures, AA and BB indicate gametophyte pairs from the same sporophyte, AB indicates gametophyte pairs from two different sporophytes * Genetic load was estimated b duce sporophytes.
* The same letter in parentheses for each item for eac in Duncan’s multiple test (95% confidence limits).
y counting the percentage of bisexual gametophytes failing to pro-
h species indicates no significant difference
paired cultures. However, single plants isolated as spores produced signifi- cantly fewer sporophytes than those isolated as young gametophytes or those grown in pairs. Thus genetic load estimates from isolated-spore and isolated- gametophyte cultures were significantly different (Table 2).
In P. pellucidum, 21% of the isolated plants produced sporophytes. How- ever, only 1% of these were produced through syngamy, the others were generated through apogamy. All of the apogamous sporophytes formed from
CHIOU ET AL: MATING SYSTEMS 71
Allele frequencies at polymorphic loci in three populations of re Soria phyllitidis. Sample sizes in each population are 10. Populations were invariable at 27
Locus Allele JD! CH’ FS! Locus Allele JD’ CH’ Fs’ Lap-b Z 1.00 0.10 0.65 Pgm-3a ab 1.00 1.00 0.90 - 0.90 a5; Z - _ 0.10
Mdh-3a s - 0.10 - Pgm-3b 1 1.00 1.00 0.80 vA 1.00 0.90 1.00 Z - = 0.20
Mdh-3b ef - 0.10 - 6Pgd-2a 1 1.00 1.00 0.40 zZ 1.00 0.90 1.00 2 - - 0.60
' JD=Jonathan Dickson State Park, CH=Castellow Hammock, FS=Fakahatchee Strand State Preserve.
IsozYMEs.—In C. phyllitidis, seventeen putative locus pairs were scored across the eleven enzyme systems (Table 3). There was no variability within or among the three populations for 12. These were fixed for the same allele at four (Fbp-1, Idh, Mdh-1, and Tpi-1) and for fixed interlocus heterozygosity at eight locus-pairs (Aco, Fbp-2, Hk, Mdh-2, 6Pgd-1, Pgi-2, Skdh, Tpi-2). Mdh- 3 was fixed for a single allele in the samples from JD and FS populations, Pgm- 2 was fixed in all populations, but for a different allele at 2b in JD, and Pgm-3 was fixed in the samples from JD and CH populations. 6Pgd-2 had fixed hetero- zygosity in populations JD and CH. The genetic similarity among the three populations was high (>0.934) for both Rogers’ (1972) and Nei’s (1978) genetic coefficient (Table 4).
In population JD, only one genotype was found at each of the locus-pairs. In population CH, Lap and Mdh-3 each had two genotypes, combining to form three multilocus genotypes. In population FS, Lap and Pgm-3 each contained three genotypes and the variable locus 6Pgd-2 contained two geno- types, combining to form six multilocus genotypes (Table 5). In combination, the three populations form 10 multilocus genotypes. Of 30 plants tested, only one displayed recombinational heterozygosity at one locus (Lap 11/23).
In P. aureum, 22 putative loci were scored among the 13 enzyme systems. There was no variability within or among the three populations for 21 locus pairs. Eleven (Ald, Fbp-1, Fbp-2, Idh-1, Idh-3, Mdh-1, Mdh-3, 6Pgd-2, Pgi-1, Skdh-1, and Skdh-2) were fixed at the same allele, and ten (Aco-1, Aco-2, Hk, Lap, Mdh-2, 6Pgd-3, Pgi-2, Pgm-1, Pgm-2, and Tpi-2) had fixed heterozygos-
TABLE 4. Matrix of Roger’s genetic similarity (above diagonal) and Nei’s unbiased genetic identity (below diagonal) among three populations of Campyloneurum phyllitidis in Florida.
Population’ JD CH FS JD ae 0.938 0.934 CCH 0.946 ae 0.951 FS 0.956 0.981 a a
1jD= — Dickson State Park, CH=Castellow Hammock, FS= Fakahatchee Strand State Preserv
v2 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
TasLe 5. Description of multilocus genotypes of two populations of Campyloneurum phyllitidis in Florida.
Locus CH! FS! Lap-a/b 11/22 11/33 11/33 11/22 41/22 11/33 11/33 11/23 11/22 Mdh-3a/b 29/99 11/11 29/22 Pgm-3a/b 47/11 11/22 ss | 22/22 as a 114/91 6Pgd-2a/b 41/22 22/22 11/22 11/22 22/22 22/22
Observed 10% 10% 80% 30% 10% 20% 10% 10% 20% ‘CH = Castellow Hammock, FS = Fakahatchee Strand State Preserve.
ity. Got displayed fixed heterozygosity for the same alleles in TS and JD pop- ulations and in 90% of the samples in FS, but had fixed heterozygosity with a different allele for Got-b in the other 10%. The genetic similarity between population FS and the two identical populations TS and JD was high, 0.998 in Rogers’ (1972) genetic similarity and 1.0 in Nei’s (1978) genetic identity. In populations TS and JD, there was only one genotype for all the locus- pairs. In population FS, only two genotypes appeared, Got-a!'/b"™ (10%) and Got-a''/b** (90%). No recombinational heterozygous individuals were de- tected among the 30 plants tested.
DISCUSSION
In multi-gametophyte cultures of all species, male and bisexual plants ap- peared only after a significant number of female plants had differentiated. This is consistent with the presence of antheridiogen systems in these spe- cies, as has been previously demonstrated (Chiou & Farrar, 1997a). Anther- idia on bisexual gametophytes grown as isolated plants were probably induced by their own gametophyte’s antheridiogen secretions. This could occur either by absence of or delayed attainment of insensitivity of a gameto- phyte to antheridiogen, or by generation of secondary lobes with reduced physiological connection to the principal antheridiogen-producing apex. A
tion from antheridiogen. Plants that remained unisexual males at eight months generally were relatively slow-growing and small.
In isolated-plant cultures, a significant difference in sexual expression between gametophytes grown from isolated spores and those grown from gametophytes isolated from multi-gametophyte cultures when they were one
pellucidum is that plants in multi-gametophyte cultures were subjected to antheridiogen before individual gametophytes were transferred to the iso-
CHIOU ET AL: MATING SYSTEMS 73
lated-gametophyte cultures. It is also possible that transplanting actively growing gametophytes might disrupt their normal developmental pattern. Neither hypothesis explains the slow rate at which bisexuality is attained by spore-isolated gametophytes of these species relative to the other species tested.
Genetic load can be estimated by the ability of isolated bisexual plants to produce sporophytes. Isolated gametophytes of Polypodium pellucidum, Microgramma heterophylla, and the B source of Campyloneurum angustifo- lium produced virtually no sporophytes through syngamy, whereas isolated gametopytes of C. phyllitidis, Phlebodium aureum, Phymatosorus scolopen- dira and the A source of C. angustifolium produced abundant sporophytes through intragametophytic selfing.
Genetic load was not significantly different between isolated-spore and isolated-gametophyte cultures, except in Campyloneurum phyllitidis where the apparent genetic load of isolated-spore cultures was much greater. The reason for this apparent elevation of genetic load in isolated-spore cultures in C. phyllitidis is not clear, but from the delayed production of bisexual plants evident in isolated spore cultures (Table 1), it is possible that anther- idia in some bisexual plants may still have been functionally immature even at eight months, falsely indicating a high genetic load. No similar delay in production of antheridia by this species was evident in multispore cultures (Chiou & Farrar, 1997a) or in isolated-gametophyte cultures. However, pair- ing of gametophytes from different sporophytes in C. phyllitidis also failed to fully relieve the sporophyte suppression observed in isolated gametophyte cultures. This suggests that some genetic load in this species is perhaps being expressed in gamete development (Klekowski, 1971), or that there was very little genetic difference between the two sporophytes, as might well be the case in a population reproducing primarily by intragametophytic selfing. In fact, most sporophytes of this species were homozygous at all loci tested.
Paired-spore and paired-gametophyte cultures allowed intergametophytic mating to relieve inbreeding depression that might prevent intragameto- phytic selfing in either gametophyte. Thus any increase in sporophyte pro- duction in paired cultures relative to isolated cultures of the same species is assumed to have resulted from intergametophytic mating.
Two spore sources were used in studies of Campyloneurum angustifolium, C. phyllitidis, and Phlebodium aureum. Genetic load estimates obtained from the two sources were not significantly different for C. phyllitidis and P. aur- eum. Estimated genetic load did differ significantly between the two sources of C. angustifolium in which it was low for source A, but very high for source B. This suggests that the B sporophyte of C. angustifolium was from a highly outcrossing population, whereas the A sporophyte was derived from a population with a higher level of inbreeding. Both diploid and tetraploid chromosome counts have been reported for C. angustifolium (n=37, Evans, 1963, from Peru; n=74, Evans, 1963, from Costa Rica; Sorsa, 1966, from Costa Rica; Knobloch, 1967, from Jamaica). Thus, judging from a correlation
74 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
143) ¥ 43. *
TABLE 6. Description of mult genoty} percentage in a combination of three Florida populations of Campyloneurum phyllitidis.
Lap-a/b Mdh-3a/b Pgm-2a/b Pgm-3a/b 6Pgd-2a/b Percentage 11/22 22/22 11/22 11/11 1/22 33.33 11/33 22/22 11/11 TN/41 11/22 26.67 11/33 14/11 14/41 14/11 11/22 3.33 11/22 22/22 71/41 11/11 41/22 3.33 41/33 22/22 11/14 11/11 11/22 6.67 11/22 APs Lies a jg 11/22 10.00 11/23 22122. 11/11 a ala 22/22 3.33 11/33 22122 11/11 22/22 Ti/22 3.33 11/22 22/22 11/11 14/22 22/22 3.33 11/22 22/22 11/11 ag 22/22 6.67
of polyploidy with low genetic load, it is possible that the A sporophyte and B sporophyte of C. angustifolium were tetraploid and diploid respectively.
Both diploid and tetraploid forms have also been reported in Phlebodium aureum (n= 37, n=74, Evans, 1963), and Phymatosorus scolopendria (2n=72, Léve et al., 1977; n=72, Tsai and Shieh, 1983). In Campyloneurum phyllitidis, only tetraploids have been found (n=74, 2n=148, Evans 1963, Nauman 1993), whereas in Polypodium pellucidum, only diploid numbers have been reported (Manton, 1951). Thus genetic load estimates indicate that the sample sporophyte spore sources of C. phyllitidis, P. aureum and P. scolopendria were probably tetraploids, whereas those of Microgramma heterophylla, and P. pellucidum were diploid. In fact, isozyme evidence also indicates that C. phyllitidis and P. aureum are polyploids.
Strong evidence for describing the mating system of diploid species in the wild can be obtained from analysis of isozyme electrophoretic patterns. Elec- trophoretic patterns for the species tested in this study, Campyloneurum phyllitidis and Phlebodium aureum, revealed a high level of fixed hetero- zygosity for both, indicating that these samples from Florida are polyploid, probably allopolyploid. Because of this, accurate counts of heterozygous in- dividuals and estimates of outcrossing from isozyme evidence are not possi- ble, although we can state that at least one putatively outcrossed individual (Lap-a'*/b**) was among the 10 samples of C. phyllitidis in the FS popula- tion. No evidence of outcrossing was present in the Florida populations of P. aureum but the extremely low level of genetic variability among sampled plants of Pz aureum (29 of 30 plants were genetically identical) would preclude isozymic detection of most recombinational heterozygotes if they were formed. However, the considerable variability among sporophytes of C. phyllitidis (10 multilocus genotypes among 30 plants tested) allows ample opportunity for detection of unbalanced heterozygotes and three-allele hetero- zygotes (at Lap) that would be produced if outcrossing was frequent (Table 6).
Assuming allopolyploidy, intragametophytic selfing, and no mutations, the maximum number of genotypes per locus-pair provides an estimate of the
CHIOU ET AL: MATING SYSTEMS 75
minimum number of hybridization events involved in producing an allote- traploid species (Ranker et al., 1994). Thus in Campyloneurum phyllitidis, the single genotype in the population JD suggests that only one hybridization event occurred in the ancestry of this population in Jonathan Dickson State Park. Two genotypes at each of Lap and Mdh-3 of population CH indicate that that population originated from at least two hybridization events. Three genotypes at Pgm-3a/b implies that at least three hybridization events pro- duced population FS. However, one of these genotypes (11/22) could have been generated (11/11 x 22/22) by the low level of outcrossing demonstrated by presence of the Lap 11/23 genotype. Because there are not great distances separating these populations in Florida and because of the high similarity of genotypes among the three populations, we can also consider them as a sin- gle population. In that case the number of hybridization events responsible for the Florida plants is at least three and may be as high as ten.
In Phlebodium aureum, the genetic similarity among sampled populations is very high. Isozyme evidence showed no genetic variation in the samples of TS and JD populations and only a single plant representing a distinct genotype in the FS population. Because Campyloneurum phyllitidis and P. aureum are widespread in tropical America and because likely parental diploids are not present in Florida, these Florida genotypes probably repre- sent three to ten separate spore introductions for C. phyllitidis and possibly only one or two for P. aureum.
In an electrophoretic study of Polypodium pellucidum, Li and Haufler (1999) found a small but significant (mean fixation index of 0.169 across populations of epiphytic P. pellucidum var. pellucidum) excess of homozy- gous individuals in most (but not all) populations sampled in the Hawaiian archipelago. Since intragametophytic selfing as a dominant breeding system would lead to much higher fixation values in relatively few generations (Hartl & Clark, 1997), the observed level of fixation likely results from a mixed mating system where intergametophytic selfing among gametophytes in populations derived from the same sporophyte is occurring. Thus electro- phoretic analysis of population structure in P. pellucidum is not inconsistent with the implications of our results that intragametophytic selfing in this species is rare, probably curtailed by genetic load. This constraint on repro- duction via single isolated spores likely contributes to the genetic differen- tiation and low values of gene flow between populations estimated by Li and Haufler (1999).
The fact that diploidy favors intergametophytic mating whereas tetraploidy favors intragametophytic selfing has been demonstrated (e.g., Masuyama and Watano, 1990). The fixation of different alleles from the different parent spe- cies of allopolyploids may mitigate the expression of recessive lethal genes caused by intragametophytic selfing. In our study, low levels of genetic load, as evidenced by production of sporophytes through intragametophytic self- ing, was well correlated with polyploidy (Table 7). Isolated gametophytes of the diploid species (Polypodium pellucidum, Microgramma heterophylla, and the B source of Campyloneurum angustifolium) produced virtually no
76 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
TaBLE 7. Genetic load, ploidy level and probable mating systems in Polypodiaceae species.
Species Genetic load’ Ploidy” Mating system?*
C. angustifolium A* 14 P (Evans 1963) Intragam. selfing C. angustifolium B* 100 D (Evans 1963) Intergam. mating C. phyllitidis 35 P (Isozyme; Evans 1963) Intergam. selfing M. heterophylla 99 D Intergam. mating hl. aureum 10 P (Isozyme; Evans 1963) Intergam. selfing Phy. scolopendria 11 P (Tsai & Shieh 1983) Intergam. selfing Pol. pellucidum 98 D (Manton 1951) Intergam. mating
‘Estimated from average sporophyte production by single gametophytes isolated as spores and as young gametophytes, except for C. phyllitidis in which the estimate is from plants isolated as young gametophytes.
* Determined from isozyme patterns (isozyme) and/or chromosome counts (reference). D=dip- loid, P= polyploid.
* Determined from isozymes and/or genetic load.
* C. angustifolium A and B plants are from two different sources.
sporophytes through Ssyngamy, whereas isolated gametopytes of the tetra- ploid species (C. phyllitidis, Phlebodium aureum, Phymatosorus scolopen- dria and the A source of C. angustifolium) produced abundant sporophytes through intragametophytic selfing.
Gametophytes derived from isolated spores resulting from long-distance
spores can produce sporophytes through either intragametophytic selfing or intergametophytic mating, the latter being augmented by antheridiogen- stimulated production of male gametophytes. Species predominantly repro- ducing by intergametophytic mating can maintain high levels of genetic variability, including a high genetic load, but may be very limited in their ability to migrate by long-distance spore dispersal (Peck et al., 1990). Thus a trade-off exists between maintenance of genetic variability on one hand and ease of migration on the other.
CHIOU ET AL: MATING SYSTEMS 77
Schneller & Hess, 1995). But, the existence of antheridiogens here could also function to promote bisexuality in isolated plants if individual thalli have reduced ability to attain bisexuality. Gametophytes of these species propa- gate vegetatively by branching (Chiou & Farrar, 1997b). Maintaining an an- theridiogen system may be adaptive in promoting antheridium formation on new vegetatively produced thalli regardless of whether the species is in- breeding or outbreeding.
Whether species are outbreeders or inbreeders, breeding behavior must be adaptive to establishment and survival of individuals of those species. In general, inbreeding may be advantageous for initiating new populations fol- lowing long-range spore dispersal where gametophytes are likely to be de- rived from single isolated spores. The opposite strategy, outbreeding, has the advantage of generating and retaining genetic diversity, and high genetic loads carried by outbreeders tend to maintain that mating system. Previous studies have demonstrated that gametophytes of the epiphytic species studied here grow perennially through branching and vegetative prolifera- tions which increase their life span and effective gametophyte size (Chiou & Farrar, 1997b) and produce antheridiogen that facilitates the production of male gametophytes (Chiou & Farrar, 1997a). Both of these characteristics have been proposed to increase the probability of intergametophytic mating (Chiou & Farrar, 1997a; b; Chiou et al., 1998). Here we demonstrate that sporophytes of these species may be produced through either outcrossing or inbreeding, as evidenced by high or low level of genetic load, respectively. Their mating systems may be controlled principally by genetic load and generally correlated with ploidy level. Interwoven with these factors are gametophyte morphology, growth habit, antheridiogen production, and envi- ronmental parameters which together maintain successful reproduction of these species. For outbreeding diploid species, genetic load is possibly the driving force leading to morphological and physiological adaptations promo- ting outcrossing.
ACKNOWLEDGMENTS
The authors thank Drs. D. B. Lellinger, C. H. Haufler, and L. Hickok for their useful comments, Dr. Paul N. Hinz for help with statistical analysis of gametophyte culture data, the Marie Selby Botanical Garden for allowing collection of spores of Campyloneurum angustifolium, Dr. H. Lu- Rivero for assisting collection of Campyloneurum angustifolium, the Florida
ia?) | 5 jor ns)
tellow Hammock, Dr. J. B. Miller and Mr. R. E. Roberts for help with collecting Campyloneurum phyllitidis and Phelebodium aureum in Jonathan Dickinson State Park, Mr. M. Owen for assis- tance in collecting Campyloneurum phyllitidis and Phelebodium aureum in Fakahatchee Strand State Preserve, Mr. Keith Fisher for help with locating Phelebodium aureum in Tosohatchee State Reserve, and Mrs. Ming-Ren Huang for help with plant cultures and data recording. This research was supported by the Taiwan Forestry Research Institute (Contribution No. 196).
78 AMERICAN FERN JOURNAL: VOLUME 92 NUMBER 2 (2002)
REFERENCES
Cuiou, W.-L. and D. R. Farrar. 1997a. Antheridiogen production and response in Polypodiaceae species. Amer. J. Bot. 84:633-640. and D. R. Farrar. 1997b. — a morphology of selected species of the wong? Polypodiaceae. Amer. Fern J. 87: , D. R. Farrar, and T. A. cai a eae eee and reproductive biol- ogy in Elaphoglossum Schott. Canad. J. Bot. 76:1967-19 CRIsT, ish Gs dees _R. Farrar. 1983. Genetic load and ue dispersal in Asplenium platy- Canad. J. Bot. 61: ab ip Dass, ee L 1995. Significance of gamatophyte form in tropical, epiphytic ferns. PhD diss. lowa ate University, Ames. —— pe D. R. Farrar. 2001. Significance . eae form in long-distance colonization by mele bigaie ferns. Brittonia 53:352-369. M.
Evans, A. 3. New peace a. in the Polypodiaceae and Grammitidaceae. Carolan i. 671-6 Farrar, D. R. 1990. heat and evolution in asexually reproducing independent fern gameto-
phytes. ak Bot. 15:98-111.
Hartt, D. L. and A. G. CLark. 1997. Principles of Population Genetics. Sinauer Associates, Sunder-
and, Massachusetts.
Haurier, C. H. and G. J. Gasrony. 1978. Antheridiogen and breeding system in the fern genus Bommeria. Canad. J. Bot. 56:1594—1601.
and D. E. Souris. 1984. Obligate mies von in a homosporous fern: field confirmation of
a laboratory prediction. Amer. J. Bot. 71:878—-881.
, M. D, WinpHaM, and T. A. Ranker. 1990. oo analysis of the Cystopteris tennes-
seensis complex. Ann. Missouri Bot. Gard. 77:314—-329.
Hooper, E. A. and C. H. HAuFer. 1997. Genetic Hii and ues system in a group of neo- tropical Kae ferns (Pleopeltis; Polypodiaceae). Amer. J. Bot. 84:1664—-1674
K.exowsKkI, E. J. JR. 1968. Reproductive biology and evolution in the pteridophyta. PhD diss. Uni- versity of Sateen Berkeley.
———. 1971. Ferns and genetics. BioScience 21:317-322.
. 1979. The genetics and reproductive biology of ferns. Pp 133-170 in A. F. Dyer, ed. The
Experimental ain of Ferns. Academic Press, Lon
966. Evolutionary significance of polyploidy i in the pteridophyta. Sci-
ence 153: 305-307. Knosiocu, I. W. 1967. Chromosome numbers in Cheilanthes and Polypodium. Amer. J. Bot. 54:461 ae
Li, J., and C . 1999. Genetic variation, breeding apres and patterns of diversifica- tion in poantonanp: Polypodinan (Polypodiaceae). Syst. Bot. 2
Love, A., D, Love, and R. E. G. oo SERMOLLI. 1977. eset renait Atlas of Pteridophyta. mer, Vaduz, Liechtenstei
Cra Manton, I. 1951. ee of Pabeodieae’ in America. Nature 167:3 Masuyama, S Y. Watano. 1990. Trends for inbreeding in cas pteridophytes. Pl. Sp.
Antheridiogens and antheridia development. Pp 435-470, in A. F. Dyer, ed. The enc ldesit regen of ferns. Academic Press, London. , C. E. 1993. Campyloneurum. Pp 328 in Flora of North America Editorial Committee,
add. Flora of eet America. Oxford Univ. Press, New York, NY.
Nei, M. 1978. Estimation of seu anes and genetic distance from a small number of individuals. Genetics 89:583—
Peck, J. H., C. J. Peck, and D. R. viciieae 90. Influences bs life history attributes on formation of local and distant fern populations. Amer Fern. J. 8
Ranker, T. A., LER, P. S. Soitis, and D, E aay he 1989. Genetic evidence for allopoly- ploidy in the Neotropical fern esis eee (Adiantaceae) and the reconstruction of an ancestral genome. Syst. Bot. 14:439-447
CHIOU ET AL: MATING SYSTEMS 79
a, : K. FLoyp, and P. G. Trapp. 1994. Multiple colonization of Asplenium adiantum-nigrum o the ea eget Archipelago. Evolution 48:1364—1370. Roc, i S. 1972. Measures of genetic similarity and genetic distance. Studies in Genetics, + Toss ate 7213:145-153 Petia J. J., C. H. Haurter, and T. A. Ranker. 1990. Antheridiogen and natural gametophyte populations. Amer. Fern J. 80:143-15 and A. Hess, 1995. Antheridiogen syatem in the fern Asplenium ruta-muraria (Asplenia-
Soitis, D. E. and P. S. Soitis. 1986a. Electrophoretic evidence for inbreeding in the fern Botrychium asl (Ophioglossaceae). Amer. J. Bot. 74:504—509. and P. S. TIs. 1986b. Polyploidy and breeding systems in homosporous pteridophyta: a eeecnhs preci Naturalist 130:219- d, P.:S, oo 1990. Genetic i pa and among populations of ferns. Amer.
Fern J. 84:161— Sorsa, V. 1966. ae studies in the rcs come Amer. Fern J. 56:113-119. Sworrorp, D. L. and R. B. SELANDER. 1989 -1. A computer program for the analysis of
allelic variation in acenuanl cites or biochemical systematics, release 1.7. Illinois Natural History Survey. Urban
Tsal, J.-L. and W.-C. SHIEH. 1983. A ptenicoienhe survey of the pie e se in Taiwan. (1) Material collection and chromosome observation. J. Sci. Engin. 20:1
VoELLER, B. 1971. Developmental adehes of fern gametophytes: enitian hr biology. Bio- Science 5 267-270.
Wertu, C. R. 1989. The use of — data for inferring ancestry of polyploid pteridophytes. 2 lid eo Ecol. 17:117—
——., and M. I. Cousens. 1990. yen the contribution of population studies on ferns. Amer. Fern J. 80:183-190.
American Fern Journal 92(2):80—92 (2002)
Belowground Distribution and Abundance of Botrychium Gametophytes and Juvenile Sporophytes
CINDY JOHNSON-GROH'’, CHANDRA RIEDEL?, LAURA SCHOESSLER®, and KrissA SKOGEN* Biology Department, Gustavus Adolphus College, 800 W. College Ave., St. Peter, MN 56082
phytes m_ * respectively. Botrychium hesperium also has a relatively high density of 478 gameto- [ ith ar
The importance of propagule banks (also referred to as seed, spore, or dia- spore banks) in community dynamics has long been recognized (Leck et al., 1989) for flowering plants. Propagules may persist belowground for many years, creating a secure reservoir from which aboveground plants can be
density of fern gametophytes resulting from cultivation of spore banks is high, ranging from 57,000 spores m ? (Milberg, 1991) to 5,000,000 m~2
y Hall, Iowa State University, Ames, IA 50011. Evolution and Behavior, University of Minnesota, 100 Ecology Bldg., 1987 Upper Buford Circle, St. Paul, MN 55108
Current address: Argonne Na | ry, ’ . ° ’ a .
JOHNSON-GROH ET AL.: BELOWGROUND DISTRIBUTION 81
of propagules (represented by germinated gametophytes) to the aboveground density of sporophytes, all previous studies have reported the remarkable lack of ferns present in the aboveground vegetation despite a belowground reservoir of spores. During and ter Horst (1983), Leck and Simpson (1987), Milberg (1991), Raffaele (1996), Rydgren and Hestmark (1997) and Strickler and Edgerton (1976) all found evidence of high densities of fern propagules where ferns were entirely absent from the aboveground vegetation.
Botrychium presents two problems in applying conventional propagule bank techniques. First is the difficulty and length of time required to culture Botrychium beyond germination (Whittier and Thomas, 1993). Botrychium spores require darkness for germination, and in nature Botrychium gameto- phytes require mycorrhizae for growth beyond the two- or three-celled stage. Therefore, it is not possible to culture and quantify the propagules using standard seed bank techniques.
The second and more significant problem relates to the different life cycle of Botrychium, relatively little of which is visible aboveground (Fig. 1). Plants produce spores that filter into the soil and germinate in darkness. Fol- lowing germination, a belowground achlorophyllous, fleshy gametophyte is produced. The gametophyte produces gametangia and sexual reproduction occurs, resulting in a belowground juvenile sporophyte. The belowground rhizome is upright and short with mycorrhizal roots